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