Phoxokalliapseudes gobinae ( Bamber, 1998 ) DRUMM, DAVID T. & HEARD, RICHARD W., 2011

DRUMM, DAVID T. & HEARD, RICHARD W., 2011, Systematic revision of the family Kalliapseudidae (Crustacea: Tanaidacea), Zootaxa 3142 (1), pp. 1-172 : 38-50

Phoxokalliapseudes gobinae ( Bamber, 1998) n. comb.

(Figs 22–24)

Kalliapseudes gobinae Bamber, 1998: 181–184 , Figs. 8–9. Guţu, 2006: 127. Drumm, 2007: 18.

Material examined. 6 “form A” males (4.1–5.3 mm) (1 dissected), 6 “form B” males (4.1–4.8 mm) (1 dissected), 4 ovigerous females (5.7–5.9 mm) (1 dissected), 5 females with oostegites (5.4–7 mm), 8 juveniles, NHM 254– 263, Indo–Pacific , Sabah, North Borneo, 20 km west of Pulau Tiga Island, 35 m, coll. Roger Bamber , 2003.

Diagnosis (adult female and male). Rostrum triangular, rounded tip. Pleotelson tapering posteriorly to an indented tip with two long terminal plumose setae. Antennule first peduncle article with ventral spiniform setae; accessory flagellum of 4–6 articles; main flagellum of female with three aesthetascs, male with clusters of aes- Labrum complex with four strong cusps. Terminal spiniform seta of mandibular palp approximately 3.5 times as long as broad. Female cheliped fixed finger cutting edge with spinules interspersed with one or two round tubercles; male with two forms of cheliped. Cheliped and pereopod 1 exopodite with two plumose setae. Pereopods 2 and 3 dactylus lacking digitiform lobe. Pereopod 5 propodus with three short bipinnate setae on inner surface. Male pereopod 6 dactylus longer than ischium, merus, carpus and propodus combined, without any subterminal setae. Uropod basal article approximately 2.8 times as long as broad, with spiniform seta on inner distal corner; exopodite with three articles, last article approximately 2.3 times as long as second article.

Description. Adult male (form A). Body (Fig. 22A): length approximately 5 mm, 4.8 times as long as broad.

Carapace (Fig. 22A). Slightly broader than long, one pair mid–lateral and dorsal setae; rostrum rounded, pronounced.

Pereonites (Fig. 22A). Pereonites 4 and 5 longer than others and pereonite 6 shortest; pereon narrowing posteriorly; at least one pair anterolateral and one pair dorsal setae; hyposphaenia present on all pereonites.

Pleon (Fig. 22A). Pleonites subequal; rounded epimera, with several plumose setae, one pair dorsolateral and dorsal simple setae; hyposphaenia present on all pleonites. Pleotelson shorter than last three pleonites combined, tapering posteriorly to indented tip, with several short dorsal simple setae and two terminal plumose setae.

Antennule (Fig. 22B). First peduncle article approximately 2.6 times as long as second and third articles com- 3.5 times shorter than first article, with several simple and broom setae. Third and fourth articles subequal (including common article). Outer flagellum approximately same length as first peduncle article, with ten articles, clusters of aesthetascs on proximal articles, decreasing in number distally. Inner flagellum with 4 articles, last article with four terminal setae (three simple and one broom).

Antenna (Fig. 22C). First peduncle article with medial extension bearing five plumose setae and one simple seta on outer subdistal corner. Second peduncle article naked and the squama with five simple setae. Third peduncle article with two plumose setae on inner margin. Peduncle article 4 (last) shorter than flagellum, with double row plumose setae, one proximal and distal broom seta. Flagellum with 6 articles, one or two plumose setae on each article and articles 2–5 with several bipectinate setae.

Labrum (Fig. 22D) with numerous posterior hair–like setae and four strong anterior cusps.

Mandibles (Fig. 22E). Left mandible incisor process with approximately eight and lacinia mobilis with approximately seven teeth; spine row with five serrate spiniform setae. Right mandible not examined. Palp with short terminal spiniform seta, approximately 3.5 times as long as broad.

Other mouthparts not examined.

Cheliped (Fig. 22F). Basis with two long simple setae on outer surface and two short spiniform setae on ventral margin. Merus as long as broad, with three distal simple setae one short simple seta midway on ventral margin. Carpus approximately 2.3 times as long as broad, with double row of long, plumose setae. Propodus with diagonal row of long, plumose setae on inner face; fixed finger with three simple setae just proximal to distal unguis; cutting edge with large proximal triangular tooth and several short spinules interspersed with round tubercle between the tooth and distal unguis; palm with several distal simple setae and a large spine on inner surface near dactylus insertion. Dactylus with three simple setae on inner surface; cutting edge with several spinules increasing in length distally (> 10). Exopodite with two plumose setae

Pereopod 1 ( Fig. 23A ). Basis approximately 2.3 times as long as broad, with one spiniform and one simple seta ventrally. Ischium with two simple setae. Merus shorter than and as broad as basis, with several simple setae distally and on ventral margin, and one spiniform seta on each distal corner. Carpus approximately 2.5 times shorter than merus, with several simple setae and two ventrodistal and one dorsodistal spiniform setae. Propodus shorter than carpus, with several simple setae and six ventral and two dorsal serrate spiniform setae, one broom seta on dorsal margin. Dactylus represented by a sensory organ, shorter than propodus, with several long, terminal sensory setae with two setae on inner surface; unguis absent. Exopodite with two plumose distal setae.

Pereopod 2 ( Figs. 23B, C ). Basis approximately 3.2 times as long as broad, with two broom and two short simple setae on dorsal margin, two simple setae (one long and one short) on outer surface near ventral margin and one ventrodistal spiniform seta. Ischium with two ventrodistal simple setae (one long and one short). Merus approximately same length as carpus, with three simple setae and one spiniform seta ventrodistally. Carpus approximately 1.7 times as long as broad, with several simple setae distally, and five spiniform setae (four ventral and one on subdistal outer surface). Propodus approximately 2.5 times shorter than basis, with several distal simple setae, one dorsal broom seta, four ventral and one dorsodistal spiniform setae, and three spiniform setae on outer surface. Dactylus shorter than basis, slightly longer than carpus and propodus combined; digitiform lobe absent; unguis fused with dactylus.

Pereopod 3 (not illustrated). Similar to pereopod 2.

Pereopod 4 ( Figs. 23E, F ). Basis approximately 2.6 times as long as broad, with two proximal broom setae, two simple dorsodistal setae (one long and one short) and one short simple ventrodistal seta. Ischium with two ventrodistal simple setae. Merus shorter than carpus, with two simple and three spiniform setae. Carpus with six spiniform setae on inner surface and seven spiniform setae on outer surface. Propodus with seven serrate spiniform setae on outer surface, five serrate spiniform setae on inner surface, one dorsodistal serrate spiniform seta, three short bipinnate setae on inner surface, and one proximal broom seta on dorsal margin. Dactylus shorter than propodus, with tuft of four aesthetascs; unguis absent.

Pereopod 5 ( Figs. 23F, G ). Similar to pereopod 4. Carpus with seven spiniform setae on inner surface and seven spiniform setae on outer surface. Dactylus with a tuft of five aesthetascs.

Pereopod 6 (Fig. 24A). Basis slender, approximately 4.5 times as long as broad, with six plumose setae on dorsal margin, four plumose and two simple setae on ventral margin and one proximal broom seta on outer surface. simple setae on ventral margin, six plumose setae on dorsal margin, and one simple seta subdistally on outer surface. Propodus shorter than the carpus, with four long spiniform setae on ventral margin and 20 short bipinnate setae. Dactylus longer than ischium, merus, carpus and propodus combined, without any subterminal setae; unguis absent.

Pleopods (not illustrated). Basal article with three long plumose setae. Exopodite with 15–16 plumose setae and endopodite with 18–20 plumose setae.

Uropods (Fig. 24B). Basal article approximately 2.8 times as long as broad, with several subdistal simple setae and one spiniform seta on inner distal corner. Exopodite with three articles, last article approximately 2.2 times as long as second article, with four terminal simple setae. Endopodite (exact number of articles difficult to determine due to incomplete fusion in some of the articles).

Ovigerous Female. Very similar to male but with the following differences:

Antennule (not illustrated). Main flagellum with three aesthetascs.

Cheliped (Figs. 24D, E). Less robust. Cutting edge of fixed finger with numerous spinules interspersed with one or two round tubercles.

Pereopod 6 (not illustrated). Dactylus shorter than carpus and propodus combined.

Type Locality. Brunei, northwest coast of Borneo , 05°21'N 114°40'E, Indo–Pacific (Fig. 1, number 9) .

Geographic distribution. Sabah, Malaysia, Indo–Pacific.

Remarks. A full description of this species is included here to expand on Bamber’s (1998) partial description. This species differs from the two other congeners by lacking sensory lobes on the dactylus of pereopods 2 and 3 and lacking a subterminal seta on the pereopod 6 dactylus. The males of this species are unique in having the pereopod 6 dactylus longer than usual (longer than the ischium, merus, carpus and propodus combined).

Two male forms were detected in the material examined (one of the male forms, “form B”, has a cheliped as in Fig. 24C). This is usually interpreted as a possibility of protogynous hermaphroditism (Mañe–Garzon, 1949), but developmental studies are needed to determine if one or both of these “forms” represent non–hermaphroditic primary or secondary (protogynous) males, or simply subterminal and terminal male stages. The males of both forms overlap in size range, making it difficult to determine which of these options is in play, but it is probable that the cheliped in Fig. 22F represents the terminal male form because it is more robust than the other form and the spine near the dactylus insertion and the tooth on the fixed finger cutting edge are better developed.

Phoxokalliapseudes multiarticulus (Gu ţ u, 2006) n. comb.

( Figs 25 –29)

Kalliapseudes multiarticulus Guţu, 2006: 127 , 138–142, Figs. 201–208.

Type material. Holotype female, Museum and Art Gallery of the Northern Territory, Darwin ( Australia), No. Cr 14429.

Material examined. 1 ovigerous female and 1 adult male, ZMUC Gal. #503, Arafura Sea , central Indo – Pacific, N Australia, 10°37'S, 139°19'E, 57 m, coral–sand and gravel, many shells, Sept. 27, 1951 .

Diagnosis (adult female). Rostrum tapering to rounded tip. Pleotelson tapering posteriorly to indented tip and with two terminal plumose setae. Antennule first peduncle article with several ventral spiniform setae; accessory flagellum of 6 or 7 articles. Antenna first peduncle article with one simple seta on outer subdistal corner; third peduncle article with three plumose setae on inner margin; flagellum with six articles, second and third articles with several pectinate setae on outer margin. Labrum complex with four strong cusps. Terminal spiniform seta of mandibular palp approximately five times as long as broad. Cutting edge of cheliped fixed finger with numerous spinules along entire margin interspersed with round protuberances. Cutting edge of cheliped dactylus with numerous (≥ 10) long spinules along entire margin. Cheliped and pereopod 1 exopodite with two plumose setae. Pereopod 5 propodus with short bipinnate setae on inner and outer surfaces. Pereopod 6 dactylus with one subterminal seta. Uropod basal article approximately 2.5 times as long as broad; exopodite with three articles, last article approximately three times as long as second article.

Description. Ovigerous female. Body ( Fig. 25A ): length approximately 9 mm, 5.7 times as long as broad.

Carapace ( Fig. 25A ). As long as broad, one pair mid–lateral setae; rostrum tapering to a rounded tip.

Pereonites ( Fig. 25A ). Pereonites 4 and 5 longer than others and pereonite 6 shortest; at least one pair anterolateral setae; hyposphaenia present on all pereonites.

Pleon ( Fig. 25A ). Pleonites subequal; rounded epimera, with several plumose setae, one pair dorsolateral simple setae; hyposphaenia present on all pleonites. Pleotelson subequal in length to last three pleonites combined, narrowing posteriorly to indented tip, with two terminal plumose setae.

Antennule ( Figs. 25B, C ). First peduncle article approximately 2.5 times as long as second and third articles combined and approximately 2.6 times as long as maximum width, with some simple setae on inner margin and several simple and broom setae on outer margin. Second peduncle article approximately four times shorter than first article, with several simple and broom setae. Third article slightly longer than fourth article (including common article). Outer flagellum shorter than first peduncle article, with 11 articles, one aesthetasc each on articles 7, 8, and 9. Inner flagellum with 6 articles, last article with three terminal setae (two simple and one broom).

Antenna ( Fig. 25D ). First peduncle article with one simple seta on outer subdistal corner and medial extension bearing eight plumose setae. Second peduncle article naked and squama with four simple setae. Third peduncle article with three plumose setae on inner margin. Peduncle article 4 (last) longer than flagellum, with double row plumose setae, one proximal broom seta. Flagellum with 6 articles, second and third articles with several pectinate setae and one or two plumose setae on each article, distal article with four terminal simple setae.

Labrum ( Fig. 25E ) rounded posteriorly, finely setose and with four strong anterior cusps.

Mandibles ( Figs. 25F–I ). Left mandible ( Fig. 25H ): incisor process with approximately 14 and lacinia mobilis with approximately 10 teeth; spine row with five spiniform setae. Right mandible: incisor process with two large and three small teeth; spine row with five spiniform setae. Palp ( Fig. 25F ) with a short terminal spiniform seta, approximately five times as long as broad.

Labium ( Fig. 25J ) with short hairs on anterior margin. Palp with long hairs on margins; ending in an acuminate inner tip, approximately 2.5 times as long as broad.

Maxillule ( Fig. 25K ). Inner endite bearing four terminally setulate setae and dense rows of hairs on outer margin. Outer endite with eleven long and one short spiniform setae, two subterminal setae and dense rows of hairs on outer margin.

Maxilla ( Figs. 26A, B ). Inner lobe of fixed endite with posterior row of five serrate spiniform setae and with long anterior row of filter setae. Outer lobe of fixed endite with four distal serrate setae, several simple and pectinate setae, and one serrate spiniform seta on posterior face. Inner lobe of moveable endite with several simple blunt–tipped setae. Outer lobe of moveable endite with two pectinate and two plumodenticulate (bearing proximal setules and distal denticles). Inner margin spinulate.

Maxilliped ( Figs. 26C, D ). Basal article fringed with plumose setae on outer margin, inner margin naked. First article of palp with five inner simple setae and one outer simple seta. Last three articles of palp with double row of long plumose setae on inner margin; second article with one simple seta on outer distal corner. Endite ( Fig. 26D ) with nine pappose setae along margin and one long simple and several pappose distal setae; two coupling hooks.

Cheliped ( Figs. 26E, F ). Basis with four simple setae (two long and two short) on ventral margin. Merus longer than broad, with three distal simple setae and one short simple seta midway on ventral margin. Carpus approximately four times as long as broad, with double row of long, plumose setae ventrally and three simple setae on dorsodistal corner. Propodus with diagonal row of long, plumose setae on inner face; fixed finger with several simple setae just proximal to distal unguis; cutting edge with short spinules interspersed with round protuberance; palm with several simple setae. Dactylus with several simple setae on inner surface; cutting edge with several spinules (> 10) increasing in length distally. Exopodite with two plumose setae.

Pereopod 1 ( Figs. 26G, H ). Basis approximately 2.5 times as long as broad, with one proximal broom seta and four simple ventrodistal setae. Ischium naked. Merus shorter than and as broad as basis, with several simple setae distally and on ventral margin, and one spiniform seta on each distal corner. Carpus approximately 2.3 times shorter than merus, with several simple setae and two ventrodistal and one dorsodistal serrate spiniform setae. Propodus shorter than carpus, with several simple setae and five ventral and two dorsal serrate spiniform setae, one broom seta on dorsal margin, and one pectinate seta distally on the inner surface. Dactylus represented by a sensory organ, shorter than propodus, with several long, terminal sensory setae with two setae on inner surface; unguis absent. Exopodite with two plumose distal setae.

Pereopod 2 ( Fig. 26I ). Basis approximately three times as long as broad, with two broom setae (broken off) on dorsal margin, one proximal and one distal simple seta on ventral margin and two shorter simple setae on ventrodistal corner. Ischium with three simple setae on ventrodistal corner. Merus approximately same length as carpus, with four simple and one spiniform setae on ventral margin. Carpus approximately 1.6 times as long as broad, with several simple setae distally and on ventral margin, and five spiniform setae. Propodus approximately three times shorter than basis, with several distal simple setae, one middorsal broom seta, six ventral and one dorsodistal spiniform setae, and three spiniform setae on outer surface. Dactylus shorter than basis, approximately as long as carpus and propodus combined; sensory organ present near base, with eight aesthetascs; unguis fused with dactylus.

Pereopod 3 ( Figs. 27A, B ). Similar to pereopod 2. Carpus with 12 spiniform setae. Propodus with seven ventral and two dorsodistal serrate spiniform setae, and four spiniform setae on outer surface. Dactylus sensory organ with approximately 13 aesthetascs.

Pereopod 4 ( Figs. 27C, D ). Basis approximately 2.4 times as long as broad, with two short ventrodistal setae, and two proximal broom setae and one simple seta near dorsal margin. Ischium with five ventrodistal simple setae. Merus shorter than carpus, with five simple setae on inner surface, one simple seta on ventral margin, and one simple and one spiniform seta on outer surface. Carpus with ten spiniform setae on outer surface and seven spiniform setae on inner surface. Propodus with nine serrate spiniform setae on outer surface, nine serrate spiniform setae on inner surface, one terminal serrate spiniform seta, three short bipinnate setae on outer surface and seven on inner surface, and one proximal broom seta on dorsal margin. Dactylus shorter than propodus, with tuft of approximately 13 sensory setae; unguis absent.

C, antennule 1 st peduncle article (ventral); D, antenna; E, labrum complex; F, right mandible; G, right mandible incisor process; H, left mandible; E, left mandibular palp; J, labium; K, maxillule. Scale bars: A = 0.5 mm; B, D, E = 0.2 mm; F = 0.1 mm; J, K = 0.05 mm.

Pereopod 5 ( Figs. 27E, F ). Similar to pereopod 4. Carpus with six spiniform setae on inner surface. Propodus with one short bipinnate seta on outer surface and five on inner surface. Dactylus with approximately 16–18 sensory setae.

Pereopod 6 ( Fig. 27G ). Basis slender, approximately 4.4 times as long as broad, with nine plumose setae on dorsal margin, five plumose and three simple setae on ventral margin. Ischium with five simple setae on ventral margin. Merus longer than ischium, with one plumose seta on dorsal margin and four simple setae on ventral margin. Carpus approximately 2.5 times as long as merus, with eight simple setae on ventral margin, seven plumose setae on dorsal margin, and four simple setae distally on outer surface. Propodus shorter than the carpus, with five long spiniform setae on ventral margin and 29 short bipinnate setae. Dactylus shorter than carpus and propodus combined, with one subdistal seta; unguis absent.

FIGURE 29. Phoxokalliapseudes multiarticulus comb. nov., adult male. A, pereopod 2 (outer); B, pereopod 3 (outer); C, pere-

Pleopods ( Fig. 28A ). Basal article with six long plumose setae. Exopodite with 27 plumose setae and endopodite with 31 plumose setae.

Uropods ( Fig. 28B ). Basal article approximately 2.5 times as long as broad, with one simple seta on outer margin (only base illustrated) and four distal simple setae. Exopodite with three articles, distal article longest, approximately three times as long as second article, with four simple setae. Endopodite with approximately 26 articles (exact number difficult to determine due to incomplete fusion in some of the articles), as long as last two pereonites and pleonites and pleotelson combined.

Male. Very similar to female but with the following differences:

Rostrum ( Fig. 28C ) rounded, not tapering.

Antennule ( Fig. 28D ). Proximal articles of outer flagellum with cluster of aesthetascs, decreasing in number distally.

Antenna ( Fig. 28E ). First peduncle article medial extension with six plumose setae. Squama with three simple setae.

Cheliped ( Fig. 28F ). Basis robust with one short spiniform seta on ventral margin and ventrodistal corner. Carpus approximately 2.6 times as long as broad. Propodus robust; fixed finger cutting edge with proximal tooth proximal to spinules; palm with one large spine on inner surface. Dactylus with proximal hook–shaped tooth on inner surface.

Pereopod 2 (Fig. 29A). Carpus with eight spiniform setae.

Pereopod 3 (Fig. 29B). Carpus with nine spiniform setae. Dactylus sensory organ with eight sensory setae.

Pereopod 4 (Figs. 29C, D). Merus with three spiniform setae. Carpus with eight spiniform setae on outer surface.

Pereopod 5 (Figs. 29E, F). Carpus with seven spiniform setae on inner surface and eight spiniform setae on outer surface.

Pereopod 6 (Fig. 29G). Basis with 10 plumose setae on dorsal margin and seven plumose setae on ventral margin. Merus with three plumose setae on dorsal margin. Carpus with eight plumose setae on dorsal margin. Propodus with three long spiniform setae and approximately 24 bipinnate setae. Dactylus longer than ischium, merus, carpus, and propodus combined.

Type locality. Dudley Point , Darwin, Northern Territory, Australia ( Fig. 25 ) .

Geographic distribution. Darwin, Northern Territory, Australia; Arafura Sea, Central Indo–Pacific (Fig. 1, number 10).

Remarks. The original description of Phoxokalliapseudes (= Kalliapseudes ) multiarticulus was based on an adult female, thus only a partial description was given ( Guţu, 2006). A full supplemental description including the male is presented here to expand on the original description. Because the holotype was unavailable for study and the material examined by us was not topotypic, we could not determine with certainty that this species is conspecific with multiarticulus . However, the material examined came from near the type locality and based on the similarity to Guţu’s description, we tentatively consider this species conspecific with P. multiarticulus until topotypic material can be examined.

This is a large species (9–9.6 mm) characterized by having a multi–segmented antennule inner flagellum (6 or 7 articles) and two plumose setae on the cheliped and pereopod 1 exopodite. The holotype has very small ventral spiniform setae on the pereopod 1 propodus and we attribute this to aberrancy because no other species of Kalliapseudinae has such small setae. It also must be pointed out that the left mandibular palp in the female described here has a very unusual terminal spiniform seta: it has a small spiniform seta coming off the base of it ( Fig. 25I ) and this is attributed to aberrancy. Guţu (2006) described this species has having two articles in the uropod exopodite but we noticed three articles. This character has caused confusion in the past (see the “remarks” section under Mesokalliapseudes crassus ) because the first article is usually very small and can be missed if the uropod is viewed dorsally because it attaches to the basal article ventrally.

Figures

FIGURE 23. Phoxokalliapseudes gobinae comb. nov., adult male. A, pereopod 1 (outer); B, pereopod 2 (outer); C, pereopod 2 (distal); D, pereopod 4 (outer); E, pereopod 4 (inner); F, pereopod 5 (inner); G, pereopod 5 (outer). Scale bars = 0.2 mm.

FIGURE 25. Phoxokalliapseudes multiarticulus comb. nov., ovigerous female. A, dorsal view of body; B, antennule (dorsal);

FIGURE 26. Phoxokalliapseudes multiarticulus comb. nov., ovigerous female. A, maxilla; B, outer lobe of fixed endite; C, maxilliped; D, maxillipedal endite; E, cheliped (inner); F, chela (inner); G, pereopod 1 (inner); H, pereopod 1 (outer); I, pereopod 2 (outer). Scale bars: A = 0.05 mm; C, E, G, I = 0.2 mm.

FIGURE 27. Phoxokalliapseudes multiarticulus comb. nov., ovigerous female. A, pereopod 3 (outer); B, pereopod 3 dactylus (outer); C, pereopod 4 (inner); D, pereopod 4 (outer); E, pereopod 5 (outer); F, pereopod 5 (inner); G, pereopod 6 (inner). Scale bars = 0.2 mm.

FIGURE 28. Phoxokalliapseudes multiarticulus comb. nov., ovigerous female (A, B); adult male (C–F). A, pleopod; B, uropod; C, dorsal view of body; D, antennule; E, antenna; F, cheliped (inner). Scale bars: A, D, F = 0.2 mm; B, E = 0.1 mm; C = 0.4 mm.