Periglypta listeri (J. E. Gray, 1838) (Bivalvia: Veneridae) in the western Atlantic: taxonomy, anatomy, life habits, and distribution

Rüdiger Bieler

Periglypta listeri (J. E. Gray, 1838), one of the largest and most distinctive western Atlantic venerids, and the only Atlantic member of the genus, is redescribed based on original mate-rial from the Florida Keys, museum specimens, and literature records. Conchologically, this species agrees with previously described venerids in having a well-developed escutcheon and lunule, and a hinge with three cardinal teeth in each valve. Within the genus, it is unique in having internal purplish brown coloration, and in the frequent presence of a purplish brown "hinge dot" on the anterior lateral tooth. This is the first anatomical study for any species in the genus Periglypta, and the most complete so far for any member of Venerinae. Periglypta listeri agrees with previously described venerids in most anatomical characteristics, and notably features an undulating mantle edge that can close in "zipper" fashion, tentacles at the anterior mantle edge, and branching tentacles at the tips of the unfused siphons, type B mantle fusion, type C(2) ctenidia, and a type V stomach. Although empty shells are commonly collected, P. listeri unusually (for venerids) lives cryptically in rubble or sand among rocks, and/or in reef settings. Thus far, the presence of an anterior lateral hinge tooth is the sole morphological feature separating the subfamily Venerinae from the closely allied Chioninae.

MALACOLOGIA, 2004, 46(2): 427−458 PERIGLYPTA LISTERI (J. E. GRAY, 1838) (BIVALVIA: VENERIDAE) IN THE WESTERN ATLANTIC: TAXONOMY, ANATOMY, LIFE HABITS, AND DISTRIBUTION Rüdiger Bieler1*, Isabella Kappner1 & Paula M. Mikkelsen2 ABSTRACT Periglypta listeri (J. E. Gray, 1838), one of the largest and most distinctive western Atlantic venerids, and the only Atlantic member of the genus, is redescribed based on original material from the Florida Keys, museum specimens, and literature records. Conchologically, this species agrees with previously described venerids in having a well-developed escutcheon and lunule, and a hinge with three cardinal teeth in each valve. Within the genus, it is unique in having internal purplish brown coloration, and in the frequent presence of a purplish brown “hinge dot” on the anterior lateral tooth. This is the first anatomical study for any species in the genus Periglypta, and the most complete so far for any member of Venerinae. Periglypta listeri agrees with previously described venerids in most anatomical characteristics, and notably features an undulating mantle edge that can close in “zipper” fashion, tentacles at the anterior mantle edge, and branching tentacles at the tips of the unfused siphons, type B mantle fusion, type C(2) ctenidia, and a type V stomach. Although empty shells are commonly collected, P. listeri unusually (for venerids) lives cryptically in rubble or sand among rocks, and/or in reef settings. Thus far, the presence of an anterior lateral hinge tooth is the sole morphological feature separating the subfamily Venerinae from the closely allied Chioninae. Key Words: Florida Keys, Mollusca, Caribbean, infaunal, clam, sanctuary. INTRODUCTION remain unresolved, due to a surprising paucity of comparative morphological work. Despite their relative abundance and commercial importance, only about 50 venerid species have some published anatomical data. Most publications focus on a few species traditionally grouped in the Chioninae, such as representatives of Mercenaria (Kellogg, 1892, 1903, 1915; Morse, 1919; Jones, 1979), Chione (Kellogg, 1915; Jones, 1979; Narchi & Gabrieli, 1980), Timoclea (Ansell, 1961; Narchi, 1980), Lirophora (Jones, 1979), Tawera (Burne, 1920), Chamelea (Odhner, 1912; Ansell, 1961), Anomalocardia (Narchi, 1972; Purchon, 1985), Bassina (Morton, 1985; Purchon, 1985), Protothaca (Guerón & Narchi, 2000), and Clausinella (Ansell, 1961). Anatomical details for members of other nominal venerid subfamilies are much sparser, with data available for individual species of Gouldiinae [= “Circinae”] (Pelseneer, 1911; Ansell, 1961; Fishelson, 2000), Cyclininae (Purchon, 1985), Dosiniinae (Thiele, 1886; Ansell, 1961; Guéron & Coelho, 1989; Fishelson, 2000); Gemminae (Morse, Veneridae is the largest marine family of bivalves, with many of the more than 500 living species forming key components in the world’s clam fisheries. The nominate subfamily Venerinae currently comprises 14 genus-group taxa (e. g., Venus, Periglypta, Globivenus, Ventricoloidea; Keen, 1969) with more than 140 nominal extant and fossil species. Members of this subfamily live in a wide range of benthic habitats, in coarse sand, mud, or gravel between tide lines to depths over 150 m, and from temperate to tropical seas. Delimiting shell characteristics are the presence of both radial and concentric sculpture and an anterior lateral tooth in the left valve (Keen, 1969; see hinge discussion below). The nominal subfamily Chioninae, another large group comprising such genera as Chione, Mercenaria, and Protothaca, has recently been synonymized with Venerinae by some authors (Coan & Scott, 1997; Coan et al., 2000). Relationships between these and among other venerid subfamilies Department of Zoology, Division of Invertebrates, Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, Illinois 60605-2496, U.S.A. Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192, U.S.A. *Corresponding author: bieler@fieldmuseum.org 1 2 427 428 BIELER ET AL. 1919; Sellmer, 1967; Narchi, 1971), Meretricinae (Kellogg, 1915; Narchi, 1972; S. Gray, 1982; Narchi & Dario, 2002), Pitarinae (G. B. Sowerby II, 1854; Thiele, 1886; Pelseneer, 1911; Kellogg, 1915; Morse, 1919; Narchi, 1971; F. R. Bernard, 1982; S. Gray, 1982; Fishelson, 2000; Morton, 2000), and Tapetinae (G. B. Sowerby II, 1854; Carrière, 1879; Pelseneer, 1894, 1897, 1911, 1923, 1931; Berkeley, 1959; Ansell, 1961; Nielsen, 1963; Joshi & Bal, 1965a, b; Morton, 1985; Fishelson, 2000). The morphological diversity of the nominate subfamily, Venerinae, remains largely unexplored, with published anatomical data restricted to Circomphalus casina (Linnaeus, 1758) (Ansell, 1961), Venus verrucosa Linnaeus, 1758 (type species of Venus Linnaeus, 1758, and of Clausina Brown, 1827; Pelseneer, 1894, 1897), and Globivenus toreuma (Gould, 1850) (Pelseneer, 1911). The current paper focuses on a venerine species currently classified in the genus Periglypta Jukes-Brown, 1914, a group not previously studied anatomically. Periglypta listeri (J. E. Gray, 1838), also known as “Lister’s venus” or “princess venus”, is the second largest Caribbean venerid species, only exceeded in shell size by Mercenaria campechiensis (Gmelin, 1791), which ranges from the mid-Atlantic coast to the Gulf of Mexico and extends into the Caribbean. A shallow-water species with a very conspicuous shell, P. listeri had at one point even been declared the type species of the genus Venus (Stoliczka, 1871: xvii; Venus verrucosa Linnaeus, 1758, was subsequently fixed as the type by ICZN Opinion 195, 1954). Empty shells of P. listeri are commonly collected, but living specimens are less frequently encountered, due in part to their relatively cryptic infaunal habitat in seagrass areas and algae-covered rubble near reefs. This paper reviews the taxonomy and geographic distribution of this species, its anatomy, and life habits, based on original information from living specimens from the Florida Keys, together with a re-evaluation of existing literature and selected museum data. Comparisons are drawn with sympatric large-bodied venerids in the western Atlantic, with selected worldwide species of Periglypta, and with known anatomical data for the family. MATERIALS AND METHODS This study is part of an ongoing investigation of marine molluscan biodiversity in peninsular Florida and the Florida Keys, formally initiated by RB and PMM in 1994. Consecutively numbered stations comprising these collections are preceded by an “FK” acronym in the following text. Living animals and empty shells were collected by hand mainly during scuba diving on coral reefs and shallow-water (2−10 m) patch reefs, rubble areas, and ledges. The majority of live observations were made on specimens from the “Horseshoe” site, bayside of West Summerland Key (Spanish Harbor Keys), comparable to stations IMBW-FK-629 and -637 reported by Mikkelsen & Bieler (2004). Interpretations of distribution records based Henderson’s Eolis expeditions are taken from the recent compilation by Bieler & Mikkelsen (2003). Specimen photography used a variety of equipment and techniques. In situ photographs of living animals (Fig. 24) were taken using a Nikonos V underwater camera with close-up lens. Other living animals (Figs. 25, 26) were photographed in aquaria using standard 35 mm single-lens reflex or electronic cameras. Wholevalve and detail light micrography (Figs. 3−12, 31, 32) used a Microptics® micro/macro imaging system based on a high-resolution Nikon® single-lens reflex digital camera. Excised preserved tissues were prepared for scanning electron microscopy (SEM) by critical point drying and gold sputter coating, then viewed at beam acceleration voltages of 10kV on a Amray 1810 scanning electron microscope at FMNH. For anatomical observation, specimens were relaxed by chilling in a household refrigerator assisted by the addition of magnesium sulfate crystals (Epsom salts) to their seawater supply, or in an isotonic aqueous magnesium chloride solution. Ciliary currents were studied using carmine particles. Anatomy was observed under a dissecting microscope; preserved tissues were dyed for better contrast with neutral red or methylene blue. Voucher FK specimens were fixed in 5% formalin, later transferred to 70% ethanol, and are deposited in the Field Museum of Natural History (FMNH), Chicago, and the American Museum of Natural History (AMNH), New York. All measurements and meristics were taken from the left valve whenever possible. Shell measurements, taken with calipers or with ocular micrometer on a stereomicroscope, include: maximum height from umbo to farthest distal point on free edge, and maximum length (= width) perpendicular to axis of height. Size is expressed as shell length unless otherwise noted. Radial ribs were counted at the growth PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC edge on the main body of the shell. For morphometric analyses (length to height ratio), 32 shells were measured and the linear regression calculated using Microsoft® Excel 2000. Other cited repositories include: ANSP BMNH BMSM CMNH DMNH MCZ MNHN MTD NCSM NTM SBMNH UMML USNM ZMB Academy of Natural Sciences of Philadelphia, Pennsylvania, U.S.A. The Natural History Museum [= British Museum (Natural History)], London, United Kingdom Bailey-Matthews Shell Museum, Sanibel Island, Florida, U.S.A. Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, U.S.A. Delaware Museum of Natural History, Wilmington, U.S.A. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A. Muséum National d’Histoire Naturelle, Paris, France Staatliche Naturhistorische Sammlungen, Museum für Tierkunde, Dresden, Germany North Carolina State Museum of Natural Sciences, Raleigh, North Carolina, U.S.A. Northern Territory Museum of Arts and Sciences, Darwin, Australia Santa Barbara Museum of Natural History, Santa Barbara, California, U.S.A. Rosenstiel School of Marine and Atmospheric Science [= University of Miami Marine Laboratory], University of Miami, Florida, U.S.A. National Museum of Natural History. [= United States National Museum], Smithsonian Institution, Washington, DC., U.S.A. Museum für Naturkunde [= Zoologisches Museum, Berlin], HumboldtUniversität, Berlin, Germany Other abbreviations: alc frag juv LV pair RV spm valve fluid-preserved (alcohol) specimen shell fragment juvenile or subadult left valve an empty (dead) complete shell (2 valves) right valve a live-collected specimen an empty (dead) single valve 429 RESULTS Veneroidea: Veneridae: Venerinae Rafinesque, 1815: 146 (as Veneridia) Periglypta Jukes-Brown, 1914: 72 Type species, by original designation: Venus puerpera Linnaeus, 1758. Periglypta was introduced as a subgenus of Antigona Schumacher, 1817, by Jukes-Brown (1914: 72). Cytherea Röding (1798), non Fabricius, 1794 (Diptera: Bombylidae). Type species (subsequent designation of Dall, 1902): Venus puerpera Linnaeus. Periglypta listeri (J. E. Gray, 1838) Selected synonymy: ?”pectunculus admodum crassus …” Lister, 1687: Liber III, fig. 178 [pl. 341, fig. 178 in later editions]. [unlabelled figure] − Lamarck, 1797: pl. 378, fig. 2a−b. Venus puerpera (2) Var. − Lamarck, 1818: 584−585 [referring to Lister (1687) and Lamarck (1797) figures; non Venus puerpera Linnaeus, 1771]. Venus puerpera Var. 2 − Deshayes, 1832: 1112 [referring to Lister (1687) and Lamarck (1797) figs.]; − Deshayes, 1835: 335. Venus puerpera − Schramm, 1869: 20. Dosina Listeri J. E. Gray, 1838: 308. Venus listeri − Hanley, 1843, in 1842−1856: 110; − Deshayes, 1853a: 106 [distribution (Philippines, Australia) erroneous]; − G. B. Sowerby II, 1853 [in part]: 705, pl. 152, fig. 8 [excluding figs. 7, 9; distribution (Philippines, Australia) erroneous]; − Reeve, 1863: no. 14, pl. 5, fig. 14 [“Hab. Philippine Islands; Cuming” erroneous]; − Krebs, 1864: 97; − Pfeiffer, 1869: 141, pl. 8, figs. 8, 9 [distribution (Nicobares, Philippines) erroneous]; − Simpson, 1889: 64; − F. C. Baker, 1891: 47; − Cockerell, 1894: 118; − Benthem Jutting, 1927: 34; − McLean, 1936b: 119; − Dance, 1974: 263−264, fig.; − FischerPiette, 1975: 36−37; − Dance, 1977: 264, fig. Venus (Periglypta) listeri − Lamy, 1929: 205. Omphaloclathrum listeri − Mörch, 1853: 24. Venus (Chione [Omphaloclathrum]) listeri − Römer, 1867: 32 [“Insulae Philippinae” in error]. Venus crispata − Dall, 1889: 54 [non Venus crispata Deshayes, 1853b]. Cytherea (Cytherea) listeri − Dall, 1902: 372. Cytherea listeri − Dall, 1903: 1275−1276, 1279; − Maury, 1920: 103. 430 BIELER ET AL. Antigona (Periglypta) listeri − Jukes-Brown, 1914: 72; − Warmke & Abbott, 1961: 185, pl. 38 fig. L; − Humfrey, 1975: 248, pl. 30, fig. 3. Antigona (Dosina) listeri − Palmer, 1927: 337 [129]; − Palmer, 1929: pl. 28, figs. 2, 11; − Abbott, 1954: 404, pl. 32, fig. m; − Abbott, 1958: 129. Antigona listeri − Weisbord, 1926: 83; − Johnson, 1934: 48; − Lermond, 1936: 6; − Clench & McLean, 1936: 166; − McLean, 1936a: 41; − Clench & McLean, 1937: 39− 40; − M. Smith, 1937: 53, pl. 21, fig. 11; − M. Smith, 1940: 110, fig. 1445; Jaume & Perez Farfante, 1942: 39 [Pleistocene]; − Jaume, 1946: 101; − Aguayo & Jaume, 1949: 1; − Pulley, 1952: 150, pl. 13, fig. 7; − NowellUsticke, 1959: 14; − Abbott, 1961: 162; − Weber, 1961: 58; − Rice & Kornicker, 1962: 382, pl. 7, fig. 6a−b; − Moulding, 1967: 83; − Brooks, 1968: 8; − J. A. Baker, 1969: 3−4; − Ross, 1969: 8; − Voss et al., 1969: 71; − Abbott, 1970: 162; − Stanley, 1970: 160, pl. 21, figs. 12, 13; − McGinty, 1970: 58 [Lower Pleistocene]; − Magnotte, [1970−1979]: 63, fig. 12; − Woods, 1970: 2−3; − McGinty & Nelson, 1972: 13; − Godcharles & Jaap, 1973: 37; − Zischke, 1973: 35; − Zischke, 1977a: 29; − Zischke, 1977b: 338, fig. A.14− 67; − Romashko, 1974: 49, fig. 17; − Ekdale, 1974: 657; − Eisenberg, 1981: 169, pl. 151, fig. 16; − Abbott, 1986: 230, fig. 5; − Sutty, 1990: 92, fig.; − Prieto et al., 2001: 593. Antigona (Antigona) listeri − McLean, 1951: 82, pl. 15, fig. 5. Periglypta aff. listeri – Weisbord, 1964: 300− 302, pl. 43, figs. 7, 8 [Pliocene]. Periglypta listeri − Morris, 1973: 58, pl. 24, fig. 13; − Abbott, 1974: 521, color pl. 24, fig. 5852; − Emerson & Jacobson, 1976: 429, pl. 42, fig. 18 [AMNH 106142; seen by authors]; − Parodiz, 1976: 20 [Mayan ruins]; − Lozet & Pétron, 1977: 129, fig. 247; − Edwards, 1980: 3; − Theroux & Wigley, 1983: 47, fig. 85 (map); − Voss et al., 1983: 316, 429; − Romashko, 1984: 96, fig. p. 97; − H. E. Vokes & E. H. Vokes, 1984: 43, pl. 45, fig. 9; − Abbott, 1984: 54, fig. 9; − Lipe & Abbott, 1991: 76, fig. 9; − Lawson, 1993: 53; − Espinosa et al., 1994: 123; − Díaz M. & Puyana H., 1994: 78, pl. 18, fig. 170; − Abbott & Morris, 1995: 60, pl. 30; − Lyons & Quinn, 1995: J-13; − Alvarez, 1998: 103 ff.; Pointier & Lamy, 1998: 214, fig.; − Tremor, 1998: 7; − Turgeon et al., 1998: 48; − Mikkelsen & Bieler, 2000: 379; − Redfern, 2001: 236, pl. 101, fig. 965. Periglyphus (sic) listeri − Voss et al., 1983: 79, 183. Distribution (Figs. 1, 2) North Carolina, southeastern and western Florida, including the Florida Keys, Texas, the West Indies, Caribbean Central America and northern South America; apparently not reaching Brazil. It appears to have a largely Caribbean island (as opposed to Gulf of Mexico or mainland North or South American) distribution. It is rare off Texas and other Gulf of Mexico locations. It extends along the entire island chain of the Florida Keys, from Key Largo to the Dry Tortugas. Localities (*= unverified): *North Carolina (Pulley, 1952). Florida: east coast (AMNH, ANSP, FMNH, USNM; Dall, 1902; M. Smith, 1937; Pulley, 1952; Stanley, 1970; McGinty & Nelson, 1972), Florida Keys (AMNH, ANSP, BMSM, CMNH, DMNH, FMNH, USNM, this study; Simpson, 1889; Dall, 1902; Palmer, 1927; Lermond, 1936; M. Smith, 1937; Pulley, 1952; Abbott, 1961; Brooks, 1968; Ross, 1969; Abbott, 1970; Magnotte, 1970−1979; Woods, 1970; Godcharles & Jaap, 1973; Zischke, 1973, 1977a, b; Edwards, 1980; Voss et al., 1983; Lyons & Quinn, 1995; Tremor, 1998; W. G. Lyons, pers. comm.), Dry Tortugas (USNM, this study), *west coast (M. Smith, 1937; Pulley, 1952); Texas (ANSP); Caribbean: Bahamas (AMNH, ANSP, FMNH, USNM; Clench & McLean, 1936; McLean, 1936b; Clench & McLean, 1937; McLean, 1938; Moulding, 1967; J. A. Baker, 1969; Lawson, 1993; Redfern, 2001), Cuba (ANSP, FMNH, MTD, USNM); McLean, 1936a; Aguayo & Jaume, 1949; Pulley, 1952), Cayman Islands (ANSP; Abbott, 1958), Jamaica (USNM), Hispaniola − Haiti and Dominican Republic (AMNH, ANSP, USNM; Palmer, 1927), Puerto Rico (ANSP, USNM; Clench & McLean, 1937; McLean, 1951; Warmke & Abbott, 1961), Virgin Islands (AMNH, ANSP, FMNH, USNM; Krebs, 1864; Dall, 1902; McLean, 1951; Nowell-Usticke, 1959; Weber, 1961), Antigua (USNM), *Martinique (Lamy, 1929; Fischer-Piette, 1975); *Guadeloupe (Schramm, 1869; Pointier & Lamy, 1998); Grenada (ANSP); Netherlands Antilles (DMNH, FMNH, USNM; Benthem Jutting, 1927); Central America: Mexico (AMNH; F. C. Baker, 1891; Weisbord, 1926; Jaume, 1946; Rice & Kornicker, 1962; Ekdale, 1974; Parodiz, 1976 [Mayan ruins]; H. E. Vokes & E. H. Vokes, 1984); Belize (ANSP, USNM), Honduras (USNM; Alvarez, 1998); Costa Rica (USNM); Panama (USNM); South America: Colombia (AMNH, FMNH, USNM; Díaz M. & Puyana H., 1994), *Venezuela (Prieto et al., 2001); *French Guyana (Femorale, 2003). PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC FIG. 1. Distribution of Periglypta listeri; * denotes record for region, without details. FIG. 2. Distribution of Periglypta listeri in the Florida Keys; (•) living records, () dead records. 431 432 BIELER ET AL. Fossil Record Diagnosis Lower Pleistocene (McGinty, 1970: 58) and Upper Pleistocene (M. Campbell, in lit., Feb. 2003) of southern Florida; Pleistocene of Cuba (Jaume & Perez Farfante, 1942) and Santo Domingo [Hispaniola] (Dall, 1903). It was collected from Mayan archaeological sites (but not from concurrent Recent collections) in Yucatan, Mexico, by Parodiz (1976). Weisbord (1964: 302) noted that the species was “reported from the Pleistocene of … Barbados, and the Island of Tortuga, Venezuela”, but did not include source citations for these records; Weisbord examined additional fragments from the Lower Mare formation (Pliocene, possibly Lower Pliocene) at Quebrada Mare Abajo, northern Venezuela. Cassab (1984) recorded the eastern Pacific P. multicostata (G. B. Sowerby I, 1835) from several Tertiary western Atlantic locations − the Chipola Formation (Lower Miocene) of Florida, the Pirabas Formation (Lower Miocene) of Para State, Brazil, and the Middle Miocene of Costa Rica; verification of the species-level identity of these materials, which date prior to the closure of the Panamanian landbridge, is warranted. All post-closure western Atlantic Periglypta records should most definitely be reconsidered against P. listeri; Dall (1903: 1279) noted that P. multicostata “has been enumerated as one of the reef Pleistocene fossils of St. Domingo, but doubtless through a misidentification, perhaps of [P.] listeri.” A largeshelled form described as P. tamiamensis Olsson & Petit, 1964, from Florida’s Late Miocene and Pliocene Tamiami formation seems closely related. The relationship of P. listeri to extinct western Atlantic species from older geological formations [e.g., Antigona dominica Palmer, 1928 (= A. caribbeana Anderson, 1927), Miocene of Santo Domingo, fide Hertlein & Strong, 1948; P. tarquinia (Dall, 1900), Oligocene of western Florida and Santo Domingo, called a small “precursor” of P. listeri by Dall, 1903; P. mauryae (H. E. Vokes, 1938), Upper Miocene of Trinidad], awaits a more comprehensive review of the genus. Large western Atlantic venerid, with thickwalled, inflated, trapezoid shell, with external sculpture of erect commarginal ridges crenulated by underlying radial sculpture, and with posterior end vertically truncated. Exterior cream-colored, with scattered brown speckles, blotches, or flames. Interior yellowish white with more-or-less strongly developed purplish brown stain around posterior adductor muscle scar, posterior margin, and above the pallial line. Anterior lateral hinge tooth often with a purplish brown “hinge dot”. Type Material No original material located (see Taxonomic Remarks, below). Material Examined See Appendix 1. Description Shell relatively heavy, equivalve, longer than high, trapezoid, with nearly straight dorsal margin and bluntly truncated posterior margin; inequilateral with low, rounded umbones approximately 1/3 of the shell length from the anterior end (Figs. 3, 4). Length to height ratio very regular throughout lifespan (R2 = 0.9929, n = 32). Inflated, with posterodorsal slope somewhat concave. External sculpture consisting of prominent erect commarginal ridges, regularly spaced, reflected umbonally, those along posterior margin higher and not reflected; ridges more-or-less alternating in strength at anterior and posterior margins as well as on later growth of shells (beginning at 2.5−3 cm or after first 15−25 commarginal ridges); ridges crenulated by underlying radial sculpture consisting of uniform flattened ribs separated by narrow grooves approximately 1/2−1/3 width of ribs (Fig. 5). Lunule broadly spindle- to teardrop-shaped (Figs. 6, 13), with deeply incised margins, asymmetric, right half slightly larger than left, with many very fine commarginal lamellae (continuing the much coarser ridge pattern of the anterior shell), without radial elements. Escutcheon distinct (Fig. 6), delimited by a marginal groove, right half overlapping left in the posteriormost third, both halves finely obliquely grooved. Externally cream-colored with brown speckles, blotches, or flames, sometimes darker posterodorsally, occasionally radially merging into (often three) broadening radial stripes; escutcheon and lunule with color of surrounding shell. Internal margin finely crenulated, continuing onto lunular margin. Anterior adductor muscle scar oval; posterior adductor muscle scar beanshaped, somewhat flattened dorsally; posterior scar larger and more curved than anterior; PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC 433 FIGS. 3−9. Periglypta listeri, Florida Keys specimens. FIGS. 3, 4: External shell; FMNH 176372, Little Duck Key, 75 mm; FIG. 5: Sculptural detail of shell in Fig. 3. Scale bar = 5 mm; FIG. 6: Umbonal aspect; FMNH 296695, Rachel Bank, 66 mm; FIGS. 7, 8: Internal shell; FMNH 296695, Rachel Bank, 62 mm; FIG. 9: Loose pearls from 78 mm shell; AMNH 295199, Spanish Harbor Keys, largest pearl with maximum dimension of 5.8 mm. (AMMS, accessory mantle muscle scars; E, escutcheon; GrL, pigmented growth lines on posterior adductor muscle scar; HD, hinge dot; L, lunule). 434 BIELER ET AL. FIGS. 13, 14. Periglypta listeri, details of umbo and prodissoconch; AMNH 296531, Looe Key reef, Florida Keys, 9.5 mm. FIG. 13: Anterior aspect with lunule. Arrows point to transition from wide commarginal lamellae on surface to fine striae on lunule; FIG. 14: Transition between prodissoconch I and II (arrow). Scale bars = 1 mm (Fig. 13), 100 µm (Fig. 14). FIGS. 10−12. Periglypta listeri, details of specimen in Figs. 7, 8. FIGS. 10, 11: Close-up of hinge teeth; FIG. 12: Articulated shells. (1, right middle cardinal tooth; 2a, left anterior cardinal tooth; 2b, left middle cardinal tooth; 3a, right anterior cardinal tooth; 3b, right posterior cardinal tooth; 4b, left posterior cardinal tooth; HD, hinge dot; AII, left anterior lateral tooth). Scale bars = 5 mm. dorsalmost portion of posterior scar formed by pedal retractor muscle scar, separated by faint demarcation. Anterior pedal retractor muscle scar on ventral side of hinge plate, dorsomedial to anterior adductor muscle scar, just below anterior cardinal tooth (Fig. 12: 3a). Pallial line entire (Figs. 7, 8); pallial sinus wide, roundly pointed anteriorly. Accessory pallial muscle scars just inside pallial line, more-or-less regularly spaced. Internal color yellowish white with purplish brown stain around posterior adduc- tor muscle scar and posterior margin, extending dorsally above the pallial line, sometimes also ventrally below, occasionally also surrounding anterior adductor and pedal retractor muscle scars; additional faint purplish brown “lines” often extending toward umbo from anterior limits of pallial sinus and anterior pedal retractor muscle scar (Figs. 7, 8). Live-collected specimens that are pure white internally also noted (e.g., USNM 464243, 890543, 890776). Color pattern within posterior adductor muscle scar sometimes showing distinct growth lines (Fig. 7). Hinge teeth of left valve (Figs. 11, 12) comprising minute anterior lateral [AII] (see Discussion below), prominent triangular anterior cardinal (2a), slightly smaller bifid middle cardinal (2b; with posterior part 1/4−1/3 less prominent), and lamellate posterior cardinal (4b). Hinge teeth of right valve (Figs. 10, 12) comprising well-defined, relatively small anterior cardinal (3a), larger bifid middle cardinal (1; with posterior part narrower and smaller), PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC 435 FIGS. 15−18. Diagrammatic anatomy of Periglypta listeri. FIG. 15: Organs of the mantle cavity, with RV and most of right mantle removed. Arrows on gills and labial palps indicate direction of particle flow. Outer labial palp (OLP) is reflected to show structure and particle flow over palps; FIG. 16: General dissection of alimentary system, with RV and right mantle, siphons, gills and labial palps removed; FIG. 17: Gut-loop variation in two additional dissected specimens (FK-273, FK-357); FIG. 18: Vertical section through gill, illustrating direction of food currents along inner and outer demibranchs. (A, anus; AAM, anterior adductor muscle; APRM, anterior pedal retractor muscle; AU, auricle of heart; BA, bulbus arteriosus; CG, cerebral ganglia; Cut M, cut mantle edge; DG, digestive gland; E, esophagus; EX, excurrent siphon; F, foot; GL, gut loop; I, intestine; IDB, inner demibranch; ILP, inner labial palp; IN, incurrent siphon; K, kidney; M, mantle edge; MO, mouth; ODB, outer demibranch; OLP, outer labial palp; PAM, posterior adductor muscle; PG, pedal ganglia; PPRM, posterior pedal retractor muscle; S, stomach; SS, style sack; V, ventricle of heart; VG, visceral ganglia). 436 BIELER ET AL. and prominent, wide, equally bifid posterior cardinal (3b). Anterior lateral tooth at base of anterior cardinal of LV, and its corresponding socket in RV, often each with purplish brown “hinge dot” (Figs. 8, 10, 11). Prodissoconch I visible at tip of umbo (Figs. 13, 14), 155 µm in length (n = 1; AMNH 296531, FK-269). Uncertain border between prodissoconch II and juvenile shells (at about 1.3 mm). Juvenile sculpture (Figs. 13, 23) initially smooth, followed by fine commarginal ridges with wide bands of radial ribs between. Surface pattern of brown to orange speckles particularly noticeable on juvenile shell (Fig. 23). Foot large (Fig. 15), with brown pigment (in living specimens) between muscular part and visceral mass, with pedal gland opening slightly anterior of midpoint; pedal groove extending from ventral posterior end toward anterior third. Mantle muscles attaching mantle to shell, some extending dorsalward to produce so-called accessory muscle scars dorsal to pallial line (approx. 1.5 mm dorsal, in larger specimens). Attachment of anteriorly pointed siphonal retractor muscles producing (and reflecting shape of) pallial sinus on shell. Anterior and posterior adductor muscles (AAM, PAM, respectively) oval; PAM slightly larger than AAM. Posterior pedal retractor muscle (PPRM) round, arising anterodorsal to PAM. Fibers of PAM differently colored in living material, with posterior third darker than remainder. Anterior pedal retractor muscle (APRM) attaching to underside of anterior hinge plate and inserted into anterodorsal part of foot a short distance posterior to AAM. Visceral retractor muscle attached to inner surface of umbo behind hinge plate, inserting into roof of visceral mass. Mantle with four mantle folds as previously described for other venerids (Ansell, 1961; Yonge, 1957). Ventral mantle margins distinctly wavy (Figs. 15, 20, 26) in living and preserved specimens; capable of closing in “zipper” fashion and held closely appressed in living indi- FIGS. 19−22. Periglypta listeri, scanning electron micrographs of critical-point dried tissues; AMNH 295199, West Summerland Key, 73.2 mm. FIG. 19: Mid-ventral portion of the outer demibranch, with food groove (arrows); FIG. 20: Ventral mantle margin with wavy inner median mantle fold; FIG. 21: Anteriormost portion of mantle margin with all four mantle folds; FIG. 22: Detail of anterior mantle margin with triangular tentacles. (IF, inner mantle fold; M1, inner middle mantle fold; M2, outer middle mantle fold; OF, outer mantle fold. Scale bars = 100 µm (Figs. 20, 22), 1 mm (Figs. 19, 21). PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC viduals at rest. Pedal gape extending ventrally from base of incurrent siphon to AAM. Anterior 1/8 of inner median mantle fold with small triangular tentacles (Figs. 21, 22). Siphons separated (Figs. 15, 26), each with distinct black pigment (persisting in preserved specimens) between tentacles internally and externally; pigment stronger at terminal margin and fading toward basal region; internally with densely placed yellow-white surface papillae; each siphon with terminal digitate tentacles and basal 437 siphonal membrane consisting of a thin tissue flap narrowing the lumen, with membrane of incurrent siphon forming a double ridge. Siphonal mantle fusion of type B (Yonge, 1957, 1982); union of inner and middle folds exposing outer surface of middle folds, with common outer ring of sensory tentacles. Incurrent (ventral) siphon slightly larger in diameter and length than excurrent (dorsal) siphon (Figs. 24−26); excurrent siphon tapering to narrower terminal diameter than wider, slightly flaring incurrent FIGS. 23−26. Periglypta listeri, juvenile shell and living specimens. FIG. 23: Juvenile shell with color pattern; FMNH 296719, West Summerland Key, Florida Keys, 7.2 mm; FIG. 24: Siphons of living specimen in situ, muddy sand and algal cover, 2 m depth; West Summerland Key, not collected; FIG. 25: Siphons of living specimen in sand in laboratory tank; FMNH 301448, West Summerland Key, 56 mm; FIG. 26: Living specimen in laboratory tank; FMNH 295706, off Stirrup Key, Florida Keys, 59 mm. 438 BIELER ET AL. siphon. Siphonal tentacles digitate, those on incurrent siphon with 5−11, on excurrent with 3−5, lateral papillae, interspersed with additional small, simple tentacles; both kinds of tentacles on excurrent siphon with distal opening of unknown function (Figs. 29, 30); complex tentacles on incurrent siphon (Figs. 24−28) 30− 75% larger, slightly more pointed and more numerous (approx. 40 tentacles of various sizes in 73 mm specimen, FK-273; AMNH 295199) than those on smaller excurrent siphon (approx. 30 tentacles, same specimen; Figs. 24−26, 29− 30). Distal end of excurrent siphon with pointed dome-shaped valve surrounded by ring of tentacles, presumably allowing control of current (Figs. 24−26, 30). Demibranchs smooth (not plicated; Fig. 15), with axes nearly vertical dorsoventrally; inner demibranch slightly larger than outer, each with numerous interlamellar junctions. Surface currents moving particles ventrally on the outer surface of each demibranch; currents on inner surfaces uncertain. Food grooves at distal edges of inner and outer demibranchs (Figs. 18, 19), with oralward currents, indicating gills and ciliation of type C(2) (Atkins, 1937); longitudinal oralward current also found between bases of adjacent demibranchs. Triangular palps with narrow lamellae on inner surface (33−37 lamellae, n = 2; FK-273, AMNH 295199, 73 mm; FK-352, FMNH 283534, 67 mm); margins smooth at ventral and dorsal side; outer surface smooth. Acceptance currents on palps along lamellae directed ventrally, and on ventral boundary oralward. Rejection currents counter-oralward on smooth ventral and dorsal margins of palps. Ctenidial/labial palp association of type II (Stasek, 1963); anteroventral tips of inner demibranch inserted and fused to distal oral groove between labial palps. Esophagus relatively short, with 7−8 longitudinal rugae, leading into anterior part of stomach (Fig. 16); stomach embedded in di- FIGS. 27−30. Periglypta listeri, scanning electron micrographs of critical point dried siphonal papillae; AMNH 295199, West Summerland Key, 73.2 mm. FIG. 27: Digitate tentacles on incurrent siphon, exterior aspect of siphon; FIG. 28: Same, viewed from interior of siphon; FIG. 29: Detail of simple tentacle of excurrent siphon with orifice of unknown function; FIG. 30: Digitate tentacles on excurrent siphon in front of flap-like valve (arrow). Scale bars = 1 mm (Fig. 27), 100 µm (Figs. 28, 30), 10 µm (Fig. 29). PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC gestive diverticula, located posterior to labial palps. Stomach of type V (Purchon, 1985), with two caeca. Right caecum with seven ducts; left caecum with 11 ducts opening into digestive diverticula; four ducts of digestive diverticula opening into left pouch. Major typhlosole (MT) and intestinal groove extending from intestine into stomach, penetrating right caecum; MT crossing stomach floor beneath esophageal opening into left caecum; MT continuing from stomach into intestine, ending just after gut loop. Minor typhlosole projecting into stomach but ending close to midgut opening. Gastric shield located opposite crystalline style sac at roof and left side of stomach, with flanges that pass into dorsal hood and left pouch. Dorsal hood with left and right sorting areas, located over esophageal opening. Style sac combined with midgut, together exiting posteroventrally from posterior end of stomach, then turning anteriorly and coiling on ventral side of stomach, turning again posteriorly and crossing style sac, then ascending (as hindgut, penetrating pericardium, ventricle and bulbus arteriosus) to continue dorsally and posteriorly across surface of PAM (Fig. 16). Gut loops varying somewhat (Fig. 17) in configuration between individuals, but always including a single ventral loop. Ventricle of heart surrounding intestine, with lateral auricles connecting to outer limbs of kidney (Fig. 16). Kidney positioned along ventral edges of pericardium and anterior to PAM (Fig. 16). Three pairs of ganglia (Fig. 16), with cerebral ganglia between APRM and AAM, joined by supraesophagal commissure; visceral ganglia ventral and slightly anterior to PAM; pedal ganglia extensively fused, anteroventral to the ventral gut loop. Gonad surrounding stomach and intestine within visceral mass; reproductive system not otherwise investigated. Dimensions and Maximum Recorded Size Median length approximately 65 mm, ranging 13−100 mm (mean 63.3 ± 14.7 mm SD, n = 103). Maximum 100.2 mm, St. Thomas, Virgin Islands; AMNH 31868 [registered specimen, K. Hutsell (San Diego, California), Registry of World Record Size Shells]. Largest Florida Keys specimen, 96 mm, FK-601, FMNH 296718. Habitat and Ecology Based on observations in southeastern Florida’s Biscayne Bay, Stanley (1970: 160) 439 described this species as “widespread in intertidal and shallow subtidal settings, in large numbers; it is generally restricted to coarse substrate and grassy areas”. Voss et al. (1969: 71) reported the species, for the same region, from the highly unlikely habitat of “rocks; pilings; seawalls” as well as from seagrass beds, open sand, and lagoonal patch reefs. Other habitat records include sand (FMNH, this study; Abbott, 1974; Humfrey, 1975; Dance, 1977; Tremor, 1998), mud (Humfrey, 1975), loose rock/rubble (this study; Godcharles & Jaap, 1973; Zischke, 1973; Voss et al., 1983), and seagrass (this study; Clench & McLean, 1937; Zischke, 1973; H. E. Vokes & E. H. Vokes, 1984). Clench & McLean (1937: 39) noted that P. listeri was “exceedingly abundant” in Savannah Sound (Eleuthera, Bahamas) but generally uncommon elsewhere. At Savannah Sound, they lived buried 5−10 cm below the surface of seagrass-covered sand bars; native Bahamians were observed using “a short stick as a probe [to locate the clams] ... [they are not] eaten by the natives though they are gathered in this way for fish bait” (Clench & McLean, 1937: 37). In the present study, one living specimen was found nestled in red algae within the cavity of a large barrel sponge at 8 m depth (FK-395), although the favored habitat for the species seems to be sand with loose rubble. It undoubtedly is a shallow-water species, in contrast to the analysis by Theroux & Wigley (1983: 47) who placed this species based on two dredge records “in the 50−99 m depth range grouping”. Collecting records of all (living and dead) material range from shallow water to 84 m (Theroux & Wigley, 1983), with the deepest confirmed livecollected specimen in this study from 8 m (FK395). Stanley (1970: 160) described the slow burrowing process of this species and noted that (in Biscayne Bay) the depth of burial is to some extent dependent upon interference by the subsurface rhizomes of surrounding turtlegrass (Thalassia testudinum König). In the laboratory, when permitted to burrow in its native sediment with the seagrass removed, a 5.8 cm-long animal assumed a position with the posterior shell margin 4.5 cm beneath the sediment surface. Specimens observed in the present study were often found hindered from deep burrowing by rubble, and had their shells barely covered by substratum. In addition to an herbivorous diet, evidenced through the presence of a crystalline style, P. listeri appears to opportunistically ingest zooplankton: two copepods were found in the stomach of 440 BIELER ET AL. one individual (FK-352). In Puerto Rico, this species is a “favourite food of the Slipper Lobster” (Sutty, 1990: 92). In the Florida Keys, empty shells have been frequently found associated with octopus middens at scuba depths, or with beveled drill holes, the latter indicative of predation by naticid gastropods. Pearls A single living animal was found with a series of loose pearls lining the center of the inside shell, apparently in response to the remains of an intruding worm-shaped organism (Fig. 9). The 78 mm specimen (FK-273, AMNH 295199) contained 11 irregular pearls, the largest with a maximum dimension of 5.8 mm. This is the first record of pearls from Periglypta; both free and attached pearls have previously been reported from several other venerids, especially Mercenaria spp. (Haas, 1931; Shirai, 1994; Hill, 1996; Landman et al., 2001; Mienis, 2001). Taxonomic Remarks Although there is general consensus in the recent literature of applying the species name “listeri” to this western Atlantic taxon, the taxonomic history of that name is not without complications. Dosina listeri was introduced by J. E. Gray (1838: 308). He placed it in an unnamed section of Dosina J. E. Gray, 1835, characterized by “anterior lateral tooth small, sometimes obliterated”, together with three previously described species: Venus verrucosa Linnaeus, 1758 (as “Dosina veerrucosa [sic], Venus veerruicosa [sic], Linn.”); Venus reticulata Linnaeus, 1758; and Venus puerpera Linnaeus, 1771. While other species in the same article were expressly identified as new species descriptions (labeled as “n.s.”, accompanied by a textual description and including an indication of the originating collection; e.g., as done for Grateloupea cuneata J. E. Gray, 1838: 304), the name D. listeri appears as a new name referring to prior literature data (1838: 308). The complete “description”, cryptic by today’s standards, introducing the name listeri for a previously unnamed variety of Venus puerpera Linnaeus, 1771, reads as follows: “Dosina Listeri, V. puerpura [sic] var., Linn. Sow. Gen. f. Ency. Meth. t. 278, f. 2”. Venus puerpera was introduced by Linnaeus (1771: 545), in the Mantissa. Linnaeus himself did not mention varieties in the original description and referred to two prior illustrations − Gualtieri (1742: pl. 83, fig. F) and Argenville (1742: pl. 26, fig. F). In Hanley’s words (1855: 453): “Neither of the very dissimilar figures referred to bears the least resemblance to the shell which has been universally accepted for the species.” In any case, Venus puerpera clearly is an Indo-Pacific, not Atlantic, species (Fig. 31). Lamarck (1818: 584−585) and Deshayes (1832: 1112) distinguished two varieties of V. puerpera; their “Venus puerpera var. 2” referred to the cited figure of Lister (1687), as well as to an illustration by Lamarck (1797: pl. 278, fig. 2a, b) in the Encyclopédie Méthodique. J. E. Gray (1838) had named Dosina listeri for Martin Lister, author of the Historia Conchyliorum (1685−1692). The first reference to a figure by Lister in conjunction with Venus puerpera, and the first reference to a distinct variety of this species (as γ), appeared in the 13 th edition of Linnaeus’ Systema naturae. The latter was authored by Gmelin (1791: sp. 3276), and it is the entry in this work that Gray seemed to have meant with his “var., Linn.”. The indicated Lister figure (1687: pl. 341, fig. 178) is in agreement with today’s concept of Periglypta listeri, but lacks detail to distinguish it from similar bivalve shells. J. E. Gray’s (1838) introduction of the new name Dosina listeri included references to two works. As outlined above, he did not seem to have actual specimens before him, and it appears that only the specimens referred to in these two works (and, possibly, Lister’s original material) qualify as the type series (ICZN, 1999: Art. 72.4). One reference, and the only one indicated by Gray with actual plate and figure numbers, was to Lamarck’s figures (1797: pl. 278, fig. 2a, b) as cited earlier by Lamarck (1818) and Deshayes (1832) for “variety 2” of Venus puerpera. The latter are excellent illustrations of an external right valve and of the external hinge area of an articulated specimen. Lamarck’s illustration matches today’s concept of western Atlantic Periglypta listeri. Somewhat confusingly, Deshayes (1835: 334) stated that Lamarck’s specimen of “var. 2” and the figure 2a, b of the Encyclopédie fit well with the typical V. puerpera of Linnaeus. Lamy & Fischer-Piette (1938: 292) agreed and considered all such Lamarck material in the MNHN collection to represent V. puerpera [as well as (from a specimen agreeing with figs. 1a, b of Encyclopédie pl. 278) Venus magnifica Hanley, 1845]. The interior features of the shell, particularly its coloration, are unknown; the specimen on which Lamarck’s excellent illustrations were PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC based cannot currently be located (B. Metivier, 10/2002 in lit.). The other work, cited by J. E. Gray as “Sow. Gen. f.”, is G. B. Sowerby I’s Genera of Recent and fossil shells (1834). In it, figure 1 of the Venus plate shows detailed color illustrations of the inside valves of a species identified as V. puerpera. However, the illustrated valves lack the brown or purple stains usually present in Periglypta puerpera and P. listeri and instead are shown with extensive orange coloration below the umbo. The illustrated specimen or specimens (considerable differences in the shape of mantle line and muscle scars indicate that these drawings of valves might originate from different individuals) have not been located in the BMNH collection. The orange interior coloration is matched by a specimen of P. puerpera from J. E. Gray’s collection (BMNH 1991135, unknown locality, 49 mm; seen by authors). However, this shell differs considerably from Sowerby’s illustrations in having dark brown markings at the shell margin, oval (not angular) anterior muscle scars, and a much more angular posterior shoulder. Another specimen at BMNH, furnished as a potential syntype of Dosinia listeri (K. Way, 02/2003 in lit.; BMNH 1840.7.1.41, 76 mm, no locality, old British Museum collection; see Appendix), is a member of P. listeri in today’s sense, but likewise not a shell illustrated by G. B. Sowerby I (1834). It is here, for reasons outlined above, not considered part of the syntypic series. Because the identity of P. listeri is not currently in dispute, and the figured specimens of Lamarck and Sowerby cannot be located at present, we refrain from designating a lectotype (or neotype) in the context of this study and defer to a future genus-wide revision. Subsequent British authors, particularly Hanley (1843, in: 1842−1856: 110) and G. B. Sowerby II (1853: 705), referred to Venus listeri as a species distinct from V. puerpera. However, their concept of this nominal species was broader and included differently colored forms with dark rayed patterns on the shells, as evidenced by the range of included illustrations. Venus listeri was thought to hail from the “Indian Seas”, from the Philippines and Australia. Nevertheless, G. B. Sowerby II (1853: 705) expressly endorsed Lamarck’s oft-cited illustration for V. listeri by stating “The figure in the Encyclopaedia is very exact for the type.” Reeve (1863: pl. 5, no. 14) narrowed the interpretation of V. listeri to shells that are “fulvous-white, obscurely freckled with flesh- 441 brown”, but still assumed the waters near the Philippines Islands as its home. It was in the context of an Indo-Pacific interpretation of V. listeri that Reeve (1863: pl. 3) stated that he doubted that eastern Pacific Periglypta multicostata “is anything more than a variety of V. listeri, in which the ribs are more timidly thickened and recurved.” Fischer-Piette (1975) assigned most of the Indo-Pacific records of Venus/Dosinia/Chione/Cytherea/Antigona listeri to P. puerpera and restricted P. listeri to Atlantic records. Comparative Remarks Periglypta listeri is one of the largest western Atlantic venerids, second only to Mercenaria campechiensis in maximum adult size. Its sculpture, of flattened radial ribs between sharp commarginal ridges (which are further crenulated by the radials), renders it distinct from other sympatric venerids: M. campechiensis lacks discernible sculpture between the commarginals, whereas Globivenus (formerly Ventricolaria) rugatina (Heilprin, 1886), G. rigida (Dillwyn, 1817), and Circomphalus strigillinus (Dall, 1902) have fine concentric striae between the commarginal ridges and are also rounder in general shell shape. Periglypta listeri is part of a worldwide complex of morphologically similar species, the relationships of which far exceed the scope of this paper. Römer (1867: 32, here translated) referred to similarities so great “that the differences often can barely be put into words” when comparing “Venus” listeri to other nominal species of Venus − Venus lacerata Hanley, 1845; V. clathrata Deshayes, 1853; V. crispata Deshayes, 1853; V. multicostata G. B. Sowerby II, 1853; V. laqueata G. B. Sowerby II, 1853; V. resticulata G. B. Sowerby II, 1853; V. chemnitzii Hanley, 1844; V. sowerbyi Deshayes, 1853; V. reticulata Linnaeus, 1758; and V. monilifera G. B. Sowerby II, 1851. Citing a high degree of variability, E. A. Smith (1885: 120−121) considered “Venus” resticulata, V. aegrota Reeve, 1863, V. lacerata Hanley, V. sowerbyi, V. clathrata, V. crispata, V. puerpera, V. listeri, and probably also V. multicostata and V. magnifica Hanley, 1845, as “races” of a single biological species, although he fell short of formalizing this action in not placing these names in synonymy under V. puerpera. Modern revisionary treatment of the Indo-Pacific taxa is urgently needed. Periglypta listeri is the only living member of the genus in the Atlantic Ocean. It differs from 442 BIELER ET AL. FIGS. 31, 32. Contrasting shell morphology in Periglypta spp. FIG. 31: P. puerpera; FMNH 82478, Mindanao, Philippines, 59 mm; FIG. 32: P. multicostata; FMNH 165936, Pacific Panama, 111 mm. the Indo-Pacific P. puerpera (Fig. 31), type species of the genus, in general shell shape and coloration. Periglypta puerpera is less anteroposteriorly elongated, and generally more rounded in outline. Its external sculpture appears smoother, a result of its less prominent commarginal ridges. Externally it is creamcolored, with or without scattered radial brown flecks, often with 1−3 darker brown rays, one almost always extensively covering the posterior third. Internally it is white with a bright purple (not purplish brown) stain at the posterior margin below the PAM, sometimes also with a yellow or peach-colored flush at the center (although internally pure-white specimens also occur). As earlier noted by E. A. Smith (1885: 120−121), P. puerpera has a “V-shaped purple mark upon the apex of the umbones”, referring to two radial color bands on the prodissoconch and earliest juvenile stage; this is lacking in examined specimens of P. listeri. Weisbord (1964: 302) called Periglypta listeri the western Atlantic analog of Panamic P. multicostata, citing a difference in the outline of the posterior end (obliquely vertical and truncated in P. listeri, subtruncated and more rounded in P. multicostata; see Fig. 32); although based on many museum specimens (AMNH, n = 31), this difference is only obvious among the largest specimens. Periglypta multicostata attains a larger size (maximum observed 120 mm, AMNH 248600, Baja California Norte, Mexico) than P. listeri. Externally, the commarginal ridges of P. multicostata are decidedly coarser and more prominently dorsally reflected than those of P. listeri, rendering the crenulations and radial ribs less obvious. Internally, P. multicostata is white (or very occasionally flushed with pink, not purplish brown, either centrally or in an oblique posterior streak from umbo to margin, but not prominently surrounding the posterior muscle scars) and marginal crenulations are less noticeable or absent in the largest specimens (although the latter are quite prominent in specimens < 65 mm). DISCUSSION Periglypta listeri agrees with previously described venerids in conchological features, such as a well-developed escutcheon and lunule, and a hinge with three cardinal teeth in each valve (Keen, 1969). Although Lamprell (1998) cited differences in the shape of the pallial sinus, we found that distinguishing characters at the species level reside in sculpture and color/color pattern. So far as is known, the pattern of the internal purplish brown coloration and the purplish brown “hinge dot” are unique to P. listeri within the genus. PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC This paper presents the first anatomical study for any species in the genus Periglypta, and is arguably the most extensive anatomical description yet available for any member of the subfamily Venerinae. Anatomical information is published for only three other venerines. Pelseneer (1894, 1897) provided minor details about the anatomy of Venus verrucosa (tongue-shaped foot without byssus, very small labial palps, siphons more or less fused). He later (Pelseneer, 1911) presented similarly cursory details (siphons short and united, with retractors; short “byssal” groove on posterior foot; very narrow external demibranch) for Venus (now Globivenus) toreuma. Ansell (1961: fig. 8) illustrated (but did not extensively discuss) the gross anatomy and stomach structure of Circomphalus casina (Linnaeus, 1758), which are closely similar to those in P. listeri with respect to the gills, labial palps, foot, stomach, and adductor muscles, as well as the flow of particles over the gills and labial palps. Like P. listeri, C. casina features a structurally complex ventral mantle edge, but its function was not mentioned. Its siphons appear short, unlike the longer, separated siphons of P. listeri. Ansell (1961; also Pelseneer, 1911) considered the degree of siphonal fusion to vary greatly within Veneridae, and from these minimal data this could also be true within Venerinae. The anatomy of Periglypta listeri agrees with these and other previously described venerids in most features. As elaborated by Yonge (1957), Ansell (1961), and Narchi (1971), the mantle edge has four folds (outer, OF; inner middle, M1; outer middle, M2; and inner, IF). The periostracum is secreted between OF and M2. In P. listeri, M1 is undulating, with its anterior part elaborated into distinctly triangular tentacles. The wavy structure and tentacles can close in “zipper” fashion (Fig. 26), presumably to protect the organs of the mantle cavity from intrusion of particles and small organisms. At the anterior end of the pedal gape, OF fuses with M2, and M1 fuses with IF. At the posterior end, M1 and IF are fused to form the siphons; this configuration typifies venerids of type B fusion (Yonge, 1948, 1957, 1982; Ansell, 1961). The conical valve on the tip of the excurrent siphon is the elaborated IF, while the ring of tentacles on the siphon represents M1 (Yonge, 1957; Ansell, 1961; Jones, 1979). The tentacles on the incurrent siphon are not distinguishable into inner and outer rings of tentacles, in contrast to Jones’ (1979) findings in members of Chione, Mercenaria, and Austrovenus. The siphonal tentacles of P. 443 listeri are long and digitate, serving as an effective screen to block intrusion of particles. Ansell (1961) contrasted the digitate tentacles of Circomphalus, Timoclea, Clausinella, Venerupis, and Gafrarium spp., which live in gravelly and stony bottom habitats, against the simple tentacles of Chamelea and Dosinia spp., which live in cleaner sand or gravelly bottoms. Periglypta listeri is not congruent with this pattern; it carries digitate tentacles but lives in a gravel and sand habitat without a high amount of detritus in suspension. At the proximal end of each siphon, Periglypta listeri has a siphonal membrane or valve comprised of thin tissue flaps, as described for other venerids (Ansell, 1961; Jones, 1979). That of the incurrent siphon consists of a basal double ridge; Ansell (1961) described this double ridge in Circomphalus casina and, like others (Kellogg, 1915; Jones, 1979; Narchi & Dario, 2002) postulated that such a valve can be opened to admit water inflow, or closed to direct water ventrally and flush out accumulated pseudofeces. The anterior end of the gill in Periglypta listeri is inserted into the distal oral groove, in the midline of the labial palps to which the gill is fused (Stasek, 1963). The ctenidia, with food grooves on the edges of both demibranchs, correspond to Atkins’ (1937) type C(2) along with members of Paphia, Mercenaria, Chione, Tivela, and Saxidomus. Other venerines, such as Venus verrucosa and Circomphalus casina, as well as other venerids (in Clausinella, Dosinia, Gafrarium, Chamelea, and Paphia), lack a groove on the outer demibranch and have therefore been assigned to type C(1b). However, it must be noted that Atkins (1937) found this character variable within genera (e.g., Paphia) and species (e.g., only one of four specimens of C. casina had a groove on the outer demibranch), indicating that this character requires larger sample sizes and further research. The circulatory and nervous systems are broadly similar to the species studied by Jones (1979), as is the general plan of the digestive system and configuration of the gut loop. The type V stomach of P. listeri agrees overall with the descriptions by numerous authors (Ansell, 1961; Dinamani, 1967; Narchi, 1971, 1972; Jones, 1979; Purchon, 1987; Narchi & Dario, 2002) for other members of the family. A combined style sac and midgut is typical of venerids, except for Placamen tiara (Dillwyn, 1817), in which the openings, although close together, lead into separate tubes (Dinamani, 444 BIELER ET AL. 1967). The numbers of ducts entering the left and right caeca vary among species and range, respectively, from a minimum of one and two in Nutricola tantilla (Gould, 1853) to a maximum of 13 and seven in Venerupis pullastra (Montagu, 1803) (Purchon, 1987); P. listeri is at the high end of this range with 11 and seven ducts. Four additional ducts open into the left pouch in P. listeri, in contrast to two or three in Meretrix, Katelysia, Placamen, Sunetta, and Irus spp. (Dinamani, 1967), five ducts in Tivela and Gafrarium spp., or eight in Dosinia spp. (Purchon, 1987). No additional ducts enter the stomach near the right caecum in P. listeri, unlike in Clausinella and Callista spp. (Purchon, 1987). Periglypta listeri lives in rubble or sand among rocks, and/or in reef settings (as opposed to soft bottoms like most other venerids). This is also true for the similar eastern Pacific P. multicostata, aptly named “giant reef clam”, described as a common species in 3−6 m depth in Baja California Sur (GarcíaDomínguez et al., 1998), and noted from sand among rocks (Keen, 1971). Other Periglypta species recorded for rocky or reef habitats are P. puerpera and P. reticulata (fide Whitehead, 1983; Graham, 1995). Periglypta was originally described as a subgenus of Antigona, and Harte (1998: 358) maintained that that is its proper placement. However, although Antigona Schumacher (type species by original designation, A. lamellaris Schumacher, 1817, Indo-Pacific) shares radial threads between the commarginal shell ridges with Periglypta, its members have radial ribs extended onto the lunule, no groove around the escutcheon, a more triangular pallial sinus, and hinge teeth with a much wider 3b and smaller 2b. Periglypta also differs from Dosina J. E. Gray (type species, D. zelandica J. E. Gray, 1835, South Pacific, by subsequent designation of Frizzell, 1936), in which P. listeri was originally described, by the predominantly concentric sculpture (Keen, 1969) and a weakly or undeveloped escutcheon in that genus (pers. obs.). Most authors (e.g., Keen, 1969) refer to an anterior lateral tooth in the left valve as characteristic of venerines, but an anterior lateral tooth is also present in members of other venerid subfamilies (e.g., Gouldiinae [= “Circinae”], Sunettinae, Meretricinae, Pitarinae, and Dosiniinae; Keen, 1954). There are indications that such lateral teeth result from different ontogenetic pathways and might not be homologous. Félix Bernard (1895: 127) presented an ontogenetic series of hinge development for a Miocene species of Gouldia, which (together with its numbering system) became the model for tooth development in the “corbiculoid” hinge type sensu Cox (1969: N54). According to this model, the venerine anterior lateral tooth of the left valve (designated as AII) derives from lamella II. Whereas F. Bernard (1895: 127) noticed differing hinge morphologies of other venerids, such as Macrocallista, he still recognized the anterior lateral tooth in the latter as a lamella II derivative, and thus an AII. Marwick (1927: 598−599), in another ontogenetic comparison, showed that the Oligocene venerine Kuia vellicata (Hutton, 1873) displayed the same AII development as shown in F. Bernard’s Gouldia study. However, Marwick (1927: 598) maintained that the left anterior lateral tooth of Macrocallista, a continuation of a low ridge proceeding from below the umbo, is “in no way connected with the anterior cardinal”. Some subsequent authors (Frizzell, 1936) followed Marwick’s argument of non-homologous anterior lateral teeth in the Veneridae; Grant & Gale (1931: 316) referred to the venerine AII of Antigona as a “pseudolateral”, in contrast to the situation in groups such as Macrocallista. It appears that venerid “anterior lateral teeth” might variously be derived from either lamella II or IV and homology assumptions of these structures need closer scrutiny. Chionines do not have anterior lateral or pseudolateral teeth even though their overall shell morphology is very similar to that of venerines. Anatomical studies carried out by Jones (1979) showed that chionine siphons are usually fused along their length in contrast to the unfused siphons of Periglypta listeri seen in this study; the high degree of variability in siphonal fusion, as well as the presumably fused siphons in the venerine Circomphalus casina (see above) render this a weak distinguishing character. At present, the presence of an AII lateral hinge tooth is postulated as the only reliable morphological feature separating these two nominal subfamilies. Although recently synonymized by Coan & Scott (1997), preliminary molecular studies (16S gene; I. Kappner, unpubl.) indicate a distinct grouping that might warrant retention of these subfamilial units. Studies of additional taxa, and more anatomical and molecular characters are needed for resolution of subfamilial synapomorphies. PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC ACKNOWLEDGMENTS This work is part of an ongoing investigation of the molluscan diversity of the Florida Keys; regional surveys and collections were supported by permits from the Florida Keys National Marine Sanctuary (080-98, 2000-036, 2002-078, and 2002-079); in the vicinity of Pigeon Key (on National Register of Historic Places) under the auspices of the Pigeon Key Foundation; in the Dry Tortugas National Park, under collecting permits DRTO-19970030 and 2002-SCI-0005; in Long Key State Park (Long Key, Florida Keys) under Florida Department of Environmental Protection permit 5-02-43; in Key West National Wildlife Refuge (near Sand Key, Florida Keys) under United States Fish and Wildlife Service permit 41580-01-07. Additional collecting was sanctioned under Florida Fish and Wildlife Conservation Commission permit 99S-024 to affiliates of The Bailey-Matthews Shell Museum (Sanibel, Florida) and permit 01S-056 (as well as annual permits for prior years of this study) to affiliates of the Smithsonian Marine Station (Ft. Pierce, Florida; logistic support by Mary E. Rice and staff is much appreciated). We thank Timothy Collins, Timothy Rawlings, Roberto Cipriani, Deirdre Gonsalves-Jackson, Cecelia Miles, Louise Crowley, Daniel Miller, Jim Culter, the Smithsonian Marine Station at Fort Pierce, and the captains and crews of R/ V EUGENIE CLARK (Mote Marine Laboratory, Sarasota, Florida) and R/V CORAL REEF II (Shedd Aquarium, Chicago) for collecting assistance. Data gathering from other museum collections was facilitated by Gary Rosenberg (ANSP), José Leal and Tina Petrikas (BMSM), Charles Sturm (CMNH), Timothy Pearce, Leslie Skibinski, and Albert Chadwick (DMNH), Katrin Schniebs (MTD), Nancy Voss (UMML), Jerry Harasewych (USNM), Ronald Jansen (Senckenberg Museum, Frankfurt), Matthias Glaubrecht and Lothar Maitas (ZMB). Bernard Metivier (Museum National d’Histoire Naturelle, Paris), and Kathie Way (BMNH) provided type specimen information and loans. Richard E. Petit, as so often before, helped us disentangle obscure literature references. IK greatly appreciates Osmar Domaneschi’s advice on anatomical questions and thanks Gustav Paulay and Rudo von Cosel for very fruitful discussions. The research, in part, was supported by NSF-PEET DEB-9978119 and Comer Science and Education Foundation grants to RB and PMM. 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Revised ms. accepted 31 October 2003 APPENDIX 1: Material Examined Material examined (this study, Florida Keys) RB unnumb., N of Bahia Honda State Park, shallow water, sand, 25 Mar 1989 (1 spm alc, FMNH 288810); FK-018, “The Stakes”, off Middle Florida Keys, scuba, 20 ft (6.1 m), patch reef/ledges, sand patches, M/V SITE FINDER, 08 July 1995 (1 pair, AMNH 308088); FK-035, Indian Key Fill, 24°53’25"N, 80°40’28"W, bayside, Thalassia seagrass bed, 1 m, shovel/ sieve, 10 March 1996 (1 juv valve, AMNH 296528); FK-039, Crawl Key, 24°44’36"N, 80°58’47"W, oceanside, beach inside channel, shallow sand, seagrass, seawall, by hand and hand dredge, 12 March 1996 (1 spm alc, FMNH 301424); FK-047, Channel marker 50A off Ramrod Key, 24°35.80’N, 81°27.24’W, rubble and patch reef, 15 ft (4.6 m), scuba, M/V THE SNAIL, 21 September 1996 (1 pair, 1 valve, AMNH 295179); FK-068, bayside of West Summerland Key (Spanish Harbor Keys), 24°39’19"N, 81°18’13"W, bayside, shovel/ sieve in Thalassia + beach combing, 0.5−1 m, 18 April 1997 (3 valves, 1 frag, FMNH 279531); FK-069, Channel marker 50A off Ramrod Key, 24°35.80’N, 81°27.24’W, rubble and patch reef, 18 ft (5.5 m), scuba, R/V FLORIDAYS, 19 April 1997 (1 pair, FMNH 279521); FK-090, Garden Key, Dry Tortugas, 24°37’50"N, 8252’20"W, sand beach with stone pilings, 2.5 m, by hand and snorkeling, 23 April 1997 (1 pair, AMNH 296520), FK-091, Fort Jefferson, Garden Key, Dry Tortugas, 24°37’50"N, 82°52’24"W, beach at mote and mote wall, by hand and snorkeling, 23 April 1997 (2 pair, AMNH 290122); FK115, East Washerwoman Shoal (Channel Marker 49), off Marathon, 24°40’N, 8104.3’W, 9 ft (2.7 m), scuba, R/V FLORIDAYS, 12 July 1997 (1 valve, FMNH 279523); FK-117, Key Vaca, channel west of Stirrup Key, 24°44.19’N, 81°02.92’W, bayside, in channel, 15 ft (4.6 m), 452 BIELER ET AL. plus in surrounding shallow Thalassia/sand, scuba, 14 July 1997 (1 spm alc [photographed alive], FMNH 295706); FK-119, “The Slabs” patch reef between “outer patches” and “coral humps” off Marathon, 24°39.53’N, 81°00.90’W, scuba, 23 ft (7.0 m), R/V FLORIDAYS, 20 July 1997 (1 valve, FMNH 279528); FK-121, “The Slabs” patch reef between “outer patches” and “coral humps” off Marathon, 24°39.53’N, 8100.90’W, 23 ft (7.0 m), scuba, R/V FLORIDAYS, 21 July 1997 (1 pair, 3 valves, AMNH 296525); FK-131, off Key Vaca, oceanside, “outer patches” south of Hawk Channel, 24°39.30’N, 81°01.30’W, rubble, gorgonians, sponges, 21 ft (6.4 m), scuba, R/ V FLORIDAYS, 07 August 1997 (3 valves, AMNH 296530); FK-135, east of Bethel Bank, Florida Bay, 2443.86’N, 81°07.41’W to 24°43.60’N, 81°07.47’W, sand/sparse seagrass, 8 ft (2.4 m), dredge, R/V FLORIDAYS, 07 August 1997 (1 valve, FMNH 279525); FK149, exposed flats outside channel off Crawl Key, 24°44’29"N, 80°58’24"W, oceanside, clean sand + Thalassia/Syringodeum seagrass, Penicillus, 0.5 ft (0.15 m), by hand, R/V FLORIDAYS, 22 August 1997 (2 valves, FMNH 279530); FK-169, Tavernier Creek, near bayside entrance, west side (Plantation Key), 25°00.76’N, 80°32.68’W, sand/Thalassia, 6− 8 ft (1.8−2.4 m), ponar grab and dredge, R/V FLORIDAYS, 17 September 1998 (1 valve, AMNH 296522); FK-171, just off mouth of Tavernier Creek, oceanside, near marker #7, 24°59.67’N, 80°31.72’W, sand, Thalassia/ Syringodeum seagrass, 0−3 ft (0−0.9 m), snorkeling/sieving, R/V FLORIDAYS, 17 September 1998 (observed pair); FK-178, east end of Rodriguez Key, oceanside of Key Largo, 25°03.13’N, 80°26.49’W, Thalassia seagrass, sand, small rubble, 2−3 ft (0.6−0.9 m), snorkeling, R/V FLORIDAYS, 20 September 1998 (2 pair, 1 valve, AMNH 295178); FK-179, Lower Matecumbe Key, 24°51’24"N, 80°43’40"W, oceanside, from beach front 3 days after Hurricane Georges, in Thalassia droves (1−3 ft deep) washed ashore by storm, 28 September 1998 (1 valve, FMNH 279529); FK-205, Carysfort Reef, 25°13.25’N, 80°12.78’W, coral rubble, sand patches, coral heads, 6−15 ft (1.8−4.6 m), scuba, R/V FLORIDAYS, 10 April 1999 (1 spm alc, AMNH 298893); FK-207, east end of Rodriguez Key, 25°03.13’N, 80°26.49’W, sand, seagrass, rubble, 1−2 m, snorkeling, R/V FLORIDAYS, 11 April 1999 (3 pair, AMNH 296519); FK-228, Old Dan Bank, bayside off Long Key, 24°49.66’N, 80°50.18’W, Thalassia/Porites/ Halimeda, 2−4 ft (0.6−1.2 m), snorkeling, R/V FLORIDAYS, 31 July 1999 (2 pair, AMNH 296521); FK-233, Old Dan Bank, bayside of Long Key, north of marker 2X, 24°49’57"N, 80°49’45"W, Thalassia seagrass, Halimeda, Porites, 2−4 ft (0.6−1.2 m), snorkeling/sieving, R/V FLORIDAYS, 01 August 1999 (2 pair, AMNH 296527); FK-236, Coffins Patch, oceanside of Grassy Key, 24 °41’05"N, 80°57’28"W, algae-covered coral reef, 16 ft (4.9 m), scuba, R/V FLORIDAYS, 02 August 1999 (3 valves, FMNH 279522); FK-244, bayside of West Summerland Key (Spanish Harbor Keys), 24°39’19"N, 81°18’13"W, bayside, rock wall and sand slope, 23 ft (7.0 m), scuba, 05 August 1999 (1 valve, AMNH 296524); FK-246, bayside of West Summerland Key (Spanish Harbor Keys), along inner shore of western arm of horseshoe, 24°39’19"N, 81°18’13"W, beach, 05 August 1999 (1 frag, AMNH 296523); FK-255, Friend Key Bank, bayside of Bahia Honda Key, north side at crest of bank, 24°42.58’N, 81°16.77’W, Thalassia/Syringodeum seagrass, sand patches, 0.5−2 ft (0.15−0.6 m), snorkeling, R/V FLORIDAYS, 09 August 1999 (3 valves, FMNH 279520); FK-260, Looe Key coral reef, oceanside of Ramrod Key, 24°32.80’N, 81°24.80’W, spur and groove reef, 24−25 ft (7.3−7.6 m), scuba, R/V FLORIDAYS, 10 August 1999 (1 valve, FMNH 279526); FK268, Looe Key back reef, 24°32.79’N, 81°24.33’W, seagrass, sand patches, rubble, 2−8 ft (0.6−2.4 m), snorkeling, R/V FLORIDAYS, 19 August 1999 (1 valve, FMNH 279527); FK-269, Looe Key back reef, 24°32.79’N, 81°24.33’W, sediment sample, 8 ft (2.4 m), snorkeling, R/V FLORIDAYS (1 juv pair, AMNH 296531); FK-273, bayside of West Summerland Key (Spanish Harbor Keys), at outermost point of western arm of horseshoe, 24°39.35’N, 81°18.22’W, 1−4 ft (0.3−1.2 m), snorkeling, hand collecting, 19 August 1999 (3 spm alc [1 dissected, 1 with pearls, 1 SEM, AMNH 295199; 1 partial animal alc [body of spm with pearls, see prior; DNA], FMNH 296696; 1 juv pair, AMNH 296526); FK-275, Looe Key back reef, 24°32.87’N, 81°24.41’W, 5−10 ft (1.5−3.0 m), snorkeling and hand-collecting, R/V FLORIDAYS, 20 August 1999 (1 valve, FMNH 279524); FK-287, bayside of West Summerland Key (Spanish Harbor Keys), inside outermost arm of horseshoe, 24°39.35’N, 81°18.22’W, shallow subtidal, by hand, snorkeling, 10 April 2000 (1 spm alc, FMNH 289982; 1 live-collected pair, FMNH 301426; 9 pair, FMNH 289939); FK-298, Looe PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC Key National Marine Sanctuary, southwest, ca. 24°32.5’N, 81°24.7’W, 30 ft (9.1 m), scuba, R/ V EUGENIE CLARK, 02 July 2000 (1 juv pair; AMNH 308080); FK-350, Looe Key National Marine Sanctuary, southwest corner of core area, 24°32.61’N, 81°24.66’W, 32 ft (9.7 m), scuba, R/V EUGENIE CLARK, 07 July 2000 (1 valve, 1 juv valve, AMNH 299545); FK-351, Looe Key back reef, 24°32.87’N, 81°24.41’W, rubble, 3−7 ft (0.9−2.1 m), snorkeling, R/V FLORIDAYS, 08 July 2000 (1 pair, AMNH 299489); FK-352, bayside of West Summerland Key (Spanish Harbor Keys), south arm, 24°39’19"N, 81°18’13"W, bayside, rubble, to 1.5 m, snorkeling, 08 July 2000 (3 spm alc [dissected], FMNH 283534; 6 juv pair, 1 valve, 2 juv valves, AMNH 299467); FK-357, American Shoals, northwest of lighthouse, 2431.54’N, 81°31.26’W, Thalassia seagrass with large coral rubble, 9−11 ft (2.7−3.3 m), scuba, R/V FLORIDAYS, 09 July 2000 (3 spm alc [1 dissected], FMNH 301428); 1 frag, AMNH 299581); FK-359, American Shoals, 24°31.56’N, 81°31.10’W, Thalassia/ Syringodeum seagrass with rubble, rocks, 11− 12 ft (3.3−3.6 m), scuba, R/V FLORIDAYS, 10 July 2000 (observed spm; 2 pair, 2 valves, AMNH 299421); FK-360, coral lumps off Newfound Harbor Keys, off Big Munson Key, 24°36.96’N, 81°23.64’W, sand, seagrass, patch reef, gorgonians, 9 ft (2.7 m), scuba, R/ V FLORIDAYS, 11 July 2000 (1 pair, AMNH 299520); FK-363, east end of Rodriguez Key, oceanside of Key Largo, 25°03.27’N, 80°26.66’W, Thalassia seagrass, sand, small rubble, 6 ft (1.8 m), snorkeling, R/V FLORIDAYS, 07 October 2000 (2 pair, AMNH 307612); FK-364, Dove Key (just southwest of Rodriguez Key), oceanside of Key Largo, 25°02.94’N, 80°28.27’W, sand, algae, sponges, Sargassum, 2−6 ft (0.6−1.8 m), snorkeling, R/V FLORIDAYS, 07 October 2000 (1 pair, AMNH 307613); FK-367, northeast corner of Conch Reef, oceanside of Key Largo, 24°57.895’N, 80°27.248’W, rubble, Thalassia seagrass, 9 ft (2.7 m), snorkeling, R/V FLORIDAYS, 08 October 2000 (1 frag, AMNH 307736); FK-392, Lower Matecumbe Key, 24°51’24"N, 80°43’40"W, oceanside, by hand in wrack line, 20 October 2000 (9 valves, FMNH 301429); FK-395, Snappers Ledge reef, 24°58.88’N, 80°25.36’W, in cavity of large sponge with red algae, 26 ft (7.9 m), scuba, M/S REPUBLIC IV, 27 March 2001 (1 pair, FMNH 301430); FK-459, Ft. Zacchary Taylor State Park, Key West, beach, shells washed ashore among rubble, 02 May 2001 (1 frag, 453 AMNH 307737); FK-463, American Shoals, 24°31.541’N, 81°33.218’W, 5 ft (1.5 m), Thalassia/Syringodium seagrass and rubble, scuba, R/V FLORIDAYS, 20 July 2001 (3 pair, 4 valves, 2 frag, FMNH 301431); FK-499, Sand Key, 24°27.18’N, 81°52.79’W, beach to 16 ft (4.9 m) under ship, sand, rocks, patch reef, snorkeling, R/V EUGENIE CLARK, 24 July 2001 (2 frag, AMNH 307738); FK-539, south of Bahia Honda Key, west of Looe Key reef, 24°34.24’N, 81°16.64’W, 30.2−34.1 m (99−112 ft), sand and rubble, pipe dredge and triangle dredge, R/V EUGENIE CLARK, 28 July 2001 (1 juv valve, AMNH 308089); FK-547, Looe Key reef, 24°32.809’N, 81°24.158’W, 25 ft (7.6 m), spur and groove reef, scuba, R/V FLORIDAYS, 30 July 2001 (2 valves, AMNH 307614); FK-559, south beach of Loggerhead Key, Dry Tortugas, 24°37.790’N, 82°55.400’W, wrack line to 7 ft (2.1 m), by hand and snorkeling, R/V CORAL REEF II, 15 April 2002 (3 valves, FMNH 301432); FK-581, north shore of Loggerhead Key, Dry Tortugas, 24°37.871’N, 82°55.447’W, wrack line and shallow subtidal, by hand and snorkeling, R/V CORAL REEF II, 16 April 2002 (5 valves, FMNH 301433); FK-601, Hospital Key, Dry Tortugas, 24°38.970’N, 82°51.284’W, patch reef, sand, 16 ft (4.9 m), scuba and snorkeling, R/V CORAL REEF II, 18 April 2002 (1 valve, 3 frag, AMNH 307615; 1 pair, FMNH 296718); FK-606, southwest of Dry Tortugas, 24°30.009’N, 82°59.914’W to 24°29.970’N, 82°59.687’W, 28 m, triangle dredge, R/V CORAL REEF II, 18 April 2002 (1 frag, AMNH 307616); FK-615, Cosgrove Shoal, 24°27.486’N, 82°11.039’W, 9.7 m (32 ft), patchy rubbly reef with sponges, gorgonians, overhangs, sand flat, scuba, R/V CORAL REEF II , 20 April 2002 (2 pair, 3 valves, AMNH 307617); FK-620, Old Dan Bank, bayside of Long Key, 24°50.45’N, 80°49.63’W, Thalassia seagrass with Halimeda, Porites, sponges, hydroids, patches of sand/Halimeda hash, 1− 2 ft (0.3−0.6 m), by hand, R/V FLORIDAYS, 16 and 18 July 2002 (2 valves, FMNH 301445); FK-622, directly off Keys Marine Laboratory, bayside of Long Key, 24°49.5’N, 80°48.9’W, seagrass bed with coral rubble, snorkeling, sieving, by hand, 0−1.5 m, 20 July 2002 (valves observed); FK-624, Horseshoe Reef, off Fat Deer Key, 24°39.91’N, 80°59.56’W, patch reef with sandy bottom, 24 ft (7.3 m), scuba, M/V SHUTTERBUG II, 20 July 2002 (3 valves, FMNH 301446); FK-625, Coffins Patch Sanctuary Preservation Area, off Crawl Key, 24°40.92’N, 80°58.26’W, patch reef with 454 BIELER ET AL. sand patches, gorgonian, pillar coral, 21 ft (6.4 m), scuba, M/V SHUTTERBUG II, 20 July 2002 (5 valves, FMNH 301447); FK-629, bayside of West Summerland Key (Spanish Harbor Keys), 24°39.3’N, 8118.2’W, among rocks along arms of quarry, to ca. 1 m, by hand, snorkeling, 21 and 26 July 2002 (1 spm alc [siphons photographed in sand, dissected], FMNH 301448; 5 pair, 4 valves, 2 juv valves [incl. photo voucher], FMNH 296719); FK-639, Coral Gardens inshore patch reef, oceanside of Lower Matecumbe Key, 24°50.23’N, 80°43.77’W, snorkeling, 1215 ft (3.6−4.6 m), Keys Marine Laboratory boat, 23 July 2002 (valves observed); FK-647, west side of Pigeon Key, 24°42.2’N, 81°09.3’W, Thalassia/ Halodule/Syringodeum seagrass on sand/ rubble, concrete bridge piers, 0.5−1 m, by hand, snorkeling, shovel/sieving (1 pair, NTM); FK-649, Sprigger Bank, bayside, just W of Everglades National Park border, 24°54.75’N, 80°56.24’W, Thalassia/ Syringodeum seagrass, 1−3 ft (0.1−0.9 m), snorkeling, shovel/sieving, Keys Marine Laboratory boat, 27 July 2002 (2 pair, 1 valve, FMNH 301449); FK-659, Pigeon Key, 24°42.2’N, 81°09.3’W, seagrass, scuba, 2−4 ft (0.6−1.2 m), 28 July 2002 (valves observed); FK-660, Old Dan Bank, bayside of Long Key, 24°50.08’N, 80°49.63’W, Thalassia seagrass with Halimeda, Porites, sponges, hydroids, patches of sand/Halimeda hash, 1−5 ft (0.3− 1.5 m), snorkeling, R/V LAST MANGO, 28 July 2002 (2 pair, 5 valves, 3 frag, FMNH 301450); FK-661, Molasses Keys, south of center of Seven-Mile Bridge, north of westernmost island, 24°41.070’N, 81°11.483’W, sandy bottom, coral rubble, Thalassia seagrass, hot water (> 30°C), snorkeling, 1−6 ft (0.3−1.8 m), R/V FLORIDAYS, 04 August 2002 (5 valves [2 with breakage by predator], FMNH 301434); FK-662, Sombrero Reef, vicinity of buoy SO3, 5 nmi south of Knights Key, 24°37.619’N, 81°06.528’W, sandy bottom adjacent to coral reef; scuba, 17−23 ft (5.2−7.0 m), R/V FLORIDAYS, 05 August 2002 (1 valve, 1 frag, FMNH 301435); FK-664, Molasses Keys, south of center of Seven-Mile Bridge, north of westernmost island, 24°41.070’N, 81°11.483’W, sandy bottom, coral rubble, Thalassia seagrass, strong current, snorkeling, 2−5 ft (0.6−1.5 m), R/V FLORIDAYS, 06 August 2002 (2 valves, FMNH 301436); FK665, Coffins Patch pillars, south of Crawl Key, 24°40.899’N, 80°58.246’W, coral reef, sand plains, some Thalassia seagrass, scuba, 18− 23 ft (5.5−7.0 m), R/V FLORIDAYS, 07 August 2002 (2 valves, FMNH 301437); FK-668, Money Key, oceanside of west end of SevenMile Bridge, off north and west ends of island, 24°41.009’N, 81°12.955’W, “ironshore” beach rock, sand, Thalassia seagrass, snorkeling, 0− 5 ft (0−1.5 m), R/V FLORIDAYS, 09 August 2002 (1 pair, FMNH 301444); FK-672, John Sawyer Bank, 3 nmi north of western part of Key Vaca, bayside, 24°45.498’N, 81°06.621’W, coral rubble, Thalassia seagrass, strong current over shoal, snorkeling, 2−6 ft (0.6−1.8 m); R/V FLORIDAYS, 11 August 2002 (1 pair, 2 valves, 1 fresh frag, FMNH 301438); FK-673, Bethel Bank, 2 nmi N of Knights Key Channel, 24°43.796’N, 81°07.588’W, bayside, coral rubble, Thalassia seagrass, strong current over shoal, 3−5 ft (0.9−1.5 m), snorkeling, R/V FLORIDAYS, 11 August 2002 (1 valve, FMNH 301439); FK-674, Sombrero Reef, southwest area of lighthouse reef, 24°37.555’N, 81°06.729’W, spur and groove reef, rubble, scuba, 22−26 ft (6.7−7.9 m), R/V FLORIDAYS, 12 August 2002 (4 valves, 1 fresh frag, FMNH 301440); FK-675, Red Bay Bank, 3 nmi north of Pigeon Key, bayside, 24°45.1000’N, 81°08.647’W, very diverse habitat: coral, seagrass, many algae, snorkeling, 1−6 ft (0.3− 1.8 m), R/V FLORIDAYS, 13 August 2002 (2 spm, 2 valves, FMNH 301441); FK-676, Rachel Bank, 2 nmi north of Key Vaca, bayside, 24°44.714’N, 81°04.599’W, muddy sand, seagrass, coral rubble, snorkeling, 35 ft (0.9−1.5 m), R/V FLORIDAYS, 13 August 2002 (5 spm, 7 valves, 1 fresh hinged frag, FMNH 296695); FK-677, Doughnut Reef, oval reef mass east of Coffins Patch (off Crawl Key), 24°41.507’N, 80°56.835’W, sand flat at reef edge, scuba, 23 ft (7.0 m), M/V SHUTTERBUG II, 13 August 2002 (1 valve, FMNH 301442); FK-678, bayside of West Summerland Key (Spanish Harbor Keys), 24°39.3’N, 81°18.2’W, inside western arm, scuba, 3−10 ft (0.9−3.0 m), 14 August 2002 (1 spm alc [dissected], 4 valves, FMNH 301443); FK-688, Conch Reef, oceanside of Key Largo, 24°57.380’N, 80°29.428’W, algae-covered reef, vertical walls and adjacent sand plains, max. 28 ft (8.5 m), scuba, R/V FLORIDAYS, 05 June 2003 (1 V, FMNH 302086; 2 LV, AMNH 308081); FK689, northeast of Dove Key, oceanside of Key Largo, 25°03.055’N, 80°28.220’W, hard bottom with silty sand, sponges, gorgonians, 0.5− 1.0 m, snorkeling, R/V FLORIDAYS, 06 June 2003 (1 juv pair, AMNH 308084); FK-692, Hen and Chickens patch reef, oceanside of Plantation Key, 24°56.099’N, 80°32.920’W, max. 21 ft (6.4 m), scuba and snorkeling, R/V PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC FLORIDAYS, 08 June 2003 (1 RV, 1 LV, AMNH 308087); FK-693, off Dove Key, oceanside of Key Largo, 25°03.039’N, 80°28.151’W, silty Thalassia seagrass and sand, 3−5 ft (0.9−1.5 m), snorkeling, R/V FLORIDAYS, 08 June 2003 (3 pair, 1 RV, 1 LV, AMNH 308083); FK696, Sand Island (near Molasses Reef), 25°01.116’N, 80°22.046’W, patch reef with rubble, max. 22 ft (6.7 m), scuba, R/V FLORIDAYS, 10 June 2003 (1 pair, AMNH 308085); FK-698, Wolfe mooring buoy (near Three Sisters Reef), 25°01.311’N, 80°23.774’W, patch reef with adjacent Thalassia seagrass, max. 16 ft (4.9 m), scuba, R/V FLORIDAYS, 11 June 2003 (2 RV, 1 fragment, AMNH 308082); FK-701, off Dove Key, oceanside of Key Largo, 25 °03.011’N, 80°28.163’W, silty Thalassia seagrass and sand, 1−2 ft (0.3−0.6 m), snorkeling, R/V FLORIDAYS, 12 June 2003 (5 pair, 1 RV, 1 LV, AMNH 308086). FK-702, seagrass flat off Yellowtail Inn, oceanside of Grassy Key, mile marker 58.3, 24°45.494’N, 80°57.179’W, 1−5 ft (0.3−1.5 m), snorkeling, 5−23 August 2003 (1 pair, 1 LV, FMNH 302071); FK-703, patch reef with sand pockets in vicinity of mooring buoy near “the Stake”, Coffins Patch coral reef, oceanside of Grassy Key, 24°41.159’N, 80°57.836’W, 8−18 ft (2.4−5.5 m), snorkeling, R/V FLORIDAYS, 7 August 2003 (1 pair, 1 RV, FMNH 302079); FK-704, Sombrero Reef, vicinity of buoy SO-4, 5 nmi S of Knights Key, 2437.591’N, 81°06.563’W, sandy bottom immediately adjacent to spur & groove coral reef; scuba, 17−25 ft (5.2−7.6 m); R/V FLORIDAYS, 08 August 2003 (1 LV, FMNH 302072); FK705, Doughnut Reef, oval reef mass E of Coffins Patch (off Crawl Key); 24°41.439’N, 80°56.862’W, patch reef surrounded by sand flats, scuba, 24 ft (7.3 m); M/S SEAFARI, 09 August 2003 (1 LV, FMNH 302073); FK-706, Elbow Reef, E of Coffins Patch (off Crawl Key); 24°41.551’N, 80°56.789’W, patch reef surrounded by sand flats, scuba, 19 ft (5.8 m); M/S SEAFARI, 09 August 2003 (1 RV, FMNH 302074); FK-710, “Porkfish” and “Hammer Ledge” reefs, E of Coffins Patch (off Conch Key); 24°42.038’N, 80°53.578’W, scuba, 19− 25 ft (5.8−7.6 m); M/S SEAFARI, 11 August 2003 (2 RV, 2 frag, FMNH 302075); FK-711, patch reef with sand pockets in vicinity of mooring buoy 9, Coffins Patch coral reef, oceanside of Grassy Key, 24°41.131’N, 80°57.818’W, 18− 20 ft (5.5−6.1 m), scuba; R/V FLORIDAYS, 12 August 2003 (1 LV, FMNH 302076); FK-714, vicinity of mooring buoy 1 at “Marker 48” patch sand and seagrass; 24°41.505’N, 81°01.528’W, 455 16−24 ft (4.9−7.3 m), scuba; R/V FLORIDAYS, 17 August 2003 (1 pair, 5 LV, FMNH 302077); FK-719, beach near moat wall, Fort Jefferson, Garden Key, Dry Tortugas, 24°37’50"N, 82°52’24"W, by hand, 21 August 2003 (1 RV, FMNH 302078); Other Material Examined Florida: Boynton, Lake Worth, T. L. McGinty! ANSP 195919 (1 pair); Lake Worth Inlet, C. T. Simpson! UMML 28.1729 (1 pair); South Lake Worth near Boynton, Kline! July 1944, DMNH 72175 (1 pair); Lake Worth, Lermond! August 1941, DMNH 153846 (3 pair); Lake Worth, south end, dredged in 6 ft (1.8 m), Lermond! 14 September 1937, DMNH 153848 (2 pair); Lake Worth, south end, Doremus! 1942, DMNH 63779 (2 pair); Lake Worth, south end, F. Lyman family! Summer 1944, FMNH 144097 (1 pair); Lake Worth, South Inlet, oyster reef, F. M. Bayer! USNM 890543 (1 pair with tissue); Boynton in Lake Worth, E Fal, sand pocket in rock reef, T. L. McGinty! June 1940, USNM 599285 (1 pair); Lake Worth, White collection, USNM 153360 (1 pair); Lake Worth, Singer Bridge, F. M. Bayer! USNM 890617 (2 juv pair); opposite Lemon City, C. T. Simpson! UMML 28.1731 (1 LV); Bal Harbor, Broad Causeway, dredgings, Poh! June 1975, DMNH 118270 (1 pair, 1 valve); Coral Gables, FMNH 54675 (1 pair); off Cape Florida, Florida, Finger Channel flats, Crovo! 05 April 1970, DMNH 30152 (1 pair); Cape Florida, Biscayne Bay, T. L. McGinty! 1937, ANSP 264016 (1 pair); Miami Bay area, flats off Cape Florida, Ingalls Family! 12 March 1967, AMNH 140432 (1 pair); Miami area, Biscayne Bay, flats off Cape Florida, W. E. Old! 12 March 1967, AMNH 136053 (2 pair); Cape Florida, T. L. McGinty! FMNH 26427 (2 valves); Cape Florida, near Miami, T. L. McGinty! June 1936, USNM 599308 (1 pair); Bear Cut, Key Biscayne, Miami, W. S. Bitler! 1963, AMNH 142323 (1 pair); Bear Cut, Key Biscayne, S. Sokoloff! 18 November 1961, AMNH 261423 (1 pair); Bear Cut, April 1939, UMML 28.45 (1 pair); Bear Cut, Hepler! DMNH 45331 (2 pair); Caesar Creek, Whitney! November 1956, DMNH 97311 (1 pair); Florida Keys, Lermond! DMNH 153830 (1 pair); Angelfish Creek [north of Key Largo, connecting Card Sound and Atlantic Ocean], Wisoff Collection, AMNH 120395 (1 pair); Key Largo, Nelson collection, FMNH 155544 (1 pair); Key Largo, T. L. Moise! August 1950, ANSP 456 BIELER ET AL. 193815 (1 juv pair); Key Largo, F. M. Bayer! June 1940, USNM 890827 (1 pair); Windley Key, Whale Harbor Channel, near bridge in 5−10 ft (1.5−3.0 m), T. R. Waller! Sta. 3, 31 August 1971, USNM 707750 (1 valve); Upper Matecumbe Key, Islamorada, on beach, 13 January 1978, BMSM 26108 (5 pair); Indian Key, D. V. Stingley! May 1959, BMSM 26109 (1 pair); Lower Matecumbe Key, Hausman! AMNH 133675 (1 pair); Conch Key, south end, J. J. Parodiz & Winters! 08 July 1976, CMNH 43728 (4 pair); Grassy Key, M. & S. Snyder! July 1966, ANSP 309750 (2 pair); Grassy Key, Minzak! February 1972, DMNH 93337 (3 pair); Crawl Key, 24°44’36"N, 80°58’47"W, beach, P. M. Mikkelsen & R. Bieler! 22 September 1996, AMNH 295180 (1 pair); Crawl Key, Richardson! DMNH 85844 (1 pair); Crawl Key, shallow water, J. M. Bijur! May 1964, AMNH 248309 (1 pair); Crawl Key, bayside, on flats at 1 ft (0.3 m), Raeihle! November 1961, AMNH 106142 (1 pair, figured Emerson & Jacobson, 1976: pl. 42, fig. 18); Crawl Key, bayside, on beach, D. Raeihle! November 1973, AMNH 179278 (1 pair); Crawl Key, bayside, D. Raeihle! AMNH 116658 (1 pair); Crawl Key, bayside, November 1959, Raeihle! AMNH 307848 (3 pair, 1 valve, livecollected); Crawl Key, G. Dingerkus & L. D. Uhler! 06 January 1977, AMNH 267424 (1 spm alc); Crawl Key, bayside, edge of borrow pit ½ mi north of mile marker 56, in weeds, M. J. de Maintenon! 24 June 1985, AMNH 307571 (1 pair); Bonefish Key, A. Koto! AMNH 133664 (1 pair) and FMNH 176293 (1 pair, 1 valve); Marathon Key, Florida Straits, K. C. Vaught Collection, AMNH 250783 (1 pair); Marathon, 1959, BMSM 26110 (2 pair); Marathon Key, south end, Jensen! 1962, DMNH 42547 (1 pair); Sombrero Reef, 25−30 ft (7.6−9.1 m), P. S. Mikkelsen! 21 May 1980, DMNH 180066 (2 valves); Washerwoman’s patch reef, 24°39’54"N, 81°04’14"W, M. Snyder! August 1966, ANSP 398069 (1 pair); Pigeon Key, 24°42’N, 81°09’W, M. Snyder! July 1966, ANSP 398068 (2 pair); Little Duck Key, A. Koto! 1955, FMNH 176330 (1 pair) and FMNH 176372 (1 pair [photo voucher]); Little Duck Key, shallow water, sand, F. Schilling! July 1968, FMNH 288716 (1 pair); Missouri Key, Richardson! DMNH 85835 (1 pair), Missouri Key, snorkeling, A. D. Barlow! 05 March 1967, AMNH 243914 (2 pair); Missouri Key, sand, F. Schilling! 16 July 1970, FMNH 288718 (1 pair); Bahia Honda State Park, Germer Collection, 11 July 1973, AMNH 269541 (1 pair); north of Bahia Honda State Park, shallow water, sand, R. Bieler & P. M. Mikkelsen! 25 March 1989, FMNH 288810 (1 spm alc); Bahia Honda Key, BMSM 26111 (1 pair); Spanish Harbor Key, beach, Piech! July 1980, DMNH143838 (1 pair); West Summerland Key, at entrance to dredge hole, L. Scheu Collection, 1984, AMNH 230097 (2 pair); Newfound Harbor Keys, living in shallow water, sand, November 1968, Raeihle! AMNH 307849 (1 pair); Torch Key channel, C. T. Simpson! UMML 28.1716 (4 pair); Sugarloaf Key, J. B. Clark! 23−24 May 1921, ANSP 9634 (1 juv pair); Boca Chica Key, H. A. Pilsbry! ANSP 100273 (1 valve); Stock Island, F. R. Kirtland! 1936, ANSP 167777 (1 pair); Key West, A. Koto! FMNH 176402 (1 pair); Key West, Nelson! FMNH 166745 (1 pair); Key West, coral banks at low tide, J. W. Milner! USNM 127385 (1 pair); Key West, Hawk Channel, 3−20 ft (0.9−6.1 m), Eolis sta. 65, J. B. Henderson Jr.! 15 May 1913, USNM 448344 (2 juv valves); Key West, reefs, rare, H. Hemphill! USNM 95672 (2 pair); Key West, Smith Shoals, Eolis sta. 335, J. B. Henderson Jr.! 1916, USNM 448343 (1 juv pair); Key West, N side, beach collecting, Eolis sta. 35, J. B. Henderson Jr.! 30 May (or 6 June) 1911, USNM 48340 (1 juv pair); Key West, USNM 406825 (2 pair); Key West, H. Hemphill! ANSP 52199 (2 pair); Key West, H. A. Pilsbry! MarchApril 1940, ANSP 175943 (1 pair); Key West, Sand Point, B. R. Bales! 1946, DMNH 21225 (2 pair) and ANSP 285406 (6 pair); Key West, at low tide, 3−4 ft (0.9−1.2 m), F. Schilling! 21 June 1967, FMNH 288719 (1 pair); Key West, C. T. Simpson! UMML 28.1748 (1 pair); Sand Key, Key West, Ostheimer! DMNH110654 (3 pair); near Boca Grande Key, UMML 28.1740, C. T. Simpson! (1 pair); Tortugas, Stm.! USNM 36403 (4 pair, 1 valve); Dry Tortugas, off Loggerhead Key, north end, beach collecting, Eolis sta. 367, J. B. Henderson Jr.! 13 June 1911, USNM 448347 (1 valve); Dry Tortugas, Garden Key, 3 mi out from red sea buoy, 5 dredge hauls, 14−15 fms (25.6−27.4 m), Eolis sta. 34, J. B. Henderson Jr.! 09 June 1911, USNM 448341 (4 pair, 1 valve); Tortugas, Bush Key, F. M. Bayer! USNM 890776 (1 pair); off Carabelle [Franklin County, panhandle of Florida], 29°15’N, 84°40’W, 90−100 ft (27.4− 30.5 m), dredged, J Moore! ex M. & B. Naide, August 1966, ANSP 402130 (1 pair). Texas: off Freeport, 28°13’N, 94°51’W, 27 ft (8.2 m), dredge, A. Kight! ANSP 338392 (1 pair). PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC Bahamas: Bahamas, J. B. Henderson Jr.! USNM 448342 (2 pair); PMM-1047 (wp277R rubble), off Andros, 24°54’44.42"N, 77°53’51.80"W, rubble in back reef pavement zone with gorgonians, 7 ft (2.1 m), scuba, P. M. Mikkelsen, et al.! 29 August 2000, AMNH 305616 (1 valve); PMM-1090 (wp415-R), off Andros, 24°53’32.2"N, 77°53’51.4"W, thick Thalassia seagrass, 12 ft (3.6 m), scuba/snorkeling, P. M. Mikkelsen, et al.! 04 September 2000, AMNH 305617 (1 pair), FMNH 301425 (1 spm alc [95%]); PMM-1079 (wp427-R), off Andros, 24°55’24.8"N, 77°55’19.8"W, sand/algal plain, 5 ft (1.5 m), scuba/snorkeling, P. M. Mikkelsen, et al.! 02 September 2000, FMNH 296720 (1 pair); PMM-1063 (BH-R seagrass), oceanic blue hole off Blue Hole Cay, off Andros, 24°53’55.2"N, 77°55’12.1"W, Thalassia seagrass, 4 ft (1.2 m), scuba/snorkeling, P. M. Mikkelsen, et al.! 31 August 2000, AMNH 307572 (1 pair); East Andros Island, Calabash Bay, Abbott! February 1971, DMNH 29242 (1 pair); East Andros Island, Small Hope Bay, Abbott! DMNH 41253, March 1971 (1 valve); Chub Cay, Berry Islands, Moise! ANSP 193106 (1 pair); Chub Cay, Periwinkle Beach, K. C. Vaught Collection, April 1977, AMNH 250782 (1 pair); Grand Bahama Island, 26°31’N, 78°46’30"W, J. N. Worsfold! ANSP 375213 (2 pair); Grand Bahama Island, 26°31’00"N, 78°46’30"W, J. N. Worsfold! ANSP 375212 (1 juv valve); Grand Bahama Island, Running Mon Canal, 26°29’45"N, 78°41’45"W, J. N. Worsfold! ANSP 369788 (1 pair); Grand Bahama Island, C. C. Allen! 1922-1923, ANSP 133697 (1 pair); Grand Bahama island, Bottle Bay Canal, 26°39’30"N, 78°57’00"W, sediment, 5 ft (1.5 m), J. Worsfold! ANSP 371908 (1 pair); east end of Grand Bahama Island, Deep Water Cay, ca. 2.5 mi northwest of Sweetings Cay Light, intertidal sand and rocks, V. O. Maes! January 1965, ANSP 307688 (1 juv pair); Grand Bahama Island, F. H. Low Collection, AMNH 113790 (1 pair); Nassau, New Providence Island, Wards! (before 1893), FMNH 2741 (1 pair, 1 valve); Nassau, New Providence Island, Pope! FMNH 187147 (2 valves); Nassau, C. C. Allen! USNM 36617 (1 pair); New Providence Island, Dicks Point, McLean & Russell! July 1936, ANSP 169917 (2 valves); New Providence [Island], Lyford Cay, AMNH 80763 (2 pair); north coast of Hog Island, north of New Providence Island, 457 R. Robertson! 11 September 1955, ANSP 299657 (1 juv pair); Great Abaco Island and Green Turtle Key, Abaco Islands, Cherokee flats, Great Abaco and Mendelson’s flats, Heilman! DMNH 37728 (3 pair); Great Abaco Island and Green Turtle Cay, Cherokee flats, DMNH 37977 (5 pair); Great Abaco, Mendelson’s flats, Heilman! February 1958, DMNH 86223 (5 pair, 2 valves); Great Abaco, Parrot Cays, west of Elbow [Little Guana] Cay, near octopus hole, R. Robertson! 14 August 1953, ANSP 299066 (3 pair); Great Abaco, west coast of north end of Elbow [Little Guana] Cay, mud/sand, Halimeda remains, Thalassia seagrass, 0.5−3 ft (0.15− 0.9 m) and near octopus hole, R. Robertson! 04 September 1953, ANSP 298847 (1 pair, 1 juv pair); Abaco Island, Green Turtle Cay, Gwillim Bay, on sand on exposed sand bar at low tide, A. & A. Taxson! 10 June 1964, AMNH 111921 (4 pair) and AMNH 269540 (2 pair, ex D. Germer Collection); Abaco, Crab Cay, 2−4 ft (0.6−1.2 m), E. I. Wright! 1974, USNM 846377 (1 pair); Abaco, Marsh Harbor, O. Bryant! USNM 180541 (1 pair); east-central Eleuthera, north end of Half Sound, 25°07’45"N, 76°09’00"W, R. Robertson! 18 April 1984, ANSP 359292 (2 pair); Eleuthera, Savannah Island, Santy Point, W. J. Clench! May 1936, ANSP 173811 (3 pair); Eleuthera, Savannah Sound, Sandy Point, Cora Staples Collection, AMNH 306250 (1 pair); Eleuthera, Current, Current Club, A. Ross! 29 July 1963, AMNH 100174 (1 valve); Eleuthera Island, Doremus! DMNH 63778 (2 pair); Eleuthera Island, Sandy Point, Savannah Sound, Doremus! May 1936, DMNH 63780 (2 pair); Harbour Island, N end Eleuthera Island, Loc.114, Kline! 17 June 1949, DMNH72173 (1 pair); Eleuthera, N end Half Sound, E central Eleuthera Island, Abbott! June 1976 DMNH 115333 (1 valve); Exuma Island, Rolle Town, Loc.16, Kline! 11 July 1951, DMNH 72174 (1 pair); southern Exuma Cays, north of Leaf Cay, R. Robertson! 07 July 1957, ANSP 285738 (1 pair); Bimini, near Bailey Town, Bimini Lagoon, R. Robertson! 1957−1958, ANSP 326271 (1 valve); South Bimini, east of Nixon’s Harbour, R. Robertson! 1957−1958, ANSP 325603 (1 pair); Bimini, 1.75 mi southeast of Orange Cay, 23 ft (7.0 m), R. Robertson! 1957−1958, ANSP 325698 (1 juv valve); Bimini, South Cat Cay, grassy, T. L. Moise! ANSP 193577 (1 pair); Bimini, around Risty Causeway, Pi- 458 BIELER ET AL. geon Cay, Steger! April 1956, DMNH 107635 (1 pair); San Salvador Island, W Pigeon Creek, beach, Piech! February 1977, DMNH 143251 (1 pair). Turks and Caicos: Providenciales, Water Cay, beach, Piech! February 1978, DMNH 144123 (1 pair). Cuba: east of Tarallones de Arena, near Santiago, sand beach, R. E. Dickerson! ANSP 182932 (3 valves); Paradise Island, Oriente, Christofferson! 17 April 1949, FMNH 144071 (4 valves); west of Guardalavaca, eastern shore near Playa Esmeralda, Province Holguin, K. & Ch. Schniebs! December 2001, 1 pair, MTD 43828; Guantanamo, E. O. Mitchell et al.! 1930, USNM 405334 (1 valve); Cayo Hutia Reef, Barrera Expedition sta. 218, USNM 448345 (2 pair); Esperanza, 2−3 fms (3.6−5.5 m), Barrera Expedition sta. 210, USNM 448346 (1 pair); Varadero Beach, Barrera Expedition sta. 213, USNM 448348 (1 juv valve). Cayman Islands: Grand Cayman Island, Gun Bay, near Blakes’, mud and turtlegrass flats, A. J. Ostheimer III! ANSP 199513 (1 pair); Grand Cayman, North Sound, Jensen! August 1970, DMNH 39561 (1 pair). Jamaica: Harboreale, near Annotta Bay, St. Mary, Orcutt! USNM 440717 (1 valve); Black River, St. Elizabeth, Orcutt! USNM 441413 (1 valve). Hispaniola: Haiti, off Port-au-Prince, southeast side of Grand Bans, east side of reef, G. Goodfriend! 25 June 1972, AMNH 177703 (1 pair); Haiti, Cape Haitien, American Haitien Dev. Company, Krieger! USNM 487861 (1 valve); Santo Domingo, Monte Christi, W. J. Clench et al.! July 1937, ANSP 173105 (2 valves); Santo Domingo, Monte Cristi, Doremus! July 1937, DMNH 63777 (2 pair). Puerto Rico: Puerto Rico, Stearns! USNM 54091 (1 pair); El Deseches Island, Mayaguez B., F. A. Gallardo! USNM 464243 (1 valve); Bahia Bramadero [south of Mayaguez], G. L. Warmke! November 1956, ANSP 222755 (1 valve); Puerto Rico, Richardson, DMNH-85845 (2 pair). U.S. Virgin Islands: St. Thomas, W. A. Haines Collection, pre-1895, AMNH 31868 (9 pair, including largest recorded specimen); St. Thomas, M. Petit! USNM 250151 (1 pair); St. Thomas, Petit collection, USNM 530502 (1 pair, 2 valves, 1 juv valve); St. Thomas, Lindberg Beach, D. M. Barringer! 1936, ANSP 166919 (1 valve); St. Thomas, Swift Collection, ANSP 53580 (5 pair, 2 juv pair); St. Thomas, ZMB-104283 (1 pair); St. Thomas, ZMB-104287 (1 pair). British Virgin Islands: Peter Island, Little Harbour, R. H. Pine! July 1976, FMNH 197453 (1 valve); Seal Dog Islands, R. H. Pine! August 1976, FMNH 197476 (1 pair); Tortola, east of Roadtown, H. G. Richards! ANSP 244999 (1 juv valve); Tortola, Kjaer! USNM 3208 (1 pair); Tortola, K. Lamprell! 25 September 1980, AMNH 303476 (1 pair); Gorda Island, Colquhon Reef, BredinSmithsonian Expedition sta. 37−58, Schmitt et al.! 07 April 1958, USNM 735909 (1 pair). Antigua: Falmouth Harbor, beach, SUI Expedition, J. B. Henderson Jr.! 1918, USNM 500994 (1 pair). Grenada: South Grenada, Little Bacaye Harbor, silt, Thalassia seagrass, sand patches, R. Ostheimer and Buerk! 23 January 1964, ANSP 297064 (1 pair). Mexico: Isla Mujeres, K. C. Vaught Collection, AMNH 250781 (1 pair). Belize: east-southeast of Punta Negra, 16°16’15"N, 88°32’10"W, R. Robertson! 23 August 1961, ANSP 281836 (1 juv valve); north of Tarpon Cay, shallow Acropora cervicornis reef, 16°37’05"N, 88°09’05"W, R. Robertson! 17 August 1961, ANSP 282574 (1 valve); Glovers Reef, NE Cay, R. S. Houbrick! USNM 771205 (1 pair). Honduras: NW shore Bonacca Island, L. Kornicker! July 1963, USNM 667961 (4 valves). Costa Rica: Port Limon, Wailes! USNM 187276 (1 pair). Panama: Payardi Island, NW end, W. P. Woodring! 13 December 1959, USNM 67821 (1 valve); Payardi Island, NW end, 6 mi NE of Colon, suction dredge at refinery site, W. P. Woodring! 13 December 1959, USNM 637931 (1 valve); Payardi Island, Minas Bay, E of Colon, R. H. Stewart! USNM 734522 (3 valves). Colombia: Old Providence Island, Sid Anderson! January−March 1966, AMNH 138019 (1 pair); San Andres Island, S. Anderson! January 1966, AMNH 137541 (1 valve, 1 frag); vicinity of Cartagena, T. A. Link! USNM 364301 (3 valves); Cartagena, R. Pfaff! 1959, FMNH 78686 (1 pair). Netherlands Antilles: Aruba, M. R. Barnes! USNM 619369 (1 valve); Bonaire, Abbott! November 1972, DMNH 72707 (1 pair); Bonaire, Abbott! February 1973, DMNH 72794 (1 pair); Curaçao, N. Dearborn! 1908, FMNH 12764 (1 valve, subfossil?).

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