Descriptions of New Species of Birubius (Amphipoda: Phoxocephalidae) from Australia and Papua New Guinea with Comments on the Bi

Memoirs of Museum Victoria 58(2): 255–295 (2001)

DESCRIPTIONS OF NEW SPECIES OF BIRUBIUS (AMPHIPODA: PHOXOCEPHALIDAE) FROM AUSTRALIA AND PAPUA NEW GUINEA WITH COMMENTS ON THE BIRUBIUS-KULGAPHOXUS-TICKALERUS-YAN COMPLEX

JOANNE TAYLOR1,2 AND GARY C. B. POORE1 1Museum Victoria, GPO Box 666E, Melbourne, Victoria 3001, Australia 2Department of Zoology, University of Melbourne, Parkville, Victoria 3010, Australia ([email protected] and [email protected])

Abstract Taylor, J. and Poore, G.C.B., 2001. Descriptions of new species of Birubius (Amphipoda: Phoxocephalidae) from Australia and Papua New Guinea with comments on the Birubius-Kul- gaphoxus-Tickalerus-Yan complex. Memoirs of Museum Victoria 58(2): 255–295. Five new species of Birubius Barnard and Drummond, 1976 (Crustacea: Amphipoda: Phox- ocephalidae) are reported: B. drummondae sp. nov and B. heislersi sp. nov from Victoria, Aus- tralia; B. wallisae sp. nov. from Queensland, Australia; and B. lowryi sp. nov., and B. wilsoni sp. nov. from Papua New Guinea. The present records extend the range of the genus previously reported from Australia and Indonesia. The genus Birubius is discussed and compared with the Australian genera Tickalerus Barnard and Drummond, 1978 (monotypic), Kulgaphoxus Barnard and Drummond, 1978 (two species) and Yan Barnard and Drummond, 1978 (two species) in light of the new species exhibiting a combination of characters from all genera. Their synonymy is foreshadowed.

Introduction and Bousfield, 1994, Metharpinia Schellenberg, 1931, Microphoxus J.L. Barnard, 1960 and Rhep- Barnard and Drummond (1978) discussed the oxynius J.L. Barnard, 1979. The genus Linca was relationships between all Australian species of not included in the new subfamily and so remains the Phoxocephalidae. The subfamily Birubiinae a member of the Birubiinae. was established to include the Australian type Our preliminary cladistic analysis (work in genus Birubius Barnard and Drummond, 1976, progress) of most species of Birubiinae, Brol- Australian genera Tickalerus Barnard and ginae, Leongathinae, Metharpiniinae, Parhar- Drummond, 1978, Kulgaphoxus Barnard and piniinae and Tipimeginae has failed to support Drummond, 1978 and Yan Barnard and the monophyly of any of the subfamilies or Drummond, 1978 and the North and South genera. We are unable to identify a synapo- American genera Microphoxus J.L. Barnard, morphy for the subfamily Birubiinae, even in the 1960 and Metharpinia Schellenberg, 1931. restricted sense, nor for its type genus Birubius. Barnard and Karaman (1991) expanded the sub- The Birubiinae shares a broad form of the basis of family to include the North and South American pereopod 5 with all subfamilies except Harpini- genera Foxiphalus J.L. Barnard, 1979 and inae which exhibit a narrow basis unique to that Grandiphoxus J.L. Barnard, 1979. The genus subfamily. It shares a biarticulate palp of maxilla Linca Alonso de Pina, 1993 was erected based on 1 with all subfamilies except Phoxocephalinae a single specimen from the Argentine continental and those members of the Harpiniinae that exhibit shelf and although showing some convergence a uniarticulate palp. It differs from Brolginae, with the Brolginae, its similarity to Birubius best Harpiniinae, Phoxocephalinae and Ponthar- placed it in the Birubiinae. Linca differs from piniinae by the medium to elongate (rather than Birubius by the presence of ventral setae on short) length of peduncular article 2 of antenna 1, uropod 1 peduncle, an autapomorphy of this a state that it also shares with Tipimeginae, monotypic genus. Jarrett and Bousfield (1994) Parharpiniinae and some members of the reassessed the North and South American genera Joubinellinae. and removed them from the Birubiinae. They Examination of unidentified phoxocephalid erected the new subfamily Metharpiniinae to amphipods from the Australian Museum, include Beringiaphoxus Jarrett and Bousfield, Museum Victoria and Queensland Museum col- 1994, Foxiphalus J.L. Barnard, 1979, Grandi- lections revealed five new species belonging to phoxus J.L. Barnard, 1979, Majoxiphalus Jarrett

255 256 J. TAYLOR AND C. B. POORE the nominal subfamily Birubiinae. The generic Until the cladistic analysis is completed we are placement of some of the species was uncertain reluctant to complicate the taxonomy further by using Barnard and Karaman’s (1991) generic erecting new small genera simply because they do diagnoses. Although closely fitting the descrip- not comply with existing diagnoses. It is unclear tion for Birubius four species exhibited a large whether the minor genera in question will come to dorsal hook on urosomite 3, a character restricted encompass larger clades but it is certain that they within the Birubiinae to members of Tickalerus cannot be justified as currently defined. Therefore and Kulgaphoxus. in this contribution we describe the new taxa as Barnard and Drummond (1978) defined members of Birubius in spite of their similar- Kulgaphoxus, Tickalerus and Yan only on the basis ities to some members of Kulgaphoxus, of differences from Birubius, the largest genus. Tickalerus and Yan. The synonymy of these gen- Tickalerus differs from Birubius in the presence era with Birubius is foreshadowed. Our revised of a dorsal hook on urosomite 3, shortened outer diagnosis of Birubius is written to include all ramus of uropod 3 and rectangular coxa 4. Kul- species included in Birubius, Kulgaphoxus, gaphoxus differs from Birubius in the presence of Tickalerus and Yan. a dorsal hook on urosomite 3, shortened outer Abbreviations are: A, antenna; H, head; rLM, ramus of uropod 3, proximal placement of setae right lacinia mobilis; MD, mandible; MX, on peduncular article 2 of antenna 1 and the ves- maxilla; MP, maxilliped; GN, gnathopod; P, pere- tigial dactyl of pereopod 7. Yan differs from Biru- opod; EP, epimeron; U, uropod; PL, pleopod; T, bius in the proximal placement of setae on pedun- telson; r, right; m, male; tl., total length; MAFRI, cular article 2 of antenna 1 and the vestigial dactyl Marine and Freshwater Resources Institute, of pereopod 7. The new species share some but Queenscliff; NMV, Museum Victoria, Mel- not all of the diagnostic features of Kulgaphoxus, bourne; AM, Australian Museum, Sydney, QM, Tickalerus and Yan and could not be placed in any Queensland Museum, Brisbane. All dissections of the genera as presently diagnosed. and illustrations follow the methods of Barnard The discovery of four species exhibiting a and Drummond (1978) whereby the left side of dorsal hook on urosomite 3 is significant. Pre- the animal is illustrated unless otherwise stated. viously it was a trait observed in only five phoxo- Descriptions of the new species closely follow cephalid species belonging to Kulgaphoxus, that of other species of the genus described in Microphoxus and Tickalerus and was partly used Barnard and Drummond (1978). to split these species from Birubius. The trait was formerly believed to be sexually dimorphic and restricted to females with males having a reduced Birubius Barnard and Drummond hump at best. Males of the new species B. drum- Birubius Barnard and Drummond, 1976: 543.— mondae sp. nov. and B. wallisae sp. nov. however Barnard and Drummond, 1978: 191.—Barnard and exhibit a well developed dorsal hook as in Karaman, 1991: 635. females. It appears that sexual dimorphism is Type species. Birubius panamunus Barnard and variable but the possibility that males with- Drummond, 1976 (by original designation). out hooks belong to other species is a remote possibility. Diagnosis. Rostrum variably constricted. Eyes Barnard and Drummond did not use cladistic present. Antenna 1 peduncular article 2 length methodology to define genera. Rather, small variable, ventral setae not confined apically. genera were picked off from larger clusters on the Antenna 2 peduncular article 1 not or scarcely basis of few differences that may or may not be ensiform, article 3 with 2 facial setules, facial unique synapomorphies. The inevitable conse- robust setae on article 4 in 2+ rows, all robust quence of this is that the large genus, Birubius in setae thick, article 5 ordinary. Right mandibular this case, is paraphyletic because its numerous incisor with 3-4+ teeth, right lacinia mobilis bifid species lack a synapomorphy. We are forced to or simple, often flabellate or absent, molar not conclude that either (a) the small genera, triturative, with 4+ splayed robust setae; palpar Kulgaphoxus, Linca, Tickalerus and Yan, as hump small to medium, apex of palp article presently constituted are gradal offshoots of 3 oblique. Maxilla 1 inner plate with 3-4 setae, Birubiinae which cannot be supported in a classi- palp 2-articulate. Maxillipedal plates small to fication based on cladistic principles; or (b) the ordinary, apex of palp article 3 not strongly type species of the four genera represent much protuberant, dactyl elongate, apical nail distinct. larger clades which may be redefined using very Gnathopods small, similar, gnathopods 1-2 different character suites (synapomorphies). carpus length medium to elongate, not cryptic NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 257

(posterior margin not concealed by the abutment Birubius drummondae sp. nov. of propodus and merus), palms oblique, gnat- Figures 1–6 hopods 1–2 propodus ordinary to narrow, ovate to rectangular, poorly setiferous anteriorly. Pere- Material examined. Holotype. Australia, Victoria, opods 3–4 carpus with (rarely without) postero- Western Port (38º22´S, 145º32´E) no further data, proximal robust setae, propodus with robust setae. NMV J47227 (1 female, tl. 3.8 mm). Pereopod 5 basis of broad form (basis equal to or Allotype. Same locality as holotype, NMV J47228 (1 male, tl. 5.25 mm). greater than twice width of ischium), pereopods Paratypes. Same locality as holotype, NMV J47226 5–6 merus-carpus broad to narrow; pereopod 7 (27 females, tl. 3.0–5.7 mm). unreduced, article 3 not enlarged, dactyl well developed, vestigial or absent. Diagnosis. Rostrum constricted. Antenna 2, arti- Epimera 1–2 with or without long facial cle 4 without well developed dorsal setation. brushes of setae, without posterior setae, Right lacinia mobilis bifid, distal branch denticu- epimeron 3 bearing long setae. Urosomite 3 with late. Pereopods 3–4 carpus with 2–3 proximopos- or without dorsal hook in females, sometimes in terior robust setae. Pereopod 5 dactyl fully male only if in female. Uropod 1 peduncle with- formed. Pereopod 7 basis without long ventral out interramal robust setae, without major dis- setae. Coxa 1 not expanded distally. Coxa 4 lack- placed robust seta (seta that is shifted onto the ing long ventral setae. Epimeron 3 without large apical margin disjunctly from the true inner mar- tooth; without ventral setae; without long poste- gin), uropods 1–2 rami occasionally continuously rior seta; with oblique row of facial setae. Uro- setose to apex (thus with minute apical robust somite 3 with large dorsal hook. Uropod 1 with- setae or nails), uropod 1 inner ramus with 1 row out basofacial setae. Uropods 1–2 inner rami of marginal robust setae. Uropod 2 inner ramus lacking accessory apical nails. Uropod 3 unre- ordinary to shortened. Uropod 3 variable, either duced, outer ramus longer than peduncle. unreduced (outer ramus longer than peduncle), or Description of female. Head about 18% of total reduced (outer ramus shorter than or subequal to body length, greatest width about 100% of length; peduncle), bearing a second article on outer rostrum constricted, exceeding apex of pedunc- ramus, with 2 long apical setae. Telson ordinary ular article 1 on antenna 1. Eyes medium, clear of to elongate. pigment. Antenna 1 peduncular article 1 about 1.3 Species. Biribius batei (Haswell, 1879); B. ros- times as long as wide, about 1.8 times as wide as tratus (Dana, 1853) = B. barnardi Pirlot, 1932. peduncular article 2, ventral margin with 4 Species described by Barnard and Drummond, setules, produced dorsal apex with 1 setule; 1978: B. apari; B. babaneekus; B. booleus; peduncular article 2 about 0.8 times as long as B. cartoo; B. chintoo; B. eake; B. eleebanus; peduncular article 1, with 5 ventral setae; primary B. gallangus; B. gambodeni; B. gelarus; B. jir- flagellum with 10 articles, about 0.9 times as long randus; B. kabbulinus; B. kareus; B. karobrani; as peduncle, bearing long aesthetascs; accessory B. kinkus; B. kokorus; B. kyeemus; B. lorus; flagellum with 8 articles. Antenna 2, peduncular B. lowannus; B. maamus; B. maldus; B. maya- article 4 robust setae formula = 1-3-4-3, dorsal mayi; B. muldarpus; B. munggai; B. myallus; margin with notch bearing 3 setae, ventral margin B. nammuldus; B. narus; B. panamunus; with 4–5 groups of 1–2 long to short setae, 1 long B. quearus; B. taldeus; B. thalmus; B. ularitus; ventrodistal robust seta; peduncular article 5 B. wirakus; B. wulgaru; B. yandus; B. yorlunus. about 0.8 times as long as peduncular article 4, Species added after 1978: B. bali Ortiz and Lalana, facial robust seta formula = 1–2, dorsal margin 1999; B. drummondae sp. nov.; B. heislersi sp. naked, ventral margin with 3 sets of 1–2 long to nov.; B. lowryi sp. nov.; B. murariui Ortiz and short setae, without ventrodistal robust setae; Lalana, 1997; B. wallisae sp. nov.; B. wilsoni flagellum 1.51 times as long as peduncular arti- sp. nov. cles 4–5 combined, with 11 articles. Mandibles with medium palpar hump; right incisor with 4 Habitat and distribution. Marine 0–70 m. Aus- teeth and notch; left incisor with 2 humps in 2 tralia; Indonesia; Papua New Guinea. branches; right lacinia mobilis bifid, distal branch much shorter than proximal branch, denticulate, Remarks. Barnard and Karaman’s (1991) proximal branch simple, pointed, with facial generic diagnosis has been altered to accommo- hump; left lacinia mobilis sub-bifid; right raker 7; date the new species and to reflect the fore- left rakers 7 plus 1 rudimentary; molar in form of shadowed synonymy of Tickalerus, Kulgaphoxus bulbous hump, right molar with 4 long robust and Yan. setae, plus 1 short robust seta strongly disjunct, 258 J. TAYLOR AND C. B. POORE

Figure 1. Birubius drummondae sp. nov., holotype female, tl. 3.80 mm (m = male allotype, 5.25 mm). left molar with 5 long robust setae, plus 1 short setae. Maxilla 1 inner plate narrow, bearing 1 robust seta strongly disjunct; palp article 1 long apical seta, 1 similar apicomedial seta, 2 api- slightly elongate, article 2 with 1 medium inner colateral much shorter seta; palp article 2 with 1 apical seta and 2 other shorter inner setae, article apicomedial marginal robust seta, 3 apicomedial 3 about equal in length to article 2, apex oblique setae and 3 submarginal setae. Maxilla 2 inner with 6 robust to slender setae, with 3 basofacial and outer plates extending equally, outer not NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 259

Figure 2. Birubius drummondae sp. nov., holotype female, tl. 3.80 mm. broader than inner. Maxilliped inner plates with 2 medium, width-length ratio of coxa 4 almost = large thick apical robust seta, 3 apicofacial setae, 36:53. Long posterior setae on basis of 5 medial setae; outer plate with 7 medial and api- gnathopods 1–2 and pereopods 3–4 = 4-1-5-7, cal robust setae, 1 apicolateral seta; palp articles short posteriors = 1-0-1-0 , long anteriors = 4-8-0- 1–2 with 1 apicolateral seta, article 3 weakly pro- 0, short anteriors = 2-2-0-0. tuberant, with 3 facial setae, 1 lateral seta, nail of Gnathopods, width ratios of carpus-propodus article 4 medium length, with 1 accessory setule. on gnathopods 1–2 = 20:27 and 20:27, length Coxa 1 not expanded distally; main ventral setae ratios = 21:26 and 1:1; palmar humps ordinary, of coxae 1–4 = 5-5-5-0, posteriormost seta of palms oblique; gnathopod 1 carpus of medium coxae 1–3 shortened; anterior and posterior mar- length gnathopod 2 carpus slightly elongate. Pere- gins of coxa 4 parallel, posterior margin straight, opods 3–4 similar, facial setae on merus = 3 and posterodorsal corner sharp, posterodorsal margin 3, on carpus = 3 and 4; main spine of carpus 260 J. TAYLOR AND C. B. POORE

Figure 3. Birubius drummondae sp. nov., holotype female, tl. 3.80 mm. extending to M. 77 on propodus, carpus with 2 pereopods 5–6 broad, facial robust setae rows and 3 proximoposterior robust setae; robust setae dense, facial ridge formula on basis of pereopods formula of propodus = 4 + 5 and 4 + 5; acclivity 5–7 = 0-2-2, anterior ridge of pereopod 7 long; on inner margin of dactyls of pereopods 3–4 width ratios of basis, merus, carpus, propodus of weak, midfacial seta ordinary. Coxae 5–7 pos- pereopod 5 = 11:12:11:5, of pereopod 6 = teroventral seta formula = 2-2-1; merus-carpus of 18:13:10:5, of pereopod 7 = 107:27:23:11, length NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 261

Figure 4. Birubius drummondae sp. nov., holotype female, tl. 3.80 mm (m = male allotype, 5.25 mm). 262 J. TAYLOR AND C. B. POORE

Figure 5. Birubius drummondae sp. nov., holotype female, tl. 3.80 mm (m = male allotype, 5.25 mm).

ratios of pereopod 5 = 45:21:24:22, of pereopod 6 cally; epimeron 3 posteroventral corner barely = 58:32:24:29, of pereopod 7 = 63:15:13:13; basis protuberant, with setule sinus, posterior margin of pereopod 7 exceeding apex of merus, naked almost straight, with 2 setule notches, ventral ventrally. Pleopods 1–3 with 2 coupling hooks; margin naked, midface with oblique row of 4 rear facial setae on peduncle = 2-2-3; articles on setae near posterior margin. Urosomite 1 naked, outer rami = 11-10-11, inner rami = 7-6-6. articulation line almost complete; urosomite 3 Epimeron 1 posteroventral corner rounded, with large hook dorsally. Uropods 1–2 rami with anteroventral margin with 3 setae, posteroventral articulate enlarged apical nails, uropod 1 outer face with 2 medium setae, set vertically; ramus with 3 dorsal robust setae, inner with 1 epimeron 2 posteroventral corner rounded, with 5 dorsomedial and 1 subapical robust setae, uropod facial setae, posteriormost pair set almost verti- 2 outer ramus with 2 dorsal robust setae, inner NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 263

Figure 6. Birubius drummondae sp. nov., holotype female, tl. 3.80 mm (m = male allotype, 5.25 mm).

with 1 broad dorsomedial robust seta; uropod 1 article 1 naked, lateral margin with 1 acclivity, peduncle with 3 apicolateral robust setae, without robust setal formula = 2-2, without slender setae basofacial slender setae, with apical enlarged formula. Telson long, length-width ratio = 34:29, robust seta; uropod 2 peduncle with 5 dorsal not fully cleft, each apex wide, rounded, lateral robust setae; apicolateral corners of peduncles on acclivity broad, shallow, bearing ordinary lateral uropods 1–2 without comb. Uropod 3 unreduced, setule, robust setae next medial little longer than outer ramus longer than peduncle. Uropod 3 setule, midlateral setules diverse. peduncle with 6 ventral robust setae, dorsally with 1 lateral robust seta; rami masculine, inner Description of male. Similar to female but eyes extending to M. 100+ on article 1 of outer ramus, larger. Antenna 1 like female but with dense apex with 3 setae, medial and lateral margins medial setation on peduncular article 1; primary naked, article 2 of outer ramus elongate, 0.38, flagellum bearing calceoli. Antenna 2 elongate, bearing 2 long setae, apicomedial margin of peduncular articles 3–4 with dense dorsal seta- 264 J. TAYLOR AND C. B. POORE tion, peduncular article 5 about as long as article Birubius heislersi sp. nov. 4, dorsal margin bearing 2 calceoli and 3 groups Figures 7–11 of male setae, flagellum 28-articulate bearing calceoli. Maxilliped and maxillae 1–2 similar to Material examined. Holotype. Australia, Victoria, female. Right lacinia mobilis bifid, distal branch Ninety Mile Beach (38°30’S, 147°25.8’E), 40 m, much shorter than proximal branch, flabellate, Smith-McIntyre grab, 8 May 1998 (MAFRI stn 37C), proximal branch simple, pointed, with facial NMV J47320 (1 female, tl. 6.8 mm). Paratypes. Australia, Victoria, Apollo Bay, Skenes humps; left lacinia mobilis with 4 teeth; right Creek (38°23.4’S, 144°15.6’E), 40 m, Smith-McIntyre raker 8; left rakers 9; right and left molars with 5 grab, 3 May 1998 (MAFRI stn 18C), NMV J47321 (2 long robust setae; palp similar to female, article 3 females, tl. 6.5–9.75 mm). with 6 basofacial setae. Main ventral setae of coxae 1–4 = 7-6-7-0. Gnathopods 1–2 similar to Diagnosis. Rostrum constricted. Antenna 2, arti- female. Urosomite 3 with large hook dorsally. cle 4 without well developed dorsal setation. Uropod 1 outer ramus with 5 dorsal robust setae, Right lacinia mobilis bifid, distal branch den- inner with 1 dorsomedial and 1 subapical robust ticulate. Pereopods 3–4 carpus with 3–4 setae, uropod 2 outer ramus with 3 dorsal robust proximoposterior robust setae. Pereopod 5 dactyl setae, inner with 1 broad dorsomedial robust seta; fully formed. Pereopod 7 basis with 1 medium uropod 1 peduncle with 2 apicolateral robust ventral setae, without long ventral setae. Coxa 1 setae, with 1 basofacial slender seta; uropod 2 expanded distally. Coxa 4 lacking long ventral peduncle with 12 dorsal robust setae. Uropod 3 setae. Epimeron 3 with small tooth; without ven- with inner ramus elongate, exceeding apex of tral setae; without long posterior seta; with article 1 on outer ramus. Telson elongate, length- oblique row of facial setae. Urosomite 3 without width ratio = 7:6. dorsal hook. Uropod 1 with basofacial setae. Uropods 1–2 inner rami lacking accessory apical Etymology. For Margaret Drummond who identi- nails. Uropod 3 reduced, outer ramus shortened, fied this species as new from Museum Victoria subequal to peduncle. collections and contributed so much to knowledge of Australian amphipods. Description of female. Head about 18% of total Remarks. The following variations from the holo- body length, greatest width about 78% of length; type were observed in the paratypes. The main rostrum constricted, exceeding peduncular article ventral setae of coxae 1–4 = (4-5)-(4-6)-(4-6)-0. 1 on antenna 1. Eyes medium, clear of pigment. Uropod 1 outer ramus with 3–4 dorsal robust Antenna 1 peduncular article 1 about 1.3 times as setae, inner ramus with 1 dorsomedial and 1 sub- long as wide, about 1.6 times as wide as pedun- apical robust setae. Uropod 2 outer ramus with cular article 2, ventral margin with 10 setules, 1–2 dorsal robust setae, inner ramus with 1 dorsal weakly produced dorsal apex with 3 setules; robust setae. peduncular article 2 about 0.8 times as long as Birubius drummondae shares the dorsal hook peduncular article 1, with 8 ventral setae; primary of urosomite 3 with Tickalerus birubi, both flagellum with 15 articles, about 0.95 times as species of Kulgaphoxus and three other new long as peduncle, bearing aesthetascs; accessory species of Birubius described herein. This species flagellum with 13 articles. Antenna 2, peduncular can not be placed in the genus Tickalerus as it article 4 robust setae formula = 1-3-5-6, dorsal lacks both the well developed dorsal setation on margin with notch bearing 3 setae, ventral margin article 4 of female antenna 1 and the shortened with 6 groups of 1–2 long to medium setae, 1 ven- outer ramus of uropod 3, characters diagnostic of trodistal robust seta; peduncular article 5 about the type species, T. birubi. It remains distinct 0.7 times as long as peduncular article 4, facial from both species of Kulgaphoxus in its lack of robust seta formula = 0-3, dorsal margin naked, accessory apical nails on the inner rami of both ventral margin with 3 sets of 1–2 long to short uropods 1–2, the unreduced rostrum and the per- setae, 3 ventrodistal long to medium robust setae; fectly rectangular coxa 4, a character it shares flagellum 1.6 times as long as peduncular articles with T. birubi. Birubius drummondae differs from 4-5 combined, with 16 articles. Mandibles with previously described species of Birubius by the medium palpar hump; right incisor with 3 teeth; presence of the dorsal hook on urosomite 3, and left incisor with 3 teeth in 2 branches; right lacinia from the other new species described herein by mobilis bifid, distal branch shorter than proximal the combination of characters listed in the diag- branch, broad, denticulate, proximal branch noses. The species is number MoV3679 in simple, pointed, with marginal denticles; left Museum Victoria’s TAXA database. lacinia mobilis with 4 teeth; right raker 10; left NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 265

Figure 7. Birubius heislersi sp. nov., holotype female, tl. 6.80 mm. rakers 7; molar in form of short protrusion demar- seta strongly disjunct; palp article 1 slightly elon- cated mainly by robust setae, right and left molar gate, article 2 with 1 medium inner apical seta and with 5–6 long robust setae, plus 1 short robust 2 other shorter inner setae, article 3 about 0.8 266 J. TAYLOR AND C. B. POORE

Figure 8. Birubius heislersi sp. nov., holotype female, tl. 6.80 mm. NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 267

Figure 9. Birubius heislersi sp. nov., holotype female, tl. 6.80 mm. 268 J. TAYLOR AND C. B. POORE

Figure 10. Birubius heislersi sp. nov., holotype female, tl. 6.80 mm. NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 269

Figure 11. Birubius heislersi sp. nov., holotype female, tl. 6.80 mm. 270 J. TAYLOR AND C. B. POORE times long as article 2, apex oblique with 8 robust setae, posteroventral face with 2 long setae, pos- setae, with 6 basofacial setae. Maxilla 1 inner terior margin with 3–5 setules in sinuses; plate large, bearing 1 long apical seta, 1 similar epimeron 2 posteroventral corner rounded, with 5 apicomedial seta, 1 similar apicolateral seta; palp facial setae, posteriormost pair set vertically, pos- article 2 with 4 apicomedial marginal robust setae terior margin with 5–6 setules in sinuses; and 6 submarginal setae. Maxilla 2 inner and epimeron 3 posteroventral corner weakly pro- outer plates extending subequally, outer not tuberant, with small tooth, posterior margin broader than inner. Maxilliped inner plates with 2 straight, with setule sinuses, ventral margin large thick apical robust seta, 3 apicofacial setae, naked, face with horizontal row of 9 setae. 4 medial setae; outer plate with 6 medial and Urosomite 1 naked, articulation almost complete; apical robust setae, 2 apicolateral setae; palp urosomite 3 weakly protuberant dorsally, without article 1 with 1 apicolateral setae, article 2 with hook. Uropods 1–2 rami with articulate enlarged 3 apicolateral setae, article 3 protuberant, with apical nails, uropod 1 outer ramus with 6 dorsal 5 proximal facial setae, with 2 lateral setae, nail of robust setae, inner with 1, uropod 2 outer ramus article 4 medium length, with 1 accessory setules. with 4 dorsal robust setae, inner with 1 dorso- Coxa 1 expanded distally, anterior margin weakly medial robust seta; uropod 1 peduncle with 7 api- concave; main ventral setae of coxae 1–4 = 8-8-9- colateral robust setae and 2 basofacial slender 0, posteriormost seta of coxae 1–3 shortened; setae, apically with 2 marginal robust setae, api- anterior and posterior margins of coxa 4 diver- calmost enlarged, medially with 3 slender setae; gent, posterior margin oblique, almost straight, uropod 2 peduncle with 7 dorsal robust setae; api- posterodorsal corner sharp, posterodorsal margin colateral corners of peduncles on uropods 1–2 short, width-length ratio of coxa 4 = 59:50. Long without comb. Uropod 3 reduced, outer ramus posterior setae on basis of gnathopods 1–2 and shortened, subequal to peduncle. Uropod 3 pereopods 3–4 = 3-8-12-12, short to medium pos- peduncle with 5 ventral robust setae, dorsally teriors =2-1-1-0 , long anteriors = 3-7-2-1, short with 1 lateral robust seta; rami masculine, inner anteriors = 1-6-4-6. extending to M. 100+ on article 1 of outer ramus, Gnathopod propodus narrow; gnathopods 1–2 apex with 2 setae, medial margin with 1 seta, lat- width ratios of carpus-propodus = 5:7 and 11:15, eral margin with 4 setae, article 2 of outer ramus length ratios = 1:1 and 25:26; palmar humps short, 0.21, bearing 2 long setae, apicomedial ordinary, palms oblique; gnathopods 1–2 carpus margin of article 1 with a single seta, lateral elongate. Pereopods 3–4 similar, facial setae on margin with 2 acclivities, robust setal formula = merus = 5 and 6, on carpus = 4 and 5; main spine 1-1-2, slender setal formula = 2-1-0. Telson of carpus extending to M. 75 on propodus, carpus length-width ratio = 1:3, almost fully cleft, each with 3–4 proximoposterior robust setae; robust apex wide, rounded, lateral acclivity broad, shal- setae formula of propodus = 4 + 5; acclivity on low, bearing ordinary lateral setule, robust setae inner margin of dactyls of pereopods 3–4 weak, next medial longer than setule, midlateral setules midfacial seta ordinary. Coxae 5–7 posteroventral diverse. setule formula = 7-9-7; merus-carpus of pereopods 5–6 broad, facial robust setae rows dense, Male. Unknown. facial ridge formula on basis of pereopods 5–7 = Etymology. For Simon Heislers, Museum Victo- 0-2-2, anterior ridge of pereopod 7 very short; ria, who identified this species as new from width ratios of basis, merus, carpus, propodus of Museum Victoria collections. pereopod 5 = 29:30:26:12, of pereopod 6 = 50:33:25:13, of pereopod 7 = 65:15:14:6, length Remarks. The following variations from the holo- ratios of pereopod 5 = 55:23:26:29, of pereopod 6 type were observed in the paratypes. The main = 62:39:31:33, of pereopod 7 = 69:19:17:17; ventral setae of coxae 1–4 = (5-8)-(5-8)-(5-9)-0. pereopod 7 basis reaching or exceeding middle of Uropod 1 outer ramus with 6–8 dorsal robust carpus, with 1 medium ventral setae. Pleopods setae, inner ramus with 1 dorsal robust seta. Uro- 1–3 with 2 coupling hooks; pleopod 1 with 3 mid pod 2 outer ramus with 3–4 dorsal robust setae, and 3 proximal facial setae, pleopod 2 with inner ramus with 1 dorsal robust seta. 2 proximal, 3 mid and 2 distal facial setae, pleo- Birubius heislersi conforms well to Barnard pod 3 with 5 proximal and 5 distal facial setae; and Drummond’s (1978) diagnosis of Birubius articles on outer rami = 15-14-16, inner rami = except that it exhibits shortened rami of uropod 3 10-9-10. as seen in Tickalerus and Kulgaphoxus. It differs Epimeron 1 posteroventral corner rounded, from these genera in the lack of a dorsal hook on anteroventral margin with 8 short to medium urosomite 3. Attempts to identify this species NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 271 using Barnard and Drummond’s (1978) key with 6–7 groups of 1–2 long to short setae, with- failed. Birubius heislersi appears most similar to out ventrodistal robust seta; peduncular article 5 B. lowannus (Barnard and Drummond, 1978) but about 0.76 times as long as peduncular article 4, differs on many accounts including the denticu- facial robust seta formula = 1–2, dorsal margin late vs simple distal branch of the right lacinia naked, ventral margin with 4 sets of 1–2 long to mobilis, more ventral setae on coxae 1–3, the short setae, 2 ventrodistal long to medium robust presence of 3–4 vs 1 proximoposterior robust seta setae; flagellum 1.07 times as long as articles 4–5 on carpus of pereopods 3–4 and the naked ventral of peduncle combined, with 11 articles. margin of uropod 3. The species is number Mandibles with medium to large palpar hump; MoV3671 in Museum Victoria’s TAXA right incisor with 3 teeth; left incisor with 2 database. humps in 2 branches; right lacinia mobilis bifid, distal branch shorter than proximal branch, Birubius lowryi sp. nov. simple, pointed, proximal branch simple, pointed; left lacinia mobilis with 5 teeth; right raker 8; left Figures 12–17 rakers 7; molar in form of short protrusion demar- Material examined. Holotype. Papua New Guinea, NW cated mainly by robust setae, right molar with 6 corner of Pig I. (05°9.98’S, 145°50.45’E), 21 m, J. D. long robust setae, left molar with 6 long robust Thomas, 4 Feb 1990 (stn PNG 33K), AM P60004 (1 setae, no seta disjunct; palp article 1 slightly elon- female, tl. 4.75 mm). gate, article 2 with 2 long-medium inner apical Allotype. Papua New Guinea, Barracuda Point, E of Pig I. (05°10.26’S, 145°50.61’E), 30 m, J. D. Thomas, setae and 2 other medium inner setae, article 3 8 Feb 1990 (stn PNG 37K), AM P56151 (1 male, tl. about 0.86 times long as article 2, apex oblique 3.75 mm). with 6 robust to slender setae, without basofacial Paratypes. Same data as allotype. AM P60005 (4 setae. Maxilla 1 inner plate narrow, bearing 1 females, tl 3.0–3.75 mm). Papua New Guinea, Horse- long apical seta plus 1 shorter apicomedial seta; shoe Reef, Bootless Inlet (09°30.05’S, 147°15.50’E), palp article 2 with 1 apicomedial marginal robust 30 m, 28 Oct 1980, AM P60006 (2 females, tl. 3.75–3.9 seta, 3 apicomedial setae and 3 submarginal setae. mm). Maxilla 2 inner and outer plates extended equally. Diagnosis. Rostrum constricted. Antenna 2, arti- Maxilliped inner plates with 1 large thick apical cle 4 without well developed dorsal setation. robust seta, 3 apicofacial setae, 1 medial seta; Right lacinia mobilis bifid, distal branch simple. outer plate with 5 medial and apical robust setae; Pereopods 3–4 carpus with 1 proximoposterior palp article 1 with 1 apicolateral seta, article 2 robust seta. Pereopod 5 dactyl fully formed. with 2 apicolateral setae and 1 other lateral seta, Pereopod 7 basis with long ventral setae. Coxa article 3 unprotuberant, with 2 facial setae, nail of 1 strongly expanded distally. Coxa 4 lacking long article 4 long, with 2 accessory setules. Coxa 1 ventral setae. Epimeron 3 with small tooth; with strongly expanded distally; posterior setae of coxa ventral setae; without long posterior seta; without 1–3 = 3-4-3, main ventral setae of coxae 1–4 = facial setae. Urosomite 3 with large dorsal hook. 6-6-8-0, posteriormost seta of coxae 1–3 elon- Uropod 1 with basofacial setae. Uropods 1–2 gate; anterior and posterior margins of coxa 4 inner rami lacking accessory apical nails. Uropod strongly divergent, posterior margin oblique, pos- 3 unreduced, outer ramus longer than peduncle. terodorsal corner rounded, posterodorsal margin medium, width-length ratio of coxa 4 almost = Description of female. Head about 16% of total 1:1. Long posterior setae on basis of gnathopods body length, greatest width about 84% of length; 1–2 and pereopods 3–4 = 2-8-9-10, short posteri- rostrum constricted, narrow, elongate, reaching ors = 5-6-6-7, long anteriors = 0-10-0-0, short middle of peduncular article 2 on antenna 1. Eyes anteriors = 6-6-12-12. large, clear of pigment. Antenna 1 peduncular Gnathopods, width ratios of carpus-propodus article 1 about 1.5 times as long as wide, about 2.0 on gnathopods 1–2 = 9:11 and 10:13, length ratios times as wide as article 2, ventral margin with = 23:26 and 25:19; palmar humps ordinary, palms 8 setules, unproduced dorsal apex without setule; oblique; gnathopods 1–2 carpus of medium peduncular article 2 about 0.6 times as long as length. Pereopods 3–4 similar, facial setae on peduncular article 1, with 5 ventral setae; primary merus = 4 and 3, on carpus = 4 and 4; main spine flagellum with 10 articles, about 0.8 times as long of carpus extending to M. 91 on propodus, carpus as peduncle, lacking aesthetascs; accessory flag- with 1 proximoposterior robust seta; robust setae ellum with 8 articles. Antenna 2, peduncular formula of propodus = 2 + 4; acclivity on inner article 4 robust setae formula = 1-3-4-4, dorsal margin of dactyls of pereopods 3–4 weak, mid- margin with notch bearing 2 setae, ventral margin facial seta short. Coxae 5–7 posteroventral seta 272 J. TAYLOR AND C. B. POORE

Figure 12. Birubius lowryi sp. nov., holotype female, tl. 4.75 mm (m = male allotype, 3.75 mm). NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 273

Figure 13. Birubius lowryi sp. nov., holotype female, tl. 4.75 mm. 274 J. TAYLOR AND C. B. POORE

Figure 14. Birubius lowryi sp. nov., holotype female, tl. 4.75 mm (m = male allotype, 3.75 mm). NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 275

Figure 15. Birubius lowryi sp. nov., holotype female, tl. 4.75 mm (m = male allotype, 3.75 mm). 276 J. TAYLOR AND C. B. POORE

Figure 16. Birubius lowryi sp. nov., holotype female, tl. 4.75 mm (m = male allotype, 3.75 mm). NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 277

Figure 17. Birubius lowryi sp. nov., holotype female, tl. 4.75 mm. formula = 6-1-3; articles 4–5 of pereopods 5–6 with 3 medium setae, ventral margin with 6 medium to narrow, facial robust setae rows medium setae. Urosomite 1 naked, articulation poorly developed, facial ridge formula on basis of line almost complete; urosomite 3 with large hook pereopods 5–7 = 0-2-2, anterior ridge of pereopod dorsally. Uropods 1–2 rami with articulate 7 long; width ratios of basis, merus, carpus, enlarged apical nails, uropod 1 outer ramus with 5 propodus of pereopod 5 = 24:27:20:9, of pereo- dorsal robust setae, inner with 1 dorsomedial pod 6 = 37:23:15:8, of pereopod 7 = 25:7:6:3, robust seta, uropod 2 outer ramus with 3 dorsal length ratios of pereopod 5 = 41:17:26:24, of robust setae, inner with 2 small robust seta; uro- pereopod 6 = 49:32:26:36, of pereopod 7 = pod 1 peduncle with 4 apicolateral robust setae, 53:14:13:15; Pereopod 7 basis of reaching apex of and 5 basofacial slender setae, medially with merus, moderately setose ventrally. Pleopods 1-3 many marginal setae plus apical enlarged robust with 2 coupling hooks; pleopod 1 with 5 distal seta; peduncle of uropod 2 with 9 dorsal robust facial setae, pleopod 2 with 2 distal facial setae, setae; apicolateral corners of peduncles on pleopod 3 with 2 mid and 5 distal facial setae; uropods 1–2 without comb. Uropod 3 unreduced, articles on outer rami = 14-13-15, inner rami = outer ramus longer than peduncle. Uropod 3 10-8-10. peduncle with 5 ventral robust setae, dorsally Epimeron 1 posteroventral corner rounded, with 1 lateral seta; rami feminine, inner extending anteroventral margin with 5 setae, posteroventral to M. 41 on article 1 of outer ramus, apex with 2 face with 3 medium setae; epimeron 2 pos- setae, medial and lateral margins naked, article 2 teroventral corner rounded, with 7 facial setae; of outer ramus elongate, 0.26, bearing 2 long epimeron 3 posteroventral corner with small to setae, apicomedial margin of article 1 with 2 medium tooth, posterior margin almost straight setae, lateral margin with 3 acclivities, robust 278 J. TAYLOR AND C. B. POORE setal formula = 1-1-1-0, slender setal formula = 1- The species is number MoV3667 in Museum Vic- 1-1-1. Telson, length-width ratio = 27:26, not toria’s TAXA database. fully cleft, each apex wide, rounded, faintly setose, lateral acclivity broad, shallow, bearing Birubius wallisae sp. nov. ordinary lateral setule, robust setae next medial shorter than setule, midlateral setules diverse. Figures 18–24 Material examined. Holotype. Australia, Queensland, N Description of male. Similar to female but eyes entrance to Moreton Bay (27°02.85´S, 153°20.11É), larger. Antenna 1 like female but with dense 19 m, Smith-McIntyre grab, G.C.B. Poore, 16 Mar medial setation on peduncular article 1; primary 1998 (stn ASB/6/24), NMV J47236 (1 female, tl. 5.0 flagellum bearing calceoli. Antenna 2 elongate, mm). Allotype. Australia, Queensland, Middle Banks, peduncular articles 3–4 with dense dorsal seta- Moreton Bay (27°15´S, 153°15É), 18 November 1976 tion, peduncular article 5 about as long as article (stn 6), QM W25241 (1 male, tl 4.8 mm). 4, dorsal margin lacking calceoli bearing 2 groups Paratypes. Australia, Queensland, N entrance to of male setae, flagellum 28–articulate bearing cal- Moreton Bay, all Smith-McIntyre grab, G.C.B. Poore, ceoli. Maxilliped and maxillae 1–2 similar to 16 Mar 1998: 27°02.94´S, 153°20.04É, 24 m, (stn female. Right mandible damaged; left lacinia ASB/6/21), NMV J47237 (3 females, tl. 3.45–4.45 mobilis with 6 spines; left rakers 9, left molar mm); 27°02.34´S, 153°19.42É, 11 m, (stn ASB/6/26), with 7 long robust setae, palp similar to female, NMV J41725 (1 female, tl. 4.10 mm); 27°02.34´S, article 3 with 1 basofacial seta. Main ventral setae 153°19.47É, 13 m, (stn ASB/6/28), NMV J41726 (1 female, tl. 3.0 mm). Queensland, Middle Banks, More- of coxae 1–4 = 5-7-6-0, Gnathopods 1–2 similar ton Bay (27°15´S, 153°15É), 18 Nov 1976 (stn 6), QM to female. Urosomite 3 without large hook dor- W8682 (12 females, tl. 3.10–4.75 mm, 9 males, tl. sally. Uropod 1 outer ramus with 3 dorsal robust 4.10–4.80 mm). setae, inner with 1 dorsomedial robust seta, uropod 2 outer ramus with 3 dorsal robust setae, Diagnosis. Rostrum constricted. Antenna 2, inner with 1 dorsal robust seta; uropod 1 peduncle article 4 without well developed dorsal setation. with 3 apicolateral robust setae, with 3 basofacial Right lacinia mobilis bifid, distal branch simple. slender setae; uropod 2 peduncle with 8 dorsal Pereopods 3–4 carpus with 2–3 proximoposterior robust setae. Uropod 3 with inner ramus falling robust setae. Pereopod 5 dactyl fully formed. short of article 1 on outer ramus. Telson elongate, Pereopod 7 basis without long ventral setae. Coxa length-width ratio = 14:13. 1 not expanded distally. Coxa 4 lacking long ventral setae. Epimeron 3 without tooth; with ventral Etymology. For Dr Jim Lowry, in gratitude for his setae; without long posterior seta; with oblique assistance and advice during the first author’s row of facial setae. Urosomite 3 with large dorsal visit to examine Australia Museum collections. hook. Uropod 1 with basofacial setae. Uropods Remarks. The following variations from the holo- 1–2 inner rami lacking accessory apical nails. type were observed in the paratypes and material Uropod 3 unreduced, outer ramus longer than examined. The main ventral setae of coxae 1–4 = peduncle. (5-7)-(5-7)-(5-7)-0. Uropod 1 outer ramus with Description of female. Head about 16% of total 3–4 dorsal robust setae, inner ramus with 1 dorsal body length, greatest width about 100% of length; robust seta. Uropod 2 outer ramus with 2–3 dor- rostrum constricted, narrow, exceeding apex of sal robust setae, inner ramus with 1 dorsal robust peduncular article 1 on antenna 1. Eyes large, seta. occluded with pigment. Antenna 1 peduncular Birubius lowryi conforms well to Barnard and article 1 about 1.3 times as long as wide, about 2.1 Drummond’s (1978) diagnosis of Birubius except times as wide as peduncular article 2, ventral that it exhibits a dorsal hook on urosomite 3 as margin with 8 setules, unproduced dorsal apex seen in Tickalerus and Kulgaphoxus. It varies with 1 setule; peduncular article 2 about 0.75 from these genera in the lack of a shortened outer times as long as peduncular article 1, with 6 ven- ramus of uropod 3. It differs from the other new tral setae; primary flagellum with 12 articles, species described herein by the combination of about 0.6 times as long as peduncle, lacking characters listed in the diagnoses. It can be distin- aesthetascs; accessory flagellum with 10 articles. guished from B. wilsoni, the only other species Antenna 2, peduncular article 4 robust setae for- described from Papua New Guinea, by the mula = 1-2-4-5, dorsal margin with notch bearing absence of posterior setae on coxae 1–3, long ven- 1 robust seta and 1 slender seta, ventral margin tral setae on coxa 4 and the presence of proximo- with 7–8 groups of 1–2 long to short setae, with posterior setae on the carpus of pereopods 3–4. ventrodistal robust seta; peduncular article 5 NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 279

Figure 18. Birubius wallisae sp. nov., holotype female, tl. 5.00 mm (m = male allotype, 4.80 mm). 280 J. TAYLOR AND C. B. POORE

Figure 19. Birubius wallisae sp. nov., holotype female, tl. 5.00 mm. NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 281

Figure 20. Birubius wallisae sp. nov., holotype female, tl. 5.00 mm. about 0.7 times as long as peduncular article 4, right incisor with 3 teeth; left incisor with 2 facial robust seta formula = 0–2, dorsal margin humps in 2 branches; right lacinia mobilis bifid, naked, ventral margin with 3 sets of 1–3 long to distal branch shorter than proximal branch, sim- short setae, 2–3 ventrodistal long to medium ple, pointed, proximal branch simple, pointed; left robust setae; flagellum 1.15 times as long as lacinia mobilis with 5–6 teeth; right raker 8; left peduncular articles 4–5 combined, with 14 rakers 9; molar in form of short protrusion demar- articles. Mandibles with medium palpar hump; cated mainly by robust setae, right molar with 10 282 J. TAYLOR AND C. B. POORE

Figure 21. Birubius wallisae sp. nov., holotype female, tl. 5.00 mm. NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 283

Figure 22. Birubius wallisae sp. nov., holotype female, tl. 5.00 mm. long robust setae, left molar with 8 long robust shorter apicomedial seta plus 2 shorter apical setae, no seta disjunct; palp article 1 short, article setae; palp article 2 with 1 apicomedial marginal 2 with 1 long inner apical seta, and 2 other robust seta, 3 apicomedial setae and 3 sub- medium inner setae, article 3 about 0.91 times marginal setae. Maxilla 2 inner and outer plates long as article 2, apex oblique with 11 robust to extended equally. Maxilliped inner plates with 1 slender setae, without basofacial setae. Maxilla 1 large thick apical robust seta, 5 apicofacial setae, inner plate narrow, bearing 1 long apical seta, 1 1 medial seta; outer plate with 6 medial and api- 284 1. TAYLOR AND C. B. POORE

Figure 23. Birllbills wallisae sp. nov., holotype female, tl. 5.00 mm. cal robust setae; palp article I with I apicolateral 2 accessory setules. Coxa I unexpanded distally; seta, article 2 with 2 apicolateral setae and 2 other main ventral setae of coxae 1-4 = 9-8-7-0, pos lateral seta, article 3 slightly protuberant, with 7 teriormost seta of coxae 1-3 shortened; anterior facial setae, nail of article 4 medium length, with and posterior margins of coxa 4 divergent, pos- NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 285

Figure 24. Birubius wallisae sp. nov., allotype male, tl. 4.80 mm. terior margin oblique, posterodorsal corner 12:11 and 1:1; palmar humps ordinary, palms rounded, posterodorsal margin medium, straight, oblique; gnathopods 1–2 carpus elongate. Pere- width-length ratio of coxa 4 almost = 5:6. Long opods 3–4 similar, facial setae on merus = 7 and posterior setae on basis of gnathopods 1–2 and 5, on carpus = 5 and 7; main spine of carpus pereopods 3–4 = 6-10-11-10, short posteriors = extending to M. 100+ on propodus, carpus with 1 0-0-0-0 , long anteriors = 6-6-0-0, short anteriors proximoposterior robust seta; robust setae for- = 2-1-4-2. mula of propodus = 6 + 7; acclivity on inner Gnathopods, width ratios of carpus-propodus margin of dactyls of pereopods 3–4 weak, mid- on gnathopods 1–2 = 8:11 and 2:3, length ratios = facial seta short. Coxae 5–7 posteroventral seta 286 J. TAYLOR AND C. B. POORE formula = 6-3-4; merus-carpus of pereopods 5–6 peduncular articles 3–4 with dense dorsal seta- medium to narrow, facial robust setae rows tion, peduncular article 5 about as long as article poorly developed, facial ridge formula on basis of 4, dorsal margin bearing 1 calceolus and 2 groups pereopods 5–7 = 0-2-2, anterior ridge of pereopod of male setae, flagellum at least 19-articulate 7 short; width ratios of basis, merus, carpus, (both broken) bearing calceoli. Maxilliped and propodus of pereopod 5 = 26:28:25:11, of pereo- maxillae 1–2 similar to female. Right lacinia pod 6 = 41:31:19:8, of pereopod 7 = 50:15:12:6, mobilis bifid, distal branch much shorter than length ratios of pereopod 5 = 47:19:23:25, of proximal branch, simple; proximal branch simple, pereopod 6 = 27:15:13:16, of pereopod 7 = pointed; left lacinia mobilis with 5 teeth; right 60:16:15:18; pereopod 7 basis reaching apex of raker 6; left rakers 9; right molar with 4 long merus, with 1 medium ventral seta. Pleopods 1–3 robust setae, left molar with 6 long robust setae, with 2 coupling hooks; pleopod 1 with 3 mid and palp similar to female, article 3 with 2 basofacial 3 proximal facial setae, pleopod 2 with 2 proxi- setae. Main ventral setae of coxae 1–4 = 6-7-7-0. mal, 3 mid and 2 distal facial setae, pleopod 3 Gnathopods 1–2 similar to female. Urosomite 3 with 5 proximal and 5 distal facial setae; articles with large hook dorsally. Uropod 1 outer ramus on outer rami = 14-16-15, inner rami = 9-11-10. with 4 dorsal robust setae, inner with 1 dorso- Epimeron 1 posteroventral corner rounded, medial robust seta, uropod 2 outer ramus with 3 anteroventral margin with 3 setae, posteroventral dorsal robust setae, inner with 1 dorsal robust face with 2 medium to long setae; epimeron 2 seta; uropod 1 peduncle with 4 apicolateral robust posteroventral corner rounded, with 9 facial setae, setae, without basofacial slender seta; uropod 2 posteriormost pair set vertically; epimeron 3 pos- peduncle with 10 dorsal robust setae. Uropod 3 teroventral corner without tooth, posterior margin with inner ramus elongate, reaching article 1 on rounded, naked, midface with oblique row of 9 outer ramus. Telson elongate, length-width ratio setae near posterior margin, ventral margin with 6 = 11:9. long setae. Urosomite 1 naked, articulation line Etymology. For Dr Elycia Wallis, Museum Vic- barely present; urosomite 3 with large hook dor- toria, in appreciation of her support during the sally. Uropods 1–2 rami with articulate enlarged first author’s PhD candidature. apical nails, uropod 1 outer ramus with 6 dorsal robust setae, inner with 1 dorsomedial robust seta, Remarks. The following variations from the holo- uropod 2 outer ramus with 4 dorsal robust setae, type were observed in the paratypes and material inner with 1 robust seta; uropod 1 peduncle with examined. The main ventral setae of coxae 1–4 = 3 apicolateral robust setae, 3 smaller setae, and 3 (5-8)-(4-8)-(5-8)-0. Uropod 1 outer ramus with basofacial slender setae, medially with few 2–5 dorsal robust setae, inner ramus with 1 dorsal marginal setae plus apical enlarged robust seta; robust seta. Uropod 2 outer ramus with 1–3 dor- uropod 2 peduncle with 9 dorsal robust setae; api- sal robust setae, inner ramus with 1 dorsal robust colateral corners of peduncles on uropods 1–2 seta. without comb. Uropod 3 unreduced, outer ramus Birubius wallisae conforms well to Barnard longer than peduncle. Uropod 3 peduncle with 5 and Drummond’s (1978) diagnosis of Birubius ventral robust setae, dorsally with 1 lateral seta; except that it exhibits a dorsal hook on urosomite rami feminine, inner extending to M. 76 on article 3 as seen in Tickalerus and Kulgaphoxus. It 1 of outer ramus, apex with 2 setae, medial differs from these genera in the lack of a short- margin naked, lateral margin with 3 long setae, ened outer ramus on uropod 3 but shares with article 2 of outer ramus short, 0.15, bearing 2 long Kulgaphoxus the proximal vs widely spread setae, apicomedial margin of article 1 with 2 placement of ventral setae on antennae 1 pedun- setae, lateral margin with 4 acclivities, robust cular article 2. Birubius wallisae differs from the setal formula = 1-1-1-1-1, slender setal formula = other new species in the combination of char- 1-1-1-1-1. Telson, length-width ratio = 1:1, not acters listed in the diagnoses. The species is fully cleft, each apex wide, rounded, lateral number MoV3716 in Museum Victoria’s TAXA acclivity broad, shallow, bearing ordinary lateral database. setule, robust setae next medial shorter than setule, single midlateral setule. Birubius wilsoni sp. nov. Figures 25–30 Description of male. Similar to female but eyes larger. Antenna 1 like female but with dense Material examined. Holotype. Papua New Guinea, NW corner of Pig I. (05°9.98´S, 145°50.45´E), 21 m, J. D. medial setation on peduncular article 1; primary Thomas, 4 Feb 1990 (stn PNG 33K), AM P56149 (1 flagellum bearing calceoli. Antenna 2 elongate, female, tl. 4.20 mm). NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 287

Figure 25. Birubius wilsoni sp. nov., holotype female, tl. 4.20 mm (m = male allotype, 3.55 mm). 288 J. TAYLOR AND C. B. POORE

Figure 26. Birubius wilsoni sp. nov., holotype female, tl. 4.20 mm. NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 289

Figure 27. Birubius wilsoni sp. nov., holotype female, tl. 4.20 mm (m = male allotype, 3.55 mm). 290 J. TAYLOR AND C. B. POORE

Figure 28. Birubius wilsoni sp. nov., holotype female, tl. 4.20 mm (m = male allotype, 3.55 mm). NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 291

Figure 29. Birubius wilsoni sp. nov., holotype female, tl. 4.20 mm (m = male allotype, 3.55 mm). 292 J. TAYLOR AND C. B. POORE

Figure 30. Birubius wilsoni sp. nov., holotype female, tl. 4.20 mm (m = male allotype, 3.55 mm).

Allotype. Papua New Guinea, Madang (05°9.57´S, ventral setae. Epimeron 3 with small tooth; with 145°59.93´E), 4 m, J. D. Thomas, 20 Feb 1990 (stn ventral setae; without long posterior seta; without PNG 54), AM P56150 (1 male, tl. 3.55 mm). facial setae. Urosomite 3 with small dorsal hook. Paratypes. Same data as holotype, AM P60002 (1 Uropod 1 with basofacial setae. Uropods 1–2 male, tl. 3.7 mm). Papua New Guinea, Madang (05°9.57´S, 145°59.93´E), 4 m, J. D. Thomas, 20 Feb inner rami lacking accessory apical nails. Uropod 1990 (stn PNG 54), AM P60003 (2 females, tl. 4.2–4.8 3 unreduced, outer ramus longer than peduncle. mm, 53 males, tl. 2.7–3.3 mm). Description of female. Head about 19% of total Diagnosis. Rostrum constricted. Antenna 2, arti- body length, greatest width about equal to length; cle 4 without well developed dorsal setation. rostrum constricted, narrow, elongate, reaching Right lacinia mobilis bifid, distal branch simple. middle of peduncular article 2 on antenna 1. Eyes Pereopods 3–4 carpus without proximoposterior medium, pigmented. Antenna 1 peduncular robust setae. Pereopod 5 dactyl fully formed. article 1 about 1.6 times as long as wide, about 2.1 Pereopod 7 basis with long ventral setae. Coxa 1 times as wide as peduncular article 2, ventral strongly expanded distally. Coxa 4 with long margin with 9 setules, unproduced dorsal apex NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 293 without setule; peduncular article 2 about 0.66 length. Pereopods 3–4 similar, facial setae on times as long as peduncular article 1, with 9 ven- merus = 5 and 5, on carpus = 6 and 5; main spine tral setae; primary flagellum with 9 articles, about of carpus extending to M. 87 on propodus, carpus 0.6 times as long as peduncle, lacking aesthetascs; without proximoposterior robust setae; robust accessory flagellum with 9 articles. Antenna 2, setae formula of propodus = 2 + 4 and 2 + 5; peduncular article 4 robust setae formula = 1-2-4- acclivity on inner margin of dactyls of pereopods 3, dorsal margin with notch bearing 3 setae, ven- 3–4 weak, midfacial seta short. Coxae 5–7 pos- tral margin with 7–8 groups of 1–2 long to short teroventral seta formula = 4-3-6; merus-carpus of setae, 1 long ventrodistal robust seta; peduncular pereopods 5–6 medium to narrow, facial robust article 5 about 0.68 times as long as peduncular setae rows poorly developed, facial ridge formula article 4, facial robust seta formula = 1–2, dorsal on basis of pereopods 5–7 = 0-2-2, anterior ridge margin naked, ventral margin with 3 sets of 1–2 of pereopod 7 long; width ratios of basis, merus, long to short setae, 2 ventrodistal long to medium carpus, propodus of pereopod 5 = 24:20:14:9, of robust setae; flagellum 0.85 times as long as pereopod 6 = 34:23:14:6, of pereopod 7 = peduncular articles 4–5 combined, with 9 articles. 50:13:9:5, length ratios of pereopod 5 = Mandibles with medium palpar hump; right 38:23:22:27, of pereopod 6 = 24:15:17:18, of incisor with 3 teeth; left incisor with 2 humps in 2 pereopod 7 = 58:17:17:18; basis of pereopod 7 branches; right lacinia mobilis bifid, distal branch reaching apex of merus, heavily setose ventrally. much shorter than proximal branch, simple, Pleopods 1–3 with 2 coupling hooks; pleopod 1 pointed, proximal branch simple, pointed; left with 4 distal facial setae, pleopod 2 with 2 distal lacinia mobilis with 4–5 teeth; right raker 6; left facial setae, pleopod 3 with 6 mid facial setae; rakers 8; molar in form of short protrusion demar- articles on outer rami = 14-13-15, inner rami = cated mainly by robust setae, right molar with 6 9-8-11. long robust setae, left molar with 6 long robust Epimeron 1 posteroventral corner rounded, setae, no seta disjunct; palp article 1 slightly elon- anteroventral margin with 4 setae, posteroventral gate, article 2 with 3 long-medium inner apical face with 3 medium setae; epimeron 2 pos- setae and 1 other medium inner seta, article 3 teroventral corner rounded, with 6 facial setae, about 0.93 times long as article 2, apex oblique posteriormost pair set vertically; epimeron 3 pos- with 8 robust to slender setae, without basofacial teroventral corner with small to medium tooth, setae. Maxilla 1 inner plate narrow, bearing 1 posterior margin almost straight with 2 medium long apical seta, 1 shorter apicomedial seta, 2 setae, ventral margin with 6 medium setae. Uro- apicolateral much shorter seta; palp article 2 with somite 1 naked, articulation line absent; uro- oneapicomedial marginal robust seta, 3 api- somite 3 with small hook dorsally. Uropods 1–2 comedial setae and 3 submarginal setae. Maxilla rami with articulate enlarged apical nails, uropod 2 inner plate shorter and broader than outer. 1 outer ramus with 4 dorsal robust setae, inner Maxilliped inner plate with 1 large thick apical with 1 dorsomedial robust seta, uropod 2 outer robust seta, 2 apicofacial setae, 3 medial setae; ramus with 2 dorsal robust setae, inner without outer plate with 5 medial and apical robust setae; robust seta; uropod 1 peduncle with 4 apicolateral palp articles 1 and 2 with 2 and 3 apicolateral seta, robust setae, and 4 basofacial slender setae, medi- article 3 unprotuberant, with 2 facial setae, nail of ally with many marginal setae plus apical article 4 long, with 2 accessory setules. Coxa 1 enlarged robust seta; uropod 2 peduncle with strongly expanded distally; main ventral setae of 7 dorsal robust setae; apicolateral corners of coxae 1–4 = 9-9-10-6, posteriormost seta of coxae peduncles on uropods 1–2 with comb. Uropod 3 1–2 medium, of coxa 3 elongate; anterior and unreduced, outer ramus longer than peduncle. posterior margins of coxa 4 strongly divergent, Uropod 3 peduncle with 6 ventral robust setae, posterior margin oblique, posterodorsal corner dorsally with 1 lateral seta; rami feminine, inner rounded, posterodorsal margin medium, undulent, extending to M. 46 on article 1 of outer ramus, width-length ratio of coxa 4 almost = 29:31. Long apex with 1 seta, medial and lateral margins posterior setae on basis of gnathopods 1–2 and naked, article 2 of outer ramus short, 0.23, bear- pereopods 3–4 = 3-8-8-8, short posteriors = ing 2 long setae, apicomedial margin of article 1 2-2-4-3 , long anteriors = 4-6-0-0, short anteriors with 2 setae, lateral margin with 4 acclivities, = 2-1-1-0. robust setal formula = 1-1-1-1-0, slender Gnathopods, width ratios of carpus-propodus setal formula = 1-1-1-1-1. Telson, length-width on gnathopods 1–2 = 21:24 and 18:25, length ratio = 55:54, not fully cleft, each apex wide, ratios = 8:11 and 8:13; palmar humps ordinary, rounded, setose, lateral acclivity broad, shallow, palms oblique; gnathopods 1–2 carpus of medium bearing ordinary lateral setule, robust setae next 294 J. TAYLOR AND C. B. POORE medial shorter than setule, midlateral setules Discussion diverse. Birubius is by far the largest genus of phoxo- Description of male. Similar to female but eyes cephalid amphipods, now comprising 38 species larger. Antenna 1 like female but with dense from Australia (Barnard and Drummond, 1978), medial setation on peduncular article 1; primary three species from Indonesia (Dana, 1853, Ortiz flagellum bearing calceoli. Antenna 2 elongate, and Lalana, 1997, 1999) and two species from peduncular articles 3–4 with dense dorsal seta- Papua New Guinea. Species of Birubius occur tion, peduncular article 5 about as long as article intertidally to 70 m in benthic sandy to muddy 4, dorsal margin lacking calceoli, bearing 2 sediments. The new species expand the depth and groups of male setae, flagellum 24-articulate geographic range of the genus from that pre- bearing calceoli. Maxilliped and maxillae 1–2 viously known. The biogeographic relationships similar to female. Right mandible damaged, left between the species from Australian waters and lacinia mobilis with 5 spines; left rakers 7; molar those from Papua New Guinea and Indonesia are in form of bulbous hump, left molar with 4 long as yet unknown but could be elucidated only by robust setae, palp similar to female, article 3 with cladistic analysis of species of Birubius, 1 basofacial seta. Main ventral setae of coxae 1–4 Kulgaphoxus, Tickalerus and Yan. = 8-7-8-5, Gnathopods 1–2 similar to female. Pereopod 7 basis narrower than female. Uro- Acknowledgements somite 3 without hook dorsally Uropod 1 outer ramus with 3 dorsal robust setae, inner with 1 dor- We acknowledge the assistance and support sal robust seta, uropod 2 outer ramus with 2 dor- throughout the first author’s PhD candidature of sal robust setae, inner without robust seta; uropod those people named in the etymologies. Examina- 1 peduncle with 2 apicolateral robust setae, with 4 tion of Australian Museum collections was made basofacial slender seta; uropod 2 peduncle with 9 possible through financial assistance awarded to dorsal robust setae. Uropod 3 with inner ramus the first author in an Australian Museum Post- falling short of article 1 on outer ramus. Telson graduate Award. This project was supported by elongate, length-width ratio = 6:5. an Australian Research Council grant to G. C. B. Poore and R. S. Wilson. Etymology. For Dr Robin Wilson, Museum Vic- toria, a good friend and colleague of both authors. References Remarks. The following variations from the holo- Alonso de Pina, 1993. Linca pinata, a new phoxo- type were observed in the paratypes. The main cephalid genus and species (Crustacea: ventral setae of coxae 1–4 = (7-9)-(7-9)-(8-11)- Amphipoda) from the Argentine continental shelf. (6-9). Uropod 1 outer ramus with 1–4 dorsal Proceedings of the Biological Society of Washing- robust setae, inner ramus with 1 dorsal robust ton 106(3): 497–507. seta. Uropod 2 outer ramus with 2 dorsal robust Barnard, J.L., 1960. The amphipod family Phoxo- setae, inner ramus without dorsal robust setae. cephalidae in the eastern Pacific Ocean, with anal- Birubius wilsoni conforms well to Barnard and yses of other species and notes for a revision of the Drummond’s (1978) diagnosis of Birubius except family. Allan Hancock Pacific Expeditions 18: 175–368. that it exhibits a dorsal hook on urosomite 3 as Barnard, J.L., 1979. Littoral gammaridean Amphipoda seen in Tickalerus and Kulgaphoxus. It varies from the Gulf of California and the Galapagos from these genera in the lack of a shortened outer Islands. Smithsonian Contributions to Zoology ramus on uropod 3 but shares with Kulgaphoxus 271: vi + 149 pp, 74 figs. the proximal vs widely spread placement of ven- Barnard, J.L. and Drummond, M.M., 1976. Clarifica- tral setae on antennae 1 peduncular article 2. Biru- tion of five genera of the Phoxocephalidae (marine bius wilsoni differs from the other new species Amphipoda). Proceedings of the Biological Soci- described herein by the combination of characters ety of Washington 88: 515–547. listed in the diagnoses. It can be distinguished Barnard, J.L. and Drummond, M.M., 1978. Gam- maridean Amphipoda of Australia, Part III: The from B. lowryi, the only other species described Phoxocephalidae. Smithsonian Contributions to from Papua New Guinea, in the presence of pos- Zoology 245: 1–551. terior setae on coxae 1–3, long ventral setae on Barnard, J.L. and Karaman, G.S., 1991. The families coxa 4 and the absence of proximoposterior setae and genera of marine gammaridean Amphipoda on the carpus of pereopods 3–4. The species is (except marine gammaroids). Parts 1 and 2. number MoV3666 in Museum Victoria’s TAXA Records of the Australian Museum, Supplement database. 13: 1–866. NEW SPECIES OF PHOXOCEPHALID AMPHIPODS 295

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