Paratrypaea bouvieri ( Nobili, 1904 )

Dworschak, Peter C., 2012, On the identities of Callianassa bouvieri Nobili, 1904, C. maldivensis Borradaile, 1904, and C. gravieri Nobili, 1905 (Crustacea: Decapoda: Callianassidae): a morphometric approach, Zootaxa 3149, pp. 39-56 : 45-50

Paratrypaea bouvieri ( Nobili, 1904)

Figs 1 D, 5A–I, 6A–J, 7A, B

Callianassa (Trypaea) Bouvieri Nobili, 1904: 236 ; 1906: 101, 105, pl. 6, figs 3–3b [type locality: Djibouti]; de Man, 1928a: 22 –23; 1928b: 27, 107.

Callianassa (Trypaea) Gravieri Nobili, 1905: 396 ; 1906: 107, pl. 6 fig. 4–4d [type locality: Obock]; Balss, 1915: 2; de Man, 1928a: 23, pl. 6 figs 11–11e; 1928b: 27, 107 (new synonym).

Callianassa (Trypaea) cristata Borradaile, 1910: 263 , pl. 6 fig.7 [type locality: Salomon Atoll, Chagos Archipelago]; de Man, 1928b: 27, 107 (new synonym).

Callianassa (Trypaea) gravieri — Holthuis, 1953: 51.

Callianassa gravieri — Sakai, 1999: 43, fig. 6a,b, d–f.

Callianassa bouvieri — Holthuis, 1958: 37, 38, fig. 15; Sakai, 1970a: 46; Fishelson, 1971: table 1, 121, table 5, 130; Por et al., 1977: table 1, 309; Sakai, 1987a: 303; 1999: 40 (part), fig. 6c; Dworschak & Pervesler, 1988: 3, fig. 3; Dworschak, 1992: 192; Sakai & Apel, 2002: 276; Sakai, 2005: 78 (part); Sepahvand & Sari, 2011: 45, fig. 3.

Paratrypaea bouvieri — Komai & Tachikawa, 2008: 36, figs 10–12; Felder & Robles, 2009: 330 (list), 335(tree).

Gilvossius bouvieri — Sakai, 2011: 374 (part) [not Callianassa maldivensis Borradaile, 1904 ].

Gilvossius gravieri — Sakai, 2011: 377.

Holotype. MNHN Th-65, male, (tl 18, cl 5.3), Djibouti, H. Coutière coll. [in poor condition: cephalothorax and abdomen separated; both P1, left P3 and right P5, P3 detached; both mxp3, right P3 and right dissected mouthparts missing].

Additional material. MNHN Th-79, female, (tl 15, cl 3.8), Obock, Mission Gravier 1904 (holotype of Callianassa gravieri Nobili, 1905 ; M. de Saint Laurent redet. as C. bouvieri ) [in fairly good condition, major P1, both P2 and mxp3, one P3, P4 and P5 missing]. – ZSM 77/1, 1 male (tl 21, cl 4.1, major cheliped missing), Red Sea, Harmil, Pola-Expedition, 4 January 1896 [det. as C. gravieri Nobili by H. Balss and J.G. de Man]. – RMNH D23658, 1 female (cl 4.4, both chelipeds detached), 1 female (cl 4.2, both chelipeds missing), Ghardaqa, Rode Zee kust van Egypte, Eulitoral in gangen in Digenia zone, 1966 Volker Storch legit. – USNM 221975, 5 males, 5 females (cl 5.2–6.2), Red Sea, Sinai Peninsula, Ras Muhamed, lagoon, 0–1 m, Lewinsohn & Manning coll. 16 Oct. 1979 [det. as C. gravieri by R.B. Manning]. – NHMW 6591, MNHN Th- 1186–1194, USNM 266243–266244, SMF 17708–17715, 20 males (tl 13–24 cl 3.1–6.6) and 17 females (tl 12–25, cl 3.1–6.5) Red Sea, Egypt, from mangrove south of Safaga (material mentioned in Dworschak & Pervesler, 1988). – NHMW 19829, 1 male (cl 4.7), Red Sea, Egypt, Tubaya Al Bayda near Safaga, Egypt, intertidal, muddy sand, P. Pervesler coll. October 1992. – ULLZ 8442, 1 female, ULLZ 8443, 1 male, NHMW 25027–25059, 6 males, 8 females, 3 ovigerous female (cl 4.1– 6.4), Red Sea, Egypt, Dahab, Al-Qura, intertidal, P. Dworschak coll. 21, 26, and 31 October 2005. – NHMW 25060–25066, 2 males, 4 females, 1 ovigerous female (cl 4.3–6.0), Red Sea, Egypt, Shura Arwashi near Nabq, intertidal, P. Dworschak coll. 27 October 2005. – SMF 26513, 5 males, 4 ovigerous females (cl 3.1–5.0), Jemen, Socotra, Qalaniyah lagoon, NW coast, SOC/POL-12 (12°41.300'N 053°30.040'E), 0–1 m, Sand, Watt, Eulitoral, T. Wehle leg. 17 April 2000. – SMF 26512, 3 males, 1 female, 2 ovigerous females (cl 3.2–5.6), Jemen, Socotra, Qariyah, NE-coast, SOC/IT 154 (12°38.430'N 054°13.340'E), offene Lagune, 35‰ salinity, sandy inter- & shallow subtidal, M. Apel leg. 7 April 1999. – MNHN Th-1617, 10 males, 5 females, 2 ovigerous females (cl 3.5–6.1), Madagascar, Tuléar, B. Thomassin coll. Aug. 1962 – Sept. 1963, M. de Saint Laurent det., for details on habitats see Thomassin (1978). – ULLZ 8444, 1 female, ULLZ 8445, 1 male, NHMW 25067–25101, 20 males, 14 females (cl 2.5–5.6), Indonesia, Bali, Nusa Dua Beach, intertidal, coll. June 2005. – NHMW 19969–19983, 8 males, 2 females, 5 ovigerous females (cl 4.6–7.2), Japan, Ryukyu, Irabu Is., G. Itani coll. 2–3 June 2000. – ULLZ 6367 ( DNA voucher, GenBank EU882913 , EU875023 , EU882914 , and EU875024 ), 11 males (cl 3.9–6.4), 11 females (10 ovigerous, cl 4.3–5.9) Japan, Fukushima, Tomioka, Station 4, Sample 15, 32°31'N 130°02'E, A. Tamaki coll. 3 August 2004. – USNM 95567, 1 ovigerous female (cl 4.4), USNM 95566, 1 ovigerous female (cl 4.4, right cheliped missing), Kiribati, Onotoa, Gilbert Islands, tide flat area south of Contouma, P.E. Cloud, Geol. Survey coll. August 21, 1951, L.B. Holthuis det. (as Callianassa gravieri ), material (with somewhat different data) mentioned in Holthuis (1953). – NHMW 23020, 2 ovigerous females (cl 3.3, 3.6), Fiji, Viti Levu, Waidroka Resort (18°16.323'S 177°54.174'E), intertidal mud, S. De Grave coll. 3 October 2005.

Description (amended from Komai & Tachikawa, 2008). Carapace 0.24 of total body length; rostrum spiniform, directed slightly downward, falling slightly short of or nearly reaching midlength of eyestalk, anterolateral projection obtuse; anterolateral concavity moderately deep, with tiny cleft; dorsal oval well defined, smooth, cervical groove across about posterior 0.2 of carapace length; linea thalassinica extending to posterolateral margin of carapace.

Length ratio of first to sixth pleonal somites measured along midline 1.0: 1.53: 1.0: 0.9: 1.13: 1.4. First pleomere narrowing anteriorly in dorsal view; pleuron not distinctly delimited. Second pleomere with posterolateral margin of pleuron slightly produced, bearing tuft of few long setae. Third to fifth pleura each with prominent tuft of plumose setae; fourth and fifth pleura also each with longitudinal row of setae on ventral margin posteriorly; posteroventral margin of each pleuron rounded. Sixth pleomere slightly wider than long, subquadrate in dorsal view, bearing shallow notches on posterior one-third; no ventrolateral projection. Telson subrectangular, about as long as wide; dorsal surface nearly flat; lateral margin with two spiniform setae on posterolateral corner, with short transverse suture subproximally; posterior margin convex, median tooth absent or minute ( Fig. 7 A, B).

Eyestalks contiguous, flattened, each reaching distal margin of first segment of antennular peduncle, terminating in rounded projection anteromesially; lateral margin convex; cornea medial, disk-shaped, corneal width 1/2 to 2/3 of peduncular width.

Antennular peduncle as long or slightly longer than antennal peduncle; first segment short, partially concealed by eye-stalk in dorsal view; second segment slightly longer than first segment; third segment more than twice length of second segment, moderately slender, slightly tapering distally; second and third segments with row of long setae on ventral surfaces; antennular flagella both 1/2 to 2/3 as long as peduncle; dorsal flagellum slightly thicker, but shorter than ventral flagellum; ventral flagellum with tufts of long setae on ventral margin. Antennal peduncle reaching distal 0.15–0.2 of third segment of antennular peduncle; distal two segments subcylindrical; scaphocerite rudimentary, subovate; flagellum distinctly longer than carapace, with some setae on every 1 or 2 articles.

Epistome devoid of prominent tuft of setae.

Sternal shield on seventh thoracic somite trapezoidal, broadened anteriorly, divided by deep median groove; no conspicuous grooves delimiting anterolateral lobes.

Mandibles with large, 3-segmented palp, elongated third article of palp slightly tapered and terminally rounded, concave on proximal external surface; incisor process with row of rounded calcareous teeth on mesial margin, with deeply concave inner surface; molar process thin, with two small marginal teeth; paragnath rounded.

Maxillula with long, narrow endopod deflected proximally at articulation; proximal endite with very dense fine setation on lower mesial margin, terminal lobe with large setae; distal endite elongate, proximally narrow, and broadening terminally, where it is armed with short stiff bristles; exopodite low and setose.

Maxilla with stout, unsegmented endopod tapering distally and bearing subterminal tuft of long setae on lateral margin; scaphognathite moderately large, anterior lobe not reaching distal margin of basial endite; coxal endite with rounded plate on outer surface; anterior lobe of basial endite subtriangular.

First maxilliped with endopod reduced to rudimentary bud, still visible in outer view; exopod curved mesially, slightly overreaching distal margin of basial endite, weakly bilobed, with submarginal cluster of very long setae on outer surface mesially; epipod unilobed. Second maxilliped with moderately slender endopod; dactylus slightly longer than wide; exopod falling far short of distal margin of merus, hardly visible in outer view; epipod greatly reduced as rudimentary bud; podobranch absent. Third maxilliped without exopod; ischium-merus operculiform, 1.5 to 1.6 times longer than wide, with dense setation on ventral margin; ischium slightly longer than broad, distinctly widened distally, crista dentata consisting of row of small acute spines arranged in sinuous row on midline; merus about twice wider than long, distinctly shorter than ischium, unarmed on distomesial margin, ventrodistal angle not produced; carpus longer than wide, subequal in length to merus; propodus about twice longer than wide, slightly longer than carpus; dactylus slender, digitiform, shorter than propodus.

Chelipeds (first pereopods) greatly dissimilar in males and females ( Fig. 2 A, B).

Major cheliped ( Fig. 5 A, F, H) located on either right or left side of body (total: 74 left, 75 right), variable in length, armament and setation of dactylus, sometimes greatly elongate in males. Ischium moderately slender, becoming wider distally in general contour, dorsal margin sinuous, unarmed; lateral surface convex; ventral margin with row of 1–7 small teeth or denticles. Merus subequal in length to carpus; dorsal margin with row of small tubercles in proximal 0.3–0.4 ( Fig. 6 A–J); lateral surface generally convex, ventral half forming shallow concavity accommodating proximal part of carpus; mesial surface slightly concave in general; ventral margin variable, from median spine to broadly subtriangular, strongly compressed, marginally denticulate projection, often notched medially in large males ( Fig. 6 A, J). Carpus quadrate or subquadrate, sometimes elongate in males; dorsomesial and ventromesial margins sharply ridged, each edge smooth ( Fig. 5 C) or slightly denticulated; lateral surface smooth, convex; mesial surface medially convex, margins strongly upturned. Palm with dorsal and ventral margin parallel or slightly diverging proximally, 1.0–1.1 times longer than wide; lateral surface smooth, convex, with tufts of short setae adjacent to margins; palmar process absent; mesial surface convex medially, without sculpture or armament, margins somewhat upturned, dorsomesial and ventromesial margins sharply ridged, smooth ( Fig. 5 B). Fixed finger about half length of palm, clearly overreaching midlength of dactylus, nearly straight to noticeably curved, terminating in subacute or acute tip; cutting edge unarmed or minutely dentate; lateral surface convex, with tufts of short setae adjacent to cutting edge; ventral margin with several long setae; concavity on mesiodorsal part occasionally with patch of dense setae. Dactylus subequal in length to or shorter than palm, weakly hooked, terminating in acute or subacute tip; dorsal and lateral surfaces occasionally with mat of dense, soft setae; cutting edge variable, from sinuous ( Fig. 5 H) to bearing 1 or 2 molar-like teeth ( Fig. 5 A, F).

Minor cheliped ( Fig. 5 D, E, G, I) rather slender, about 0.6–0.7 length of major cheliped, also showing considerable variation in stoutness and proportion of segments. Ischium with margins unarmed, ranging from slightly shorter to longer than merus; dorsal margin nearly straight; ventral margin weakly concave to straight, unarmed. Merus varying from shorter to slightly longer than carpus; dorsal margin convex, unarmed; lateral surface weakly convex, smooth; ventral margin usually with 1 small spine at midlength. Carpus 1.5–2.0 times longer than wide; dorsal and ventral margins bluntly ridged, smooth, with row of short setae. Palm as long as wide or slightly wider than long; dorsal margin bluntly ridged, smooth; ventral margin sharply ridged, with row of tufts or individual setae; lateral and mesial surfaces weakly convex, smooth. Fixed finger triangular, terminating in acute tip; cutting edge with small denticles. Dactylus longer than palm, as long as fixed finger, terminating in acute tip; dorsal margin rounded, with row of tufts of setae; cutting edge with small denticles; hiatus between fingers prominent, sometimes filled with dense setation ( Fig. 5 E).

Second pereopod chelate, moderately long and slender; ischium with numerous setae along ventral margin; merus with dorsal margin smooth, nearly straight, ventral margin sinuous, with row of numerous long setae; carpus subtriangular; chela triangular, with scattered tufts of short setae on lateral surface; palm much shorter than fingers; both fingers triangular, terminating in small corneous tip, cutting edges bordered by thin corneous ridge; carpus and chela fringed with short to long setae along margins.

Third pereopod moderately stout; ischium with weakly produced ventrodistal angle; merus about 2.7 times longer than wide, with small spine on ventral margin proximally; carpus subtriangular, unarmed; propodus roundly subrectangular, about 1.5 times wider than long, with numerous tufts of short setae on lateral surface and row of numerous long setae along dorsal and ventral margins; ventral margin armed with 1 small subdistal spiniform seta practically obscured by simple setae; posterior margin not forming conspicuous heel; dactylus triangular, terminating in short, ventrolaterally directed corneous tip, lateral surface covered with short setae, dorsal and ventral margins with numerous short setae.

Fourth pereopod moderately stout, all segments unarmed; coxa flattened ventrally, unarmed, movable; propodus 2.4–2.6 times longer than wide, slightly longer than carpus, lateral surface and ventral margin densely setose; dactylus elongate oval, about 2.5 times longer than wide, terminating in small, corneous tip.

Fifth pereopod chelate, moderately stout; propodus shorter than carpus, somewhat broadened distally.

First and second pleopods absent in males. Female first pleopod with ramus slightly longer than protopod, abruptly tapering at midlength. Female second pleopod with exopod noticeably curved mesially, slightly shorter than endopod; endopod with shallow concavity on mesial margin slightly distal to midlength; protopod strongly curved. Third to fifth pleopods biramous, rami broad; appendices internae stubby, arising at about proximal 0.3 of endopod, distinctly projecting beyond margin of endopod, bearing numerous small adhesive hooks along subtruncate mesial margin.

Uropod overreaching posterior margin of telson. Endopod distinctly longer than wide, with obsolete middorsal carina and small dorsal plate, bordered with row of corneous spinules or stiff setae subterminally; posterior margin of endopod unarmed. Exopod nearly as long as wide, with broad middorsal carina, unarmed on posterior margin; dorsal plate conspicuous, bordered with mixture of spiniform setae.

Size. Males from 2.5 to 6.8 mm cl, females (ovigerous) from 3.1 to 6.8 mm cl.

Colour. Chelipeds with a touch of pink on chelipeds dorsally, whitish ventrally, and red chromatophores on dorsal abdomen. Yellow to orange hepatopancreas and orange ovaries in mature females visible in abdomen through transparent integument. Hemolymph of body partially or entirely blue or green in many specimens ( Fig. 1 D).

Parasites. Two males from Safaga (1986), 1 male from Ras Muhamed, 1 female each from Socotra and Dahab (Al Qura) were parasitised by a bopyrid isopod; one male from Bali had a cryptoniscid.

Habitat. Tidal flats consisting of fine to muddy fine sand, often between or near mangroves ( Fig. 1 A, B). Here, P. bouvieri occurs in high densities of up to 454 individuals m -2; their burrow openings are characterised by small mounds and funnels ( Fig. 1 C) ( Dworschak & Pervesler, 1988).

Distribution. Red Sea, Egypt, Sudan; Gulf of Aden, Djibouti (type locality); Socotra; Persian Gulf; Madagascar; Chagos Archipelago ( Borradaile, 1910 as C. cristata ), Indonesia; Japan; Kiribati; Fiji.

Remarks. For P. bouvieri , the male:female ratio of individuals in samples with more than 20 specimens (Safaga May 1986, Nusa Dua, Tomioka) ranged from 1:1 to 1.46:1, the overall ratio for the studied specimens is 1.14:1. The percentage of ovigerous females in these three samples ranged from 0 (Nusa Dua) to 70% (Safaga) and 91% (Tomioka); altogether 39% of the females studied were ovigerous.

The differences between C. bouvieri and C. gravieri as outlined by Nobili (1906) —propodus of P3 not projecting on its lower anterior border and telson with crests dorsally in the former versus anteriorly projecting P3 propodus (pl. 6 fig. 3a, b) and dorsally not crested telson in the latter (pl. 6 fig. 4b, d)—are rather subjective and could not be detected in the type material. Sakai (1999: 36) gives in the key as the difference between the two species the relative length of the antennular and antennal peduncles: C. bouvieri with A1 peduncle longer than A2 peduncle and C. gravieri with A2 peduncle longer than A1 peduncle, but reported the other way round in the diagnoses (p. 41 and 43), both of which comply neither with the descriptions by Nobili (1906) —A1 peduncle longer than A2 peduncle—nor with the figures presented by Sakai (1999: fig. 6 a and b), which show clearly that A1 peduncle is longer than A2 peduncle in both species he considered as such.

Michèle de Saint Laurent (N. Ngoc-Ho, pers. comm. 2001) considered C. gravieri synonymous with C. bouvieri . The denticulation on the ischium of both P1 and the three spines on the merus of major P1 seem to confirm this view, as the P 1 in C. bouvieri shows great variations (compare Dworschak & Pervesler, 1988) and the type specimen of C. gravieri is very similar to small females of C. bouvieri (NHMW 6591/24). Unfortunately, the major cheliped is now missing. The missing telson spine and the ratio telson length/telson width points to P. bouvieri . Assuming a serrated upper border of the major cheliped, the specimen falls into P. bouvieri when included in the MDS and cluster analyses (not shown).

The specimen mentioned by Balss (1915) and re-investigated by de Man (1928a) is clearly a male C. bouvieri with the major cheliped missing.

The two females from Kiribati mentioned by Holthuis (1953) as well as the two females from the Red Sea listed by Sakai (1999) are considered here to belong to P. bouvieri , because the chelipeds (as far as present on both sides) are unequal (see Fig. 2 B), the ischium of the major one shows spines on the lower margin and the telson lacks a spine.

De Man (1928b) speculated on the synonymy of C. cristata with C. bouvieri ; this was accepted by Sakai (1999) without further investigation of the type (which was lost according to de Man, 1928b).

Figures

FIGURE 1. Paratrypaea bouvieri (A – D) and P. maldivensis (E, F), habitat (A, B), burrow openings (C, E) and live specimens (D, F). A, C, Dahab, El Qura; B, mangrove at Shura Arwashi near Nabq; D, male specimen (NHMW 25048, cl 6.8) from Dahab; E, sediment surface in 10 m depth at " Laguna " (photograph: Philip Hofer); F, male (top, NHMW 25115, cl 3.9) and female (bottom, NHMW 25125, cl 3.6) specimen from Aqaba. Scale is 5 cm in C and E, 1 cm in D and F.

FIGURE 2. Plots of propodus height of major (phma, filled symbols) and minor (phmi, open symbols) chelipeds of males (A, C, squares) and females (B, D, circles) of P. bouvieri (A, B) and P. maldivensis (C, D). Triangles represent data for the holotypes of C. bouvieri and C. maldivensis, inverted triangles data for holotypes of C. gravieri and C. rectangularis.

FIGURE 5. Paratrypaea bouvieri (A – I) and P. maldivensis (J – U), major and minor chelipeds. A – D, MNHN Th- 65 (male holotype of C. bouvieri, cl 5.3); E, MNHN Th- 79 (female holotype of C. gravieri, cl 3.8, minor cheliped only); F, G, NHMW 6591 / 2 (male cl 4.4); H, I, NHMW 6591 / 15 (ovigerous female cl 5.2); J – M, UMC I. 58505 (male holotype of C. maldivensis, cl 4.8); N – Q, MNHN Th- 1069 (male holotype of C. rectangularis, cl 4.5); R, S, NHMW 25120 (male cl 3.5); T, U, NHMW 25128 (female cl 4.0). Mesial aspects except H, I, M; B, K, N, lower border of propodus in dorsomesial view; C, L, O, carpus in dorsal view. Most setation omitted. Scale is 1 mm.

FIGURE 6. Outline of selected major cheliped merus in mesial view (partially mirrored for uniform orientation). A – E, P. b o u - vieri males; F – J, P. bouvieri females; K – P, P. maldivensis males; Q – U, P. maldivensis females. A, NHMW 25098; B, NHMW 25086; C, NHMW 19829; D, NHMW 25076; E, NHMW 19983; F, NHMW 6591 / 24; G, NHMW 25050; H, NHMW 25073; I, NHMW 25085; J, NHMW 19980; K, NHMW 25135; L, NHMW 25102; M, ULLZ 8428; N, NHMW 25120; O, MNHN Th- 1069 (holotype of C. rectangularis); P, UMC I. 58505 (holotype of C. maldivensis); Q, NHMW 25117; R, NHMW 25107; S, NHMW 25129; T, NHMW 25128; U, NHMW 25105. Scale is 1 mm. Numbers to the left indicate carapace length of the respective specimen in mm.

FIGURE 7. Posterior border of telson. A, no median spine (P. bouvieri, NHMW 25076, male cl 5.3); B, tiny median spine (P. bouvieri, NHMW 25065, male cl 6.6); C, prominent median spine (P. maldivensis, NHMW 25145, female cl 4.4). Scale is 100 µm.