Periglypta listeri (J. E. Gray, 1838) (Bivalvia: Veneridae) in the western Atlantic: taxonomy,...

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 Atlanticvenerids, 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, thisspecies agrees with previously described venerids in having a well-developed escutcheonand lunule, and a hinge with three cardinal teeth in each valve. Within the genus, it is uniquein 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 thegenus Periglypta, and the most complete so far for any member of Venerinae. Periglypta

listeri agrees with previously described venerids in most anatomical characteristics, and no-tably features an undulating mantle edge that can close in “zipper” fashion, tentacles at theanterior mantle edge, and branching tentacles at the tips of the unfused siphons, type Bmantle fusion, type C(2) ctenidia, and a type V stomach. Although empty shells are commonlycollected, 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 solemorphological feature separating the subfamily Venerinae from the closely allied Chioninae.

Key Words: Florida Keys, Mollusca, Caribbean, infaunal, clam, sanctuary.

427

MALACOLOGIA, 2004, 46(2): 427−458

1Department of Zoology, Division of Invertebrates, Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, Illinois 60605-2496, U.S.A.2Division 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: [email protected]

INTRODUCTION

Veneridae is the largest marine family ofbivalves, with many of the more than 500 livingspecies forming key components in the world’sclam fisheries. The nominate subfamilyVenerinae currently comprises 14 genus-grouptaxa (e. g., Venus, Periglypta, Globivenus,Ventricoloidea; Keen, 1969) with more than 140nominal extant and fossil species. Members ofthis subfamily live in a wide range of benthichabitats, in coarse sand, mud, or gravel be-tween tide lines to depths over 150 m, and fromtemperate to tropical seas. Delimiting shell char-acteristics are the presence of both radial andconcentric sculpture and an anterior lateraltooth in the left valve (Keen, 1969; see hingediscussion below). The nominal subfamilyChioninae, another large group comprisingsuch genera as Chione, Mercenaria, and Proto-thaca, has recently been synonymized withVenerinae by some authors (Coan & Scott,1997; Coan et al., 2000). Relationships betweenthese and among other venerid subfamilies

remain unresolved, due to a surprising paucityof comparative morphological work. Despitetheir relative abundance and commercial im-portance, only about 50 venerid species havesome published anatomical data. Most publi-cations focus on a few species traditionallygrouped in the Chioninae, such as representa-tives 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), andClausinella (Ansell, 1961). Anatomical detailsfor members of other nominal venerid subfami-lies are much sparser, with data available forindividual 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,

BIELER ET AL.428

1919; Sellmer, 1967; Narchi, 1971), Meretricinae(Kellogg, 1915; Narchi, 1972; S. Gray, 1982;Narchi & Dario, 2002), Pitarinae (G. B. SowerbyII, 1854; Thiele, 1886; Pelseneer, 1911; Kellogg,1915; Morse, 1919; Narchi, 1971; F. R. Ber-nard, 1982; S. Gray, 1982; Fishelson, 2000;Morton, 2000), and Tapetinae (G. B. SowerbyII, 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 nominatesubfamily, Venerinae, remains largely unex-plored, with published anatomical data re-stricted to Circomphalus casina (Linnaeus,1758) (Ansell, 1961), Venus verrucosa Linna-eus, 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 fo-cuses on a venerine species currently classi-fied in the genus Periglypta Jukes-Brown, 1914,a group not previously studied anatomically.Periglypta listeri (J. E. Gray, 1838), also knownas “Lister’s venus” or “princess venus”, is thesecond largest Caribbean venerid species,only exceeded in shell size by Mercenariacampechiensis (Gmelin, 1791), which rangesfrom the mid-Atlantic coast to the Gulf of Mexicoand extends into the Caribbean. A shallow-wa-ter species with a very conspicuous shell, P.listeri had at one point even been declared thetype species of the genus Venus (Stoliczka,1871: xvii; Venus verrucosa Linnaeus, 1758,was subsequently fixed as the type by ICZNOpinion 195, 1954). Empty shells of P. listeriare commonly collected, but living specimensare less frequently encountered, due in part totheir relatively cryptic infaunal habitat inseagrass areas and algae-covered rubble nearreefs.

This paper reviews the taxonomy and geo-graphic distribution of this species, its anatomy,and life habits, based on original informationfrom living specimens from the Florida Keys,together with a re-evaluation of existing litera-ture and selected museum data. Comparisonsare drawn with sympatric large-bodied veneridsin the western Atlantic, with selected worldwidespecies of Periglypta, and with known anatomi-cal data for the family.

MATERIALS AND METHODS

This study is part of an ongoing investigationof marine molluscan biodiversity in peninsular

Florida and the Florida Keys, formally initiatedby RB and PMM in 1994. Consecutively num-bered stations comprising these collections arepreceded by an “FK” acronym in the followingtext. Living animals and empty shells were col-lected by hand mainly during scuba diving oncoral reefs and shallow-water (2−10 m) patchreefs, rubble areas, and ledges. The majorityof live observations were made on specimensfrom the “Horseshoe” site, bayside of WestSummerland Key (Spanish Harbor Keys), com-parable to stations IMBW-FK-629 and -637 re-ported by Mikkelsen & Bieler (2004). Interpre-tations of distribution records basedHenderson’s Eolis expeditions are taken fromthe recent compilation by Bieler & Mikkelsen(2003).

Specimen photography used a variety ofequipment and techniques. In situ photographsof living animals (Fig. 24) were taken using aNikonos V underwater camera with close-uplens. Other living animals (Figs. 25, 26) werephotographed in aquaria using standard 35 mmsingle-lens reflex or electronic cameras. Whole-valve and detail light micrography (Figs. 3−12,31, 32) used a Microptics® micro/macro imag-ing system based on a high-resolution Nikon®single-lens reflex digital camera. Excised pre-served tissues were prepared for scanning elec-tron microscopy (SEM) by critical point dryingand gold sputter coating, then viewed at beamacceleration voltages of 10kV on a Amray 1810scanning electron microscope at FMNH.

For anatomical observation, specimens wererelaxed by chilling in a household refrigeratorassisted by the addition of magnesium sulfatecrystals (Epsom salts) to their seawater sup-ply, or in an isotonic aqueous magnesium chlo-ride solution. Ciliary currents were studied usingcarmine particles. Anatomy was observed un-der a dissecting microscope; preserved tissueswere dyed for better contrast with neutral redor methylene blue. Voucher FK specimens werefixed in 5% formalin, later transferred to 70%ethanol, and are deposited in the Field Museumof Natural History (FMNH), Chicago, and theAmerican Museum of Natural History (AMNH),New York.

All measurements and meristics were takenfrom the left valve whenever possible. Shellmeasurements, taken with calipers or with ocu-lar micrometer on a stereomicroscope, include:maximum height from umbo to farthest distalpoint on free edge, and maximum length (=width) perpendicular to axis of height. Size isexpressed as shell length unless otherwisenoted. Radial ribs were counted at the growth

PERIGLYPTA LISTERI IN THE WESTERN ATLANTIC 429

edge on the main body of the shell. For mor-phometric analyses (length to height ratio), 32shells were measured and the linear regres-sion calculated using Microsoft® Excel 2000.

Other cited repositories include:

ANSP Academy of Natural Sciences ofPhiladelphia, Pennsylvania, U.S.A.

BMNH The Natural History Museum [= Brit-ish Museum (Natural History)], Lon-don, United Kingdom

BMSM Bailey-Matthews Shell Museum,Sanibel Island, Florida, U.S.A.

CMNH Carnegie Museum of Natural History,Pittsburgh, Pennsylvania, U.S.A.

DMNH Delaware Museum of Natural History,Wilmington, U.S.A.

MCZ Museum of Comparative Zoology,Harvard University, Cambridge, Mas-sachusetts, U.S.A.

MNHN Muséum National d’Histoire Natu-relle, Paris, France

MTD Staatliche Naturhistorische Sammlun-gen, Museum für Tierkunde, Dresden,Germany

NCSM North Carolina State Museum ofNatural Sciences, Raleigh, NorthCarolina, U.S.A.

NTM Northern Territory Museum of Arts andSciences, Darwin, Australia

SBMNH Santa Barbara Museum of NaturalHistory, Santa Barbara, California,U.S.A.

UMML Rosenstiel School of Marine and At-mospheric Science [= University ofMiami Marine Laboratory], Universityof Miami, Florida, U.S.A.

USNM National Museum of Natural History.[= United States National Museum],Smithsonian Institution, Washington,DC., U.S.A.

ZMB Museum für Naturkunde [= Zoolo-gisches Museum, Berlin], Humboldt-Universität, Berlin, Germany

Other abbreviations:

alc fluid-preserved (alcohol) specimenfrag shell fragmentjuv juvenile or subadultLV left valvepair an empty (dead) complete shell (2

valves)RV right valvespm a live-collected specimenvalve an empty (dead) single valve

RESULTS

Veneroidea: Veneridae: VenerinaeRafinesque, 1815: 146 (as Veneridia)

Periglypta Jukes-Brown, 1914: 72

Type species, by original designation: Venuspuerpera Linnaeus, 1758. Periglypta was in-troduced as a subgenus of Antigona Schu-macher, 1817, by Jukes-Brown (1914: 72).

Cytherea Röding (1798), non Fabricius, 1794(Diptera: Bombylidae). Type species (subse-quent designation of Dall, 1902): Venus puer-pera Linnaeus.

Periglypta listeri (J. E. Gray, 1838)

Selected synonymy:?”pectunculus admodum crassus …” Lister,

1687: Liber III, fig. 178 [pl. 341, fig. 178 inlater editions].

[unlabelled figure] − Lamarck, 1797: pl. 378,fig. 2a−b.

Venus puerpera (2) Var. − Lamarck, 1818:584−585 [referring to Lister (1687) andLamarck (1797) figures; non Venus puerperaLinnaeus, 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 (Philip-pines, Australia) erroneous]; − G. B. SowerbyII, 1853 [in part]: 705, pl. 152, fig. 8 [excludingfigs. 7, 9; distribution (Philippines, Australia)erroneous]; − Reeve, 1863: no. 14, pl. 5, fig.14 [“Hab. Philippine Islands; Cuming” errone-ous]; − Krebs, 1864: 97; − Pfeiffer, 1869: 141,pl. 8, figs. 8, 9 [distribution (Nicobares, Philip-pines) erroneous]; − Simpson, 1889: 64; − F.C. Baker, 1891: 47; − Cockerell, 1894: 118; −Benthem Jutting, 1927: 34; − McLean, 1936b:119; − Dance, 1974: 263−264, fig.; − Fischer-Piette, 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” inerror].

Venus crispata − Dall, 1889: 54 [non Venuscrispata Deshayes, 1853b].

Cytherea (Cytherea) listeri − Dall, 1902: 372.Cytherea listeri − Dall, 1903: 1275−1276,

1279; − Maury, 1920: 103.

BIELER ET AL.430

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 & PerezFarfante, 1942: 39 [Pleistocene]; − Jaume,1946: 101; − Aguayo & Jaume, 1949: 1; −Pulley, 1952: 150, pl. 13, fig. 7; − Nowell-Usticke, 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 [LowerPleistocene]; − 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 etal., 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 westernFlorida, including the Florida Keys, Texas, theWest Indies, Caribbean Central America andnorthern South America; apparently not reach-ing Brazil. It appears to have a largely Carib-bean island (as opposed to Gulf of Mexico ormainland North or South American) distribu-tion. It is rare off Texas and other Gulf of Mexicolocations. It extends along the entire islandchain of the Florida Keys, from Key Largo tothe Dry Tortugas.

Localities (*= unverified): *North Carolina (Pul-ley, 1952). Florida: east coast (AMNH, ANSP,FMNH, USNM; Dall, 1902; M. Smith, 1937; Pul-ley, 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), *westcoast (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), CaymanIslands (ANSP; Abbott, 1958), Jamaica (USNM),Hispaniola − Haiti and Dominican Republic(AMNH, ANSP, USNM; Palmer, 1927), PuertoRico (ANSP, USNM; Clench & McLean, 1937;McLean, 1951; Warmke & Abbott, 1961), VirginIslands (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).


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.

BIELER ET AL.432

Fossil Record

Lower Pleistocene (McGinty, 1970: 58) andUpper Pleistocene (M. Campbell, in lit., Feb.2003) of southern Florida; Pleistocene of Cuba(Jaume & Perez Farfante, 1942) and SantoDomingo [Hispaniola] (Dall, 1903). It was col-lected from Mayan archaeological sites (but notfrom concurrent Recent collections) in Yucatan,Mexico, by Parodiz (1976). Weisbord (1964: 302)noted that the species was “reported from thePleistocene of … Barbados, and the Island ofTortuga, Venezuela”, but did not include sourcecitations for these records; Weisbord examinedadditional fragments from the Lower Mare for-mation (Pliocene, possibly Lower Pliocene) atQuebrada Mare Abajo, northern Venezuela.

Cassab (1984) recorded the eastern PacificP. multicostata (G. B. Sowerby I, 1835) fromseveral Tertiary western Atlantic locations − theChipola Formation (Lower Miocene) of Florida,the Pirabas Formation (Lower Miocene) of ParaState, Brazil, and the Middle Miocene of CostaRica; verification of the species-level identityof these materials, which date prior to the clo-sure of the Panamanian landbridge, is war-ranted. All post-closure western AtlanticPeriglypta records should most definitely bereconsidered against P. listeri; Dall (1903: 1279)noted that P. multicostata “has been enumer-ated as one of the reef Pleistocene fossils ofSt. Domingo, but doubtless through a mis-identification, perhaps of [P.] listeri.” A large-shelled form described as P. tamiamensisOlsson & Petit, 1964, from Florida’s Late Mi-ocene and Pliocene Tamiami formation seemsclosely related. The relationship of P. listeri toextinct western Atlantic species from older geo-logical formations [e.g., Antigona dominicaPalmer, 1928 (= A. caribbeana Anderson,1927), Miocene of Santo Domingo, fide Hertlein& Strong, 1948; P. tarquinia (Dall, 1900), Oli-gocene of western Florida and Santo Domingo,called a small “precursor” of P. listeri by Dall,1903; P. mauryae (H. E. Vokes, 1938), UpperMiocene of Trinidad], awaits a more compre-hensive review of the genus.

Type Material

No original material located (see TaxonomicRemarks, below).

Material Examined

See Appendix 1.

Diagnosis

Large western Atlantic venerid, with thick-walled, inflated, trapezoid shell, with externalsculpture of erect commarginal ridges crenu-lated by underlying radial sculpture, and withposterior end vertically truncated. Exteriorcream-colored, with scattered brown speck-les, blotches, or flames. Interior yellowish whitewith more-or-less strongly developed purplishbrown stain around posterior adductor musclescar, posterior margin, and above the pallialline. Anterior lateral hinge tooth often with apurplish brown “hinge dot”.

Description

Shell relatively heavy, equivalve, longer thanhigh, trapezoid, with nearly straight dorsalmargin and bluntly truncated posterior margin;inequilateral with low, rounded umbones ap-proximately 1/3 of the shell length from theanterior end (Figs. 3, 4). Length to height ratiovery regular throughout lifespan (R2 = 0.9929,n = 32). Inflated, with posterodorsal slopesomewhat concave. External sculpture con-sisting of prominent erect commarginal ridges,regularly spaced, reflected umbonally, thosealong posterior margin higher and not re-flected; ridges more-or-less alternating instrength at anterior and posterior margins aswell as on later growth of shells (beginning at2.5−3 cm or after first 15−25 commarginalridges); ridges crenulated by underlying radialsculpture consisting of uniform flattened ribsseparated by narrow grooves approximately1/2−1/3 width of ribs (Fig. 5). Lunule broadlyspindle- to teardrop-shaped (Figs. 6, 13), withdeeply incised margins, asymmetric, right halfslightly larger than left, with many very finecommarginal lamellae (continuing the muchcoarser ridge pattern of the anterior shell),without radial elements. Escutcheon distinct(Fig. 6), delimited by a marginal groove, righthalf overlapping left in the posteriormost third,both halves finely obliquely grooved. Externallycream-colored with brown speckles, blotches,or flames, sometimes darker posterodorsally,occasionally radially merging into (often three)broadening radial stripes; escutcheon andlunule with color of surrounding shell. Internalmargin finely crenulated, continuing ontolunular margin. Anterior adductor muscle scaroval; posterior adductor muscle scar bean-shaped, somewhat flattened dorsally; poste-rior scar larger and more curved than anterior;


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: Umbonalaspect; 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 pearlwith 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).

BIELER ET AL.434

tor muscle scar and posterior margin, extend-ing dorsally above the pallial line, sometimesalso ventrally below, occasionally also sur-rounding anterior adductor and pedal retrac-tor muscle scars; additional faint purplish brown“lines” often extending toward umbo from an-terior limits of pallial sinus and anterior pedalretractor muscle scar (Figs. 7, 8). Live-collectedspecimens that are pure white internally alsonoted (e.g., USNM 464243, 890543, 890776).Color pattern within posterior adductor musclescar sometimes showing distinct growth lines(Fig. 7). Hinge teeth of left valve (Figs. 11, 12)comprising minute anterior lateral [AII] (seeDiscussion below), prominent triangular ante-rior cardinal (2a), slightly smaller bifid middlecardinal (2b; with posterior part 1/4−1/3 lessprominent), and lamellate posterior cardinal(4b). Hinge teeth of right valve (Figs. 10, 12)comprising well-defined, relatively small ante-rior cardinal (3a), larger bifid middle cardinal(1; with posterior part narrower and smaller),

FIGS. 10−12. Periglypta listeri, details of specimenin Figs. 7, 8. FIGS. 10, 11: Close-up of hinge teeth;FIG. 12: Articulated shells. (1, right middle cardinaltooth; 2a, left anterior cardinal tooth; 2b, left middlecardinal tooth; 3a, right anterior cardinal tooth; 3b,right posterior cardinal tooth; 4b, left posteriorcardinal tooth; HD, hinge dot; AII, left anteriorlateral tooth). Scale bars = 5 mm.

FIGS. 13, 14. Periglypta listeri, details of umboand prodissoconch; AMNH 296531, Looe Keyreef, Florida Keys, 9.5 mm. FIG. 13: Anterioraspect with lunule. Arrows point to transition fromwide commarginal lamellae on surface to finestriae on lunule; FIG. 14: Transition betweenprodissoconch I and II (arrow). Scale bars = 1mm (Fig. 13), 100 µm (Fig. 14).

dorsalmost portion of posterior scar formed bypedal retractor muscle scar, separated by faintdemarcation. Anterior pedal retractor musclescar on ventral side of hinge plate, dorsomedialto anterior adductor muscle scar, just belowanterior cardinal tooth (Fig. 12: 3a). Pallial lineentire (Figs. 7, 8); pallial sinus wide, roundlypointed anteriorly. Accessory pallial musclescars just inside pallial line, more-or-less regu-larly spaced. Internal color yellowish white withpurplish brown stain around posterior adduc-


FIGS. 15−18. Diagrammatic anatomy of Periglypta listeri. FIG. 15: Organs of the mantle cavity, withRV and most of right mantle removed. Arrows on gills and labial palps indicate direction of particleflow. 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 palpsremoved; 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 ofheart; 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, innerlabial 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 pedalretractor muscle; S, stomach; SS, style sack; V, ventricle of heart; VG, visceral ganglia).

BIELER ET AL.436

and prominent, wide, equally bifid posteriorcardinal (3b). Anterior lateral tooth at base ofanterior cardinal of LV, and its correspondingsocket 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 prodisso-conch II and juvenile shells (at about 1.3 mm).Juvenile sculpture (Figs. 13, 23) initiallysmooth, followed by fine commarginal ridgeswith wide bands of radial ribs between. Sur-face pattern of brown to orange speckles par-ticularly noticeable on juvenile shell (Fig. 23).

Foot large (Fig. 15), with brown pigment (inliving specimens) between muscular part andvisceral mass, with pedal gland opening slightlyanterior of midpoint; pedal groove extendingfrom ventral posterior end toward anterior third.Mantle muscles attaching mantle to shell, someextending dorsalward to produce so-calledaccessory muscle scars dorsal to pallial line

(approx. 1.5 mm dorsal, in larger specimens).Attachment of anteriorly pointed siphonal re-tractor muscles producing (and reflectingshape of) pallial sinus on shell. Anterior andposterior adductor muscles (AAM, PAM, re-spectively) oval; PAM slightly larger than AAM.Posterior pedal retractor muscle (PPRM)round, arising anterodorsal to PAM. Fibers ofPAM differently colored in living material, withposterior third darker than remainder. Anteriorpedal retractor muscle (APRM) attaching tounderside of anterior hinge plate and insertedinto anterodorsal part of foot a short distanceposterior to AAM. Visceral retractor muscleattached to inner surface of umbo behind hingeplate, inserting into roof of visceral mass.

Mantle with four mantle folds as previouslydescribed for other venerids (Ansell, 1961;Yonge, 1957). Ventral mantle margins distinctlywavy (Figs. 15, 20, 26) in living and preservedspecimens; capable of closing in “zipper” fash-ion and held closely appressed in living indi-

FIGS. 19−22. Periglypta listeri, scanning electron micrographs of critical-point dried tissues; AMNH295199, West Summerland Key, 73.2 mm. FIG. 19: Mid-ventral portion of the outer demibranch, withfood 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 mantlemargin with triangular tentacles. (IF, inner mantle fold; M1, inner middle mantle fold; M2, outer middlemantle fold; OF, outer mantle fold. Scale bars = 100 µm (Figs. 20, 22), 1 mm (Figs. 19, 21).


viduals at rest. Pedal gape extending ventrallyfrom base of incurrent siphon to AAM. Anterior1/8 of inner median mantle fold with small tri-angular tentacles (Figs. 21, 22). Siphons sepa-rated (Figs. 15, 26), each with distinct blackpigment (persisting in preserved specimens)between tentacles internally and externally; pig-ment stronger at terminal margin and fadingtoward basal region; internally with denselyplaced yellow-white surface papillae; each si-phon with terminal digitate tentacles and basal

siphonal membrane consisting of a thin tissueflap narrowing the lumen, with membrane ofincurrent siphon forming a double ridge.Siphonal mantle fusion of type B (Yonge, 1957,1982); union of inner and middle folds expos-ing outer surface of middle folds, with commonouter ring of sensory tentacles. Incurrent (ven-tral) siphon slightly larger in diameter and lengththan excurrent (dorsal) siphon (Figs. 24−26);excurrent siphon tapering to narrower terminaldiameter than wider, slightly flaring incurrent

FIGS. 23−26. Periglypta listeri, juvenile shell and living specimens. FIG. 23: Juvenile shell with colorpattern; FMNH 296719, West Summerland Key, Florida Keys, 7.2 mm; FIG. 24: Siphons of livingspecimen 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, 56mm; FIG. 26: Living specimen in laboratory tank; FMNH 295706, off Stirrup Key, Florida Keys, 59 mm.

BIELER ET AL.438

siphon. Siphonal tentacles digitate, those onincurrent siphon with 5−11, on excurrent with3−5, lateral papillae, interspersed with addi-tional small, simple tentacles; both kinds of ten-tacles on excurrent siphon with distal openingof unknown function (Figs. 29, 30); complextentacles on incurrent siphon (Figs. 24−28) 30−75% larger, slightly more pointed and morenumerous (approx. 40 tentacles of various sizesin 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 pointeddome-shaped valve surrounded by ring of ten-tacles, presumably allowing control of current(Figs. 24−26, 30).

Demibranchs smooth (not plicated; Fig. 15),with axes nearly vertical dorsoventrally; innerdemibranch slightly larger than outer, each withnumerous interlamellar junctions. Surface cur-rents moving particles ventrally on the outersurface of each demibranch; currents on inner

surfaces uncertain. Food grooves at distaledges of inner and outer demibranchs (Figs.18, 19), with oralward currents, indicating gillsand ciliation of type C(2) (Atkins, 1937); longi-tudinal oralward current also found betweenbases of adjacent demibranchs. Triangularpalps with narrow lamellae on inner surface(33−37 lamellae, n = 2; FK-273, AMNH295199, 73 mm; FK-352, FMNH 283534, 67mm); margins smooth at ventral and dorsalside; outer surface smooth. Acceptance cur-rents on palps along lamellae directed ven-trally, and on ventral boundary oralward.Rejection currents counter-oralward onsmooth ventral and dorsal margins of palps.Ctenidial/labial palp association of type II(Stasek, 1963); anteroventral tips of innerdemibranch inserted and fused to distal oralgroove between labial palps.

Esophagus relatively short, with 7−8 longi-tudinal rugae, leading into anterior part ofstomach (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 simpletentacle of excurrent siphon with orifice of unknown function; FIG. 30: Digitate tentacles on excurrentsiphon in front of flap-like valve (arrow). Scale bars = 1 mm (Fig. 27), 100 µm (Figs. 28, 30), 10 µm(Fig. 29).


gestive diverticula, located posterior to labialpalps. Stomach of type V (Purchon, 1985), withtwo caeca. Right caecum with seven ducts;left caecum with 11 ducts opening into diges-tive diverticula; four ducts of digestive diver-ticula opening into left pouch. Major typhlosole(MT) and intestinal groove extending from in-testine into stomach, penetrating right caecum;MT crossing stomach floor beneath esoph-ageal opening into left caecum; MT continu-ing from stomach into intestine, ending justafter gut loop. Minor typhlosole projecting intostomach but ending close to midgut opening.Gastric shield located opposite crystalline stylesac at roof and left side of stomach, withflanges that pass into dorsal hood and leftpouch. Dorsal hood with left and right sortingareas, located over esophageal opening. Stylesac combined with midgut, together exitingposteroventrally from posterior end of stom-ach, then turning anteriorly and coiling on ven-tral side of stomach, turning again posteriorlyand crossing style sac, then ascending (ashindgut, penetrating pericardium, ventricle andbulbus arteriosus) to continue dorsally andposteriorly across surface of PAM (Fig. 16).Gut loops varying somewhat (Fig. 17) in con-figuration between individuals, but always in-cluding a single ventral loop.

Ventricle of heart surrounding intestine, withlateral auricles connecting to outer limbs ofkidney (Fig. 16). Kidney positioned along ven-tral edges of pericardium and anterior to PAM(Fig. 16). Three pairs of ganglia (Fig. 16), withcerebral ganglia between APRM and AAM,joined by supraesophagal commissure; vis-ceral ganglia ventral and slightly anterior toPAM; pedal ganglia extensively fused,anteroventral to the ventral gut loop. Gonadsurrounding stomach and intestine within vis-ceral mass; reproductive system not otherwiseinvestigated.

Dimensions and Maximum Recorded Size

Median length approximately 65 mm, rang-ing 13−100 mm (mean 63.3 ± 14.7 mm SD, n =103). Maximum 100.2 mm, St. Thomas, VirginIslands; AMNH 31868 [registered specimen, K.Hutsell (San Diego, California), Registry ofWorld Record Size Shells]. Largest Florida Keysspecimen, 96 mm, FK-601, FMNH 296718.

Habitat and Ecology

Based on observations in southeasternFlorida’s Biscayne Bay, Stanley (1970: 160)

described this species as “widespread in in-tertidal and shallow subtidal settings, in largenumbers; it is generally restricted to coarsesubstrate and grassy areas”. Voss et al. (1969:71) reported the species, for the same region,from the highly unlikely habitat of “rocks; pil-ings; seawalls” as well as from seagrass beds,open sand, and lagoonal patch reefs. Otherhabitat records include sand (FMNH, thisstudy; 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 abun-dant” in Savannah Sound (Eleuthera, Baha-mas) but generally uncommon elsewhere. AtSavannah Sound, they lived buried 5−10 cmbelow the surface of seagrass-covered sandbars; native Bahamians were observed using“a short stick as a probe [to locate the clams]... [they are not] eaten by the natives thoughthey are gathered in this way for fish bait”(Clench & McLean, 1937: 37). In the presentstudy, one living specimen was found nestledin red algae within the cavity of a large barrelsponge at 8 m depth (FK-395), although thefavored habitat for the species seems to besand with loose rubble. It undoubtedly is ashallow-water species, in contrast to the analy-sis by Theroux & Wigley (1983: 47) who placedthis species based on two dredge records “inthe 50−99 m depth range grouping”. Collect-ing records of all (living and dead) materialrange from shallow water to 84 m (Theroux &Wigley, 1983), with the deepest confirmed live-collected specimen in this study from 8 m (FK-395). Stanley (1970: 160) described the slowburrowing process of this species and notedthat (in Biscayne Bay) the depth of burial is tosome extent dependent upon interference bythe subsurface rhizomes of surroundingturtlegrass (Thalassia testudinum König). Inthe laboratory, when permitted to burrow in itsnative sediment with the seagrass removed,a 5.8 cm-long animal assumed a position withthe posterior shell margin 4.5 cm beneath thesediment surface. Specimens observed in thepresent study were often found hindered fromdeep burrowing by rubble, and had their shellsbarely covered by substratum. In addition toan herbivorous diet, evidenced through thepresence of a crystalline style, P. listeri ap-pears to opportunistically ingest zooplankton:two copepods were found in the stomach of

BIELER ET AL.440

one individual (FK-352). In Puerto Rico, thisspecies is a “favourite food of the Slipper Lob-ster” (Sutty, 1990: 92). In the Florida Keys,empty shells have been frequently found as-sociated with octopus middens at scubadepths, or with beveled drill holes, the latterindicative of predation by naticid gastropods.

Pearls

A single living animal was found with a se-ries of loose pearls lining the center of the in-side shell, apparently in response to theremains of an intruding worm-shaped organ-ism (Fig. 9). The 78 mm specimen (FK-273,AMNH 295199) contained 11 irregular pearls,the largest with a maximum dimension of 5.8mm. This is the first record of pearls fromPeriglypta; both free and attached pearls havepreviously been reported from several othervenerids, especially Mercenaria spp. (Haas,1931; Shirai, 1994; Hill, 1996; Landman et al.,2001; Mienis, 2001).

Taxonomic Remarks

Although there is general consensus in therecent literature of applying the species name“listeri” to this western Atlantic taxon, the taxo-nomic history of that name is not without com-plications. Dosina listeri was introduced by J.E. Gray (1838: 308). He placed it in an un-named section of Dosina J. E. Gray, 1835,characterized by “anterior lateral tooth small,sometimes obliterated”, together with three pre-viously described species: Venus verrucosaLinnaeus, 1758 (as “Dosina veerrucosa [sic],Venus veerruicosa [sic], Linn.”); Venusreticulata Linnaeus, 1758; and Venus puerperaLinnaeus, 1771. While other species in thesame article were expressly identified as newspecies descriptions (labeled as “n.s.”, accom-panied by a textual description and includingan indication of the originating collection; e.g.,as done for Grateloupea cuneata J. E. Gray,1838: 304), the name D. listeri appears as anew name referring to prior literature data(1838: 308). The complete “description”, cryp-tic by today’s standards, introducing the namelisteri for a previously unnamed variety of Ve-nus puerpera Linnaeus, 1771, reads as follows:“Dosina Listeri, V. puerpura [sic] var., Linn. Sow.Gen. f. Ency. Meth. t. 278, f. 2”. Venus puer-pera was introduced by Linnaeus (1771: 545),in the Mantissa. Linnaeus himself did not men-tion varieties in the original description and re-ferred to two prior illustrations − Gualtieri (1742:

pl. 83, fig. F) and Argenville (1742: pl. 26, fig.F). In Hanley’s words (1855: 453): “Neither ofthe very dissimilar figures referred to bears theleast resemblance to the shell which has beenuniversally accepted for the species.” In anycase, Venus puerpera clearly is an Indo-Pa-cific, 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 citedfigure of Lister (1687), as well as to an illustra-tion by Lamarck (1797: pl. 278, fig. 2a, b) inthe Encyclopédie Méthodique.

J. E. Gray (1838) had named Dosina listerifor Martin Lister, author of the HistoriaConchyliorum (1685−1692). The first refer-ence to a figure by Lister in conjunction withVenus puerpera, and the first reference to adistinct variety of this species (as γ), appearedin the 13 th edition of Linnaeus’ Systemanaturae. The latter was authored by Gmelin(1791: sp. 3276), and it is the entry in this workthat Gray seemed to have meant with his “var.,Linn.”. The indicated Lister figure (1687: pl.341, fig. 178) is in agreement with today’s con-cept of Periglypta listeri, but lacks detail to dis-tinguish it from similar bivalve shells.

J. E. Gray’s (1838) introduction of the newname Dosina listeri included references to twoworks. As outlined above, he did not seem tohave actual specimens before him, and it ap-pears that only the specimens referred to inthese two works (and, possibly, Lister’s origi-nal material) qualify as the type series (ICZN,1999: Art. 72.4). One reference, and the onlyone indicated by Gray with actual plate andfigure numbers, was to Lamarck’s figures(1797: pl. 278, fig. 2a, b) as cited earlier byLamarck (1818) and Deshayes (1832) for “va-riety 2” of Venus puerpera. The latter are ex-cellent illustrations of an external right valveand of the external hinge area of an articu-lated specimen. Lamarck’s illustration matchestoday’s concept of western Atlantic Periglyptalisteri. Somewhat confusingly, Deshayes(1835: 334) stated that Lamarck’s specimenof “var. 2” and the figure 2a, b of theEncyclopédie fit well with the typical V. puer-pera of Linnaeus. Lamy & Fischer-Piette(1938: 292) agreed and considered all suchLamarck material in the MNHN collection torepresent V. puerpera [as well as (from a speci-men agreeing with figs. 1a, b of Encyclopédiepl. 278) Venus magnifica Hanley, 1845]. Theinterior features of the shell, particularly itscoloration, are unknown; the specimen onwhich Lamarck’s excellent illustrations were


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 Re-cent and fossil shells (1834). In it, figure 1 ofthe Venus plate shows detailed color illustra-tions of the inside valves of a species identi-fied as V. puerpera. However, the illustratedvalves lack the brown or purple stains usuallypresent in Periglypta puerpera and P. listeriand instead are shown with extensive orangecoloration below the umbo. The illustratedspecimen or specimens (considerable differ-ences in the shape of mantle line and musclescars indicate that these drawings of valvesmight originate from different individuals) havenot been located in the BMNH collection. Theorange interior coloration is matched by aspecimen of P. puerpera from J. E. Gray’s col-lection (BMNH 1991135, unknown locality, 49mm; seen by authors). However, this shell dif-fers considerably from Sowerby’s illustrationsin having dark brown markings at the shellmargin, oval (not angular) anterior musclescars, and a much more angular posteriorshoulder. Another specimen at BMNH, fur-nished 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 collec-tion; see Appendix), is a member of P. listeriin today’s sense, but likewise not a shell illus-trated by G. B. Sowerby I (1834). It is here, forreasons outlined above, not considered partof the syntypic series. Because the identity ofP. listeri is not currently in dispute, and the fig-ured specimens of Lamarck and Sowerby can-not be located at present, we refrain fromdesignating a lectotype (or neotype) in thecontext of this study and defer to a future ge-nus-wide revision.

Subsequent British authors, particularlyHanley (1843, in: 1842−1856: 110) and G. B.Sowerby II (1853: 705), referred to Venus listerias a species distinct from V. puerpera. How-ever, their concept of this nominal species wasbroader and included differently colored formswith dark rayed patterns on the shells, as evi-denced by the range of included illustrations.Venus listeri was thought to hail from the “In-dian Seas”, from the Philippines and Austra-lia. Nevertheless, G. B. Sowerby II (1853: 705)expressly endorsed Lamarck’s oft-cited illus-tration for V. listeri by stating “The figure in theEncyclopaedia is very exact for the type.”Reeve (1863: pl. 5, no. 14) narrowed the in-terpretation of V. listeri to shells that are“fulvous-white, obscurely freckled with flesh-

brown”, but still assumed the waters near thePhilippines Islands as its home. It was in thecontext of an Indo-Pacific interpretation of V.listeri that Reeve (1863: pl. 3) stated that hedoubted that eastern Pacific Periglyptamulticostata “is anything more than a varietyof V. listeri, in which the ribs are more timidlythickened and recurved.” Fischer-Piette (1975)assigned most of the Indo-Pacific records ofVenus/Dosinia/Chione/Cytherea/Antigonalisteri to P. puerpera and restricted P. listeri toAtlantic records.

Comparative Remarks

Periglypta listeri is one of the largest west-ern Atlantic venerids, second only to Merce-naria campechiensis in maximum adult size.Its sculpture, of flattened radial ribs betweensharp commarginal ridges (which are furthercrenulated by the radials), renders it distinctfrom other sympatric venerids: M. campe-chiensis lacks discernible sculpture betweenthe commarginals, whereas Globivenus (for-merly Ventricolaria) rugatina (Heilprin, 1886),G. rigida (Dillwyn, 1817), and Circomphalusstrigillinus (Dall, 1902) have fine concentricstriae between the commarginal ridges and arealso rounder in general shell shape.

Periglypta listeri is part of a worldwide com-plex of morphologically similar species, therelationships of which far exceed the scope ofthis paper. Römer (1867: 32, here translated)referred to similarities so great “that the differ-ences often can barely be put into words” whencomparing “Venus” listeri to other nominal spe-cies of Venus − Venus lacerata Hanley, 1845;V. clathrata Deshayes, 1853; V. crispataDeshayes, 1853; V. multicostata G. B. SowerbyII, 1853; V. laqueata G. B. Sowerby II, 1853;V. resticulata G. B. Sowerby II, 1853; V.chemnitzii Hanley, 1844; V. sowerbyiDeshayes, 1853; V. reticulata Linnaeus, 1758;and V. monilifera G. B. Sowerby II, 1851. Cit-ing a high degree of variability, E. A. Smith(1885: 120−121) considered “Venus” resti-culata, V. aegrota Reeve, 1863, V. lacerataHanley, 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, al-though he fell short of formalizing this actionin not placing these names in synonymy un-der V. puerpera. Modern revisionary treatmentof the Indo-Pacific taxa is urgently needed.

Periglypta listeri is the only living member ofthe genus in the Atlantic Ocean. It differs from

BIELER ET AL.442

the Indo-Pacific P. puerpera (Fig. 31), typespecies of the genus, in general shell shapeand coloration. Periglypta puerpera is lessanteroposteriorly elongated, and generallymore rounded in outline. Its external sculptureappears smoother, a result of its less promi-nent commarginal ridges. Externally it is cream-colored, with or without scattered radial brownflecks, often with 1−3 darker brown rays, onealmost always extensively covering the poste-rior third. Internally it is white with a bright purple(not purplish brown) stain at the posterior mar-gin below the PAM, sometimes also with a yel-low or peach-colored flush at the center(although internally pure-white specimens alsooccur). As earlier noted by E. A. Smith (1885:120−121), P. puerpera has a “V-shaped purplemark upon the apex of the umbones”, refer-ring to two radial color bands on theprodissoconch and earliest juvenile stage; thisis lacking in examined specimens of P. listeri.

Weisbord (1964: 302) called Periglypta listerithe western Atlantic analog of Panamic P.multicostata, citing a difference in the outlineof the posterior end (obliquely vertical and trun-cated in P. listeri, subtruncated and morerounded in P. multicostata; see Fig. 32); al-though based on many museum specimens(AMNH, n = 31), this difference is only obvi-ous among the largest specimens. Periglyptamulticostata attains a larger size (maximum

observed 120 mm, AMNH 248600, Baja Cali-fornia Norte, Mexico) than P. listeri. Externally,the commarginal ridges of P. multicostata aredecidedly coarser and more prominently dor-sally reflected than those of P. listeri, render-ing the crenulations and radial ribs lessobvious. Internally, P. multicostata is white (orvery occasionally flushed with pink, not pur-plish brown, either centrally or in an obliqueposterior streak from umbo to margin, but notprominently surrounding the posterior musclescars) and marginal crenulations are less no-ticeable or absent in the largest specimens(although the latter are quite prominent inspecimens < 65 mm).

DISCUSSION

Periglypta listeri agrees with previously de-scribed venerids in conchological features,such as a well-developed escutcheon andlunule, and a hinge with three cardinal teethin each valve (Keen, 1969). Although Lamprell(1998) cited differences in the shape of thepallial sinus, we found that distinguishing char-acters at the species level reside in sculptureand color/color pattern. So far as is known,the pattern of the internal purplish brown col-oration and the purplish brown “hinge dot” areunique to P. listeri within the genus.

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.


This paper presents the first anatomical studyfor any species in the genus Periglypta, and isarguably the most extensive anatomical de-scription yet available for any member of thesubfamily Venerinae. Anatomical informationis published for only three other venerines.Pelseneer (1894, 1897) provided minor detailsabout the anatomy of Venus verrucosa(tongue-shaped foot without byssus, very smalllabial palps, siphons more or less fused). Helater (Pelseneer, 1911) presented similarly cur-sory details (siphons short and united, withretractors; short “byssal” groove on posteriorfoot; very narrow external demibranch) forVenus (now Globivenus) toreuma. Ansell(1961: fig. 8) illustrated (but did not extensivelydiscuss) the gross anatomy and stomach struc-ture of Circomphalus casina (Linnaeus, 1758),which are closely similar to those in P. listeriwith respect to the gills, labial palps, foot, stom-ach, and adductor muscles, as well as the flowof particles over the gills and labial palps. LikeP. listeri, C. casina features a structurally com-plex ventral mantle edge, but its function wasnot mentioned. Its siphons appear short, un-like the longer, separated siphons of P. listeri.Ansell (1961; also Pelseneer, 1911) consideredthe degree of siphonal fusion to vary greatlywithin Veneridae, and from these minimal datathis could also be true within Venerinae.

The anatomy of Periglypta listeri agrees withthese and other previously described veneridsin most features. As elaborated by Yonge(1957), Ansell (1961), and Narchi (1971), themantle edge has four folds (outer, OF; innermiddle, M1; outer middle, M2; and inner, IF).The periostracum is secreted between OF andM2. In P. listeri, M1 is undulating, with its an-terior part elaborated into distinctly triangulartentacles. The wavy structure and tentaclescan close in “zipper” fashion (Fig. 26), presum-ably to protect the organs of the mantle cavityfrom intrusion of particles and small organisms.At the anterior end of the pedal gape, OF fuseswith M2, and M1 fuses with IF. At the posteriorend, M1 and IF are fused to form the siphons;this configuration typifies venerids of type Bfusion (Yonge, 1948, 1957, 1982; Ansell,1961). The conical valve on the tip of the ex-current siphon is the elaborated IF, while thering of tentacles on the siphon represents M1(Yonge, 1957; Ansell, 1961; Jones, 1979). Thetentacles on the incurrent siphon are not dis-tinguishable into inner and outer rings of ten-tacles, in contrast to Jones’ (1979) findings inmembers of Chione, Mercenaria, andAustrovenus. The siphonal tentacles of P.

listeri are long and digitate, serving as an ef-fective screen to block intrusion of particles.Ansell (1961) contrasted the digitate tentaclesof Circomphalus, Timoclea, Clausinella,Venerupis, and Gafrarium spp., which live ingravelly and stony bottom habitats, against thesimple tentacles of Chamelea and Dosiniaspp., which live in cleaner sand or gravellybottoms. Periglypta listeri is not congruent withthis pattern; it carries digitate tentacles but livesin a gravel and sand habitat without a highamount of detritus in suspension.

At the proximal end of each siphon,Periglypta listeri has a siphonal membrane orvalve comprised of thin tissue flaps, as de-scribed for other venerids (Ansell, 1961; Jones,1979). That of the incurrent siphon consists ofa basal double ridge; Ansell (1961) describedthis double ridge in Circomphalus casina and,like others (Kellogg, 1915; Jones, 1979; Narchi& Dario, 2002) postulated that such a valvecan be opened to admit water inflow, or closedto direct water ventrally and flush out accu-mulated pseudofeces.

The anterior end of the gill in Periglypta listeriis inserted into the distal oral groove, in themidline of the labial palps to which the gill isfused (Stasek, 1963). The ctenidia, with foodgrooves on the edges of both demibranchs,correspond to Atkins’ (1937) type C(2) alongwith members of Paphia, Mercenaria, Chione,Tivela, and Saxidomus. Other venerines, suchas Venus verrucosa and Circomphalus casina,as well as other venerids (in Clausinella,Dosinia, Gafrarium, Chamelea, and Paphia),lack a groove on the outer demibranch andhave 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 offour specimens of C. casina had a groove onthe outer demibranch), indicating that this char-acter requires larger sample sizes and furtherresearch.

The circulatory and nervous systems arebroadly similar to the species studied by Jones(1979), as is the general plan of the digestivesystem and configuration of the gut loop. Thetype V stomach of P. listeri agrees overall withthe 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 com-bined style sac and midgut is typical ofvenerids, except for Placamen tiara (Dillwyn,1817), in which the openings, although closetogether, lead into separate tubes (Dinamani,

BIELER ET AL.444

1967). The numbers of ducts entering the leftand right caeca vary among species andrange, respectively, from a minimum of oneand two in Nutricola tantilla (Gould, 1853) to amaximum of 13 and seven in Venerupispullastra (Montagu, 1803) (Purchon, 1987); P.listeri is at the high end of this range with 11and seven ducts. Four additional ducts openinto the left pouch in P. listeri, in contrast totwo or three in Meretrix, Katelysia, Placamen,Sunetta, and Irus spp. (Dinamani, 1967), fiveducts in Tivela and Gafrarium spp., or eight inDosinia spp. (Purchon, 1987). No additionalducts enter the stomach near the right cae-cum in P. listeri, unlike in Clausinella andCallista spp. (Purchon, 1987).

Periglypta listeri lives in rubble or sandamong rocks, and/or in reef settings (as op-posed to soft bottoms like most othervenerids). This is also true for the similar east-ern Pacific P. multicostata, aptly named “giantreef clam”, described as a common speciesin 3−6 m depth in Baja California Sur (García-Domínguez et al., 1998), and noted from sandamong rocks (Keen, 1971). Other Periglyptaspecies recorded for rocky or reef habitats areP. puerpera and P. reticulata (fide Whitehead,1983; Graham, 1995).

Periglypta was originally described as a sub-genus 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 commar-ginal shell ridges with Periglypta, its membershave radial ribs extended onto the lunule, nogroove around the escutcheon, a more trian-gular pallial sinus, and hinge teeth with a muchwider 3b and smaller 2b. Periglypta also dif-fers from Dosina J. E. Gray (type species, D.zelandica J. E. Gray, 1835, South Pacific, bysubsequent designation of Frizzell, 1936), inwhich P. listeri was originally described, by thepredominantly concentric sculpture (Keen,1969) and a weakly or undeveloped escutch-eon in that genus (pers. obs.).

Most authors (e.g., Keen, 1969) refer to ananterior lateral tooth in the left valve as char-acteristic of venerines, but an anterior lateraltooth is also present in members of othervenerid subfamilies (e.g., Gouldiinae [=“Circinae”], Sunettinae, Meretricinae, Pitarinae,and Dosiniinae; Keen, 1954). There are indi-cations that such lateral teeth result from dif-ferent ontogenetic pathways and might not be

homologous. Félix Bernard (1895: 127) pre-sented an ontogenetic series of hinge devel-opment for a Miocene species of Gouldia,which (together with its numbering system)became the model for tooth development inthe “corbiculoid” hinge type sensu Cox (1969:N54). According to this model, the venerineanterior lateral tooth of the left valve (desig-nated as AII) derives from lamella II. WhereasF. Bernard (1895: 127) noticed differing hingemorphologies of other venerids, such asMacrocallista, he still recognized the anteriorlateral tooth in the latter as a lamella II deriva-tive, and thus an AII. Marwick (1927: 598−599),in another ontogenetic comparison, showedthat the Oligocene venerine Kuia vellicata(Hutton, 1873) displayed the same AII devel-opment as shown in F. Bernard’s Gouldiastudy. However, Marwick (1927: 598) main-tained that the left anterior lateral tooth ofMacrocallista, a continuation of a low ridgeproceeding from below the umbo, is “in no wayconnected with the anterior cardinal”. Somesubsequent authors (Frizzell, 1936) followedMarwick’s argument of non-homologous an-terior lateral teeth in the Veneridae; Grant &Gale (1931: 316) referred to the venerine AIIof Antigona as a “pseudolateral”, in contrastto the situation in groups such as Macro-callista. It appears that venerid “anterior lat-eral teeth” might variously be derived fromeither lamella II or IV and homology assump-tions of these structures need closer scrutiny.

Chionines do not have anterior lateral orpseudolateral teeth even though their overallshell morphology is very similar to that ofvenerines. Anatomical studies carried out byJones (1979) showed that chionine siphonsare usually fused along their length in con-trast to the unfused siphons of Periglyptalisteri seen in this study; the high degree ofvariability in siphonal fusion, as well as thepresumably fused siphons in the venerineCircomphalus casina (see above) render thisa weak distinguishing character. At present,the presence of an AII lateral hinge tooth ispostulated as the only reliable morphologicalfeature separating these two nominal subfami-lies. Although recently synonymized by Coan& Scott (1997), preliminary molecular studies(16S gene; I. Kappner, unpubl.) indicate a dis-tinct grouping that might warrant retention ofthese subfamilial units. Studies of additionaltaxa, and more anatomical and molecularcharacters are needed for resolution ofsubfamilial synapomorphies.


ACKNOWLEDGMENTS

This work is part of an ongoing investigationof the molluscan diversity of the Florida Keys;regional surveys and collections were sup-ported by permits from the Florida Keys Na-tional Marine Sanctuary (080-98, 2000-036,2002-078, and 2002-079); in the vicinity ofPigeon Key (on National Register of HistoricPlaces) under the auspices of the Pigeon KeyFoundation; in the Dry Tortugas National Park,under collecting permits DRTO-19970030 and2002-SCI-0005; in Long Key State Park (LongKey, Florida Keys) under Florida Departmentof Environmental Protection permit 5-02-43;in Key West National Wildlife Refuge (nearSand Key, Florida Keys) under United StatesFish and Wildlife Service permit 41580-01-07.Additional collecting was sanctioned underFlorida Fish and Wildlife Conservation Com-mission permit 99S-024 to affiliates of TheBailey-Matthews Shell Museum (Sanibel,Florida) and permit 01S-056 (as well as an-nual permits for prior years of this study) toaffiliates 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 atFort 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 as-sistance. Data gathering from other museumcollections 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), MatthiasGlaubrecht and Lothar Maitas (ZMB). BernardMetivier (Museum National d’HistoireNaturelle, Paris), and Kathie Way (BMNH) pro-vided type specimen information and loans.Richard E. Petit, as so often before, helpedus disentangle obscure literature references.IK greatly appreciates Osmar Domaneschi’sadvice on anatomical questions and thanksGustav Paulay and Rudo von Cosel for veryfruitful discussions. The research, in part, wassupported by NSF-PEET DEB-9978119 andComer Science and Education Foundationgrants to RB and PMM. Additional fieldworksupport from the Bertha LeBus Charitable

Trust, Field Museum’s Womens Board, FieldMuseum’s Zoology Department’s MarshallField Fund and AMNH’s Proctor-Old-SageMalacology Fund is also gratefully acknowl-edged. Two anonymous reviewers providedhelpful comments on an earlier draft. This isSmithsonian Marine Station, Ft. Pierce, con-tribution number 603.

LITERATURE CITED

ABBOTT, R. T., 1954, American seashells. D. vanNostrand, Princeton, New Jersey. xiv + 541 pp.,40 pls.

ABBOTT, R. T., 1958, The marine mollusks ofGrand Cayman Island, British West Indies.Monographs of the Academy of Natural Sci-ences of Philadelphia, no. 11: 1−138, pls. 1−5.

ABBOTT, R. T., 1961, How to know the Ameri-can marine shells. New American Library, NewYork. 222 pp., 12 color pls.

ABBOTT, R. T., 1970, How to know the Ameri-can marine shells, revised ed. The New Ameri-can Library, New York. 222 pp., 12 pls.

ABBOTT, R. T., 1974, American seashells: themarine Mollusca of the Atlantic and Pacificcoasts of North America, 2nd ed. Van NostrandReinhold, New York. 663 pp., 24 pls.

ABBOTT, R. T., 1984, Collectible Florida shells.American Malacologists, Melbourne, Florida.64 pp.

ABBOTT, R. T., 1986, A guide to field identifica-tion: seashells of North America, rev. ed.Golden Press, New York. 280 pp.

ABBOTT, R. T. & P. A. MORRIS, 1995, A fieldguide to shells: Atlantic and Gulf coasts andthe West Indies, 4th ed. Houghton Mifflin Com-pany, Boston & New York. xxxiii + 350 pp., 74pls.

AGUAYO, C. G. & M. L. JAUME, 1949,Pelecypoda − Veneridae. Catalogo Moluscosde Cuba, no. 566, 1 p.

ALVAREZ, F. M. A., 1998, Shallow water mol-luscs (Gastropoda y Pelecypoda) from CayosCochinos Biological Reserve, Honduras.Moluscos (Gastropoda y Pelecypoda) deaguas someras, Reserva Biológica de CayosCochinos, Honduras. Revista de Biologia Tropi-cal , 46(suppl. 4): 103−107 [also http://www.ots.duke.edu/tropibiojnl/tbonline/Fascicul/r9846.htm; last accessed 07 January 2003].

ANSELL, A. D., 1961, The functional morphol-ogy of the British species of Veneracea(Eulamellibranchia). Journal of the Marine Bio-logical Association of the United Kingdom,41(1): 489−515.

ARGENVILLE, A. J. D. d’, 1742, L’histoirenaturelle éclaircie dans deux de ses partiesprincipales, la lithologie et la conchyliologie,don’t l’une traite des pierres et l’autre descoquillages … De Bure, Paris. [viii] + 491 pp.,36 pls.

ATKINS, D., 1937, On the ciliary mechanismsand interrelationships of lamellibranches. Part

BIELER ET AL.446

III. Types of lamellibranch gills and their foodcurrents. Quarterly Journal of MicroscopicalScience, 79: 375−421.

BAKER, F. C., 1891, Notes on a collection ofshells from southern Mexico. Proceedings ofthe Academy of Natural Sciences of Philadel-phia, 1891(1): 45−55.

BAKER, J. A., 1969, Shelling at St. George Cay,Bahamas. Seafari [Palm Beach County ShellClub newsletter], 11(11): 3−4.

BENTHEM JUTTING, T. VAN, 1927, Marinemolluscs of the Island of Curaçao. Bijdragentot de Dierkunde, 25: 1−36.

BERKELEY, C., 1959, Some observations ofcristispira in the crystalline style of Saxidomusgiganteus Deshayes and in that of some otherLamellibranchiata. Canadian Journal of Zool-ogy, 37(1): 53−58.

BERNARD, F., 1895, Première note sur ledéveloppement et la morphologie de la coquillechez les lamellibranches. Bulletin de la SociétéGéologique de France, (3)23: 104−154.

BERNARD, F. R., 1982, Nutricola n. gen. forTransennella tantilla (Gould) from the north-eastern Pacific (Bivalvia: Veneridae). Venus,41(2): 146−149.

BIELER, R. & P. M. MIKKELSEN, 2003, Thecruises of the Eolis − John B. Henderson’smollusc collections off the Florida Keys, 1910−1916. American Malacological Bulletin, 17(1−2) “2002”: 125−140.

BROOKS, J., 1968, Keys after “Abby”. Seafari[Palm Beach County Shell Club newsletter],10(7): 8.

BURNE, R. H., 1920, Mollusca Part IV. − Anatomyof Pelecypoda. British Antarctic (“Terra Nova”)Expedition, 1910. Natural History Report. Zo-ology, 2(10): 233−256, pls. 1−4.

CARRIÈRE, J., 1879, Die Drüsen im Fusse derLamellibranchiaten. Arbeiten aus demZoologisch-Zootomischen Institut in Würzburg,5: 56−92, pls. 5−6.

CASSAB, R. DE CASSIA TARDIN, 1984, Sobrea ocorrência de Periglypta multicostata naFormação Pirabas, Oligoceno-Mioceno doBrasil. Anais da Academia Brasileira deCiências, 56(3): 283−286.

CLENCH, W. J. & R. A. McLEAN, 1936, Marinebivalves collected by the Harvard-BahamaExpedition of 1935. Memorias de la SociedadCubana de Historia Natural “Felipe Poey”,10(3): 157−168.

CLENCH, W. J. & R. A. McLEAN, 1937, Marinebivalves from Little and Great Abaco, GrandBahama, and Eleuthera, Bahama Islands.Memorias de la Sociedad de Historia Natural,11(1): 31−42, pls. 5−6.

COAN, E. V. & P. H. SCOTT, 1997, Checklist ofthe marine bivalves of the northeastern PacificOcean. Santa Barbara Museum of Natural His-tory Contributions in Science, 1: (ii +) 28 pp.

COAN, E. V., P. VALENTICH SCOTT & F. R.BERNARD, 2000, Bivalve seashells of west-ern North America: marine bivalve mollusksfrom Arctic Alaska to Baja California. SantaBarbara Museum of Natural History Mono-graphs, no. 2: viii + 764.

COCKERELL, T. D. A., 1894, A list of the Brachio-poda, Pelecypoda, Pteropoda, and Nudibran-chiata of Jamaica, living and fossil. The Nauti-lus, 7(10): 103−107, 113−118.

COX, L. R. (with additions by C. P. NUTTALL &E. R. TRUEMAN), 1969, General features ofthe Bivalvia. Part N [Bivalvia], Mollusca 6, vol.1: N2−N129, in: R. C. MOORE, ed., Treatise onInvertebrate Paleontology. Geological Societyof America and University of Kansas,Lawrence, Kansas.

DALL, W. H., 1889, A preliminary catalogue ofthe shell-bearing marine mollusks and brachio-pods of the south-eastern coast of the UnitedStates, with illustrations of many of the species.Bulletin of the United States National Museum,no. 37: 221 pp., 74 pls. [December 26].

DALL, W. H., 1902, Synopsis of the familyVeneridae and of the North American Recentspecies. Proceedings of the United States Na-tional Museum, 26(1312): 335−412, pls. 12−16 [December 29].

DALL, W. H., 1903, Contributions to the Tertiaryfauna of Florida, with especial reference to thesilex-beds of Tampa and the Pliocene beds ofthe Caloosahatchie River, including in manycases a complete revision of the generic groupstreated of and their American Tertiary species.Part VI. Concluding the work. Transactions ofthe Wagner Free Institute of Science, Phila-delphia, 3(6): 1219−1654, pls. 48−60.

DANCE, S. P., 1974, The collector’s encyclope-dia of shells. McGraw-Hill Book Company, NewYork, etc. 288 pp.

DANCE, S. P. [German edition by R. VONCOSEL], 1977, Das große Buch derMeeresmuscheln . Verlag Eugen Ulmer,Stuttgart. 304 pp.

DESHAYES, G. P., 1832, Encylopédieméthodique. Histoire naturelle des vers. Vol-ume 3. Agasse, Paris. Pp. 595−1152 [for colla-tion, see Coan et al., 2000: 597, underBruguière].

DESHAYES, G. P., 1835, Histoire naturelle desanimaux sans vertebres, 2nd ed. Volume 6,Histoire des mollusques. J. B. Baillière, Paris.iv + 600 pp.

DESHAYES, G. P., 1853a, Catalogue of theConchifera or bivalve shells in the collection ofthe British Museum. Part I. Veneridae, Cyprin-idae and Glauconomidae. British Museum,London. 216 pp.

DESHAYES, G. P., 1853b, Descriptions of newspecies of shells in the collection of Mr.Cuming. Proceedings of the Zoological Soci-ety of London, 1853: 1−11.

DÍAZ MERLANO, J. M. & M. PUYANAHEGEDUS, 1994, Moluscos del CaribeColombiano: un catálogo ilustrado. Colcienciasy Fundación Natura Colombia, Santafe deBogota. 291 pp., 74 pls.

DINAMANI, P., 1967, Variation in the stomachstructure of the Bivalvia. Malacologia, 5(2):225−268.

EDWARDS, C. [E.], 1980, Convention shellingand other thoughts … Conchologists ofAmerica Bulletin, (21): 3, 8, 13.


EISENBERG, J. M., 1981, A collector’s guide toseashells of the world. McGraw-Hill Book Com-pany, New York, London. 239 pp., incl. 158 pls.

EKDALE, A. A., 1974, Marine molluscs from shal-low-water environments (0 to 60 meters) offthe northeast Yucatan coast, Mexico. Bulletinof Marine Science, 24(3): 638−668.

EMERSON, W. K. & M. K. JACOBSON, 1976,The American Museum of Natural History guideto shells; land freshwater, and marine, fromNova Scotia to Florida. Alfred Knopf, New York.482 + xviii pp., 47 pls.

ESPINOSA, J., J. ORTEA & Á. VALDÉS, 1994,Catálogo de los moluscos bivals [sic] recientesdel Archipiélago cubano. Avicennia, 2: 109−129.

FEMORALE, LTDA. [COLTRO, J., JR. & M.COLTRO], 2003, Shell photos: Veneridae −Periglypta listeri (Gray, 1838). http://w w w. f e m o r a l e . c o m . b r / s h e l l p h o t o s /detail.asp?photo=25542.jpg, last accessed 13January 2003.

FISCHER-PIETTE, E., 1975, Révision duVenerinae s. s. (Mollusques Lamellibranches).Mémoires du Muséum National d’HistoireNaturelle, Nouvelle Série, Série A, Zoologie,93: 1−64, pls. 1−8.

FISHELSON, L., 2000, Comparative morphologyand cytology of siphons and siphonal sensoryorgans in selected bivalve molluscs. MarineBiology, 137: 497−509.

FRIZZELL, D. L., 1936, Preliminary reclassifica-tion of veneracean pelecypods (1). Bulletin duMusée royal d’Histoire naturelle de Belgique,12(34): 1−84.

GARCÍA-DOMÍNGUEZ, F., B. P. CEBALLOS-VÁZQUEZ, M. VILLALEJO-FUERTE & M.ARELLANO-MARTÍNEZ, 1998, Reproductivecycle of the giant reef clam Periglypta multi-costata (Sowerby, 1835) (Pelecypoda:Veneridae) at Isla Espiritu Santo, Baja Califor-nia Sur, Mexico. Journal of Shellfish Research,17(4): 1009−1013.

GMELIN, J. F., 1791, Caroli a Linné … Systemanaturae per regna tria naturae, secundumclasses, ordines, genera, species, cumcharacteribus, differentiis, synonymis, locis.Editio Decima Tertia, Aucta, Reformata. G. E.Beer, Leipzig, 1(6): 3021−3909.

GODCHARLES, M. F. & W. C. JAAP, 1973,Fauna and flora in hydraulic clam dredge col-lections from Florida west and southeastcoasts. Florida Department of Natural Re-sources, Marine Research Laboratory, St. Pe-tersburg, Florida, Special Scientific Reports,40: 1−89.

GRANT, U. S., IV. & H. R. GALE, 1931, Cata-logue of the marine Pliocene and PleistoceneMollusca of California and adjacent regions;with notes on their morphology, classification,and nomenclature and a special treatment ofthe Pectinidae and the Turridae (including afew Miocene and Recent species). Memoirsof the San Diego Society of Natural History, 1:1036 pp., 32 pls.

GRAY, J. E., 1838, Catalogue of the species ofthe genus Cytherea, of Lamarck, with the de-

scription of some new genera and species. TheAnalyst; a Quarterly Journal of Science, Lit-erature, Natural History, and the Fine Arts, VIII(24), July 1838: 302−308.

GRAY, S., 1982, Morphology and taxonomy oftwo species of the genus Transennella (Bi-valvia: Veneridae) from western North Americaand a description of T. confusa sp. nov. Mala-cological Review, 15(1−2): 107−117.

GUALTIERI, N., 1742, Index TestarumConchyliorum quae adservantur in Museo N.Gualtieri ... Albizzini, Florence. xxiii pp., 110 pls.

GUÉRON, C. DE OLIVEIRA CASTRO & A. C.DOS SANTOS COELHO, 1989, Consi-deracoes taxonomicas e morfologia de Dosinia(Dosinia) concentrica (Born, 1778) (Mollusca,Bivalvia, Veneridae). Boletim Museu NacionalRio de Janeiro Zoologia, 334: 1−19.

GUERÓN, C. DE OLIVEIRA CASTRO & W.NARCHI, 2000, Anatomia funcional deProtothaca (Leukoma) pectorina (Lamarck)(Bivalvia: Veneridae). Revista Brasileira deZoologia, 17(4): 1007−1039.

HAAS, F., 1931, Bau und Bildung der Perlen.Academische Verlagsgesellschaft, Leipzig. 116pp.

HANLEY, S., 1842−1856, An illustrated and de-scriptive catalogue of Recent bivalve shells …with 960 figures by Wood and Sowerby, form-ing an appendix to the Index Testaceologicus.Williams & Norgate, London. xviii + 392 + 24pp., pls. 9−24.

HANLEY, S., 1855, Ipsa Linnaei Conchylia; theshells of Linnaeus, determined from his manu-scripts and collections ... Also, an exact reprintof the Vermes Testacea of the ‘SystemaNaturae’ and ‘Mantissa’. Williams & Norgate,London. 556 pp., 5 pls.

HARTE, M. E., 1998, Superfamily Veneroidea.Pp. 355−362, in: P. L. BEESLEY, G. J. B. ROSS & A.WELLS, eds., Mollusca: the southern synthe-sis. Fauna of Australia, Vol. 5, Part B. CSIRO,Melbourne.

HERTLEIN, L. G. & A. M. STRONG, 1948, East-ern Pacific expeditions of the New York Zoo-logical Society. XXXIX. Mollusks from the westcoast of Mexico and Central America. Part VI.Zoologica, Scientific Contributions of the NewYork Zoological Society, 33(13): 163−198, pls.1−2.

HILL, L., 1996, Shells: treasures of the sea. HughLauter Levin Associates, Hong Kong. 311 pp.

HUMFREY, M., 1975, Sea shells of the WestIndies: a guide to the marine molluscs of theCaribbean. Taplinger, New York. 351 pp., 32pls.

INTERNATIONAL COMMISSION ON ZOOLOGI-CAL NOMENCLATURE (ICZN), 1954, Opinion195: Designation, under the Plenary Powers,of a type species for the genus “Venus”Linnaeus, 1758 (Class Pelecypoda) in harmonywith accustomed usage. Opinions and Decla-rations Rendered by the International Commis-sion on Zoological Nomenclature, 3(pt. 14):191−198.

INTERNATIONAL COMMISSION ON ZOOLOGI-CAL NOMENCLATURE (ICZN), 1999, Interna-

BIELER ET AL.448

tional Code of Zoological Nomenclature, 4th ed.International Trust for Zoological Nomencla-ture, London. xxix + 306 pp.

JAUME, M. L., 1946, Moluscos marinos litoralesdel Cabo Catoche, Yucatán, México. Revistade la Sociedad Malacólogia “Carlos de laTorre”, 4(3): 95−110.

JAUME, M. L. & I. PÉREZ FARFANTE, 1942,Moluscos Pleistocenicos de la zona franca deMatanzas. Memorias de la Sociedad Cubanade Historia Natural “Felipe Poey”, 16(1): 37−44.

JOHNSON, C. W., 1934, List of marine Molluscaof the Atlantic coast from Labrador to Texas.Proceedings of the Boston Society of NaturalHistory, 40(1): 1−204.

JONES, C. C., 1979, Anatomy of Chionecancellata and some other chionines (Bivalvia:Veneridae). Malacologia, 19(1): 157−199.

JOSHI, M. C. & D. V. BAL, 1965a, Nervous sys-tem of clam Katelysia marmorata (Lam.). Jour-nal of the University of Bombay, 33(3/5): 1−7.

JOSHI, M. C. & D. V. BAL, 1965b, The circula-tory system of clam Katelysia marmorata(Lam.). Journal of the University of Bombay,33(3/5): 13−23.

JUKES-BROWN, A. J., 1914, A synopsis of thefamily Veneridae. Part I. Proceedings of theMalacological Society of London, 11: 58−74.

KEEN, A. M., 1969, Superfamily VeneraceaRafinesque, 1815. Part N [Bivalvia], Mollusca6, vol. 2: N670−N690, in: R. C. MOORE, ed.,Treatise on invertebrate paleontology. Geologi-cal Society of America and University of Kan-sas: Lawrence, Kansas.

KEEN, A. M., 1971, Sea shells of tropical WestAmerica. Stanford University Press, Stanford,California. 1064 pp.

KELLOGG, J. L., 1892, A contribution to ourknowledge of the morphology of lamelli-branchiate mollusks. Bulletin of the U.S. FishCommission for 1890: 389−436, pls. 7994.

KELLOGG, J. L., 1903, Feeding habits andgrowth of Venus mercenaria. New York StateMuseum Bulletin 71, Zoology 10: 28 pp., 4 pls.

KELLOGG, J. L., 1915, Ciliary mechanisms oflamellibranches with description of anatomy.Journal of Morphology, 26(4): 625−701.

KREBS, H., 1864, The West-Indian marine shellswith some remarks [a manuscript printed forcirculation among collectors, by H. Krebs,1864]. A republication (1947−1948) by W. J.Clench, C. G. Aguayo and R. D. Turner, withremarks and a brief account of the life of H.Krebs. Revista de la Sociedad Malacólogia“Carlos de la Torre”, 5(1): 23−40, 5(2): 59−80,5(3): 91−116, 6(1): 11−43, 6(2): 45−48.

LAMARCK, J. B. P. A., 1797, [plates 190−286] inTableu Encyclopédie et Methodique des triosRègnes de la Nature. Vers, Coquilles,Mollusques et Polypiers. Pp. I−viii, 1−83 [83−84], 85−180. Agasse, Paris [for collation, seeCoan et al., 2000: 597, under Bruguière].

LAMARCK, J. B. P. A., 1818, Histoire naturelledes animaux sans vertèbres, 5: 612 pp.

LAMPRELL, K., 1998, Periglypta revisited.Australasian Shell News, 98: 6.

LAMY, E., 1929, Notes sur quelques Lamellibran-ches de la Martinique. Bulletin du Muséum Na-tional d’Histoire Naturelle, Paris, ser. 2, 1(3):201−208.

LAMY, E. & E. FISCHER-PIETTE, 1938, Notessur les espèces Lamarckiennes d’Antigona(Moll. Lamellibr.). Bulletin du Muséum Nationald’Histoire Naturelle (2), 10(3): 292−297.

LANDMAN, N. H., P. M. MIKKELSEN, R. BIELER& B. BRONSON, 2001, Pearls: a natural his-tory. Harry N. Abrams Inc., New York. 232 pp.

LAWSON, B., 1993, Shelling San Sal. BahamianField Station, San Salvador, Bahamas. 63 pp.

LERMOND, N. W., 1936, Check list of Floridamarine shells. Norman W. Lermond, Gulfport,Florida. 56 pp.

LINNAEUS, C., 1771, Mantissa plantarum alteragenerum editionis VI. & specierum editionis II.Holmiae [Stockholm], L. Salvii, 1771. [i−vi] +143−588. (containing also “Regni AnimalisAppendix”, pp. 521−552).

LIPE, R. E. & R. T. ABBOTT, 1991, Living shellsof the Caribbean and Florida Keys. AmericanMalacologists, Melbourne, Florida. 80 pp.

LISTER, M., 1685−1692, Historiae Conch-yliorum. M. Lister, London [only figures num-bered; pages and plates unnumbered in 1stedition; for a collation see: G. L. Wilkins, 1957].

LOZET, J. B. & C. PÉTRON (English version: W.REED & S. REED), 1977, Shells of the Carib-bean. Les Éditions du Pacifique, Tahiti. 138 pp.

LYONS, W. G. & J. F. QUINN, JR., 1995, Appen-dix J. Marine and terrestrial species and algae:Phylum Mollusca. Pp. J10−J26, in: Florida KeysNational Marine Sanctuary draft managementplan/environmental impact statement, Vol. III.Appendices. National Oceanographic and At-mospheric Administration, Silver Spring, Mary-land.

MAGNOTTE, G., 1970−1979? (various versions,all undated), Shelling & beachcombing insouthern & Caribbean waters. InternationalGraphics, Hollywood, Florida, etc. 96 pp.

MARWICK, J., 1927, The Veneridae of NewZealand. Transactions and Proceedings of theNew Zealand Institute, 57: 567−636.

MAURY, C. J., 1920, Recent molluscs of the Gulfof Mexico and Pleistocene and Pliocene spe-cies from the Gulf States. Part I: Pelecypoda.Bulletins of American Paleontology, 8(34): 35−147, pl. 1.

McGINTY, T. L., 1970, Mollusca of the “Glades”unit of southern Florida: Part I. Introduction andobservations. Tulane Studies in Geology andPaleontology, 8(2): 53−68.

McGINTY, T. L. & M. M. NELSON, 1972, Mol-lusca dredged off Pompano Beach, Florida, in10 fathoms. Unpublished report, 14 pp.; cop-ies on file in AMNH and FMNH Division of In-vertebrate Zoology libraries.

McLEAN, R. A., 1936a, A list of bivalves fromnorthern Cuba. Memorias de la SociedadCubana de Historia Natural, 10(1): 39−42.

McLEAN, R. A., 1936b, Some marine bivalvesfrom the Bahama Islands. The Nautilus, 49(4):116−119.


McLEAN, R. A., 1951, The Pelecypoda or bivalvemollusks of Porto Rico, and the Virgin Islands.Scientific Survey Puerto Rico, New York Acad-emy of Sciences, 17(1): 1−183, 26 pls.

MIENIS, H. K., 2001, A blister pearl formation inPaphia cor (Sowerby, 1853) from the ArabianSea off Pakistan. De Kreukel, 37(6): 108.

MIKKELSEN, P. M. & R. BIELER, 2000, Marinebivalves of the Florida Keys: discoveredbiodiversity. In: E. M. HARPER, J. D. TAYLOR & J.A. CRAME, eds., The evolutionary biology of theBivalvia [Proceedings of “Biology & Evolutionof the Bivalvia”, an international symposiumorganized by the Malacological Society of Lon-don, 14−17 September 1999, Cambridge, UK],Geological Society, London, Special Publica-tion 177: 367−387.

MIKKELSEN, P. M & R. BIELER, 2004, Interna-tional Marine Bivalve Workshop 2002: Intro-duction and summary. In: R. BIELER & P. M.MIKKELSEN, eds., Bivalve studies in the FloridaKeys, Proceedings of the International MarineBivalve Workshop, Long Key, Florida, July2002. Malacologia, 46(2): 241−248.

MÖRCH, O. A. L., 1853, Catalogus conchyliorumquae reliquit D. Alphonso d’Aguirra & GadeaComes de Yoldi, Regis Daniae CubiculariorumPrinceps, Ordinis Dannebrogici in PrimaClasse & Ordinis Caroli Tertii Eques. Fascicu-lus secundus. Acephala. Annualata Cirrepedia.Echinodermata. L. Klein, Copenhagen. [iv +]74 [+ ii] pp.

MORRIS, P. A., 1973, A field guide to shells ofthe Atlantic and Gulf coasts and the WestIndies, 3rd ed. Houghton Mifflin, Boston. 330pp., 76 pls.

MORSE, E. S., 1919, Observations on livinglamellibranchs of New England. Proceedingsof the Boston Society of Natural History, 35(5):139−196.

MORTON, B., 1985, Aspects of the biology andfunctional morphology of Irus irus (Bivalvia:Veneridae: Tapetinae) with a comparison ofBassina calophylla (Chioninae). Pp. 321−336,in: B. S. MORTON & D. DUDGEON, eds., Themalacofauna of Hong Kong and southernChina. II, Vol. 1. Hong Kong University, HongKong. viii + 361 pp.

MORTON, B., 2000, The anatomy of Callocardiahungerfordi (Bivalvia: Veneridae) and the ori-gin of its shell camouflage. Journal of Mollus-can Studies, 66: 2130.

MOULDING, M. B., 1967, Shells at our feet: anintroduction to shelling in the Bahamas. SeaScapers, Chicago. 102 pp.

NARCHI, W., 1971, Structure and adaptation inTransennella tantilla (Gould) and Gemmagemma (Totten) (Bivalvia: Veneridae). Bulletinof Marine Science, 21(4): 866−885.

NARCHI, W., 1972, Comparative study of thefunctional morphology of Anomalocardiabrasiliana (Gmelin, 1791) and Tivelamactroides (Born, 1778) (Bivalvia, Veneridae).Bulletin of Marine Science, 22(3): 643−670.

NARCHI, W., 1980, On the biology of Veremolpascabra (Hanley 1845) (Bivalvia: Veneridae)from the South China Sea. Pp. 277−289, in: B.

MORTON, ed., The malacofauna of Hong Kongand southern China. Proceedings, First Inter-national Workshop on the Malacofauna ofHong Kong and Southern China, 23 March−8April 1977. Hong Kong University Press for theDepartment of Zoology, University of HongKong, Hong Kong.

NARCHI, W. & F. DI DARIO, 2002, The anatomyand functional morphology of Tivela ventricosa(Gray, 1838) (Bivalvia: Veneridae). The Nauti-lus, 116(1): 13−24.

NARCHI, W. & M. A. GABRIELI, 1980, Sobreanatomia funcional de Chione subrostrata(Lamarck, 1818). Revista Nordestina deBiologia, 3(Especial): 25−46.

NIELSEN, B. J., 1963, Studies on the genusKatelysia Römer 1857 (Mollusca, Lamellibran-chiata). Memoirs of the National Museum ofVictoria, 26: 219−257.

NOWELL-USTICKE, G. W., 1959, A check list ofthe marine shells of St. Croix U.S. Virgin Is-lands with random annotations. G. W. Nowell-Usticke, Christiansted, St. Croix. vi + 90 pp.

ODHNER, N., 1912, Morphologische undphylogenetische Untersuchungen über dieNephridien der Lamellibranchien. Zeitschrift fürWissenschaftliche Zoologie, 100: 287−391.

PALMER, K. V. W., 1927, 1929, The Veneridaeof eastern America, Cenozoic and Recent.Palaeontographica Americana, 1(5): 209−522(1927), pls. 32−76 (1929).

PARODIZ, J. J., ca. 1976, Marine Mollusca col-lected in Yucatan. Unpublished report, 20 pp.,map; copies on file in AMNH and FMNH Divi-sion of Invertebrate Zoology libraries.

PELSENEER, P., 1894, Introduction a l’étude desmollusques. Henri Lamertin, Brussels. 216 pp.[+ 1 p. errata].

PELSENEER, P., 1897, Fascicule XVI.Mollusques. 187 pp., in: R. BLANCHARD, ed.,Traité de Zoologie. Rueff et Cie, Paris.

PELSENEER, P., 1911, Les lamellibranches del’expedition du Siboga. Partie anatomique.Siboga-Expeditie, 53a. E. J. Brill, Leiden. 125pp., 26 pls.

PELSENEER, P., 1923, Variations dans lesmollusques. Annales de la Société RoyaleZoologique de Belgique, 54: 68−78.

PELSENEER, P., 1931, Quelques particularitésd’organisation chez des Pectinacea. Annalesde la Société Royale Zoologique de Belgique,61: 12−17.

PFEIFFER, L., 1869, Die Familie derVenusmuscheln, Veneracea. Nebst einemAnhange, enthaltend die Chemnitz’schenLucinen, Galateen und Corbis. In: H. C. KÜSTER,ed., Systematisches Conchilien-Cabinet vonMartini und Chemnitz, XI(1), 302 pp., 42 pls.

POINTIER, J.-P. & D. LAMY, 1998, Guide descoquillages des Antilles. PLB Editions,Guadeloupe. 225 pp.

PRIETO, A. S., L. J. RUIZ, N. GARCÍA & M.ALVAREZ, 2001, Diversidad malacológica enuna comunidad de Arca zebra (Mollusca:Bivalvia) en Chacopata, Estado Sucre, Ven-ezuela. Revista de Biología Tropical, 49(2):591−598 [also http://rbt.ots.ac.cr/revistas/49-2/

BIELER ET AL.450

prietor/prietor.html, last accessed 07 January2003] + online supplement at http://rbt.ots.ac.cr/revistas/49-2/prietor/cuadro1.html[last accessed 07 January 2003].

PULLEY, T. E., 1952, A zoogeographic studybased on the bivalves of the Gulf of Mexico.Ph.D. Dissertation, Harvard University, Cam-bridge, Massachusetts. 215 pp., 19 pls.

PURCHON, R. D., 1985, Studies on the internalstructure and function of the stomachs of bi-valve molluscs: stomach types III, IV and V.Pp. 337−361, in: B. S. MORTON & D. DUDGEON,eds., The malacofauna of Hong Kong andsouthern China. II, Vol. 1. Hong Kong Univer-sity, Hong Kong. viii + 361 pp.

PURCHON, R. D., 1987, The stomach in theBivalvia. Philosophical Transactions of theRoyal Society of London, series B, BiologicalSciences, 316: 183−276.

RAFINESQUE, C. S., 1815, Analyse de la na-ture, ou tableau de l’univers et des corpsorganisés. Palermo. 224 pp.

REDFERN, C., 2001, Bahamian seashells: athousand species from Abaco, Bahamas.Bahamianseashells.com, Boca Raton, Florida.ix + 280 pp.

REEVE, L. A., 1863−1864, Monograph of thegenus Venus. Pls. 1−26 (with captions and in-dex), in: L. A. REEVE, ed., Conchologia iconica;or, illustrations of the shells of molluscous ani-mals, vol. 14 [pls. 2−11: April 1863; pls. 12−23:June; pl. 1: Feb. 1864; pls. 24−26: March 1864].

RICE, W. & L. S. KORNICKER, 1962, Mollusksof Alacran Reef, Campeche, Mexico. Publica-tions of the Institute of Marine Science, Uni-versity of Texas, 8: 366−403, 9 pls.

ROMASHKO, S., 1974, The shell book, 2nd ed.Windward Publishing, Miami, Florida. 64 pp.

ROMASHKO, S., 1984, The complete collector’sguide to shells & shelling. Windward Publish-ing, Miami, Florida. 112 pp.

RÖMER, E., 1867, Kritische Uebersicht aller zumSubgenus Chione gehörenden Arten von Ve-nus. Malakozoologische Blätter, 14: 28−62.

ROSS, B., 1969, Field trip to the Keys. Seafari[Palm Beach County Shell Club newsletter],11(7): 8−10.

SCHRAMM, A., 1869, Catalogues des coquilleset des crustacés de la Guadeloupe envoyés al’Exposition Universelle de 1867 parl’Administration de la Colonie. CollectionsCaillet et I. Desbonne. Déterminations etClassements, 2nd ed. Basse-Terre. 27 pp.

SELLMER, G. P., 1967, Functional morphologyand ecological life history of the gem clam,Gemma gemma (Eulamellibranchia:Veneridae). Malacologia, 5(2): 137−223.

SHIRAI, S., 1994, Pearls and pearl oysters ofthe world. Marine Planning Company, Okinawa,Japan. 108 pp.

SIMPSON, C. T., 1889, Contributions to the Mol-lusca of Florida. Proceedings of the DavenportAcademy of Natural Sciences, 5: 45−72.

SMITH, E. A., 1885, XXXV. Report on theLamellibranchiata. Report of the Scientific Re-sults of the Voyage of H. M. S. Challenger dur-

ing the years 1873−76 … Zoology, Vol. XIII,part 1, 341 pp., 25 pls.

SMITH, M., 1937, East coast marine shells.Edwards Brothers, Ann Arbor, Michigan. 308pp., 74 pls., 1 map.

SMITH, M., 1940, World-wide sea shells. Tropi-cal Photographic Laboratory, Lantana, Florida.xviii + 139 pp. [pp. xi−xviii by J. L. Baily].

SOWERBY, G. B. [I], 1834, Venus. 4 pp., 2 pls.In: The Genera of Recent and fossil shells, forthe use of students in conchology and geol-ogy. Part 41 [text and plates unnumbered; re-issued in 1875 with title page, index, and withmost of the plates numbered; collation inpreparation by R. E. Petit, February 2003, inlit.].

SOWERBY, G. B. [II], 1853, Monograph of thegenus Venus ; Supplement; index [toVeneridae]. In: G. B. SOWERBY II, ed., Thesau-rus conchyliorum; or, monographs of generaof shells, 2(14): 703−762, pls. 152−163.

SOWERBY, G. B. [II], 1854, Popular British con-chology. A familiar history of the molluscs in-habiting the British Isles. Lovell Reeve, London.304 pp., 20 pls.

STANLEY, S. M., 1970, Relation of shell form tolife habits of the Bivalvia (Mollusca). The Geo-logical Society of America, Memoir, 125: xiii +296 pp.

STASEK, C. R., 1963, Synopsis and discussionof the association of ctenidia and labial palpsin the bivalved Mollusca. The Veliger, 6(2): 91−97.

STOLICZKA, F., 1870−1871, Cretaceous faunaof southern India, Vol. III: The Pelecypoda,with a review of all known genera of this class,fossil and recent, … Memoirs of the Geologi-cal Survey of India, Palaeontologica Indica,xxii + 538 pp., 50 + 4 pls. (1−222, pls. 1−12 +4, 1 Sept. 1870; 223−408, pls. 13−28, 1 March1871; xxii + 409−538, pls. 29−50, 1 Aug.1871).

SUTTY, L., 1990, Seashells of the Caribbean.MacMillan Press, London. vi + 106 pp., figs.

THIELE, J., 1886, Die Mundlappen derLamellibranchiaten. Zeitschrift für Wissen-schaftliche Zoologie, 44: 239−272, pls. 17−18.

THEROUX, R. B. & R. L. WIGLEY, 1983, Distri-bution and abundance of east coast bivalvemollusks based on specimens in the NationalMarine Fisheries Service Woods Hole Collec-tion. NOAA Technical Report NMFS SSRF-768:i−xvi, 1−172.

TREMOR, M. E., JR., 1998, A shelling trip to KeyWest. Shell-o-gram [Jacksonville Shell Club,Jacksonville, Florida], 39(5): 7−9 [orig. pub-lished in Tidelines (St. Petersburg Shell Club),Sept. 1998, not seen].

TURGEON, D. D., J. F. QUINN, JR., A. E.BOGAN, E. V. COAN, F. G. HOCHBERG, W.G. LYONS, P. M. MIKKELSEN, R. J. NEVES,C. F. E. ROPER, G. ROSENBERG, B. ROTH,A. SCHELTEMA, F. G. THOMPSON, M.VECCHIONE & J. D. WILLIAMS, 1998, Com-mon and scientific names of aquatic inverte-brates from the United States and Canada:


Mollusks, 2nd ed. American Fisheries Society,Special Publication 26, Bethesda, Maryland.526 pp.

VOKES, H. E. & E. H. VOKES, 1984 (“1983”),Distribution of shallow-water marine Mollusca,Yucatan Peninsula, Mexico. MesoamericanEcology Institute Monograph 1. Middle Ameri-can Research Institute, Publication 54, 183 pp.,50 pls.

VOSS, G. L., F. M. BAYER, C. R. ROBINS, M.GOMON & E. T. LAROE, 1969, The marineecology of the Biscayne National Monument[Report to the National Park Service, Depart-ment of the Interior]. Institute of Marine andAtmospheric Sciences, University of Miami,Miami, Florida. iv + 128 pp., 40 figs.

VOSS, G. L., N. A. VOSS, A. Y. CANTILLO & M.J. BELLO, 1983, An environmental assessmentof the John Pennekamp Coral Reef State Parkand the Key Largo Coral Reef Marine Sanctu-ary. Joint NOAA/University of Miami Report.NOAA Technical Memorandum NOS NCCOSCCMA 161. NOAA LISD Current References2002-6. Rosenstiel School of Marine and At-mospheric Science, University of Miami, Mi-ami, Florida. 452 pp. [2002 edited versionavailable at http://www.aoml.noaa.gov/general/lib/cedardoc.html, last accessed 04 April 2003].

WARMKE, G. L. & R. T. ABBOTT, 1961, Carib-bean seashells. Dover Publications, New York.348 pp., 44 pls.

WEBER, J. A., 1961, Marine shells of Water Is-land, Virgin Is. The Nautilus, 75(2): 55−60.

WEISBORD, N. E., 1926, Notes on marine mol-lusks from the Yucatan Peninsula, Mexico. TheNautilus, 39(3): 81−87.

WEISBORD, N. E., 1964, Late Cenozoic pele-cypods from northern Venezuela. Bulletins ofAmerican Paleontology, 45(204): 564 pp., 59pls.

WHITEHEAD, T., 1983, The genus PeriglyptaJukes-Brown 1914 in Australia. Australian ShellNews, no. 41: 4−5.

WILKINS, G. L., 1957, Notes on the HistoriaeConchyliorum of Martin Lister (1638−1712).The Journal of the Society for the Bibliographyof Natural History, 3(4) 196−205.

WOODS, E., 1970, June Keys field trip. Seafari[Palm Beach County Shell Club newsletter],12(10): 2−4.

YONGE, C. M., 1948, Formation of siphons inLamellibranchia. Nature, 161(4084): 198−199.

YONGE, C. M., 1957, Mantle fusion in theLamellibranchia. Pubblicazioni della StazioneZoologica di Napoli, 29: 151−171.

YONGE, C. M., 1982, Mantle margins with a re-vision of siphonal types in the Bivalvia. Jour-nal of Molluscan Studies, 48(1): 102−103.

ZISCHKE, J. A., 1973, An ecological guide to theshallow-water marine communities of PigeonKey, Florida. Privately published, St. Olaf Col-lege, Northfield, Minnesota. [vi +] 44 pp.

ZISCHKE, J. A., 1977a, Checklist of macroflora,invertebrates and fishes of Pigeon Key. Pp.2730, in: H. G. MULTER, ed., Field guide to somecarbonate rock environments − Florida Keys

and western Bahamas, new ed. Kendall/HuntPublishing Company: Dubuque, Iowa. 415 pp.,10 maps.

ZISCHKE, J. A., 1977b, Some common inverte-brates of Pigeon Key, Florida. Pp. 338−347,fig. A.14, in: H. G. MULTER, 1977, Field guide tosome carbonate rock environments − FloridaKeys and western Bahamas, new ed. Kendall/Hunt Publishing Company: Dubuque, Iowa.415 pp., 10 maps.

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”, offMiddle Florida Keys, scuba, 20 ft (6.1 m), patchreef/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, AMNH296528); FK-039, Crawl Key, 24°44’36"N,80°58’47"W, oceanside, beach inside channel,shallow sand, seagrass, seawall, by hand andhand dredge, 12 March 1996 (1 spm alc, FMNH301424); FK-047, Channel marker 50A offRamrod Key, 24°35.80’N, 81°27.24’W, rubbleand patch reef, 15 ft (4.6 m), scuba, M/V THESNAIL, 21 September 1996 (1 pair, 1 valve,AMNH 295179); FK-068, bayside of WestSummerland 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 April1997 (1 pair, FMNH 279521); FK-090, GardenKey, Dry Tortugas, 24°37’50"N, 8252’20"W,sand beach with stone pilings, 2.5 m, by handand snorkeling, 23 April 1997 (1 pair, AMNH296520), FK-091, Fort Jefferson, Garden Key,Dry Tortugas, 24°37’50"N, 82°52’24"W, beachat mote and mote wall, by hand and snorkel-ing, 23 April 1997 (2 pair, AMNH 290122); FK-115, East Washerwoman Shoal (ChannelMarker 49), off Marathon, 24°40’N, 8104.3’W,9 ft (2.7 m), scuba, R/V FLORIDAYS, 12 July1997 (1 valve, FMNH 279523); FK-117, KeyVaca, channel west of Stirrup Key, 24°44.19’N,81°02.92’W, bayside, in channel, 15 ft (4.6 m),

BIELER ET AL.452

plus in surrounding shallow Thalassia/sand,scuba, 14 July 1997 (1 spm alc [photographedalive], FMNH 295706); FK-119, “The Slabs”patch reef between “outer patches” and “coralhumps” off Marathon, 24°39.53’N, 81°00.90’W,scuba, 23 ft (7.0 m), R/V FLORIDAYS, 20 July1997 (1 valve, FMNH 279528); FK-121, “TheSlabs” patch reef between “outer patches” and“coral humps” off Marathon, 24°39.53’N,8100.90’W, 23 ft (7.0 m), scuba, R/VFLORIDAYS, 21 July 1997 (1 pair, 3 valves,AMNH 296525); FK-131, off Key Vaca,oceanside, “outer patches” south of HawkChannel, 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 to24°43.60’N, 81°07.47’W, sand/sparse sea-grass, 8 ft (2.4 m), dredge, R/V FLORIDAYS,07 August 1997 (1 valve, FMNH 279525); FK-149, exposed flats outside channel off CrawlKey, 24°44’29"N, 80°58’24"W, oceanside,clean sand + Thalassia/Syringodeum sea-grass, Penicillus, 0.5 ft (0.15 m), by hand, R/VFLORIDAYS, 22 August 1997 (2 valves, FMNH279530); FK-169, Tavernier Creek, nearbayside 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/VFLORIDAYS, 17 September 1998 (1 valve,AMNH 296522); FK-171, just off mouth ofTavernier Creek, oceanside, near marker #7,24°59.67’N, 80°31.72’W, sand, Thalassia/Syringodeum seagrass, 0−3 ft (0−0.9 m), snor-keling/sieving, R/V FLORIDAYS, 17 Septem-ber 1998 (observed pair); FK-178, east end ofRodriguez 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), snor-keling, 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 3days after Hurricane Georges, in Thalassiadroves (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, coralheads, 6−15 ft (1.8−4.6 m), scuba, R/VFLORIDAYS, 10 April 1999 (1 spm alc, AMNH298893); 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/VFLORIDAYS, 31 July 1999 (2 pair, AMNH296521); FK-233, Old Dan Bank, bayside ofLong 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 August1999 (3 valves, FMNH 279522); FK-244,bayside of West Summerland Key (SpanishHarbor Keys), 24°39’19"N, 81°18’13"W,bayside, rock wall and sand slope, 23 ft (7.0m), scuba, 05 August 1999 (1 valve, AMNH296524); FK-246, bayside of WestSummerland Key (Spanish Harbor Keys),along inner shore of western arm of horse-shoe, 24°39’19"N, 81°18’13"W, beach, 05August 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 sea-grass, 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 Keycoral 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); FK-268, 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 FLORI-DAYS, 19 August 1999 (1 valve, FMNH279527); FK-269, Looe Key back reef,24°32.79’N, 81°24.33’W, sediment sample, 8ft (2.4 m), snorkeling, R/V FLORIDAYS (1 juvpair, AMNH 296531); FK-273, bayside of WestSummerland Key (Spanish Harbor Keys), atoutermost 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 ofspm with pearls, see prior; DNA], FMNH296696; 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-col-lecting, R/V FLORIDAYS, 20 August 1999 (1valve, FMNH 279524); FK-287, bayside ofWest Summerland Key (Spanish HarborKeys), inside outermost arm of horseshoe,24°39.35’N, 81°18.22’W, shallow subtidal, byhand, snorkeling, 10 April 2000 (1 spm alc,FMNH 289982; 1 live-collected pair, FMNH301426; 9 pair, FMNH 289939); FK-298, Looe


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 NationalMarine Sanctuary, southwest corner of corearea, 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/VFLORIDAYS, 08 July 2000 (1 pair, AMNH299489); FK-352, bayside of WestSummerland Key (Spanish Harbor Keys),south arm, 24°39’19"N, 81°18’13"W, bayside,rubble, to 1.5 m, snorkeling, 08 July 2000 (3spm 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 seagrasswith large coral rubble, 9−11 ft (2.7−3.3 m),scuba, R/V FLORIDAYS, 09 July 2000 (3 spmalc [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, 10July 2000 (observed spm; 2 pair, 2 valves,AMNH 299421); FK-360, coral lumps offNewfound 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, AMNH299520); FK-363, east end of Rodriguez Key,oceanside of Key Largo, 25°03.27’N,80°26.66’W, Thalassia seagrass, sand, smallrubble, 6 ft (1.8 m), snorkeling, R/VFLORIDAYS, 07 October 2000 (2 pair, AMNH307612); FK-364, Dove Key (just southwestof 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), snor-keling, R/V FLORIDAYS, 07 October 2000 (1pair, AMNH 307613); FK-367, northeast cor-ner of Conch Reef, oceanside of Key Largo,24°57.895’N, 80°27.248’W, rubble, Thalassiaseagrass, 9 ft (2.7 m), snorkeling, R/VFLORIDAYS, 08 October 2000 (1 frag, AMNH307736); FK-392, Lower Matecumbe Key,24°51’24"N, 80°43’40"W, oceanside, by handin wrack line, 20 October 2000 (9 valves,FMNH 301429); FK-395, Snappers Ledgereef, 24°58.88’N, 80°25.36’W, in cavity of largesponge with red algae, 26 ft (7.9 m), scuba,M/S REPUBLIC IV, 27 March 2001 (1 pair,FMNH 301430); FK-459, Ft. Zacchary TaylorState Park, Key West, beach, shells washedashore among rubble, 02 May 2001 (1 frag,

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, SandKey, 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 July2001 (2 frag, AMNH 307738); FK-539, southof Bahia Honda Key, west of Looe Key reef,24°34.24’N, 81°16.64’W, 30.2−34.1 m (99−112ft), sand and rubble, pipe dredge and triangledredge, R/V EUGENIE CLARK, 28 July 2001(1 juv valve, AMNH 308089); FK-547, LooeKey reef, 24°32.809’N, 81°24.158’W, 25 ft (7.6m), spur and groove reef, scuba, R/VFLORIDAYS, 30 July 2001 (2 valves, AMNH307614); FK-559, south beach of LoggerheadKey, Dry Tortugas, 24°37.790’N, 82°55.400’W,wrack line to 7 ft (2.1 m), by hand and snor-keling, R/V CORAL REEF II, 15 April 2002 (3valves, FMNH 301432); FK-581, north shoreof Loggerhead Key, Dry Tortugas,24°37.871’N, 82°55.447’W, wrack line andshallow subtidal, by hand and snorkeling, R/VCORAL REEF II, 16 April 2002 (5 valves,FMNH 301433); FK-601, Hospital Key, DryTortugas, 24°38.970’N, 82°51.284’W, patchreef, sand, 16 ft (4.9 m), scuba and snorkel-ing, R/V CORAL REEF II, 18 April 2002 (1valve, 3 frag, AMNH 307615; 1 pair, FMNH296718); 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/VCORAL REEF II, 18 April 2002 (1 frag, AMNH307616); 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 CORALREEF II , 20 April 2002 (2 pair, 3 valves, AMNH307617); FK-620, Old Dan Bank, bayside ofLong Key, 24°50.45’N, 80°49.63’W, Thalassiaseagrass 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, CoffinsPatch Sanctuary Preservation Area, off CrawlKey, 24°40.92’N, 80°58.26’W, patch reef with

BIELER ET AL.454

sand patches, gorgonian, pillar coral, 21 ft (6.4m), scuba, M/V SHUTTERBUG II, 20 July2002 (5 valves, FMNH 301447); FK-629,bayside of West Summerland Key (SpanishHarbor Keys), 24°39.3’N, 8118.2’W, amongrocks 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, oceansideof 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 Pi-geon Key, 24°42.2’N, 81°09.3’W, Thalassia/Halodule/Syringodeum seagrass on sand/rubble, concrete bridge piers, 0.5−1 m, byhand, snorkeling, shovel/sieving (1 pair,NTM); FK-649, Sprigger Bank, bayside, justW 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 Labo-ratory boat, 27 July 2002 (2 pair, 1 valve,FMNH 301449); FK-659, Pigeon Key,24°42.2’N, 81°09.3’W, seagrass, scuba, 2−4ft (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 seagrasswith Halimeda, Porites, sponges, hydroids,patches of sand/Halimeda hash, 1−5 ft (0.3−1.5 m), snorkeling, R/V LAST MANGO, 28 July2002 (2 pair, 5 valves, 3 frag, FMNH 301450);FK-661, Molasses Keys, south of center ofSeven-Mile Bridge, north of westernmost is-land, 24°41.070’N, 81°11.483’W, sandy bot-tom, coral rubble, Thalassia seagrass, hotwater (> 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 SO-3, 5 nmi south of Knights Key, 24°37.619’N,81°06.528’W, sandy bottom adjacent to coralreef; scuba, 17−23 ft (5.2−7.0 m), R/VFLORIDAYS, 05 August 2002 (1 valve, 1 frag,FMNH 301435); FK-664, Molasses Keys,south of center of Seven-Mile Bridge, northof westernmost island, 24°41.070’N,81°11.483’W, sandy bottom, coral rubble,Thalassia seagrass, strong current, snorkel-ing, 2−5 ft (0.6−1.5 m), R/V FLORIDAYS, 06August 2002 (2 valves, FMNH 301436); FK-665, Coffins Patch pillars, south of Crawl Key,24°40.899’N, 80°58.246’W, coral reef, sandplains, 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 Seven-Mile Bridge, off north and west ends of island,24°41.009’N, 81°12.955’W, “ironshore” beachrock, sand, Thalassia seagrass, snorkeling, 0−5 ft (0−1.5 m), R/V FLORIDAYS, 09 August2002 (1 pair, FMNH 301444); FK-672, JohnSawyer Bank, 3 nmi north of western part ofKey Vaca, bayside, 24°45.498’N, 81°06.621’W,coral rubble, Thalassia seagrass, strong cur-rent over shoal, snorkeling, 2−6 ft (0.6−1.8 m);R/V FLORIDAYS, 11 August 2002 (1 pair, 2valves, 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, coralrubble, Thalassia seagrass, strong current overshoal, 3−5 ft (0.9−1.5 m), snorkeling, R/VFLORIDAYS, 11 August 2002 (1 valve, FMNH301439); FK-674, Sombrero Reef, southwestarea 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, FMNH301440); FK-675, Red Bay Bank, 3 nmi northof 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 (2spm, 2 valves, FMNH 301441); FK-676,Rachel Bank, 2 nmi north of Key Vaca,bayside, 24°44.714’N, 81°04.599’W, muddysand, 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, FMNH296695); FK-677, Doughnut Reef, oval reefmass east of Coffins Patch (off Crawl Key),24°41.507’N, 80°56.835’W, sand flat at reefedge, scuba, 23 ft (7.0 m), M/V SHUTTERBUGII, 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 (0.9−3.0m), 14 August 2002 (1 spm alc [dissected], 4valves, FMNH 301443); FK-688, Conch Reef,oceanside of Key Largo, 24°57.380’N,80°29.428’W, algae-covered reef, verticalwalls and adjacent sand plains, max. 28 ft (8.5m), scuba, R/V FLORIDAYS, 05 June 2003 (1V, FMNH 302086; 2 LV, AMNH 308081); FK-689, northeast of Dove Key, oceanside of KeyLargo, 25°03.055’N, 80°28.220’W, hard bot-tom with silty sand, sponges, gorgonians, 0.5−1.0 m, snorkeling, R/V FLORIDAYS, 06 June2003 (1 juv pair, AMNH 308084); FK-692, Henand Chickens patch reef, oceanside of Plan-tation Key, 24°56.099’N, 80°32.920’W, max.21 ft (6.4 m), scuba and snorkeling, R/V


FLORIDAYS, 08 June 2003 (1 RV, 1 LV, AMNH308087); FK-693, off Dove Key, oceanside ofKey Largo, 25°03.039’N, 80°28.151’W, siltyThalassia seagrass and sand, 3−5 ft (0.9−1.5m), snorkeling, R/V FLORIDAYS, 08 June2003 (3 pair, 1 RV, 1 LV, AMNH 308083); FK-696, Sand Island (near Molasses Reef),25°01.116’N, 80°22.046’W, patch reef withrubble, max. 22 ft (6.7 m), scuba, R/VFLORIDAYS, 10 June 2003 (1 pair, AMNH308085); FK-698, Wolfe mooring buoy (nearThree Sisters Reef), 25°01.311’N,80°23.774’W, patch reef with adjacentThalassia seagrass, max. 16 ft (4.9 m), scuba,R/V FLORIDAYS, 11 June 2003 (2 RV, 1 frag-ment, AMNH 308082); FK-701, off Dove Key,oceanside of Key Largo, 25°03.011’N,80°28.163’W, silty Thalassia seagrass andsand, 1−2 ft (0.3−0.6 m), snorkeling, R/VFLORIDAYS, 12 June 2003 (5 pair, 1 RV, 1LV, AMNH 308086). FK-702, seagrass flat offYellowtail Inn, oceanside of Grassy Key, milemarker 58.3, 24°45.494’N, 80°57.179’W, 1−5ft (0.3−1.5 m), snorkeling, 5−23 August 2003(1 pair, 1 LV, FMNH 302071); FK-703, patchreef with sand pockets in vicinity of mooringbuoy 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, vi-cinity of buoy SO-4, 5 nmi S of Knights Key,2437.591’N, 81°06.563’W, sandy bottom im-mediately 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); FK-705, Doughnut Reef, oval reef mass E of Cof-fins Patch (off Crawl Key); 24°41.439’N,80°56.862’W, patch reef surrounded by sandflats, scuba, 24 ft (7.3 m); M/S SEAFARI, 09August 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 sur-rounded by sand flats, scuba, 19 ft (5.8 m);M/S SEAFARI, 09 August 2003 (1 RV, FMNH302074); FK-710, “Porkfish” and “HammerLedge” reefs, E of Coffins Patch (off ConchKey); 24°42.038’N, 80°53.578’W, scuba, 19−25 ft (5.8−7.6 m); M/S SEAFARI, 11 August2003 (2 RV, 2 frag, FMNH 302075); FK-711,patch reef with sand pockets in vicinity of moor-ing buoy 9, Coffins Patch coral reef, oceansideof Grassy Key, 24°41.131’N, 80°57.818’W, 18−20 ft (5.5−6.1 m), scuba; R/V FLORIDAYS, 12August 2003 (1 LV, FMNH 302076); FK-714,vicinity of mooring buoy 1 at “Marker 48” patchsand and seagrass; 24°41.505’N, 81°01.528’W,

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); SouthLake Worth near Boynton, Kline! July 1944,DMNH 72175 (1 pair); Lake Worth, Lermond!August 1941, DMNH 153846 (3 pair); LakeWorth, south end, dredged in 6 ft (1.8 m),Lermond! 14 September 1937, DMNH153848 (2 pair); Lake Worth, south end,Doremus! 1942, DMNH 63779 (2 pair); LakeWorth, south end, F. Lyman family! Summer1944, FMNH 144097 (1 pair); Lake Worth,South Inlet, oyster reef, F. M. Bayer! USNM890543 (1 pair with tissue); Boynton in LakeWorth, 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); oppositeLemon 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, FingerChannel flats, Crovo! 05 April 1970, DMNH30152 (1 pair); Cape Florida, Biscayne Bay,T. L. McGinty! 1937, ANSP 264016 (1 pair);Miami Bay area, flats off Cape Florida, IngallsFamily! 12 March 1967, AMNH 140432 (1pair); Miami area, Biscayne Bay, flats offCape Florida, W. E. Old! 12 March 1967,AMNH 136053 (2 pair); Cape Florida, T. L.McGinty! FMNH 26427 (2 valves); CapeFlorida, near Miami, T. L. McGinty! June1936, USNM 599308 (1 pair); Bear Cut, KeyBiscayne, Miami, W. S. Bitler! 1963, AMNH142323 (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 (2pair); Caesar Creek, Whitney! November1956, DMNH 97311 (1 pair); Florida Keys,Lermond! DMNH 153830 (1 pair); AngelfishCreek [north of Key Largo, connecting CardSound and Atlantic Ocean], Wisoff Collec-tion, AMNH 120395 (1 pair); Key Largo,Nelson collection, FMNH 155544 (1 pair);Key Largo, T. L. Moise! August 1950, ANSP

BIELER ET AL.456

193815 (1 juv pair); Key Largo, F. M. Bayer!June 1940, USNM 890827 (1 pair); WindleyKey, Whale Harbor Channel, near bridge in5−10 ft (1.5−3.0 m), T. R. Waller! Sta. 3, 31August 1971, USNM 707750 (1 valve); Up-per Matecumbe Key, Islamorada, on beach,13 January 1978, BMSM 26108 (5 pair); In-dian Key, D. V. Stingley! May 1959, BMSM26109 (1 pair); Lower Matecumbe Key,Hausman! AMNH 133675 (1 pair); ConchKey, south end, J. J. Parodiz & Winters! 08July 1976, CMNH 43728 (4 pair); Grassy Key,M. & S. Snyder! July 1966, ANSP 309750 (2pair); 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); CrawlKey, shallow water, J. M. Bijur! May 1964,AMNH 248309 (1 pair); Crawl Key, bayside,on flats at 1 ft (0.3 m), Raeihle! November1961, AMNH 106142 (1 pair, figured Emerson& Jacobson, 1976: pl. 42, fig. 18); Crawl Key,bayside, on beach, D. Raeihle! November1973, 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, live-collected); Crawl Key, G. Dingerkus & L. D.Uhler! 06 January 1977, AMNH 267424 (1spm alc); Crawl Key, bayside, edge of bor-row pit ½ mi north of mile marker 56, inweeds, M. J. de Maintenon! 24 June 1985,AMNH 307571 (1 pair); Bonefish Key, A.Koto! AMNH 133664 (1 pair) and FMNH176293 (1 pair, 1 valve); Marathon Key,Florida Straits, K. C. Vaught Collection,AMNH 250783 (1 pair); Marathon, 1959,BMSM 26110 (2 pair); Marathon Key, southend, Jensen! 1962, DMNH 42547 (1 pair);Sombrero Reef, 25−30 ft (7.6−9.1 m), P. S.Mikkelsen! 21 May 1980, DMNH 180066 (2valves); Washerwoman’s patch reef,24°39’54"N, 81°04’14"W, M. Snyder! August1966, 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) andFMNH 176372 (1 pair [photo voucher]); LittleDuck Key, shallow water, sand, F. Schilling!July 1968, FMNH 288716 (1 pair); MissouriKey, Richardson! DMNH 85835 (1 pair), Mis-souri Key, snorkeling, A. D. Barlow! 05 March1967, AMNH 243914 (2 pair); Missouri Key,sand, F. Schilling! 16 July 1970, FMNH288718 (1 pair); Bahia Honda State Park,

Germer Collection, 11 July 1973, AMNH269541 (1 pair); north of Bahia Honda StatePark, shallow water, sand, R. Bieler & P. M.Mikkelsen! 25 March 1989, FMNH 288810 (1spm alc); Bahia Honda Key, BMSM 26111 (1pair); Spanish Harbor Key, beach, Piech! July1980, DMNH143838 (1 pair); WestSummerland Key, at entrance to dredge hole,L. Scheu Collection, 1984, AMNH 230097 (2pair); Newfound Harbor Keys, living in shal-low water, sand, November 1968, Raeihle!AMNH 307849 (1 pair); Torch Key channel,C. T. Simpson! UMML 28.1716 (4 pair); Sugar-loaf Key, J. B. Clark! 23−24 May 1921, ANSP9634 (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, HawkChannel, 3−20 ft (0.9−6.1 m), Eolis sta. 65, J.B. Henderson Jr.! 15 May 1913, USNM448344 (2 juv valves); Key West, reefs, rare,H. Hemphill! USNM 95672 (2 pair); Key West,Smith Shoals, Eolis sta. 335, J. B. HendersonJr.! 1916, USNM 448343 (1 juv pair); KeyWest, N side, beach collecting, Eolis sta. 35,J. B. Henderson Jr.! 30 May (or 6 June) 1911,USNM 48340 (1 juv pair); Key West, USNM406825 (2 pair); Key West, H. Hemphill! ANSP52199 (2 pair); Key West, H. A. Pilsbry! March-April 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! 21June 1967, FMNH 288719 (1 pair); Key West,C. T. Simpson! UMML 28.1748 (1 pair); SandKey, Key West, Ostheimer! DMNH110654 (3pair); near Boca Grande Key, UMML 28.1740,C. T. Simpson! (1 pair); Tortugas, Stm.! USNM36403 (4 pair, 1 valve); Dry Tortugas, off Log-gerhead Key, north end, beach collecting,Eolis sta. 367, J. B. Henderson Jr.! 13 June1911, USNM 448347 (1 valve); Dry Tortugas,Garden Key, 3 mi out from red sea buoy, 5dredge hauls, 14−15 fms (25.6−27.4 m), Eolissta. 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 ofFlorida], 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 (1pair).


Bahamas: Bahamas, J. B. Henderson Jr.!USNM 448342 (2 pair); PMM-1047 (wp277-R rubble), off Andros, 24°54’44.42"N,77°53’51.80"W, rubble in back reef pave-ment zone with gorgonians, 7 ft (2.1 m),scuba, P. M. Mikkelsen, et al.! 29 August2000, AMNH 305616 (1 valve); PMM-1090(wp415-R), off Andros, 24°53’32.2"N,77°53’51.4"W, thick Thalassia seagrass, 12ft (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/algalplain, 5 ft (1.5 m), scuba/snorkeling, P. M.Mikkelsen, et al.! 02 September 2000, FMNH296720 (1 pair); PMM-1063 (BH-Rseagrass), oceanic blue hole off Blue HoleCay, off Andros, 24°53’55.2"N,77°55’12.1"W, Thalassia seagrass, 4 ft (1.2m), scuba/snorkeling, P. M. Mikkelsen, et al.!31 August 2000, AMNH 307572 (1 pair); EastAndros Island, Calabash Bay, Abbott! Feb-ruary 1971, DMNH 29242 (1 pair); EastAndros Island, Small Hope Bay, Abbott!DMNH 41253, March 1971 (1 valve); ChubCay, Berry Islands, Moise! ANSP 193106 (1pair); 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, Run-ning Mon Canal, 26°29’45"N, 78°41’45"W,J. N. Worsfold! ANSP 369788 (1 pair); GrandBahama Island, C. C. Allen! 1922-1923,ANSP 133697 (1 pair); Grand Bahama is-land, Bottle Bay Canal, 26°39’30"N,78°57’00"W, sediment, 5 ft (1.5 m), J.Worsfold! ANSP 371908 (1 pair); east endof Grand Bahama Island, Deep Water Cay,ca. 2.5 mi northwest of Sweetings Cay Light,intertidal sand and rocks, V. O. Maes! Janu-ary 1965, ANSP 307688 (1 juv pair); GrandBahama Island, F. H. Low Collection, AMNH113790 (1 pair); Nassau, New ProvidenceIsland, Wards! (before 1893), FMNH 2741(1 pair, 1 valve); Nassau, New ProvidenceIsland, Pope! FMNH 187147 (2 valves);Nassau, C. C. Allen! USNM 36617 (1 pair);New Providence Island, Dicks Point, McLean& Russell! July 1936, ANSP 169917 (2valves); New Providence [Island], LyfordCay, AMNH 80763 (2 pair); north coast ofHog Island, north of New Providence Island,

R. Robertson! 11 September 1955, ANSP299657 (1 juv pair); Great Abaco Island andGreen Turtle Key, Abaco Islands, Cherokeeflats, Great Abaco and Mendelson’s flats,Heilman! DMNH 37728 (3 pair); Great AbacoIsland 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! 14August 1953, ANSP 299066 (3 pair); GreatAbaco, west coast of north end of Elbow[Little Guana] Cay, mud/sand, Halimeda re-mains, 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 barat 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, MarshHarbor, O. Bryant! USNM 180541 (1 pair);east-central Eleuthera, north end of HalfSound, 25°07’45"N, 76°09’00"W, R.Robertson! 18 April 1984, ANSP 359292 (2pair); Eleuthera, Savannah Island, SantyPoint, W. J. Clench! May 1936, ANSP173811 (3 pair); Eleuthera, SavannahSound, Sandy Point, Cora Staples Collec-tion, AMNH 306250 (1 pair); Eleuthera, Cur-rent, Current Club, A. Ross! 29 July 1963,AMNH 100174 (1 valve); Eleuthera Island,Doremus! DMNH 63778 (2 pair); EleutheraIsland, 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 cen-tral Eleuthera Island, Abbott! June 1976DMNH 115333 (1 valve); Exuma Island,Rolle Town, Loc.16, Kline! 11 July 1951,DMNH 72174 (1 pair); southern ExumaCays, north of Leaf Cay, R. Robertson! 07July 1957, ANSP 285738 (1 pair); Bimini,near Bailey Town, Bimini Lagoon, R.Robertson! 1957−1958, ANSP 326271 (1valve); South Bimini, east of Nixon’s Harbour,R. Robertson! 1957−1958, ANSP 325603 (1pair); Bimini, 1.75 mi southeast of OrangeCay, 23 ft (7.0 m), R. Robertson! 1957−1958,ANSP 325698 (1 juv valve); Bimini, SouthCat Cay, grassy, T. L. Moise! ANSP 193577(1 pair); Bimini, around Risty Causeway, Pi-

BIELER ET AL.458

geon Cay, Steger! April 1956, DMNH 107635(1 pair); San Salvador Island, W PigeonCreek, beach, Piech! February 1977, DMNH143251 (1 pair).

Turks and Caicos: Providenciales, Water Cay,beach, Piech! February 1978, DMNH 144123(1 pair).

Cuba: east of Tarallones de Arena, nearSantiago, sand beach, R. E. Dickerson!ANSP 182932 (3 valves); Paradise Island,Oriente, Christofferson! 17 April 1949, FMNH144071 (4 valves); west of Guardalavaca,eastern shore near Playa Esmeralda, Prov-ince Holguin, K. & Ch. Schniebs! December2001, 1 pair, MTD 43828; Guantanamo, E.O. Mitchell et al.! 1930, USNM 405334 (1valve); Cayo Hutia Reef, Barrera Expeditionsta. 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, GunBay, near Blakes’, mud and turtlegrass flats,A. J. Ostheimer III! ANSP 199513 (1 pair);Grand Cayman, North Sound, Jensen! Au-gust 1970, DMNH 39561 (1 pair).

Jamaica: Harboreale, near Annotta Bay, St.Mary, Orcutt! USNM 440717 (1 valve); BlackRiver, St. Elizabeth, Orcutt! USNM 441413(1 valve).

Hispaniola: Haiti, off Port-au-Prince, southeastside of Grand Bans, east side of reef, G.Goodfriend! 25 June 1972, AMNH 177703(1 pair); Haiti, Cape Haitien, American HaitienDev. Company, Krieger! USNM 487861 (1valve); Santo Domingo, Monte Christi, W. J.Clench et al.! July 1937, ANSP 173105 (2valves); Santo Domingo, Monte Cristi,Doremus! July 1937, DMNH 63777 (2 pair).

Puerto Rico: Puerto Rico, Stearns! USNM54091 (1 pair); El Deseches Island,Mayaguez B., F. A. Gallardo! USNM 464243(1 valve); Bahia Bramadero [south ofMayaguez], G. L. Warmke! November 1956,ANSP 222755 (1 valve); Puerto Rico,Richardson, DMNH-85845 (2 pair).

U.S. Virgin Islands: St. Thomas, W. A. HainesCollection, pre-1895, AMNH 31868 (9 pair,including largest recorded specimen); St.Thomas, M. Petit! USNM 250151 (1 pair); St.Thomas, Petit collection, USNM 530502 (1pair, 2 valves, 1 juv valve); St. Thomas,Lindberg Beach, D. M. Barringer! 1936,ANSP 166919 (1 valve); St. Thomas, SwiftCollection, ANSP 53580 (5 pair, 2 juv pair);

St. Thomas, ZMB-104283 (1 pair); St. Tho-mas, ZMB-104287 (1 pair).

British Virgin Islands: Peter Island, LittleHarbour, R. H. Pine! July 1976, FMNH197453 (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! 25September 1980, AMNH 303476 (1 pair);Gorda Island, Colquhon Reef, Bredin-Smithsonian Expedition sta. 37−58, Schmittet al.! 07 April 1958, USNM 735909 (1 pair).

Antigua: Falmouth Harbor, beach, SUI Expe-dition, J. B. Henderson Jr.! 1918, USNM500994 (1 pair).

Grenada: South Grenada, Little Bacaye Har-bor, 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! 23August 1961, ANSP 281836 (1 juv valve);north of Tarpon Cay, shallow Acroporacervicornis 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 (4valves).

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 NEof Colon, suction dredge at refinery site, W.P. Woodring! 13 December 1959, USNM637931 (1 valve); Payardi Island, Minas Bay,E of Colon, R. H. Stewart! USNM 734522 (3valves).

Colombia: Old Providence Island, Sid Ander-son! January−March 1966, AMNH 138019 (1pair); San Andres Island, S. Anderson! Janu-ary 1966, AMNH 137541 (1 valve, 1 frag);vicinity of Cartagena, T. A. Link! USNM364301 (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, DMNH72794 (1 pair); Curaçao, N. Dearborn! 1908,FMNH 12764 (1 valve, subfossil?).

Periglypta listeri (J. E. Gray, 1838) (Bivalvia: Veneridae) in the western Atlantic: taxonomy,...

Documents

Transcript of Periglypta listeri (J. E. Gray, 1838) (Bivalvia: Veneridae) in the western Atlantic: taxonomy,...