Alpheus berumeni, Anker, 2024

Anker, Arthur, 2024, Preliminary revision of Alpheus djeddensis Coutière, 1897 species complex, with description of three new species of goby-associated snapping shrimps and taxonomic notes on A. macellarius Chace, 1988 and A. djiboutensis De Man, 1909 (Decapoda: Alpheidae), Zootaxa 5472 (1), pp. 1-63 : 42-50

Alpheus berumeni sp. nov.

Figs. 26A–E , 27A–F , 28 , 29

(+ Alpheus cf. berumeni : Figs. 26F–K , 27G–I , 30 , 31 )

Alpheus djiboutensis . —Karplus et al. 1974: 260 (part., A. djiboutensis 2) [not A. djiboutensis De Man, 1909 ].

Alpheus purpurilenticularis .— Karplus et al. 1979: 337; Karplus 1979: 174; Karplus 1981: 25; Karplus et al. 1981: 6, fig. 2A; Karplus 1987: 514, fig. 2A; Karplus & Thompson 2011: 564, fig. 4.4.5A [nomen nudum, see Anker 2022a].

(?) Alpheus djeddensis. — Baensch & Debelius 1992: 494, colour photograph; Gosliner et al. 1996: 209, colour photograph 753; Debelius 1998: 273, colour photograph; Minemizu 2000: 84, colour photograph; Debelius 2001: 150, colour photograph; Thompson 2004: 106, fig. 1; Jaafar & Randall 2009: 28, pl. 1, fig. H; Thompson et al. 2013: E63, fig. 2 (colour photograph insert); Burns et al. 2019: 275 [not A. djeddensis Coutière, 1897 , = A. cf. berumeni , see text].

(?) Alpheus sp. B .— Senou et al. 2004: 300, colour photograph [= A. cf. berumeni , see text].

(?) Alpheus sp. 1 .— Minemizu 2013: 101, colour photograph [= A. cf. berumeni , see text].

(?) Alpheus sp. 14.— Minemizu 2000: 86, colour photograph [= A. cf. berumeni , see text].

(?) Alpheid shrimp. — Randall et al. 1998: 531, upper photograph [= A. cf. berumeni , see text].

Type material. Holotype: male (cl 14.3 mm), FLMNH UF 36584 , Red Sea , Saudi Arabia, off Farasan Banks, Dolphen Lagoon, 19°00’19.1”N / 40°08’53.5”E, reef lagoon, barrier reef flat, depth 1–25 m, leg. M.L. Berumen (A. Anker et al.), 10.03.2013 (BDJRS-1688) . Paratypes: 1 ovig. female (cl 9.5 mm), FLMNH UF 71919 , Red Sea , Saudi Arabia, west of Yanbu, offshore reef, 24°00’13.3”N / 37°58’35.2”E, reef flat, depth 15.8 m, in burrow with Ctenogobiops crocineus , leg. V.N. Peinemann & L.P. Aroyo, 09.10.2022 (AA-22-318 / Art-35); 1 ovig. female (cl 10.4 mm), FLMNH UF 71921 , same collection data as for previous specimen, in burrow with Amblyeleotris steinitzi (AA-22-320 / Art-33); 1 ovig. female (cl 8.7 mm), RSRC , Red Sea , Saudi Arabia, north-west of Mastorah, offshore reef, 23°14’33.1”N / 38°37’33.4”E, outer reef near channel, depth 13 m, in burrow with A. steinitzi , leg. V.N. Peinemann & L.P. Aroyo, 05.10.2022 (AA-22-277 / Art-11); 1 male (cl 7.0 mm), FLMNH UF 71920 , Red Sea , Saudi Arabia, off King Fahd Industrial Port, offshore reef, 23°50’41.0”N / 38°07’54.8”E, sandflat near coral reef, depth 15.1 m, in burrow with A. steinitzi , leg. V.N. Peinemann & L.P. Aroyo, 08.10.2022 (AA-22-349 / Art-30); 1 ovig. female (cl 15.2 mm), FLMNH UF 38116 , Red Sea , Saudi Arabia, Gulf of Aqaba, Magna coast guard station, depth 0–60 m, leg. M.L. Berumen, 29- 30.09.2013 ( NORS 14 /15/16.2) .

Other material examined. Alpheus cf. berumeni . 1 male (cl 10.4 mm), FLMNH UF 16538 , French Polynesia, Society Islands, Moorea, between Papetoai and Sheraton Hotel , 17°29’22.6”S / 149°53’03.1”W, sandflat with large mounds, corals, rubble, depth 1 m, suction pump in burrow, leg. A. Anker, 23.11.2008 (BMOO-5682); 1 ovig. female (cl 14.1 mm), FLMNH UF 16539 , same collection data as for previous specimen (BMOO-5683) ; 1 female (cl 8.3 mm), FLMNH UF 16518 , French Polynesia, Society Islands, Moorea, Motu Tiahura , Fareone channel, 17°29’20.0”S / 149°54’52.6”W, sandflat with coral rubble and living corals, depth 1–2 m, suction pump, in burrow in sand, leg. A. Anker, 21.11.2008 (BMOO-5625) ; 4 males (cl 8.4–11.9 mm, all missing major and/or major cheliped), 2 females (cl 11.0, 12.1 mm, missing major or both chelipeds), MNHN-IU-2018-5649, French Polynesia, Society Islands, Moorea , sand patches between corals, depth 5–15 m, leg.A. R. Thompson, 06–08.2000 ; 1 male (cl 12.3 mm, missing minor cheliped), MNHN-IU-2022-129, same collection data as for previous specimens ; 1 female (cl 10.2 mm), MNHN-IU-2022-127, same collection data as for previous specimens ; 1 male (cl 10.1 mm), MNHN-IU-2022- 128, same collection data as for previous specimens ; 1 male (cl 11.6 mm), MNHN-IU-2022-132, same collection data as for previous specimens ; 1 male (cl 6.5 mm), 1 female (cl 6.4 mm), MNHN-IU-2014-6017, Papua New Guinea, Madang, PAPUA NIUGINI sta. PR76, Rempi, SW Hargun Island , 5°01’58.8”S / 145°48’08.6”E, depth 2–15 m, sandflat near coral reef, leg. A. Anker et al. 21.11.2012 (PR76-PZD-300L) ; 1 female (cl 8.9 mm), MNHN- IU-2014-6032, same collection data as for previous specimens (PR76-PZD-300H); 1 ovig. female (cl 8.2 mm), MNHN-IU-2014-6018, Papua New Guinea, Madang, PAPUA NIUGINI sta. PR106, Bilbil Island , 5°17’53.2”S / 145°46’42.2”E, depth 14 m, sandflat with coral heads, leg. A. Anker et al. 21.11.2012 (PR106-PZD-402) ; 1 female (cl 8.9 mm), FLMNH UF 68851 , Solomon Islands, New Georgia, Munda, Sosuhite Island , shallow sandflat with rubble around small island, in burrow under coral rubble, depth 0.5–1 m, leg. A. Anker et al., 10.09.2016 (SOL-068) ; 1 female (cl 7.6 mm), FLMNH UF 37329 , USA, Northern Mariana Islands, Guam, W coast near Fish Eye Marine Park , 13°28’33.5”N / 144°42’19.4”E, under large rocks in sand, depth 0.5–1 m, leg. A. Anker, 14.06.2010 (GUOK-0279) .

Diagnosis. Morphology largely as in A. djeddensis . Carapace strongly pitted, except for mid-dorsal portion, glabrous, sometimes with very few short setae. Rostrum well developed, reaching mid-length of first article of antennular peduncle, sometimes almost reaching its distal margin; rostral carina distinct, blunt, raising above orbital hoods, extending well beyond base of orbital hoods, not reaching mid-length of carapace; orbital hoods unarmed, rounded; adrostral furrows very shallow ( Fig. 26A, B ). Rostro-orbital region moderately protruding beyond anterolateral margin of carapace ( Fig. 26A, B ). Telson very broad, at most 1.5 times as long as wide at base, gently tapering; dorsal surface strongly pitted, with two pairs of stout spiniform setae inserted at considerable distance from lateral margins; posterior margin broadly rounded, with row of slender spiniform setae between two pairs of more robust spiniform setae at each distolateral angle, mesial much longer than lateral ( Fig. 26C ). Antennule with stylocerite slightly swollen laterally, its tip falling well short of distal margin of first article of peduncle, ventromesial carina with subtriangular tooth; second article at most 2.5 times as long as wide ( Fig. 26A–C ).Antenna with distoventral margin of basicerite armed with sharp tooth; scaphocerite with lateral margin noticeably concave; distolateral tooth by far overreaching distal margin of blade ( Fig. 26A, B ). Third maxilliped with penultimate article cup-shaped, widening distally, strongly convex ventrally, with few moderately elongate setae on distoventral margin ( Fig. 26E ). Major cheliped of brevirostris type; merus stout, without tooth on distomesial margin, ventromesial margin with some spiniform setae; chela not elongate; palm about twice as long as wide, with deep transverse groove on dorsal margin, mesial surface faintly granulated; fingers almost 0.7 length of palm; plunger of dactylus feebly bulging ( Fig. 27A–C ; see also Fig. 27G, A . cf. berumeni). Minor cheliped distinctly shorter and weaker than major cheliped; merus without tooth on distomesial margin, ventromesial margin with some spiniform setae; chela not elongate; palm about twice as long as wide, without grooves, mesial surface distinctly granulated; fingers about 1.2 times as long as palm, somewhat gaping, sexually dimorphic (but see below), with rows of balaeniceps setae on each side of dactylus and pollex in males, without or with poorly developed balaeniceps setae in females ( Figs. 27D–F , 29A ; see also Figs. 26H, I , 30 , 31 , A. cf. berumeni ). Second pereiopod with ratio of carpal subarticles equal to 3.0 / 2.5 / 1.0 / 1.0 / 1.6 ( Fig. 28B ; see also Fig. 26F, A . cf. berumeni). Third pereiopod moderately robust; ischium armed with stout spiniform seta; merus about 4.2 times as long as maximal width, unarmed; propodus with six or so spiniform setae along ventral margin and one pair of spiniform setae on distoventral angle; dactylus about 0.45 length of propodus, gradually curving distally, spatulate ( Fig. 28B ; see also Fig. 26G–I, A . cf. berumeni). Fifth pereiopod ischium armed with spiniform seta ( Fig. 26J, K, A . cf. berumeni). Appendix masculina with stiff setae on apical part and along margin opposed to endopodal ramus ( Fig. 26L ). Uropodal exopod with diaeresis almost straight, except for broad lateral lobe; endopod with row of spiniform setae on distal margin ( Fig. 26M ). Colour pattern diagnostic, as described below.

Colour pattern. Background colour whitish with yellowish tinge; carapace with pale brown-yellow or greenyellow reticulation in form of short irregular spots and blotches, interconnected or not, many forming chains and narrow bands; carapace flanks with large, conspicuous, black or dark brown patch surrounded by colourless area, stretching from about mid-length of carapace to near its posterior margin; rostrum and post-rostral area whitish with short orange-yellow bands; pleon with similar reticulation as on carapace, forming two more or less distinct transverse bands on each pleonite; saddle on first pleonite usually conspicuous, white; saddle on fifth pleonite much smaller, white; telson white with pale brown-yellow mottling; antennular and antennal peduncles white with yellowish tinge and golden brown mottling; antennular and antennal flagella yellow; chelipeds largely white with scarce brown-yellow blotches on merus, carpus and mesial face of chela; calcified distal portion of chela fingers ivory; lateral face of chelae with faint grey-yellow mottling; second to fifth pereiopods pale blue with yellowish articulations; uropods white with pale brown-yellow mottling; eggs in ovigerous females yellow ( Figs. 28 , 29 ). See also Figs. 30 , 31 for colour pattern of A. cf. berumeni .

Etymology. The new species is named after Michael L. Berumen, director of the Red Sea Research Center of KAUST, for his support and for collecting several important specimens used in the present study.

Common names. Berumen’s snapping shrimp; burnt-flanks goby shrimp.

Type locality. Farasan Banks , Saudi Arabia .

Distribution. Western Indian Ocean: presently known with certainty only from the Red Sea ( Saudi Arabia, Israel, Egypt); records from other Indo-West Pacific localities, including Seychelles ( Polunin & Lubbock 1977), Indonesia, Philippines, Japan, Australia (Great Barrier Reef), New Caledonia (photographic records in Randall et al. 1998; Minemizu 2000; Debelius 2001; Senou et al. 2004; Randall 2005; Kuiter & Debelius 2009), as well as Papua New Guinea, Solomon Islands (New Georgia), Mariana Islands ( Guam), French Polynesia (Moorea) (material identified as A. cf. berumeni ), require confirmation (see below).

Ecology. Coral reefs and sandflats near reef slopes, deeper lagoons with sand-rubble bottom, usually between 1 and 20 m, possibly deeper; in the Red Sea typically associated with Amblyeleotris steinitzi (Klausewitz) ( Fig. 29D, E ; see also Karplus 1979; Karplus et al. 1979; Jaafar & Randall 2009); occasionally associated with other gobies, e.g., Ctenogobiops maculosus (Fourmanoir) , C. crocineus (Smith) or Cryptocentrus lutheri ( Karplus & Thompson 2011; present study); in the central Indian Ocean and western Pacific ( A. cf. berumeni ) appears to be commonly associated with Ctenogobiops feroculus (Lubbock & Polunin) , as well as other species of Ctenogobiops , and Amblyeleotris ( Figs. 30F , 31G, H ; Thompson 2004; A. Anker., pers. obs.).

Remarks. The holotype from the Farasan Banks, Saudi Arabia (FLMNH UF 36584), is by far the largest male (cl 14.3 mm) within the Red Sea type material. The specimen was largely intact when it was set apart for preparation of illustrations (line drawings), with both chelipeds, as well as the left second, left third and right fifth pereiopods attached to the body ( Fig. 28 ). However, when the holotype returned from a loan in 2022, it was missing the second, third and fifth pereiopods, which were either used for molecular analyses or lost during manipulation of the specimen. Therefore, illustrations of the second, third and fifth pereiopods were prepared using a large nontype specimen of A. cf. berumeni from Moorea, French Polynesia (see below), after a comparison of the relative proportions of these appendages, which appear to be very similar (cf. Figs. 26F–K , 27B ). Most of the paratype specimens of A. berumeni from the Red Sea were collected or located after the final preparations of illustrations.

De Man (1909), while examining type material of A. djeddensis and A. djiboutensis , stated that in both species, the carapace is pubescent and illustrated this feature very clearly in his figures ( Fig. 1B, D ). In contrast, A. berumeni sp. nov. has a completely glabrous carapace, without any pubescence, although occasionally scarce setae are present near the anterolateral margin ( Fig. 26A, B ). The lack of any pubescence on the carapace allows to distinguish A. berumeni sp. nov. from A. djeddensis , A. shukran sp. nov., A. macellarius , A. tigrinus sp. nov. and A. djiboutensis , in which the carapace is pubescent. Furthermore, A. berumeni sp. nov. can be separated from A. djeddensis and A. shukran sp. nov. by the proportions of the major and minor chelipeds, with stouter meri and chelae (cf. Figs. 3F, G , 4 , 27 ); from A. macellarius and A. djiboutensis by the distomesial margin of the cheliped meri ending bluntly vs. armed with a sharp tooth on the major cheliped or on both chelipeds in the other two species, respectively (cf. Fig. 27A, D , 32G ; Chace 1988: fig. 6e, 7 h); from A. tigrinus sp. nov. by the absence of long stiff setae on the anterolateral surface of the carapace, which are present in all specimens of A. tigrinus sp. nov., and a slightly stouter telson, about 1.7 times as long as maximal width vs. twice as long as wide in A. tigrinus sp. nov. (cf. Figs. 18A–C , 26A–C ); from A. djiboutensis by the much broader telson, about 1.7 times as long as maximal with vs. 2.2 times as long as maximal width in A. djiboutensis (cf. Figs. 1F , 26C , 32C ); and from A. djeddensis , A. shukran sp. nov., A. macellarius and A. tigrinus sp. nov. by the narrower blade of the scaphocerite (cf. Figs. 3A , 4A , 18A , 26A ; Chace 1988: fig. 6a). The poor development or absence of the balaeniceps condition in females of A. berumeni sp. nov. requires confirmation since most of the females in the type series are relatively young (except for FLMNH UF 38116), but could represent an additional character distinguishing the new species from A. djeddensis , A. shukran sp. nov. and A. tigrinus sp. nov.

By far the best diagnostic feature of A. berumeni sp. nov. is its unmistakable colour pattern, consisting of a pale brown yellowish reticulation and a very large black or dark brown patch on the carapace flanks ( Figs. 28 , 29 ). This colour pattern radically differs from those of A. djeddensis , A. shukran sp. nov., A. macellarius (including A. cf. macellarius ) A. tigrinus sp. nov. (including A. cf. tigrinus and A. aff. tigrinus ), as well as specimens tentatively identified as A. djiboutensis (cf. Figs. 6–17 , 21–25 , 33–35 ).

Numerous in situ colour photographs ( Figs. 29D, E , 30F , 31G, H ; see also Randall et al. 1998; Minemizu 2000, 2013; Debelius 2001; Randall 2005; Kuiter & Debelius 2009) suggest that A. berumeni sp. nov. may have a wide distribution in the Indo-West Pacific, ranging from the Red Sea to the Philippines, Australia and French Polynesia. However, all material from outside of the Red Sea, including the material herein cautiously referred to as A. cf. berumeni ( Figs. 30 , 31 ), needs further morphometric studies and molecular comparisons. Most of the western Pacific specimens available for the present study were collected in French Polynesia ( Fig. 30 ), with a few additional, mostly relatively small specimens collected in the Mariana Islands ( Guam), Papua New Guinea and Solomon Islands ( Fig. 31A–F ). The colour pattern of the Red Sea specimens of A. berumeni sp. nov. ( Fig. 28 , 29 ) and the specimens identified as A. cf. berumeni from other Indo-West Pacific localities ( Figs. 30 , 31 ) is essentially the same. Furthermore, the material from French Polynesia, Papua New Guinea and Solomon Islands presents no obvious morphological differences with the material from the Red Sea, except perhaps for the presence of a small post-rostral tubercle in some smaller specimens (e.g., MNHN-2014-6018). Therefore, and because similar phylogeographic patterns are observed in several other goby shrimps ( Anker 2022 a, 2022b; see also discussion under A. tigrinus sp. nov.), it seems reasonable, at least for the time being, to treat all Indo-West Pacific material with the colour pattern of A. berumeni sp. nov. as A. cf. berumeni , until more material and comparative DNA data become available. The presence of a genetically distinct but morphologically cryptic sister species of A. berumeni sp. nov. outside of the Red Sea (or the western Indian Ocean), for instance, in the “Coral Triangle” and adjacent western Pacific (from Japan and Mariana Islands to Australia, New Caledonia and French Polynesia) is not unlikely.

Figures

FIGURE 1. Historical illustrations of Alpheus djeddensis Coutière, 1897 [A–C] and Alpheus djiboutensis De Man, 1909 [D–K]: A, right (minor) chela, mesial; B, carapace and frontal appendages, dorsal; C, antennal scaphocerite, dorsal; D, carapace and frontal appendages, dorsal; E, antennal scaphocerite, distal half, dorsal; F, telson, dorsal; G, left (major) cheliped, lateral; H, right (minor) cheliped, lateral; I, same, chela, dorsal; J, second pereiopod, lateral; K, third pereiopod, lateral. After Coutière (1899) [A, B] and De Man (1909) [C–K].

FIGURE 3. Alpheus djeddensis Coutière, 1897, paralectotype, male (cl 18.9 mm) from Jeddah, Saudi Arabia (MNHN-IU-2016- 9011); A, frontal region, dorsal; B, same, lateral; C, telson, dorsal; D, antennal scaphocerite, dorsal; E, third maxilliped, lateral; F, left (major) cheliped, mesial; G, same, lateral; H, same, dactylus, lateral; I; right (minor) cheliped, ischium to proximal portion of propodus (chela palm), lateral; J, same, distal portion of merus carpus and chela, lateral; K, same, chela, mesial; L, second pereiopod, lateral; M, third pereiopod, lateral; N, same, propodus and dactylus, lateral; O, same, dactylus, ventromesial; P, fourth pereiopod, lateral; Q, same, propodus and dactylus, lateral.

FIGURE 4. Alpheus djeddensis Coutière, 1897, male (cl 19.3 mm) from Farasan Islands, Saudi Arabia (FLMNH UF 36913); A, frontal region, dorsal; B, same, lateral; C, telson, dorsal; D, tooth on ventromesial carina of first article of antennular peduncle, lateral; E, third maxilliped, lateral; F, same, distal portion of antepenultimate article, and penultimate and ultimate articles, ventrolateral (setae omitted); G, second pereiopod, distal portion of merus, carpus and chela, lateral; H, third pereiopod, lateral; I, same, propodus and dactylus, lateral; J, same, dactylus, ventrolateral; K, fifth pereiopod, lateral; L, same, propodus and dactylus, lateral.

FIGURE 6. Alpheus djeddensis Coutière, 1897, male (cl 19.3 mm) from Farasan Islands, Saudi Arabia (FLMNH UF 36913); A, shrimp post-mortem (carapace damaged during collection), dorsal; B, same, lateral. Photographs by the author.

FIGURE 7. Alpheus djeddensis Coutière, 1897, male (cl indet.) from Thuwal, Saudi Arabia (specimen kept alive in RSRC); A, shrimp alive, dorsal; B, same, lateral. Photographs by the author.

FIGURE 8. Alpheus djeddensis Coutière, 1897, male (cl indet.) from Thuwal, Saudi Arabia (specimen kept alive in RSRC) [A–F]; major cheliped and third pereiopod (RSRC, used for DNA sequencing) [G]; A, frontal region, dorsal; B, tail fan, dorsal; C, left (major) chela, mesial, D, same, lateral; E, right (minor) cheliped and second pereiopod, lateral; F, right (minor) chela, lateral; G, major cheliped, mesial, and third pereiopod, lateral. Photographs by the author.

FIGURE 9. Alpheus djeddensis Coutière, 1897, female (cl indet.) from Thuwal, Saudi Arabia (specimen kept alive in RSRC); A, shrimp alive, dorsal; B, frontal region, dorsal; C, right (major) chela, mesial; D, left (minor) chela, distal portion of palm and fingers, mesial. Photographs by the author.

FIGURE 10. Alpheus djeddensis Coutière, 1897, in situ photographs; A, shrimp (ovigerous female) near burrow entrance, together with Cryptocentrus caeruleopunctatus (Rüppell) (goby not in frame and out focus), near Thuwal, Saudi Arabia; B, shrimp (sex unknown) outside of its burrow, together with C. caeruleopunctatus, southern Egypt. Photographs courtesy of Viktor Nunes Peinemann [A] and Ülar Tikk [B].

FIGURE 11. Alpheus shukran sp. nov., holotype, ovigerous female (cl 18.5 mm) from Masirah Island, Oman (FLMNH UF 71923); A, shrimp alive, dorsal; B, same, lateral. Photographs by the author.

FIGURE 12. Alpheus shukran sp. nov., holotype, ovigerous female (cl 18.5 mm) from Masirah Island, Oman (FLMNH UF 71923); A, frontal region, dorsal; B, tail fan, dorsal; C, left (major) chela, lateral; D, same, mesial; E, right (minor) chela, lateral; F, same, mesial; G, second pereiopod, lateral. Photographs by the author.

FIGURE 13. Alpheus shukran sp. nov., male (cl 11.0 mm) from Bandar Khayran, Oman (FLMNH UF 68873) [A–C]; ovigerous female (cl 13.2 mm) from the same locality (FLMNH UF 68870) [D–G]; A, shrimp alive, dorsal; B, same, lateral; C, frontal region, dorsal; D, shrimp alive, dorsal; E, same, lateral; F, carapace, frontal appendages and chelipeds, dorsal; G, frontal region, dorsal. Photographs by the author.

FIGURE 14. Alpheus shukran sp. nov., female (cl 12.7 mm) from Thuwal, Saudi Arabia (RSRC) [A–E]; aquarium-kept specimens (cl indet.) from Eilat, Israel [F, G]; male (cl 11.0 mm) from Bandar Khayran, Oman (FLMNH UF 68873) [H]; A, shrimp alive, dorsal; B, same, lateral; C, frontal region, dorsal; D, left (major) chela, mesial; E, right (minor) chela, mesial; F, shrimp alive, dorsal; G, shrimp in aquarium together with Cryptocentrus lutheri Klausewitz; H, male second pleopod, appendices masculina and interna, anterior. Photographs by the author [A–E] and Ilan Karplus [F, G, latter also published in Karplus et al. (1972a)].

FIGURE 15. Alpheus macellarius Chace, 1988, 1 male (cl 14.5 mm) from Raffles Lighthouse, Singapore (OUMNH.ZC.2014- 11-081); A, shrimp alive, dorsal; B, same lateral; C, frontal region, dorsal; D, tail fan, dorsal. Photographs by the author.

FIGURE 16. Alpheus macellarius Chace, 1988, female (cl 5.5 mm) from Nosy-Bé, Madagascar (FLMNH UF 14035) [A, B]; A, shrimp alive, dorsal; B, same, lateral. Alpheus cf. macellarius Chace, 1988, specimens from different western Pacific localities [C–H]; C, specimen (sex and cl indet.) from Koumac, New Caledonia (MNHN-IU-2018-778), shrimp alive, dorsal; B, same, lateral; E, female (cl 8.9 mm) from Madang, Papua New Guinea (MNHN-IU-2014-6031), shrimp alive, dorsal; F, same, lateral; G, ovigerous female (cl 10.2 mm) from Guam, Northern Mariana Islands (FLMNH UF 37357), shrimp alive, dorsal; H, same, lateral. Photographs by the author.

FIGURE 17. Alpheus macellarius Chace, 1988, in situ photographs; A, shrimp near burrow entrance, together with Cryptocentrus sp., Bali, Indonesia; B, shrimp and goby, Cryptocentrus caeruleomaculatus (Herre), outside of burrow, Bali, Indonesia; C, shrimp near its burrow among algae, partly exposed tide flat, Pulau Sekudu, Singapore; D, shrimp near burrow entrance, together with Cryptocentrus maudae Fowler, Sentosa, Singapore; E, F, shrimp (with bopyrid parasite) outside of burrow, together with Cryptocentrus cf. strigilliceps (Jordan & Seale), Watamu, Kenya. All identifications of shrimps and gobies tentative. Photographs courtesy of Andrey Ryanskiy (A, B), Marcus Ng (C, D), and Dawn Goebbels (E, F).

FIGURE 18. Alpheus tigrinus sp. nov., holotype, male (cl 15.3 mm) from Farasan Islands (Zahrat Durakah), Saudi Arabia (FLMNH UF 37012); A, frontal region, dorsal; B, same, lateral; C, telson, dorsal; D, tooth on ventromesial carina of first article of antennular peduncle, lateral; E, third maxilliped, lateral; F, second pereiopod, lateral; G, third pereiopod, lateral; H, same, propodus and dactylus, lateral; I, same, dactylus, ventromesial; J, fifth pereiopod, lateral; K, same, propodus and dactylus, mesial; L, male second pleopod, appendices masculina and interna, anterior; M, uropod, dorsal.

FIGURE 21. Alpheus tigrinus sp. nov., paratype, ovigerous female (cl 16.7 mm) from Farasan Islands (Tiger Head Island), Saudi Arabia (FLMNH UF 36648); A, shrimp alive, dorsal; B, same, lateral. Photographs by the author.

FIGURE 22. Alpheus tigrinus sp. nov., paratype, male (cl 13.0 mm) from Farasan Islands (Tiger Head Island), Saudi Arabia (FLMNH UF 36957) [A]; holotype, male (cl 15.3 mm) from Farasan Islands (Zahrat Durakah), Saudi Arabia (FLMNH UF 37012) [B]; A, shrimp alive, dorsal; B, shrimp post-mortem (carapace damaged during collection), dorsal. Photographs by the author.

FIGURE 23. Alpheus tigrinus sp. nov., paratype, ovigerous female (cl 16.7 mm) from Farasan Islands (Tiger Head Island), Saudi Arabia (FLMNH UF 36648) [A, B]; holotype, male (cl 15.3 mm) from Farasan Islands (Zahrat Durakah), Saudi Arabia (FLMNH UF 37012) [C]; paratype, male (cl 13.0 mm) from Farasan Islands (Tiger Head Island), Saudi Arabia (FLMNH UF 36957) [D]; male (cl 15.4 mm) from Farasan Islands (Zahrat Durakah), Saudi Arabia (FLMNH UF 37015); A, frontal region, dorsal; B, tail fan, dorsal; C, frontal region and chelipeds, dorsolateral; D, frontal region, dorsal; E, left (major) cheliped, mesial. Photographs by the author.

FIGURE 24. Alpheus tigrinus sp. nov., ovigerous female (cl 14.3 mm) from Thuwal, Saudi Arabia (RSRC) [A–D]; young female (cl indet., released) from Rabigh, Saudi Arabia [E]; A, shrimp alive, dorsal; B, same, lateral; C, frontal region, dorsal; D, tail fan, dorsal; E, shrimp alive, dorsal. Alpheus aff. tigrinus, female (cl 6.5 mm) from St. John’s Island, Singapore (OUMNH. ZC.2014-11-082) [F, G]; F, shrimp alive, dorsal; G. same, lateral. Photographs by the author.

FIGURE 25. Alpheus cf. tigrinus [A–C] and Alpheus aff. tigrinus [D–H], mostly in situ photographs (except A); A, male (cl 14.3 mm) from Abu Dhabi, UAE (FLMNH UF 71625); B, shrimp near burrow entrance, together with Cryptocentrus lutheri Klausewitz, Kubbar Island, Kuwait; C, shrimp at burrow entrance, together with C. lutheri, Qatar; D, E, G, different shrimps at burrow entrances, together with Cryptocentrus nigrocellatus (Yanagisawa), Philippines (C), Japan (D, G); F, shrimp outside of its burrow (goby not in frame), Japan; H, shrimp at burrow entrance (burrow made under massive coral), with Cryptocentrus albidorsus (Yanagisawa), Kenting, Taiwan. Identifications of gobies tentative. Photographs courtesy of FLMNH team [A], Najim Al-Misbah [B], Redha Al-Kallabi [C], Ülar Tikk [D], Tomonori Hirata [E], Shunzo [F], Tomoki Iida [G], Ming-Shiou Jeng [H].

FIGURE 26. Alpheus berumeni sp. nov., holotype, male (cl 14.3 mm) from Farasan Banks (Dolphen Lagoon), Saudi Arabia (FLMNH UF 36584) [A–E, L, M]; A, frontal region, dorsal; B, same, lateral; C, telson, dorsal; D, tooth on ventromesial carina of first article of antennular peduncle, lateral; E, third maxilliped, lateral; L, male second pleopod, appendices masculina and interna, anterior; M, uropod, dorsal. Alpheus cf. berumeni, male (cl 12.3 mm) from Moorea, Society Islands, French Polynesia (MNHN-IU-2022-129) [F–K]; F, second pereiopod, lateral; G, third pereiopod, lateral; H, same, propodus and dactylus, lateral; I, same, dactylus, ventral; J, fifth pereiopod, lateral; K, same, propodus and dactylus, mesial.

FIGURE 27. Alpheus berumeni sp. nov., holotype, male (cl 14.3 mm) from Farasan Banks (Dolphen Lagoon), Saudi Arabia (FLMNH UF 36584) [A–F]; A, right (major) cheliped, mesial; B, same, carpus and chela, lateral; C, same, dactylus, lateral; D, left (minor) cheliped, mesial; E, same, ischium to proximal portion of propodus (chela palm), lateral; F, same, carpus and chela, lateral. Alpheus cf. berumeni, two males (cl 12.3 mm, 11.6 mm) from Moorea, Society Islands, French Polynesia (MNHN-IU- 2022-129, MNHN-IU-2022-132, respectively) [G–I]; G, left (major) cheliped of larger male (cl 12.3 mm), carpus and chela, lateral; H, left (minor) chela of smaller male (cl 11.6 mm), mesial; I, same, lateral.

FIGURE 28. Alpheus berumeni sp. nov., holotype, male (cl 14.3 mm) from Farasan Banks (Dolphen Lagoon), Saudi Arabia (FLMNH UF 36584); A, shrimp alive (cephalothorax damaged during collection), dorsal; B, same, lateral. Photographs by the author.

FIGURE 29. Alpheus berumeni sp. nov., non-deposited specimens and in situ photographs; A, ovigerous female (cl indet.) from Yanbu, Saudi Arabia (released), dorsal; B, same, lateral; C, aquarium-kept specimen, Eilat, Israel; D, shrimp near burrow entrance, together with Amblyeleotris steinitzi (Klausewitz), Sharm El-Sheikh, Egypt; E, shrimp outside of burrow entrance, together with A. steinitzi, Al Lith, Saudi Arabia. Photographs courtesy of Viktor Nunes Peinemann [A, B], Ilan Karplus [C], Andrey Ryanskiy [D], Lucia Aroyo Pombo [E].

FIGURE 30. Alpheus cf. berumeni from Moorea, Society Islands, French Polynesia, material examined [A–E] and photograph in situ [F]; A, male (cl 10.4 mm) from near Papetoai (FLMNH UF 16538), shrimp alive, dorsal; B, same, lateral; C, same, ventral; D, ovigerous female (cl 14.1 mm) from the same locality (FLMNH UF 16539), shrimp alive, lateral; E, female (cl 8.3 mm) from Motu Tiahura (FLMNH UF 16518), shrimp alive, dorsal; F, shrimp outside of burrow, together with Ctenogobiops feroculus (Lubbock & Polunin). Photographs by the author [A–E] and courtesy of François Libert [F].

FIGURE 31. Alpheus cf. berumeni, specimens from different western Pacific localities; A, female (cl 8.9 mm) from New Georgia, Solomon Islands (FLMNH UF 68851), shrimp alive, dorsal; B, same, lateral; C, female (cl 7.6 mm) from Guam, Northern Mariana Islands (FLMNH UF 37329), shrimp alive, dorsal; D, same, lateral; E, ovigerous female from Madang, Papua New Guinea (MNHN-IU-2014-6018), shrimp alive, dorsal; F, same, lateral; G, shrimp outside burrow entrance, together with Amblyeleotris sp., Papua New Guinea; H, shrimp at burrow entrance, together with Ctenogobiops sp., Indonesia. Photographs by the author [A–F] and courtesy of Andrey Ryanskiy [G] and Paul Humann [H].

FIGURE 32. Alpheus cf. djiboutensis De Man, 1909, male (cl 16.7 mm) [A–I] and ovigerous female (cl 16.9 mm) [J] from Farasan Islands (Abulad), Saudi Arabia, both specimens dried out (see text) (FLMNH UF 36942); A, frontal region, dorsal; B, same, lateral; C, telson, dorsal; D, left (minor) cheliped, mesial; E, same, lateral; F, right (major) cheliped, in situ, heavily damaged due to dry condition, lateral; G, same, detached, mesial; H, second pereiopod, detached, lateral; I, second, third and fourth pereiopods, in situ, lateral; J, left (minor) cheliped, mesial. Photographs by the author.

FIGURE 33. Alpheus cf. djiboutensis De Man, 1909, male (cl 16.7 mm) from Farasan Islands (Abulad), Saudi Arabia (FLMNH UF 36942); A, shrimp post-mortem (carapace damaged during collection), dorsal; B, same, lateral; C, same, anterior region of carapace, frontal appendages and chelipeds, dorsal. Photographs by the author.

FIGURE 34. Alpheus cf. djiboutensis De Man, 1909; ovigerous female (cl 16.9 mm) from Farasan Islands (Abulad), Saudi Arabia (FLMNH UF 36942) [A, B]; A, shrimp alive, dorsal; B, same, frontal region; two shrimps photographed in situ in the Red Sea [C, D]; C, shrimp outside of burrow entrance, together with Ctenogobiops crocineus (Smith), Sharm El-Sheikh, Egypt; D, shrimp bulldozing sediments, Thuwal, Saudi Arabia. Photographs by the author [A, B] and courtesy of Andrey Ryanskiy [C] and Viktor Nunes Peinemann [D].

FIGURE 35. Alpheus cf. djiboutensis De Man, 1909, two shrimps photographed in situ near Thuwal, Saudi Arabia; A, shrimp outside of burrow entrance, together with Cryptocentrus caeruleopunctatus (Rüppell), B, shrimp bulldozing sediments, together with Cryptocentrus cryptocentrus (Valenciennes). Photographs courtesy of Viktor Nunes Peinemann.