Neocallichirus jousseaumei ( Nobili, 1904 )

Dworschak, Peter C., 2011, Redescription of Callianassa jousseaumei Nobili, 1904, a junior subjective synonym of Callianassa indica de Man, 1905 with description of a new species of Neocallichirus (Decapoda: Axiidea: Callianassidae), Zootaxa 2746, pp. 1-19 : 2-9

Neocallichirus jousseaumei ( Nobili, 1904)

( Figs 1–4 , 6 F–H)

Callianassa (Cheramus) Jousseaumei Nobili, 1904: 236 ; 1906: 101, pl. 6 fig. 2; de Man, 1928: 26, 97, 100 (key), pl. 18 fig. 27– 27a.

Callianassa (Cheramus) indica de Man, 1905: 605; 1928: 26, 100, 159, 160, pl. 17 fig. 26– 26g [Type locality: Station 16, Lat. 6°59'S long. 115°24.7' E, Bay of Kankamaraän, S. coast of Kangeang, reef].

Callianassa indica . — Kensley, 1976: 50, fig. 2A–E; Sakai, 1987: 302, 306.

Callichirus indicus . — de Saint Laurent & LeLoeuff, 1979: 97.

Callichirus jousseaumei . — de Saint Laurent & LeLoeuff, 1979: 97.

Neocallichirus taiaro Ngoc-Ho, 1995: 212, figs. 1–2. [Type locality: Taiaro atoll, Tuamotu Island, French Polynesia]; Tudge et al., 2000: 144.

Neocallichirus indicus . — Sakai, 1999: 99, fig. 23a, b, d, e (not Neocallichirus indicus ; Sakai, 1999: 99, 100, fig. 23c; 2005: 178 [part, holotype of C. natalensis ]); Sakai & Apel, 2002: 277, fig. 2; Robles et al., 2009: 314 (tree), 317 (list).

Neocallichirus jousseaumei . — Sakai, 1999: fig. 22g.

Neocallichirus indica . — Tudge et al., 2000: 144 (list).

Not Callichirus jousseaumei . — de Vaugelas, 1984: 529 [misindentification = N. vaugelasi sp. n.].

Not Callianassa jousseaumei . — Dworschak, 1992: 198, fig. 5a–d, 6a–c [misindentification = N. vaugelasi sp. n.].

Not Neocallichirus jousseaumei . — Sakai, 1999: 100, fig. 22e, f; 2005: 179; Tudge et al., 2000: 144 (list) [misindentification = N. vaugelasi sp. n.]

Type material. " Djibouti et Perim, Dr. Jousseaume coll.", 1 male (tl 57, cl 15.4, figured) LECTOTYPE (here designated to fix the concept of Callianassa jousseaumei Nobili, 1904 and to ensure the universal and consistent interpretation of the same); PARALECTOTYPES, 1 female (tl 61, cl 15, figured), 1 female (tl 61, cl 15.7, minor P1 missing, figured), 1 male (tl 56, cl 14.9), 1 female (tl 45, cl 12, minor P1 missing), 1 female (cl 10), 1 male (tl 54, cl 14.4, major P1 missing), 1 female (cl 14, both P1 missing), 1 female (cl 11, both P1 missing), 1 female (cl 13.5, both P1 missing), 1 male (damaged), 1 male (tl 46, cl 12), 1 male (tl 40, cl 10.8, both P1 missing), 1 male (tl 31, cl 8.1), 1 female (tl 40, cl 10.6, both P1 missing), 1 female (tl 38, cl 10.8, both P1 missing), 1 male (tl 48, cl 12.4, both P1 missing), 1 male (tl 34, cl 8.6, both P1 missing), 1 male (tl 19, cl 5.0, major P1 missing, figured) MNHN Th-83.

—"mer Rouge, M. Jousseaume coll. 1897, auct. det.", 2 females (not measured) MNHN Th-81. — "golf de Tadjourah, M. Faurot coll. 242.95", 1 male, 1 female (not measured) MNHN Th-82. — "Museum Paris Djibouti, H. Coutière, 109-97", 1 male (cl 8, major P1 missing, minor P1 detached), 1 female (cl 9.5, major P1 detached) MRSN Cr.460 (ex 2295).

Non-type material. " Djibouti ( Somalia) Det. Auch. 1905", 1 male (tl 60, cl 16.3, both P1 missing), 1 female (cl 14.6, broken in 2 parts), 1 male (tl 40, cl 11.7, both P1 missing), 1 male (tl 37, cl 9.4, both P1 missing), 1 male (tl 31, cl 8.3, both P1 missing), 1 vial with dissected appendages and one minor P1, MRSN Cr.467. — Red Sea, coll. E. Rüppell 1827, 1 male (tl 36, cl 10.4) SMF 4959. — Yemen, Socotra, E of Hadibo, N-coast, SOC/IT-178a (12°39.740'N 054°02.630'E) under stones in shallow water, M. Apel coll. 14 April 1999, 1 female (tl 40, cl 12.4) SMF 26515. — United Arab Emirates, Fujairah, Sandy Beach Hotel, Al Aqqa, between mainland and small island at reef near small island, UAE 95-28 (25°30.000'N 056°22.000'E), 3–4 m, in sand under stone, M. Apel coll. 4 July 1995, 1 male (tl 28, cl 9.2) SMF 26516. — Philippines, Bohol, Panglao I., Alona Beach (M1: 09°32.9'N, 123°46.6'E), intertidal: coll. June 2004, 1 male (tl 34 cl 10.2) ZRC 2010.0396; 1 female (tl 28 cl 8) NHMW 25032; –P.K.L. Ng coll 3 June 2004, 1 male (tl 35 cl 10.2) ZRC 2010.0385 (Photo sp.2); –P. Dworschak coll. 5 June 2004, 1 female (tl 74 cl 21.3) NHMW 21942 (PD030); –P. Dworschak coll. 6 June 2004, 1 male (tl 76 cl 21.4) NMCR 39021, (PD032/1); 1 female (tl 66 cl 17.7) ZRC 2010.0391 (PD032/2); 1 female (tl 59 cl 16) MNHN Th-1608 (PD032/3); 1 female (tl 51 cl 14.2) ZRC 2010.0392 (PD032/4); 1 male (tl 46 cl 12) MNHN Th-1609 (PD032/5); – P. Dworschak coll. 8 June 2004, 1 female (tl 70 cl 19) MNHN Th-1611; –P. Dworschak coll. 15 June 2004, 1 male (tl 83 cl 23.5), NHMW 25036 (PD109); 1 female (tl 73 cl 21), NMCR 39024 (PD110); 1 male (tl 66 cl 17.7), ZRC 2010.0397 (PD111); 1 male (tl 85 cl 22.8), NHMW 25033 (PD112); 1 male (tl 70 cl 18.5) ZRC 2010.0395 (PD113); 1 female (tl 58 cl 15.7) MNHN Th-1613 (PD114); 1 female (damaged tl ca 48), MNHN Th-1614 (PD115); 1 male (tl 61 cl 17), MNHN Th-1615 (PD116); 1 male (tl 65 cl 18.2); NHMW 25034 (PD117); 1 female (tl 53 cl 16), NMCR 39023 (PD118); –Joelle Lai coll. 16 June 2004, 1 male (tl 85 cl 24.4), NHMW 25041 (JL010); 1 female (tl 73 cl 20.2) NMCR 39027 (JL011); 1 female (damaged, cl 20) NMCR 39022 (JL012); 1 female (tl 55 cl 15.8) ZRC 2010.0390 (JL013); 1 female (tl 56 cl 15.9) NHMW 25031 (JL014); 1 male (damaged, cl 12.6) NMCR 39025 (JL015); –P. Dworschak coll. 18 June 2004, 1 male (tl 70 cl 20.7), 1 female (tl 35 cl 11), NHMW 25035 (PD144); –P.K.L. Ng coll. 3 July 2004, 1 female (tl 49 cl 13.5) ZRC 2010.0386 (3.7./01); 1 female (tl 50 cl 13.1) NMCR 39029 (3.7./02); 1 female (tl 60 cl 17) MNHN Th-1604 (3.7./03); 1 female (tl 75 cl 20.7) NMCR 39026 (3.7./04); 1 male (tl 52 cl 15) NHMW 25027 (3.7./05); 1 male (58 cl 16.3) MNHN Th-1605 (3.7./06); 1 female (tl 69 cl 18.6) ZRC 2010.0387 (3.7./07); 1 male (tl 57 cl 16.6) NHMW 25039 (3.7./08); 1 female (tl 55 cl 15.1) ZRC 2010.0388 (3.7./09); 1 female (tl 80 cl 22.5) NHMW 25038 (3.7./10); 1 male (tl 63 cl 16.7) NHMW 25028 (3.7./ 11); 1 male (tl 61 cl 16.6) NHMW 25040 (3.7./12); 1 female (tl 75 cl 20.8) ZRC 2010.0389 (3.7./13); 1 female (tl 63 cl 17.5) MNHN Th-1606 (3.7./14); 1 male (tl 50 cl 13.8) NHMW 25029 (3.7./15); 1 male (tl 36 cl 10.6) NHMW 25030 (3.7./16); 1 female (tl 39 cl 12) MNHN Th-1607 (3.7./17). – Philippines, Bohol, Panglao I., Sungcolan Bay (M11: 09°38.3'N, 123°49.6'E), intertidal sand, P. Dworschak coll. 7 Jun.2004, 1 female (tl 75 cl 20) ZRC 2010.0393 (PD041); 1 female (tl 83 cl 22.2) NHMW 25037 (PD042); –P. Dworschak coll. 8 June 2004, 1 male (tl 62 cl 17.2) MNHN Th-1610 (PD051); 1 female (tl 66 cl 18.3) ZRC 2010.0394 (PD052); 1 male (tl 74 cl 21.3) MNHN Th-1612 (PD054); 1 male (tl 84 cl 24.5) NMCR 39028 (PD055); 1 ovigerous female (tl 75 cl 23.2) NMCR 39030 (PD056). — Thailand, Andaman Sea, Phangnga, Laem Pakarang [8° 44'06"N 98°13'18"E], intertidal, in muddy sand at transition from beach to coral rubble, coll. July 2007: 1 female (carapace damaged) NHMW 21952; 1 female (tl 44 cl 12) NHMW 21953; 1 male (tl 37 cl 10.2) NHMW 21954; 1 female (tl 38 cl 11) NHMW 21955. – Thailand, Andaman Sea, Phangnga, Khao Lak, Sunset Beach [8°37'41" N 98°14'26" E], small beach, sand between boulders, intertidal, coll. July 2007: 1 male (tl 42 cl 12.2) NHMW 21956; 1 male (tl 57 cl 16) NHMW 21957.

Redescription. Dorsally, carapace as long as abdominal somites 1 and 2 combined, ca 1/4 of total length. Frontal margin of carapace with three anterior prominences, lateral prominences obtusely angular, sometimes appearing uncalcified; median prominence reaching beyond laterals, forming short, broadly rounded rostrum ( Figs 1 A, B, 2A, B, 3A, B, 4A, B). Carapace lacking cardiac prominence and dorsal carina, with distinct linea thalassinica; dorsal oval distinctly marked posteriorly by deep transverse cardiac furrow, the latter extending anteroventrally to either side above linea thalassinica as shallow groove marking posterior half of dorsal oval. Frontal margin of carapace continued ventrolaterally beyond intersection with linea thalassinica as thickened oblique ridge ending anteriorly at prominent hepatic boss ( Fig. 3 B). Sclerotised ridge along anterodorsal margin of anterior branchiostegal lobe articulating at junction of oblique ridge and linea thalassinica. Subantennular region of epistome bearing dense tuft of long setae.

Eyestalks reaching to or beyond basal antennular article, ca 1.5 times as long as broad; outer margins convex, terminating in a dorsoventrally flattened rounded lobe bearing one to several tubercles distally ( Figs 1 A, B, 2 A, B, M, N, 3A–D, 4A, B). Cornea black, situated dorsolaterally in distal 1/2 of eyestalk, ca 1/3 to 1/2 width of eyestalk. Black pigment filling eyestalk to variable extent proximal to cornea (extracorneal pigment) ( Figs 2 M, N, 3C, D).

Antennular peduncle ( Fig. 2 O) thicker, but shorter than antennal peduncle; second article slightly shorter than basal article; terminal article 1.5 times as long as second.

Antennal peduncle ( Fig. 2 P) with basal article with dorsolateral carina forming lip above excretory pore; second article longer than first, third article short, visible in lateral view as short triangle ventral to second article and vestigial antennal scale; fourth article elongate, as long as basal, second and third article combined; fifth article narrower, as long as fourth article.

Mouthparts as figured ( Fig. 3 E–K), typical for genus. Mandible with toothed molar process ( Fig. 3 E). Third maxilliped ( Figs 1 C, 2C, Q, 3K, J, 4C) without exopod; endopodal ischium 1.1 times as long as broad (lectotype), mesial surface with row of teeth (crista dentata); merus triangular, 1.3 times as broad as long; carpus triangular, longer than broad; propodus large, ovoid, 1.1 times as broader as long; dactylus narrow, arcuate.

Major cheliped massive, ( Figs 1 D, E, 2D, R, 3L, M, 4E, F) located on either right or left side of body (Panglao: 28 left, 22 right), shape not sexually dimorphic (see below). Ischium slender, inferior margin with row of teeth; merus with toothed blade on inferior margin, widest proximally; carpus slightly shorter than merus, with straight superior and convex inferior margin; propodus much longer than carpus (see Table 1) with low keel in proximal half of superior margin, inferior margin of palm serrated; fixed finger slightly curved, cutting edge smooth or with low tubercles proximally, dactylus curved, broad, longer than fixed finger, cutting edge variable (see below).

Minor cheliped massive ( Figs 1 F, G, 2H, 3N), about 0.7 times as long and 0.5 times as high as major cheliped, sparsely armed, ischium with row of minute denticles proximally on inferior margin; merus unarmed, with rounded superior and inferior margins; carpus as long as high; palm of propodus slightly longer than carpus (see Table 1), fixed finger shorter than palm, triangular; dactylus slightly curved; cutting edges of fixed finger and dactylus smooth. Tips of both fixed fingers and dactylus corneous.

N. jousseaumei N. jousseaumei N. jousseaumei N. vaugelasi sp. n. N. natalensis type material Panglao Khao Lak Aqaba Malindi

plma/calma 1.7 (1.35–2.07) 1.8 (1.33–2.36) 1.9 (1.45–2.22) 1.6 (1.42–1.82) 1.0 plmi/calmi 1.5 (1.27–1.74) 1.14 (0.8–1.9) 1.1 (0.89–1.52) 0.68 (0.61–0.85) 0.64 Second to fifth pereopod as figured ( Figs 1 H–K), typical for genus. Third pereopod ( Figs. 1 I, 2I, 3O) propodus rhomboidal with proximally-directed lobe of inferior margin not reaching beyond broadest part of carpus (not heeled).

Abdomen long ( Figs. 6 F–H); dorsal length ratio (along midline) of first to sixth abdominal somites 1.0: 1.47: 1.18: 1.05: 1.15: 1.57 (lectotype).

Male first pleopod consisting of two articles ( Figs 1 L, 3Q, 4G–I, L); second article same length as first, bilobed or with rounded lobe and acute hooked tip distally (see Variations below). Female first pleopod simple ( Fig. 2 J), consisting of two articles; terminal article with shoulder at midlength.

Male second pleopod ( Figs 1 M, N, 3R, 4J, K, M–O) biramous, variable with respect to shape and demarcation of appendix masculina and appendix interna (see below). Female second pleopod biramous ( Fig. 2 K), endopod with appendix interna.

Third to fifth pleopods with appendix interna embedded into mesial margin of endopod.

Telson ca 1.2 times as broad as long, broadest proximally, lateral and posterior margins convex. Uropodal endopod slightly longer than telson, rhomboidal, about as long as wide. Uropodal exopod longer than endopod, with anterodorsal plate ( Figs 2 L, 3P, 4D).

Variations. The shape of Mxp3 is variable, ischium-merus length to width ratio ranged from 1.7 to 2.5 (mean 2.1) in the specimens from Panglao; the Mxp3 differed between left and right in two specimens (see Fig. 3 J, K). The tuberculation on the eyestalks varies also, there is at least one tubercle on the distal corner, additional tubercles may be present mesiodistally. The male first pleopods, irrespective of size, may have two rounded lobes ( Figs 3 Q, 4H) (5 in type series, 7 from Panglao) or be hooked ( Figs 1 L, 4G, I) (6 in type series, 8 from Panglao); one male had an intermediate form, another one ( NHMW 25030, tl 36) a simple Plp1 terminus ( Fig. 4 L). The latter showed also an appendix interna on the endopod of Plp2 ( Figs 4 M, N). Another male ( ZRC 2010.0396, tl 34) has a hooked Plp1 and an appendix interna on Plp2 ( Figs 4 I–K). An appendix interna on the left side only was observed in another small male from the type series ( MNHN Th-83, tl 19), whereas all the larger males have no appendix interna and show a weakly demarcated appendix masculina ( Figs 1 N, 3R, 4O). The major chelipeds show no obvious sexual dimorphism in shape, but chelipeds become slightly larger in males (plma = 0.8276 cl - 2.6583, n = 22, r = 0.90) than in females (plma = 0.5652 cl + 0.5113, n = 29, r = 0.88) from Panglao. Slight variations exist in the cutting edge of the dactylus, which may be entire (8 specimens, e.g. as in the lectotype, Figs 1 D, E) or show a distal triangular and an indented proximal rectangular tooth (41 specimens, e.g. as in PD32/1, Figs 3 L, M, 4E, F). At Panglao, 28 specimens had the major cheliped on the left side, 23 had it on the right side, sex ratio (males: females) was 22: 30 for Panglao and 16: 14 for the type material, only one (tl 75) out of 30 females from Panglao was ovigerous. Embryo diameter is 714–785 µm

Size. For Panglao, tl 28–85 mm, cl 8–24.5 mm.

Colour. Transparent with a touch of pink on the dorsal abdomen and chelipeds ( Fig. 6 F) to brightly pink ( Fig. 6 G, H).

Commensals. At Panglao, many specimens had numerous clausidiid copepods on their body surface.

Habitat. Lower intertidal in coral rubble covered by fine sand or fine sand between boulders (see Figs 6 A–C).

Distribution. Djibouti, Perim, Gulf of Tadjourah, Red Sea (type locality); Socotra, Persian-Arabian Gulf ( Sakai & Apel 2002); Indonesia (type locality of C. indica de Man, 1905); Thailand, the Philippines (this study); French Polynesia (type locality of N. taiaro Ngoc-Ho, 1995).

Remarks. Present in the Muséum national d’Histoire Naturelle, Paris (as of May 2007) are the specimens from Djibouti et Perim (MNHN Th-83 collected by Jousseaume in 1891), Mer Rouge (MNHN Th-81 collected by Jousseaume in 1897) and golfe de Tadjourah (MNHN Th-82 collected by Faurot). The number of specimens agrees with that given by Nobili (1906). The seven specimens from Djibouti collected by Coutière remained in Torino, where Nobili worked; some (5 specimens) were later sent on loan to J. G. de Man. Parenti (1971) lists two lots of C. jousseaumei : MRSN Cr.460 (2 specimens) and MRSN Cr.467 (5 specimens), none indicated as type material. Both lots were found at the MRSN in September 2008; only the former is indicated as type on the label. The sizes given by de Man agree well with the two larger specimens from MRSN Cr.467, the sexes given by Nobili (1906) – 4 males, 3 females – and those found in 2008 – 5 males, 2 females – do not. The "Det. Auch. 1905" on the label, however, seems to indicate that the specimens came to the collection later. It is unknown what "Det. Auch." might mean (L. Levi, C. Froglia pers. comm. Sep. 2008), but it could be a misinterpretation of "auct. det", which can be found on the label of MNHN Th-81, meaning that the author of the species (G. Nobili) identified the specimens. The specimens of MRSN Cr.467 are here not considered as syntypes and therefore do not become paralectotypes. The type material both in MNHN and MRSN is in fairly good condition, but fewer than half of the specimens had the major cheliped still attached.

De Man (1928) compared C. indica with material of C. jousseaumei he received on loan from Turin (probably MRSN Cr.460, see above). He concluded that there are two differences, one in the shape of the male Plp1, the other in the length relation of Mxp3 ischium and merus. Edmondson (1944), in his description of C. variabilis , concluded that "Structural features selected by De Man to distinguish C. indica from C. jousseaumei are found in the Hawaiian form to have wide ranges of variation, which suggest that if sufficient material had been available de Man might have found the species from the Red Sea and Kangean to be identical". He finally gave, as a very constant character in which indica / jousseaumei and variabilis differ, the presence of a distinct anteroventral lobe on the Mxp3 propodus in the latter. The "lobe" on the Mxp3 propodus, however, was also figured for the type material of C. jousseaumei by de Man (1928: pl. 18 fig. 27). In the type series as well as in the material from Panglao, this lobe is recognisable in most specimens (Panglao: no lobe 20, with lobe 29).

According to Art.23.9.1 (International Code on Zoological Nomenclature, 1999), the use of a junior synonym is to be conserved when it has been used in 25 works by 10 different authors in the past 50 years. This is not applicable here because indicus has been published in that period in only 12 papers by 7 authors (see synonymy). Therefore, jousseaumei takes precedence over indicus .

The specimen attributed to C. jousseaumei by Dworschak (1992), subsequently by Sakai (1999) and mentioned in Tudge et al. (2000), is different from this species (see below).

Sakai (1999) synonymised several taxa with N. indicus :

1) Callianassa (Cheramus) variabilis Edmondson, 1944 from Hawaii, Hanauma Bay, Oahu, in gravel bed of the intertidal zone. As mentioned above, no essential morphological difference exists between this species and N. jousseaumei . Study of recently collected material from Hawaii (ULLZ) confirmed this (pers. obs. June 2008). Molecular studies, however, indicate that N. variabilis is sufficiently different from N. jousseaumei (as N. indicus ) (R. Robles & D.L. Felder, pers. comm. 2009). Neocallichirus variabilis is therefore considered a separate species here.

2) Callianassa natalensis Barnard, 1947 based on a single female specimen of tl 100 mm collected from the stomach of Rock Cod at the Natal coast, South Africa. Barnard (1947, 1950) mentioned that it resembles C. indica , yet without giving further details. Neocallichirus natalensis is considered different from N. jousseaumei here. Further details are given below.

3) Neocallichirus manningi Kazmi & Kazmi, 1992 known from two females with tl 35 and 22 mm collected in the lower intertidal region at Sandpit, Karachi, Pakistan. Sakai (2005) removed this species from synonymy with N. indicus , but still lists the locality among the latter's distribution. Neocallichirus manningi differs from N. jousseaumei by i) the shape of the telson, which has a median terminal spine, ii) the lower border of the major cheliped is smooth (denticulated blade in N. jousseaumei , always present even in small specimens both from the type series and Panglao). In addition, the carpus of both major and minor P1 is much longer than the palm.

Figures

FIGURE 1. Neocallichirus jousseaumei (Nobili, 1904), lectotype of Callianassa jousseaumei Nobili, 1904, MNHN Th 83, male (tl 57, cl 15.4, telson damaged): A, front in dorsal view; B, front in lateral view; C, third maxilliped in lateral view (setae omitted); major cheliped in lateral (D) and mesial (E) view; F, minor cheliped in lateral view (in situ); G, propodus and dactylus of minor cheliped in mesial view; H, second pereopod (setae omitted); I, third pereopod (setae omitted); J, fourth pereopod (setae omitted); K, fifth pereopod (setae omitted); L, first left pleopod in lateral view; M, second left pleopod in anterior view; N, same, detail of endopod. Scale is 1 mm.

FIGURE 2. Neocallichirus jousseaumei (Nobili, 1904), paralectotypes, MNHN Th 83, A – L, female (tl 61, cl 15); M – S, female (tl 61, cl 15.7). A, front in dorsal view; B, front in lateral view; C, third maxilliped in lateral view (setae omitted); D, major cheliped in mesial view; E, same, lower border of carpus and propodus in dorsomesial view; F, same, carpus in lateral view; G, dactylus and fixed finger of major cheliped in lateral view; H, minor cheliped in lateral view; I, distal articles of left third pereopod (setae omitted); J, first right pleopod in lateral view (setae omitted); K, second right pleopod in posterior view (setae omitted); L, telson and right uropod in dorsal view; eyestalk in dorsal (M) and lateral (N) view; left antennula (O) and antenna (P) in lateral view; Q, third maxilliped in lateral view (setae omitted), R, major cheliped in mesial view (setae omitted); S, same, lower border of carpus and propodus in dorsomesial view. Scale is 1 mm.

FIGURE 3. Neocallichirus jousseaumei (Nobili, 1904), NMCR 39021, male (tl 76 cl 21.4). Carapace in dorsal (A) and lateral (B) aspect; C, detail of front in lateral view; D, eyestalk in dorsal view; E, right mandible; F, right first maxilla; G, right second maxilla; H, right first maxilliped; I, right second maxilliped; right (J) and left (K) third maxilliped; L, major cheliped in mesial view; M, same, propodus and dactylus in lateral view; N, minor cheliped in mesial view; O, distal segments of third pereopod; P, telson and left uropod; right first (Q) and second (R) pleopod in posterior view. All setae omitted except for L, M, N, P. Scale is 1 mm.

FIGURE 4. Neocallichirus jousseaumei (Nobili, 1904). A – D, paralectotype, MNHN Th- 83, male (tl 19, cl 5.0); E, F, I – K, ZRC 2010.0396 (tl 34 cl 10.2); G, NHMW 25033 (tl 85 cl 22.8); H, NHMW 25039 (tl 57 cl 16.6); L – N, NHMW 25030 (tl 36 cl 10.6); O, NHMW 25027 (tl 52 cl 15). Front in dorsal (A) and lateral (B) view; C, right third maxilliped; D, telson and left uropod; major cheliped in mesial (E) and lateral (F) view; F, H, right first pleopod, lateral view; I, L, left first pleopod, lateral view; J, M, left second pleopod, anterior view; K, N, same, detail of appendix interna; O, left second pleopod, distal part of endopod, posterior view. Scale is 1 mm.

FIGURE 6. Neocallichirus jousseaumei (Nobili, 1904). Habitats: Alona Beach on Panglao, the Philippines at low tide (A); in Thailand, Leam Pakarang (B) and Sunset Beach (C); D, detail of sediment surface with burrow openings (diameter of sieve 20 cm); E, detail of one burrow opening with faecal pellets; F, male specimen NHMW 21956, (tl 42 cl 12.2) from Sunset Beach; female specimen NHMW 21942 (tl 74 cl 21.3) in dorsal (G) and lateral (H) view from Alona Beach (Photos: Tin-Yam Chan and Chia-Wei Lin). I, Neocallichirus natalensis (Barnard, 1947), female specimen NHMW 24900; major cheliped detached. Note associated red copepods on body surface in G – I. Scale is 1 cm.