Echinolittorina jamaicensis (C.B. Adams, 1850 )

Reid, David G., 2009, The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the western Atlantic Ocean 2184, Zootaxa 2184 (1), pp. 1-103 : 42-50

Echinolittorina jamaicensis (C.B. Adams, 1850)

( Figures 18–21 )

Littorina lineata —d’Orbigny, 1841: 208–209, pl. 14, fig. 25 (in part, includes E. angustior , E. lineolata ; not Lamarck, 1822 = Littoraria tessellata ).

Littorina (Melaraphis) lineata —Arango, 1880: 159 (not Lamarck, 1822).

Littorina (Melarhaphe) lineata —von Martens, 1900: 583 (in part, includes E. angustior , E. interrupta ; not Lamarck, 1822).

Littorina jamaicensis C.B. Adams, 1850: 71–72 ( Jamaica; lectotype ( Clench & Turner 1950: 380) MCZ 186133, seen, Fig. 18M herein + 9 paralectotypes MCZ 156161, 5 paralectotypes MCZ 156162, 1 paralectotype MCZ 156163, paralectotypes not seen; paralectotypes include E. interrupta according to Bandel & Kadolsky 1982, although their fig. 24 is not a paralectotype of this species as claimed (p. 24, contra caption p. 16), but is the specimen here designated neotype of Litorina ziczac var. interrupta ). Bandel, 1974a: 99, 108, figs 18–22 and 50–51 (radula) (in part, includes E. angustior ).

Littorina (Melarhaphe) jamaicensis — Mörch, 1876: 139–140 (as Melaraphe ).

Littorina (Melarhaphe) floccosa var. α Mörch, 1876: 138 (as Melaraphe ).

Littorina glaucocincta ‘Beck mss’ Mörch, 1876: 138 (name published in synonymy, unavailable).

Nodilittorina (Nodilittorina) glaucocincta — Bandel & Kadolsky, 1982: 27–28, figs 5 (shell, radula), 9 (map), 32–33 (shells and radulae).

Nodilittorina glaucocincta — Garcia & Luque, 1986: 37. Britton & Morton, 1989: 86, fig. 4- 5M.

Littorina (Melarhaphe) riisei Mörch, 1876: 140 (Havanna [La Habana], Cuba; lectotype ( Bandel & Kadolsky 1982) ZMK, not seen, specimen now lost, but figured by Bandel & Kadolsky 1982: fig. 30; as Melaraphe ).

Litorina riisei —Weinkauff, 1882: 104.

Nodilittorina (Nodilittorina) riisei — Bandel & Kadolsky, 1982: 26–27, figs 5 (shell, egg capsule, operculum, radula), 9 (map), 30–31 (shells and radulae).

Nodilittorina riisei — Britton & Morton, 1989: 86, fig. 4- 5F. Reid, 2002a: 259–281.

Nodilittorina (Echinolittorina) riisei — Reid, 1989: 99, fig. 11j (egg capsule).

Echinolittorina riisei — Williams, Reid & Littlewood, 2003: 60–86. Williams & Reid, 2004: 2227–2251, fig. 6C (map).

Litorina carinata — Weinkauff, 1878, 1882: 31, 107 (in part, includes E. angustior ; not d’Orbigny, 1841). Weinkauff, 1883: 220 (in part, includes E. angustior ; not d’Orbigny, 1841).

Litorina angustior — Weinkauff, 1883: 220 (in part, includes E. angustior ; not Mörch, 1876).

Litorina ziczac — Weinkauff, 1883: 220 (in part, includes E. angustior , E. interrupta , E. ziczac ; not Gmelin, 1791).

Littorina (Melarhaphe) ziczac — Tryon, 1887: 251 (in part, includes E. angustior , E. interrupta , E. lineolata , E. ziczac , Littoraria glabrata ; not Gmelin, 1791; as Melaraphe ).

Littorina ziczac — Bequaert, 1943: 14–18, pl. 5, figs 6–10 (in part, includes E. lineolata , E. ziczac , E. interrupta , E. angustior , E. placida ; not Gmelin, 1791). Abbott, 1954a: 132 (in part, includes E. placida , E. angustior , E. ziczac ; not Gmelin, 1791). Lewis, 1960: 415–416, fig. 11E, F (egg capsule) (in part, includes E. angustior ; not Gmelin, 1791).

Littorina floccosa — Abbott, 1968: 82–83, fig. (not Beck in Mörch, 1876 = E. interrupta ).

Littorina lineolata — Abbott, 1964: 65–66 (in part, includes E. angustior , E. placida ; not d’Orbigny, 1840). T.V. Borkowski & M.R. Borkowski, 1969: 409, fig. 1 (map), fig. 4A (egg capsule), pl. 66, figs 3, 4 (in part, includes E. lineolata ; not d’Orbigny, 1840). Borkowski, 1971: fig. 1A, B (paraspermatozoa), 2 (egg capsule) (not d’Orbigny, 1840). Bandel, 1974a: 99–100, 103–104, 108, figs 17 (egg capsule), 19–22 and 36–38 (radula) (not d’Orbigny, 1840). Borkowski, 1975: 369–377, fig. 1B (radula) (in part, includes E. lineolata ; not d’Orbigny, 1840). Harasewych, 1977: 60–62, figs 1–3 (hyperstrophic shell) (not d’Orbigny, 1840). Janson, 1985: 871–879 (not d’Orbigny, 1840).

Littorina (Austrolittorina) lineolata — Rosewater, 1970: 423 (not d’Orbigny, 1840). Abbott, 1974: 68, fig. 559 (in part, includes E. placida ; not d’Orbigny, 1840).

Nodilittorina (Nodilittorina) mordax Bandel & Kadolsky, 1982: 28–29 , figs 5 (shell, operculum, radula), 9 (map), 37–39 (shells and radulae) ( Paradise Island , Nassau, Bahamas; holotype USNM 749813, seen, + 23 paratypes USNM 749814, seen, + paratypes USNM 749815, 749816, not seen).

Littorina mordax — De Jong & Coomans, 1988: 19, pl. 22, fig. 77B.

Nodilittorina mordax — Britton & Morton, 1989: 86, fig. 4-5H. Redfern, 2001: 28–29, pl. 14, fig. 117.

Taxonomic history: Littorina jamaicensis was fully described by C.B. Adams (1850), who stressed the characteristic “very deep rather distant spiral striae”, sometimes “enlarged nearly to the width of the intervening ridges”, and a lectotype specimen confirms the identity. It seems unlikely that Adams could have included the weakly sculptured E. interrupta in his concept of the present species, as suggested by Bandel & Kadosky’s (1982: 27) identification of some paralectotypes. In their monograph Bandel & Kadolsky (1982) divided the present species into three ( N. riisei , N. glaucocincta , N. mordax ) solely on the basis of striking differences in the radula. Since there were no diagnostic differences in the shells, it was not possible to determine the identity of L. jamaicensis , which they consequently considered a nomen dubium. No anatomical differences could be found among these forms and intraspecific radular variation was discovered in other littorinids, so it seemed likely that only a single variable species was involved ( Reid 1989, 2002a; Williams & Reid 2004; under the specific name riisei ). This is confirmed here (see Remarks) and a valid name must be chosen. Littorina jamaicensis has priority, and neither of the conditions for the reversal of precedence (ICZN, 1999: Art. 23.9) is met. The name L. jamaicensis may be relatively unfamiliar, but has in fact been used as a valid name on two previous occasions ( Mörch 1876; Bandel 1974a). Furthermore, L. glaucocincta is not an available name (see below), and neither L. riisei nor N. mordax has been sufficiently widely used to justify their conservation of either. The changed name of the present species emphasizes the new taxonomic concept of a single species with no diagnostic radular form.

In his synopsis of molluscs of the West Indies, Mörch (1876) included extensive synonymy lists, brief Latin descriptions, and the Latin names were not printed in italic font. This format has led to confusion regarding usage and availability of names. Mörch (1876) listed ‘ L. glaucocincta Beck mss’ as a synonym of his ‘ Littorina floccosa Beck mss. var. Α sulcis profundioribus’. Bandel & Kadolsky (1982) located corresponding specimens in the Mörch collection in ZMK, confirming the identity of the variety, which had not been figured or adequately described. However, they proceeded to use the name as valid. Nevertheless, since the name was first published in synonymy and had not previously been considered to be available, it is not an available name (ICZN, 1999: Art. 11.6).

Under his new species L. angustior, Mörch (1876) listed a ‘var. Α fasciata’ and listed L. lineata d’Orbigny as a synonym. From the format of the work it is clear that ‘fasciata’ was merely a descriptive adjective (sometimes, as in the example discussed above, he used two or more adjectives) and not intended as a new name. Bandel & Kadolsky (1982) were incorrect to regard it as such. They listed the name in the synonymy of E. interrupta , although as a nomen dubium due to lack of radulae in the ‘type’ lot. This ‘type’ material included d’Orbigny’s (1841) figured specimen of L. lineata from Martinique (BMNH 1854.10.4.139); it and five others are the present species; the other specimen in the lot is E. angustior .

This species has had a confused history and been known under numerous names. The identity of Phasianella lineata Lamarck, 1822 and ‘ L. lineata d’Orbigny, 1840 ’ are discussed in the Taxonomic Histories of E. interrupta and E. angustior respectively. In common with all other black-and-white-striped Echinolittorina species in the western Atlantic, this has been misidentified as L. ziczac when a broad concept of that taxon prevailed ( Tryon 1887; Bequaert 1943; Abbott 1954a). Following the recognition that this was a complex of species, the present one was widely know as L. lineolata ( Abbott 1964, 1974; T.V. Borkowski & M.R. Borkowski 1969; Rosewater 1970; Bandel 1974a) until that name was correctly restricted to the Brazilian species ( Bandel & Kadolsky 1982). As discussed above, Bandel & Kadolsky (1982) divided the present species into three ( N. riisei , N. glaucocincta , N. mordax ) on the basis of radular form, but these names have not been widely used. In the recent literature this species has been most frequently referred to under the specific name riisei ( Reid 1989, 2002a; Williams, Reid & Littlewood 2003; Williams et al.).

Diagnosis: Shell medium size; 6–8 primary spiral grooves; 8–15 deep and often wide grooves above periphery of last whorl; dark oblique wavy axial stripes, most of which bifurcate as they cross a grey spiral band midway between suture and periphery (giving approximately twice as many stripes on anterior than posterior part of shell). Penis with obliquely wedge-shaped tip and subterminal opening of sperm groove. Islands of Caribbean Sea, Bahamas, E coast of Florida. COI: GenBank AJ623043 , AJ623044 , FN298405 , FN298406 , FN298407 .

Material examined: 139 lots (including 25 penes, 15 sperm samples, 11 pallial oviducts, 7 radulae).

Shell ( Fig. 18 ): Mature shell height 4.4–19.1 mm. Shape turbinate to high turbinate (H/B = 1.37–1.67, SH = 1.54–1.80); spire whorls rounded; suture distinct; spire profile slightly convex; periphery of last whorl angled; base may be slightly concave. Columella short, wide, slightly hollowed and pinched at base; no eroded parietal area. Sculpture of 6–8 primary spiral grooves on spire whorls; secondary sculpture may be absent, leaving 8–10 deep grooves above periphery on last whorl, posterior 3 ribs and peripheral rib are up to twice width of others, widest grooves may equal rib width; some secondary riblets may appear (by interpolation in grooves) near suture and periphery of last whorl, giving up to 15 grooves above periphery; spiral microstriae visible in widest grooves; base with 9–17 spiral threads. Protoconch 0.28–0.31 mm diameter, 1.7–1.9 whorls. Ground colour white; usually a spiral pale to dark grey band on anterior half of spire whorls and at mid-point between suture and periphery of last whorl (sometimes extending from shoulder to periphery); with brown to black oblique or wavy axial stripes, most of which bifurcate as they cross the grey band, giving approximately twice as many stripes on anterior as on posterior part of shell; in palest shells the stripes may be yellow-brown lines, darkening over centre of whorl ( Fig. 18B, F ); darkest shells (present at a low frequency in some populations) almost entirely blackish brown, with short whitish stripes at suture, periphery and centre of base only ( Fig. 18A ); aperture dark brown with pale band at base and rarely a second band at shoulder; columella purple brown.

Animal: Head ( Fig. 19F ) black, usually an unpigmented stripe across snout; tentacle pale around eye and across base, with two short longitudinal grey to black stripes up to half tentacle length (often fused to give a grey or black basal band) and usually a black dot at tip; sides of foot with grey or black blotches. Opercular ratio 0.36–0.40. Penis (Fig. A–E, G): filament 0.6–0.7 total length of penis, with annular wrinkles for most of its length (so not clearly differentiated from wrinkled base), sperm groove ends at about two-thirds length of filament, where its lips are broadened, filament extends distally in a smooth oblique wedge-shape; mamilliform gland approximately half size of penial glandular disc, borne together on projection of base; penis unpigmented. Euspermatozoa 53–70 µm; paraspermatozoa ( Fig. 19I, J ) containing 1–2 parallel-sided (occasionally slightly fusiform) rod-pieces 16–29 µm (exceptionally to 49 µm), with rounded ends, usually projecting slightly from cell, which is packed with round granules. Pallial oviduct ( Fig. 19H ): copulatory bursa separates at posterior end of straight section and extends back almost to albumen gland. Spawn ( Fig. 19K ): a large pelagic capsule 660–720 µm diameter by 425–440 µm high, with broad peripheral skirt bearing oblique striations overhanging base, dome-shaped upper half sculptured by 4 concentric rings, containing single ovum 135–160 µm diameter ( Lewis 1960: Barbados; Borkowski 1971: Florida).

Radula ( Fig. 20 ): Relative radula length 4.54–9.60. Rachidian: length/width 2.26–6.0, sometimes narrow ( Fig. 20E, G ); tip of major cusp pointed. Lateral and inner marginal: 4 cusps, tip of major cusp rounded; when rachidian is narrow the major cusp of both lateral and inner marginal is enlarged and more pointed, innermost and outermost cusp of each tooth are then vestigial or absent ( Fig. 20E, G ). Outer marginal: normal form with 6 cusps, but narrowed to a rod with 5 cusps in form with narrow rachidian ( Fig. 20G ).

Range ( Fig. 21 ): On islands throughout the Caribbean Sea, Bahamas, E Florida, rare in Bermuda and Gulf of Mexico. Range limits: Boca Chica Key, Florida ( USNM 820421 ) ; Cape Canaveral Inlet, Brevard Co., Florida ( USNM 820420 ) ; Bermuda ( USNM 94396 ) ; Elbow Cay , Bahamas ( BMNH) ; Mayaguana I., Bahamas ( USNM 663919 ) ; Playa Dorada , Dominican Republic ( BMNH) ; Runaway Beach , Antigua ( BMNH) ; Harrison Point , Barbados ( BMNH) ; Piedra Pretu , Bonaire ( ZMA) ; Cabo San Roman , Paraguana Pen., Venezuela ( BMNH) ; Les Cayes, Dept. du Sud, Haiti ( USNM 440369 ) ; Fort Charlotte , Lucea, Jamaica ( BMNH 20080995 ) ; Gibara , Cuba ( IRSNB) ; Cienfuegos, Cuba ( BMNH) ; La Habana, Cuba ( BMNH 20080994 ) ; Swan Is, Honduras ( USNM 853029 ) ; South Water Cay , Belize ( BMNH) ; Bahía de la Ascención, Quintana Roo, Mexico ( USNM 736255 ) ; Isla Mujeres, Quintana Roo, Mexico ( USNM 662238 ) ; Padre I., Corpus Christi, Texas ( USNM 665901 ) .

This species has a wide distribution, but is almost entirely restricted to coasts and islands with oceanic conditions. It is therefore common on the east coast of Florida, throughout the Bahamas, in northeastern Hispaniola, from the Virgin Islands to Antigua, and as far south as Barbados and St Vincent. It is likewise common in the Netherlands Antilles, northeastern Jamaica, northwestern Cuba, on the northeastern Yucatan Peninsula and on the Caribbean islands off Honduras and Belize. However, on the rest of the Central and South American mainland shore of the Caribbean there is only a single record, from the Paraguana Peninsula in Venezuela, a relatively oceanic site. A single specimen has been recorded from Texas, evidently a rare arrival, since the species is otherwise absent from the Gulf of Mexico. Its occurrence in Bermuda is also based on a single specimen. T.V. Borkowski & M.R. Borkowski (1969) likewise recorded the northern limit in Florida at Cape Canaveral, but noted that it occurred only sporadically south of Key Biscayne.

Habitat: On limestone, beachrock and concrete, rarely on driftwood, in littoral fringe and uppermost eulittoral, on sheltered and moderately exposed coasts. Only in oceanic settings with clear water.

Together with E. ziczac , this occupies the lowest zone of the larger Echinolittorina species , corresponding to the upper yellow zone of the eulittoral and the black zone of the littoral fringe on coral limestone ( Vermeij 1973a; Brattström 1999; Dobson-Moore & Britton 2001; Minton & Gochfeld 2001). On limestone platforms in Jamaica E. jamaicensis is found on surfaces wetted by waves at the seaward edge ( Lang, Britton & Metz 1998), often in pits and crevices ( Minton & Gochfeld 2001) and its erosive activity has been measured ( Dobson-Moore & Britton 2001). The heat coma temperature has been measured as 45.3°C ( McMahon 2001) and it is less resistant to heat than E. ziczac ( Fraenkel 1968) . Animals can survive emersion for at least 14 days ( Britton 1992) and show thermoregulation both above and below ambient temperature ( Lang, Britton & Metz 1998).

In Florida Borkowski (1971) found some mature animals all through the year, but peak spawning occurred from April to November. Spawning in the field occurred whenever the tide rose above MHWS and could be induced by splash in the laboratory; there was no lunar rhythm ( Borkowski 1971). In Jamaica, sizefrequency data suggest settlement from January to March ( Lang, Britton & Metz 1998). Growth rate was measured by Borkowski (1974).

Remarks: The egg capsule and contained ovum are extraordinarily large, two to three times the size of those of any other known Echinolittorina species , either tropical or warm temperate ( Reid 2002 a, b, 2007). A figure of the egg capsule was given by T.V. Borkowski & M.R. Borkowski (1969), with a scale bar erroneously labeled as 1 mm (probably 0.1 mm). Subsequently a new figure with measured dimensions was given ( Borkowski 1971). These were still so large that an error could be suspected, but this large size is confirmed by the earlier observation of Lewis (1960). The protoconch is nevertheless of normal size for the genus, although the larval shell is of only 1.7–1.9 whorls, compared with 2.3–3.0 whorls in other members ( Reid 2002 a, b, 2007). These large capsules are pelagic ( Borkowski 1971), and Lewis (1960) reported that they hatched into swimming larvae (although observations were on a mixture of E. jamaicensis and E. angustior ). Presumably the large egg size reduces the time or possibly even the necessity of planktotrophy.

Together with E. mespillum , E. jamaicensis shows the strongest association with oceanic habitats of all the Caribbean Echinolittorina species. The distributional ranges of these two species are almost identical. The apparent rarity of E. jamaicensis in Bermuda is noteworthy, since this is an oceanic island inhabited by five other Echinolittorina species. There may perhaps be a connection with the type of larval development, if the large egg size reduces the length of pelagic life and hence the potential for dispersal from source populations to the southwest.

An abnormal hyperstrophic shell was reported from Barbados by Harasewych (1977); the larval shell was normal (orthostrophic), but the axis of coiling shifted by 140° following metamorphosis. The resulting adult shell superficially appears to be sinistral, but the adapical and abapical sides of the shell (above and below the periphery) are reversed.

Radular variation in E. jamaicensis is very marked. The radular form with a narrowed rachidian ( Fig. 20G, H ) is strikingly different from the normal type ( Fig. 20A, B ), but there is a range of variation between these extremes ( Fig. 20C–F ). Surprisingly, in a study of radular variability in samples from Florida, Borkowski (1975) did not find it to be pronounced and no narrowed forms were reported. At the time when Bandel & Kadolsky (1982) revised the western Atlantic species, radular form was considered a reliable diagnostic character of many gastropods, and they distinguished three species within E. jamaicensis solely by their radulae. These were not recognized by Reid (1989, 2002a) because no differences in shells or reproductive anatomy could be found, nor consistent patterns of geographical distribution. Similar intraspecific variation is now known in two Echinolittorina from the Indo-West Pacific ( Reid 2007) and three others from the western Atlantic. Limited molecular evidence shows no consistent differentiation of the mitochondrial COI gene between the contrasting radular forms of E. jamaicensis , which all cluster tightly, with genetic distance (K2P) among five individuals in the range 0.08–0.60% (average 0.34%; Fig. 37 ). In Littoraria the radular variation appears to be an ecophenotypic effect of substrate and striking differences are found between specimens collected from wood and rocks ( Reid & Mak 1999). However, a single specimen of E. jamaicensis from driftwood, rather than the usual rock substrate, showed a radula of normal form.

Of the 139 samples examined, a few very dark or almost black shells were present in 7 samples. The occurrence of largely black shells was mentioned by Adams (1850) in his original description of this species.

The shell of this species is readily distinguished from other black-and-white congeners in the region by its broader shape, deep and wide primary grooves above the periphery, axial stripes that bifurcate towards the anterior at the mid-point of the whorl, and usual lack of a distinct spiral black band (although anastomosing of axial stripes at the point of bifurcation may sometimes give this impression; Fig. 18C, J, N ).

The fossil Littorina seminole Petuch, 1991 from the Plio-Pleistocene Caloosahatchee Formation of Florida closely resembles E. jamaicensis .

Figures

FIGURE 18. Echinolittorina jamaicensis. A–C, Eight-mile Rock, Grand Bahama I., Bahamas (BMNH 20080988). D, E, St Ann’s Bay, Jamaica (BMNH 20080989). F, Fort Pierce Inlet, St Lucie Co., Florida, USA (BMNH 20080990). G,H, Fort Pierce Inlet, St Lucie Co., Florida, USA (BMNH 20080991). I, Cape Florida, Dade Co., Florida, USA (BMNH 20080992). J, K, Little Stirrup Cay, Berry Is, Bahamas (BMNH 20080993). L, La Habana, Cuba (BMNH 20080994). M, Littorina janaicensis C.B. Adams, 1850, lectotype, Jamaica (MCZ 186133). N, Fort Charlotte, Lucea, Jamaica (BMNH 20080995). O, Sebastian Inlet, Brevard Co., Florida, USA (BMNH 20080996).

FIGURE 19. Echinolittorina jamaicensis. A–E, G, penes. F, head. H, pallial oviduct. I, J, paraspermatozoa. K, egg capsule (after Borkowski 1971). A, D, Eight-mile Rock, Grand Bahama I., Bahamas (BMNH 20080988; shell H A = 10.4 mm, D = 11.2 mm). B, Tulum, Quintana Roo, Mexico (BMNH 20080997; shell H = 12.4 mm). C, F, I, Fort Pierce Inlet, St Lucie Co., Florida, USA (BMNH 20080990; shell H C = 10.5 mm, F = 9.6 mm). E, G, Jingle Beach, Priory, Jamaica (BMNH 20080999; shell H E = 10.9 mm, G = 8.0 mm). H, Fort Pierce Inlet, St Lucie Co., Florida, USA (BMNH 20080998; shell H = 14.0 mm). J, La Toc Beach, St Lucia (BMNH 20081000). K, south Florida. Abbreviation: o, subterminal opening of penial sperm groove. Shading conventions as in Figure 15.

FIGURE 20. Radulae of Echinolittorina jamaicensis (two views of each radula, flat and at 45°). A, B, normal form; Little Stirrup Cay, Berry Is, Bahamas (BMNH 20080993; shell H = 12.5 mm). C, D, intermediate form; Fort Pierce Inlet, St Lucie Co., Florida, USA (BMNH 20080990; shell H = 12.0 mm). E, F, intermediate form. G,H, narrowed form. E–H, Jingle Beach, Priory, Jamaica (BMNH 20080999; shell H E, F = 13.1 mm, G,H = 8.5 mm). Scale bars = 50 µm.

FIGURE 21. Distribution of Echinolittorina jamaicensis.

FIGURE 37. Molecular phylogeny of Echinolittorina species generated by Bayesian analysis of COI sequence. Outgroups and 26 Indo-West Pacific species of Echinolittorina not shown (for these see Williams & Reid 2004: fig. 2B). Numbers adjacent to branches are posterior probabilities. Radular types (normal or narrowed) of sequenced individuals are shown for E. jamaicensis and E. tuberculata only. This tree does not resolve basal relationships (for these see Fig. 1), but shows high support for species-level groupings. The two radular types do not define distinct species groups. Numbers adjacent to species names indicate the localities, using the same codes as Williams & Reid (2004: appendix). Additional localities are: E. placida: 1, 20 km S Campeche, Mexico; 2, Dauphin I., Alabama, USA; 3, Sebastian Inlet, Florida, USA; E. jamaicensis: 2, Eight-mile Rock, Grand Bahama, Bahamas (incorrect locality was given in Williams & Reid 2004); 3, Playa Dorada, Dominican Republic; 4, Fort Pierce, Florida, USA; 5, Harrison Point Lighthouse, Barbados; E. pulchella: 2, Carajamba, Angola; 3, São Tiago, Angola; 4, Old Ningo, 30 km E Accra, Ghana; 5, Aboadze, 10 km E Takoradi, Ghana; E. tuberculata: 3, Church Bay, Bermuda; 4 and 5, Harrison Point, Barbados; E. meleagris B: 3, Takoradi, Ghana; 4, Old Ningo, 30 km E Accra, Ghana. Voucher specimens are depositied in BMNH. Sequences are available from Genbank.