.

PROC. BIOL. SOC. WASH. 104(2), 1991, pp. 253-268 REVISION OF IPHIMEDIIDAE AND SIMILAR FAMILIES (: )

C. O. Coleman and J. L. Barnard

Abstract.— The amphipod families Acanthonotozomatidae, Amathillopsi- dae, Astryidae, Epimeriidae (=Paramphithoidae), Iphimediidae, Ochlesidae and are rediagnosed and three new families, Acanthonotozomel- lidae, Dikwidae, and Odiidae are described. New keys distinguish these fam- ilies; other similar families, Lafystiidae and Laphystiopsidae are diagnosed and included in the keys.

In the process of rediagnosing the Och- more discretely from Iphimediidae, without lesidae Stebbing, 1910 (see Coleman & resorting to the difficult judgment about the Barnard 1991a), as a family distinct from conical or quadrate shapes of the mouthpart Iphimediidae, we determined that other al- group. Within the Iphimediidae two prin- terations to the of the Iphime- cipal groups remain: those genera similar to diidae and Epimeriidae were necessary. Our Iphimedia and those genera similar to Acan- contribution here offers a different division thonotozoma. These two groups are distin- than used in the past based on characters guished and the Acanthonotozomatidae are not previously used or inconsistently used revived. The new family Acanthonotozo- to define these groups. For example, we mellidae are split away from Acanthono- found relatively good consistency of char- tozomatidae. acters on incisors, rakers, gnathopods and coxae in proposing this new division of the Methods taxa. All diagnoses presented herein follow the Discussion format of comparison to the model amphi- pod known as the "basic gammaridean" By using characters other than reduction pubUshed by Barnard (1969:50, figs. 1, 2). of maxillipedal palp, we are able to tighten The formula for setae on the mandibular the diagnosis of Ochlesidae, but this re- palp was invented by Stock (1974). Two quires, at the same time, the removal of the keys with different starting points are pre- genera and Postodius from the Iphi- sented. To account for possible differing in- mediidae to form the contents ofa new fam- terpretations of certain characters some taxa ily Odiidae, the removal of the peculiar Di- are entered twice in the keys. For example, kwa from its paramphithoid-iphimediid Dikwidae are entered twice to treat the al- associations to form a unique family Dikwi- ternatives of coxa 1 acuminate and tapering, dae, and the reinstatement of the Amathil- or subtruncate and not tapering. lopsidae (an old family synonymized with Epimeriidae some years ago). The genera Key I to the Families of the Parepimeria Chevreux (1911) and Parepi- Iphimediidae and Similar Taxa meriella Schellenberg (1931) are removed to the Pleustidae. By removing the above 1 Mandibular rakers present 2 contents from the two families, the diag- - Mandibular rakers absent 10

nosis of Epimeriidae becomes strengthened 2. Coxa 1 truncate, not pointed or ta- and these amphipods can be distinguished pering 3 .

254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

- Coxa 1 tapering or pointed 7 - Carpus of gnathopod 2 not pro-

3. Gnathopod 1 propodochelate, duced, molar absent head deformed Dikwidae Acanthonotozomatidae

- Gnathopod 1 not propodochelate 4 Key 2 to the Families of the Iphimediidae and Similar Taxa 4. Molar triturative .. . Amathillopsidae

- Molar not triturative, or absent . . 5 1. Gnathopod 1 propodochelate .... 2

5. Neither molar present nor incisor - Gnathopod 1 subchelate or simple

broadened . . Acanthonotozomellidae 4 - Either molar present or incisor 2. Gnathopod 2 enlarged, subchelate, broadened 6 carpus lobate, (molar triturative, 6. Small setose molar present, inci- rakers present) Odiidae sors narrow Astryidae - Gnathopod 2 feeble, propodoche- - Molar absent, incisors very broad late or simple, carpus not lobate 3 certain Stilipedidae 3. Rakers present, gnathopod 2 sim- 7. Gnathopod 2 much broader than ple Dikwidae

gnathopod 1 , carpus lobate . . Odiidae - Rakers absent, gnathopod 2 pro- - Gnathopod 2 almost as slender as podochelate Iphimediidae

gnathopod 1 , carpus not lobate 8 4. Carpus of gnathopod 2 produced

8. Gnathopod 1 propodochelate, Ochlesidae head deformed and reduced .... - Carpus of gnathopod 2 not pro- Dikwidae duced 5

- Gnathopod 1 simple, head formed 5. Coxa 1 tapering or pointed 6

normally 9 - Coxa 1 not tapering or pointed . . 8 9. Molar absent 6. Molar produced, triturative or se- Acanthonotozomellidae tose Epimeriidae - Molar present Epimeriidae - Molar reduced or absent, not tritu-

10. Coxa 1 truncate below 11 rative or setose 7

- Coxa 1 tapering below 14 7. Rakers absent 11. Gnathopods 1-2 propodochelate Acanthonotozomatidae Iphimediidae - Rakers present - Gnathopods 1-2 not propodoche- Acanthonotozomellidae late 12 8. Incisors extremely broad and flat 12. Body and incisors ofmandible very Stilipedidae broad and flat Stilipedidae - Incisors of ordinary width and not - Body and incisors of mandible or- extremely flat 9 dinary 13 9. Rakers present 10 13. Gnathopod 2 subchelate, molar - Rakers absent 11

absent, coxa 4 not reduced, point- 1 0. Molar only setose, gnathopods fee- ed Lafystiidae ble, propodus smaller than carpus, - Gnathopod 2 simple, molar pres- rectangular or ovate Astryidae ent, all coxae 1-4 reduced, trun- - Molar triturative, gnathopods en- cate Laphystiopsidae larged, propodus larger than car-

14. Gnathopod 2 propodochelate . . pus, almost almond-shaped Iphimediidae Amathillopsidae - Gnathopod 2 not propodochelate - Molar absent, gnathopods feeble, 15 propodus dominated by carpus, 15. Carpus of gnathopod 2 produced, subrectangular molar present Ochlesidae Acanthonotozomellidae VOLUME 104, NUMBER 2 255

1 1 . Gnathopod 2 subchelate, molar least slightly shorter than 4. Gnathopods of absent, coxa 4 not reduced, taper- moderate size and with short articles 5-6, ing Lafystiidae not flagellar, gnathopod 2 usually slightly

- Gnathopod 2 simple, molar pres- longer than gnathopod 1; gnathopods 1-2 ent, all coxae 1-4 reduced, trun- propodosubchelate, carpi lobed, propodi cate Laphystiopsidae enlarged and almost almond-shaped. Arti- cle 2 of pereopods 5-7 occasionally with Special Couplets Distinguishing posterior cusps or teeth. Epimeron 3 lacking Ochlesidae and Odiidae 2 large cusps. Urosomites free. Uropods 1- 3 biramous. Rami of uropod 3 flattened, A. Pereonites 1-6 flat and keel-like lanceolate, 1 -articulate. Telson entire or dorsally, flagella of both antennae weakly incised, generally not longer than 1-2 reduced, merus and carpus of peduncle of uropod 3. gnathopod 2 produced, palp of Relationship.— T>\^Qnng from the Epi- maxilla 1 reduced, inner plate of meriidae in the weaker rostrum, slightly maxilla 1 reduced B shortened and unpointed coxa 1 , lobate car- B. Gnathopod 1 simple, gnathopod 2 pi and enlarged almond-shaped propodi of simple, coxae 1 and 4 shorter than gnathopods, and presence of large inner coxae 2-3 Ochlesidae lobes on the lower lip. - 1 propodochelate, Gnathopod List of genera.— Amathillopsis Heller, coxae 1 gnathopod 2 subchelate, and 1875 (=Acanthopleustes Holmes, 1908). 4 ordinary Odiidae Epimeriidae Amathillopsidae Epimerinae [sic] Boeck, 1871:183. Amathillopsidae Pirlot, 1934:201. Paramphithoidae Stebbing, 1906:320.

Diagnosis.— Body compressed. Rostrum Diagnosis. —Body compressed, with dor- of medium size or very small (not "well sal teeth sometimes confined to metasome, developed"). Antennae elongate, flagella or urosome, and sometimes weak (Epimeri- with 5+ articles; accessory flagellum 1-2 ella). Rostrum well developed. Antennae articulate. Mouthpart part field quadrately elongate, flagella with 5 + articles; accessory developed (box-like). Epistome and labrum flagellum 0-1 articulate. Mouthpart part broad, short, entire. Incisor of mandible or- field quadrately developed (box-like). Ep- dinary, toothed; raker row strong; molar istome and labrum broad, short, incised or large and triturative, palp always present, entire. Incisor of mandible ordinary, 3-articulate. Lower lip with large fleshy in- toothed; raker row strong; molar large and ner lobes, without distinct inner notches. triturative or reduced, conical and pubes-

Inner plate of maxilla 1 ordinary, apically cent {Epimeriella and Epimeria victoria and somewhat medially setose; outer plate Hurley, 1957); palp always present, 3-artic- oblique, normally spinose; palp large, 2-ar- ulate. Lower lip without inner lobes, with- ticulate. Inner plate of maxilla 2 without out distinct inner notches. Inner plate of

facial setae. Palp of maxilliped 4-articulate, maxilla 1 ordinary, medially setose (except article 2 not produced medially. Coxae 2- Uschakoviella); outer plate oblique, nor- 3 more or less acuminate, ventral margins mally spinose; palp large, 2-articulate. Inner fitting normal ventral parabolic curve of an- plate of maxilla 2 without facial setae. Palp

terior coxae; coxa 1 but not 4 shortened, of maxilliped 3-4-articulate, article 2 not

coxa 1 scarcely to not acuminate, mostly produced medially. Coxae \-^ acuminate, truncate, coxa 4 not strictly acuminate, with ventral margins fitting normal ventral par-

weak or no posteroventral lobe, coxa 5 at abolic curve of anterior coxae; coxae 1 and ;

256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

4 not shortened, coxa 4 with large postero- gellum. Most Pardaliscidae have evenly ex- ventral lobe, coxa 5 at least slightly shorter tending coxae 1-7, there being little differ- than 4. Gnathopods feeble and with elon- ence in the ventral extension of these plates. gate articles 5-6 but not flagellar, gnathopod The Stilipedidae have an expanded, non-

2 usually slightly longer than gnathopod 1 acuminate coxa 1, and foliaceous mandi- gnathopods 1-2 propodosubchelate, or sim- bles. The Astryidae have a lower lip with ple, merus and carpus not produced. Article widely separated outer lobes and the 2 ofpereopods 5-7 often with posterior cusps gnathopods have a characteristic pattern, the or teeth. Epimeron 3 lacking 2 large cusps carpus being weakly lobate but much wider (though occasional with very small than the thinner, simple propodus. The Sti- epimeral bicuspidation), occasionally body lipedidae have an exaggerated version of this or coxae with surficial cusps or rarely artic- kind of gnathopod but the carpus is not so ulate spines (Uschakoviella). Urosomites strongly lobate as it is strongly enlarged. free. Uropods 1-3 biramous. Rami of uro- Stegocephalidae lack a mandibular palp, pod 3 longer than peduncle, flattened, lan- Dexaminidae have fusion of certain uro- ceolate, usually 1 -articulate. Telson entire somites and normal maxillipeds. or weakly incised, generally not longer than Pleustidae have characteristic lower lips. peduncle of uropod 3. Gammaridans with acuminate coxae have Relationship.— Difkring from the Acan- a large accessory flagellum and normal max- thonotozomatidae in the box-like field of illipeds. mouthparts and the well developed man- Within Pontoporeiidae Priscillina also dibular molar and the non-flagellar gnatho- bears acuminate coxae but differs otherwise pod 1. from Epimeriidae in the multiarticulate ac- Differing from the Odiidae in the well de- cessory flagellum and short inner ramus of veloped flagella of antenna 1 , well devel- uropod 3 (gammaroid facies). oped medial setae on the first maxillae, ab- Rhachotropis is distinguished from Epi- sence of distinct medial notches on the main meriidae (except Epimeria yaquinae) in the lobes of the lower lip, strongly acuminate large gnathopods with expanded propodus coxae 1-3, and well developed (versus re- and short deeply lobate carpus; the telson duced) D-E setae on the mandibular palp is elongate. Oradarea in the Eusiridae has (an accessory character not otherwise used an occasional species with incipiently acu- in diagnoses). minate coxae. Cleippides quadricuspis has The maxillipedal palp of Ochlesidae is 0- points on some anterior coxae, but no coxa 2 articulate, coxa 4 is shortened, the merus tapers to a single point downward. and carpus of gnathopod 2 is otherwise pro- Amathillopsidae appear similar to eusi- duced, and the pereon has an anterior keel, rid-calliopiid groups, but differ in the short except Meraldia which has a series of dorsal telson. The ratio of size between the plates plaques. and the palps of the maxilliped in Amathil- Differing from the Acanthonotozomelli- lopsidae are not typical for Epimeriidae (a dae in the presence of a molar. character not otherwise used in the diag- The Iphimediidae have chelate gnatho- noses). The shape of the gnathopods differs pods and have a severely reduced molar. strongly from all Epimeriidae. The Dikwidae have a chelate gnathopod Removals.— Parepimeria Chevreux

1, flagellar gnathopod 2, and reduced head. (1911) and Parepimeriella Schellenberg The Pardaliscidae have poorly developed (1931) are removed to the Pleustidae be- inner plates on the maxillipeds and one of cause of the condition of lower lips and the mandibles is heavily flattened; usually maxillipeds. pardaliscids have an elongate accessory fla- Eclysis and Epimeriella were removed to VOLUME 104, NUMBER 2 257 the Astryidae by Andres (1985). Walker the weakness of carpal expansion on the (1907) originally differentiated Epimeriella gnathopods. The latter character is a prob- from Epimeria on the lack of significant lem of context and needs much better doc- dorsal body cuspidation, lack of a point on umentation in the literature than heretofore coxa 5, the elongate pereopods 5-6, small- available. ness of rostrum, and the imperfect molar. List ofgenera. —Paramphithoe Bruzelius,

His two figures of coxa 1 on his plate 9 are 1859 (=Acanthosoma Ross, 1835, hom- slightly different views but show some ta- onym, Hemiptera) (=type ); Actina- pering. Gnathopod 1 is less typical of As- canthus Stebbing, 1888; Epimeria Costa, tryidae than of Epimeriidae. Andres (1985) 1851, 1851-1853 (=Pseudepimeria Chev- partially described some non-type speci- reux, 1912a, 1912b, =Subepimeria BeWan- mens of the type species. Epimeriella mac- Santini, 1972); Epimeriella Walker, 1906, ronyx, but no full analysis and figures of a 1907; Metepimeria Schellenberg, 1931; single specimen, nor comparison with types Uschakoviella Gurjanova, 1955. appears to be present in the literature. Gnathopods have not been reillustrated. Acanthonotozomellidae, new family Andres addresses dactylar lengths of per- eopod 5 but not overall lengths relative to Type genus. —Acanthonotozomella Schel- pereopod 7. Coxae 1-2 of the type species lenberg, 1926. are not illustrated but coxae 1-2 of Epi- Diagnosis. —Body compressed, with dor- meriella truncata show coxa 1 definitely sal teeth (except Acanthonotozomopsis). blunt distally but not as greatly and dispro- Rostrum well developed (except Acanthono- portionately widened as in species ofAstrya tozomopsis). Antennae elongate or short, and Stilipes. The gnathopodal carpi are flagella with 5 + articles, rarely with 2 (Ama- much less dominant than in species of ^45- tiquakius); accessory flagellum absent. trya and Stilipes, and scarcely differ from Mouthpart part field conical. Epistome and the classic Epimeria cornigera shown by Sars labrum narrow, long, incised. Incisor of (1895:pl. 128). mandible ordinary, toothed; raker row Epimeriella victoria (Hurley 1957; and see strong; molar reduced or absent; palp al-

Moore 1985) has a narrow pointed coxa 1, ways present, 3 -articulate. Lower lip with- unlike other supposed members of the ge- out inner lobes, without distinct inner nus (but coxa 1 of the type-species of Epi- notches. Inner plate of maxilla 1 ordinary, meriella is still not well documented). The medially setose or setation reduced; outer close congruency of mandibular molar of £". plate oblique, normally spinose; palp large, victoria to the other species of Epimeriella 2-articulate. Inner plate of maxilla 2 with- suggests there might be a close relationship; out facial or medial setae. Palp of maxilli- one may establish a transformation series ped 4-articulate, article 2 often produced between the least cuspidate species of Epi- medially. Coxae 2—4 more or less acumi- meriella and E. victoria to account for the nate, ventral margins fitting normal ventral very cuspidate conditions in the latter. For parabolic curve of anterior coxae or in type the moment we have removed E. victoria genus coxa 2 shortened; coxae 1 and 4 not to Epimeria. It fits into Epimeriidae as well shortened, coxa 1 widened in 2 genera but as it does into Astryidae; it differs from As- not significantly wider than coxa 2 (versus tryidae in its tapering coxa 1 and from Epi- Stilipedidae), subtruncate or rounded (ex- meriidae in the reduced molar and slightly cept Acanthonotozomoides with concave more opened outer lobes of the lower lip. It margin and anteroventral tooth), coxa 4 with fits Epimeriidae better because of the cus- large posteroventral lobe (except Acantho- pidation character of its body and coxae and notozomopsis and small in Amatiquakius). 258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Coxa 5 shorter than posteroventral lobe of separate Acanthonotozomellidae from 4. Gnathopods feeble, subequal; gnatho- Iphimediidae, because Pseudiphimediella pods 1-2 simple or weakly parachelate, me- bears the plesiomorphic mandibular con- rus and carpus not produced, carpi slender. ditions. Article 2 of pereopods 5-7 often with pos- Maxilliphimedia bears the fully plesio- terior cusps or teeth. Epimeron 3 often with morphic mandibular conditions and An- 2 large cusps posteroventrally. Urosomites chiphimedia intermediates between the ple- free. Uropods 1-3 biramous. Rami of uro- siomorphic state and the apomorphic frontal pod 3 longer than peduncle, flattened, lan- mandibular cutting plane (compare with ceolate, 1 -articulate. Telson entire or weak- Watling & Thurston 1989). These two gen- ly incised, generally not longer than peduncle era differ from Stilipedidae in the chelate

of uropod 3. gnathopods with otherwise completely dif- Relationship. —See the Epimeriidae, Odi- ferent form than in Stilipedidae and in the idae, Ochlesidae, for distinctions from those paired dorsal cusps typical of most iphi- families and see Epimeriidae for mention mediids. of families outside of the iphimedioid group Acanthonotozomellidae are similar to

of families included herein, which differ in Stilipedidae and Astryidae in that coxa 1 is the same way as from Acanthonotozomel- expanded and apically truncate or rounded lidae. in most species of Acanthonotozomellidae.

Differing from the Acanthonotozomati- For the most part coxa 1 of Acanthonoto- dae in the presence of rakers on the man- zomellidae is not broader than coxa 2 as in dible. Astryidae and Stilipedidae. Astryidae have Differing from the Iphimediidae in the a molar and Stilipedidae have broadened presence of rakers, the simple gnathopods mandibular bodies and incisors. The 1-2, and in the absence of medial setae on gnathopods of Acanthonotozomellidae are the inner plate of maxilla 2. subtly distinct from those of Astryidae and Differing from the Dikwidae in the ab- Stilipedidae where the carpus significantly sence of a molar, the normal head, simple dominates the propodus in its breadth.

gnathopod 1 , and the non-ffagellar gnatho- Composition. —Acanthonotozomella pod 2. Schellenberg, 1926 {=Paracanthonotozoma Acanthonotozomella and Acanthonoto- Bellan-Santini, 1972); Acantonotozomoides zomopsis share with Stilipedidae a slightly Schellenberg, 1931; Acanthonotozomopsis

expanded coxa 1 , and in most species coxae Watling & Holman, 1980; Amatiquakius 1-3 are not sharply pointed but rather trun- Coleman & Barnard, 1991b. cate or rounded. The Stilipedidae are dis- tinguished by the immensely broadened Acanthonotozomatidae Stebbing mandibles. Acanthonotozomatidae Stebbing, 1906:210. The mandibular incisors in Acanthono- tozomellidae move plesiomorphically in the Diagnosis.— ^ody compressed, with or "horizontal transverse plane of the body and without dorsal teeth. Rostrum well devel- in a plane perpendicular to the long axis of oped. Antennae elongate, flagella with 5 + the mandible" (Watling & Thurston 1989), articles; uniarticulate accessory flagellum whereas in the Iphimediidae there occurs present. Mouthpart field conical. Epistome gradual change of the cutting plane towards and labrum narrow, long, incised. Incisor the "vertical frontal plane of the body and of mandible narrow, toothed; raker row ab- in the medial-lateral plane through the long sent; molar reduced or absent; palp always axis of the mandible." This transition of the present, 3-articulate. Lower lip apically cutting plane, however, cannot be used to pointed, without inner lobes, without dis- VOLUME 104, NUMBER 2 259 tinct inner notches. Inner plate of maxilla cisor of mandible ordinary, toothed, or nee-

1 ordinary, medially setose; outer plate dle-shaped or blunt or scarcely toothed; oblique, normally spinose; palp large raker row absent; molar small and weakly (somewhat shortened in A. cristatum), triturative or reduced and simple; palp al- 2-articulate. Inner plate of maxilla 2 with- ways present, 3-articulate, D-E setae re- out facial setae. Palp of maxilliped 4-artic- duced. Lower lip without inner lobes, outer ulate, article 2 not produced medially. Cox- lobes very thin, with inner notches only at ae 1-4 acuminate, ventral margins fitting apices. Inner plate of maxilla 1 minute, of- even curve or straight line of anterior coxae; ten medially setose, setae usually very small; coxae 1 and 4 not shortened, coxa 4 with outer plate coniform, spinal margin very large posteroventral lobe, coxa 5 slightly oblique, spines few and mostly fused; palp shorter than 4. Gnathopod 1 feeble, slender, tiny, 1 -articulate. Inner plate of maxilla 2 articles 5-6 weakly elongate, gnathopods 1- with only terminal setae. Plates of maxilli- 2 propodosubchelate, merus and carpus not ped very thin, palp 0-2-articulate. Coxae 1- produced. Article 2 of pereopods 5-7 often 3 more or less acuminate, at least one coxa with posterior cusps or teeth. Epimeron 3 tapering, ventral margins not fitting normal with 2 large cusps posteromarginally (except ventral parabolic curve of anterior coxae;

Acanthonotozoma cristatum), occasionally coxae 1 and 4 shortened, coxa 3 usually with body or coxae with surficial cusps. Urosom- posterodorsal buttress, coxa 4 with peculiar ites free. Uropods 1-3 biramous. Rami of shape (see figures in Coleman and Barnard uropod 3 longer than peduncle, flattened, 1991a), with posteroventral lobe, coxa 5 lanceolate, 1 -articulate. Telson weakly in- about as long as 4. Gnathopods feeble and cised, generally not longer than peduncle of with elongate articles 5-6 (article 3 of uropod 3. gnathopod 1 mostly elongate, versus Odi-

Relationship. —The Epimeriidae differ in idae), gnathopod 1 weakly flagellar, gnatho- the well developed mandibular molar. See pod 2 usually slightly broader and shorter

Epimeriidae for mention ofdifferences from than gnathopod 1; gnathopods 1-2 simple, families outside of the families included merus and carpus of gnathopod 2 produced. herein. Article 2 of pereopods 5-7 lacking posterior See the Odiidae, Ochlesidae, Acanthono- cusps or teeth. Epimeron 3 rarely with 2 tozomellidae, Iphimediidae, and the Dikwi- large cusps. Urosomites free. Uropods 1-3 dae for distinctions from those families. biramous. Rami of uropod 3 subequal to or List ofgenera. —Acanthonotozoma Boeck, longer than peduncle, flattened, lanceolate, 1876 (=Acanthonotus Ross, 1835 [=hom- usually 1 -articulate. Telson entire, generally onym, Pisces], =Panoploeopsis Kunkel, not longer than peduncle of uropod 3. 1910). Relationship.— Differing from the Epi- meriidae in the thin dorsal keel or plaques Ochlesidae Stebbing on the pereon, short antennal ffagella, short

coxae 1 and 4, in the simple gnathopod 1 Ochlesidae Stebbing, 1910:581. and produced merus and carpus of gnatho- Diagnosis.— Body compressed, at least pod 2, reduction of D-E setae on the man- pereonites 1-6 projecting dorsalwards as thin dibular palp, reduction of medial maxillary ffat keel or formed in to ffat plaques {Meral- setae and loss of at least 2 articles on the dia). Rostrum well developed. Antennae maxillipedal palp. See Epimeriidae for poorly developed, short, flagella with 4 or mention of differences from families out- fewer articles; accessory ffagellum absent. side of the families included herein. Mouthpart field conically developed. Epi- Differing from the Odiidae in the reduc- stome and labrum narrow, long, entire. In- tion of coxae 1 and 4, absence of rakers, in .

260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the produced carpus of gnathopod 2 and the large and triturative; palp 3 -articulate (D- simple gnathopod 1 E setae well developed, versus Odiidae), Differing from the Acanthonotozomati- Lower lip without inner lobes, without inner dae in the presence of a molar, and from notches. Inner plate of maxilla 1 slender, that family and Acanthonotozomellidae in only apically setose; outer plate truncate, the reduction of the maxillipedal palp, re- with 13 spines; palp large, 2-articulate. In- duction of D-E setae on the mandibular ner plate of maxilla 2 [unknown]. Palp of palp, reduced coxae 1 and 4, and the pro- maxilliped 4-articulate, article 2 not pro- duced carpus of gnathopod 2. duced medially. Coxae 3-4 more or less acu- Differing from the Iphimediidae in the minate, coxa 2 shortened and blunt below, produced carpus of gnathopod 2, simple ventral margins not fitting normal ventral gnathopods 1-2, short coxae 1 and 4, keel- parabolic curve of anterior coxae; coxae 1 like dorsal margin of pereonites 5-6, and and 4 not shortened, coxa 1 truncate-con- the presence of a mandibular molar. cave below, with marginal points and an- Differing from the Dikwidae in the more teroventral lobule, coxa 4 with small pos- or less well developed head, short coxa 4, teroventral lobe, coxa 5 shorter than simple gnathopod 1, produced carpus of posteroventral lobe of 4. Gnathopods feeble gnathopod 2, conical outer plate of maxilla and with elongate articles 5-6 on gnathopod

1 , reduced palp of maxilliped, and the re- 2, and article 6 on gnathopod 1 gnathopod , duced D-E setae of the mandibular palp. 2 flagellar and much longer than gnathopod

1 Like the Ochlesidae, the Lafystiidae have 1 ; gnathopod propodochelate, gnathopod a 2 -articulate palp on the maxilliped; oth- 2 simple, merus and carpus not produced. erwise they differ from Ochlesidae in the Article 2 of pereopods 5-7 with posterior long coxa 4, normal lower lip, broad, un- cusps or teeth. Epimeron 3 lacking 2 large keeled body, parachelate gnathopod 2, lack cusps posteroventrally. Urosomites free. of meral and carpal lobe on gnathopod 2 Uropods 1-3 biramous. Rami of uropod 3 and well developed laciniae mobiles on both longer than peduncle, though peduncle mandibles. elongate, flattened, lanceolate, 1 -articulate. List of genera. — Ochlesis Stebbing, 1910 Telson weakly incised, not longer than pe- (=type genus); Curidia Thomas, 1983; Me- duncle of uropod 3. raldia Barnard & Karaman, 1987; Ochle- Relationship.— Uiffering from Epimeri- sodius Ledoyer, 1982. idae in the severely reduced head, chelate

gnathopod 1 , simple gnathopod 2, and elon- gate articles of the gnathopods. Dikwidae, new family Differing from Odiidae in the acuminate

Type genus. —Dikwa Griffiths, 1 974:266. coxae 3-4, simple gnathopod 2, conical out-

Diagnosis. —Body compressed, with thin er plate of maxilla 1, broad (ordinary) lower middorsal keel on pereonites 1-5, dorsal ca- lip, and broad prebuccal parts. rina on pereonites 6-7 and pleonites 1-3 Differing from Ochlesidae in the reduced

(based on new observation of material from head, chelate gnathopod 1, well developed

South African Museum). Head poorly and extraordinary maxilla 1, 4-articulate formed, rostrum absent (our interpretation maxillipedal palp, well developed antennal despite Griffiths 1977). Antennae elongate, flagella, and unshortened coxae 1 and 4. flagella with 5 + articles; accessory flagellum Differing from the Acanthonotozomati- absent. Mouthpart part field ?quadrately de- dae and Acanthonotozomellidae in having veloped (box-like). Epistome and labrum a mandibular molar, chelate gnathopod 1, broad, short, incised. Incisor of mandible flagellar gnathopod 2 and reduced head. ordinary, toothed; raker row strong; molar Differing from Iphimediidae in the sim- VOLUME 104, NUMBER 2 261 pie gnathopod 2, presence of mandibular somites free. Uropods 1-3 biramous. Rami rakers, triturative molar, normal lower lip, of uropod 3 longer than peduncle, flattened, and severely reduced head. lanceolate, 1 -articulate. Telson entire or See Epimeriidae for mention of differ- weakly incised, generally not longer than ences from families outside of the families peduncle of uropod 3. included herein. i^^mflrA:^.— Gnathopod 1 of Nodotergum List ofgenera.— Dikwa Griffiths, 1974. is simple, and because rakers and notches on lower lip are absent, Nodotergum might Iphimediiae be assigned to Acanthonotozomatidae, but the chelate gnathopod 2 and 3 -articulate Iphimedinae Boeck, 1871:178. maxillipedal palp suggest closer affinities to Diagnosis. —Body compressed, with dor- Iphimediidae than to Acanthonotozomati- sal teeth (except minutely in Iphimedia gib- dae. The outer plate of maxilla 1 in Nodo- ba and absent in Pariphimedia normani). tergum is reduced and poorly spinose. Max- Rostrum well developed. Antennae elon- illiphimedia and Paranchiphimedia differ gate, flagella with 5 + articles; accessory fla- from typical Iphimediidae in the untapered gellum 0-1 articulate. Mouthpart part field coxa 1 and broad, flattened mandible. conically developed. Epistome and labrum Relationship.— T>\f^Qr\ng from the Epi- highly variable, broad or narrow, short or meriidae in the absence of raker spines and long, incised or entire. Incisor of mandible the presence of at least one pair of chelate variable, broad and toothed in transition to gnathopods. needle shaped or blunt, untoothed, callused Differing from the Odiidae in the chelate or hollowed out; raker row absent; molar gnathopod 2, reduction or absence of man- absent or reduced {Stegopanoploea with dibular molar, and absence of rakers. conical molar); palp always present, 3-ar- The maxillipedal palp of Ochlesidae is 0- ticulate. Lower lip without inner lobes, with 2 articulate, the antennal flagella are re- or without distinct inner notches. Inner plate duced, coxae 1 and 4 are shortened, the ofmaxilla 1 ordinary or minute, usually me- propodi of the gnathopods are simple, and dially setose; outer plate oblique, normally gnathopod 2 otherwise has lobate merus and spinose or reduced (Nodotergum); palp large carpus. or small, 1-2-articulate. Inner plate of max- Differing from the Acanthonotozomati- illa 2 rarely with facial setae. Palp of max- dae in the presence of at least one pair of illiped 3-4-articulate, article 2 produced chelate gnathopods. medially in several genera. Coxae 2-4 more Differing from the Acanthonotozomelli- or less acuminate, ventral margins fitting dae in the presence of at least one pair of normal ventral parabolic curve of anterior chelate gnathopods and the absence of rak- coxae; coxae 1 and 4 not shortened, coxa 1 ers. variable, broad or narrow, acuminate or not, Differing from the Dikwidae in the che- coxa 4 with large posteroventral lobe, coxa late gnathopod 2, unreduced head and the 5 shorter than 4. Gnathopods feeble and reduced or absent mandibular molar and with elongate articles 5-6, gnathopod 1 of- rakers. ten flagellar, gnathopod 2 usually slightly Differing from Stilipedidae in the chelate longer or stouter than gnathopod 1 gnatho- gnathopod 2. ; pods 1-2 propodochelate, or gnathopod 1 See Epimeriidae for mention of differ- simple in Nodotergum, merus and carpus ences from families outside of the families not produced. Article 2 of pereopods 5-7 included herein. often with posterior cusps or teeth. Epi- Removals.— Bathypanoploea and its syn- meron 3 usually with 2 large cusps. Uro- onyms are removed to the Stilipedidae. ;

262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

List ofgenera.— Iphimedia Rathke, 1843 teroventral lobe, coxa 5 shorter than 4. (type genus) =Microcheles Kroyer, 1846, Gnathopod 1 feeble and with elongate ar- =Panoploea Thomson, 1880, =Iphime- ticles 5-6, weakly flagellar, gnathopod 2

fifzop5Z5 Delia Valle, 1893, =Cypsiphimedia much larger and broader than gnathopod 1

K. H. Barnard, 1955; Anchiphimedia K. H. gnathopod 1 propodochelate or with spine Barnard, 1930; Anisoiphimedia Karaman, forming chela; 2 propodosubchelate, merus 1980; Coboldus Krapp-Schickel, 1974; and carpus produced. Article 2 of pereo- Echiniphimedia K. H. Barnard, 1930; pods 5-7 with or without posterior cusps or Gnathiphimedia K. H. Barnard, 1930; Iphi- teeth. Epimeron 3 with 2 large cusps pos-

mediella Chevreux, 1911, 1 9 1 2b; Labriphi- teroventrally. Urosomites free. Uropods 1- media K. H. Barnard, 1931, 1932 (=Maor- 3 biramous. Rami of uropod 3 longer than iphimedia Hurley, 1954); Maxilliphimedia peduncle, flattened, lanceolate, usually K. H. Barnard, 1930; Nodotergum Bellan- 1 -articulate. Telson weakly incised, not lon- Santini, 1972; Paranchiphimedia RufFo, ger than peduncle of uropod 3. 1949; Parapanoploea Nicholls, 1938; Par- Remarks. —We have examined the types iphimedia Chevreux, 1906; Pseudiphime- of O. antarcticus which have a thin but weak diella Schellenberg, 1931; Stegopanoploea dorsal keel from pereonite 5 to pleonite 2. Karaman, 1980. Relationship.— Xyi^Qving from Epimeri- idae in the diverse gnathopods, with chelate

gnathopod 1 and produced merus and car- Odiidae, new family pus on gnathopod 2, conically arranged Type genus. — Odius Liljeborg, 1865:11. mouthparts and generally reduced subsid- Diagnosis.— ^ody compressed, all pere- iary features on mouthparts, the presence onites dorsally flush, in most species pro- of distinct notches or medial excavations on jecting dorsalwards as thin flat keel (see the lobes of the lower lip, and a flush dorsal "Remarks" below), teeth present only on keel on the pereon. pleon. Rostrum well developed. Antennae Odius has many of the same body for- poorly developed, short, flagella with 6-8 mations as found in Ochlesidae. For ex- short articles; accessory flagellum absent. ample, its basic members, albeit weakly in Mouthpart part field conically developed. O. antarcticus, have the compressed body Epistome and labrum narrow, elongate, mi- formed into a dorsal keel and have a dorsal nutely incised. Incisor of mandible narrow, tooth on pleonite 3 characteristic of prim- scarcely toothed; raker row strong; molar itive ochlesids (compare O. carinatus and small and triturative; palp present, 3 -artic- O. eridunda). The head is tall dorsoventrally ulate (D-E setae reduced, versus Dikwidae). and short anteroposteriorly and incipiently

Lower lip without inner lobes, outer lobes appears to be telescoped into pereonite 1. thin, with inner notches or excavations. In- The antennae are short and the flagella

ner plate of maxilla 1 small, apically with poorly articulate. The labrum is slender, 1-3 setules; outer plate oblique ("conical"), elongate and triangular. The mandibular normally spinose; palp 1 -articulate. Inner molar is strongly processed, and the number plate of maxilla 2 without facial or medial of D and E setae on the third palp article is setae. Palp of maxilliped 4-articulate, article reduced (a character not otherwise used in 2 not produced medially. Coxae 1-3 weakly diagnoses except for Odiidae and Ochlesi-

tapering but coxae 2-4 remaining quadrate dae). Odius, is, however, not an ochlesid below, ventral margins fitting normal ven- because articles 4—6 ofgnathopod 2 are short tral parabolic curve of anterior coxae; coxae and broad, and the propodus is broadly sub-

1 and 4 not shortened, coxa 3 lacking pos- chelate; article 3 of gnathopod 1 is not elon- terodorsal buttress, coxa 4 with large pos- gate and the propodus is cheliform; coxa 4 .

VOLUME 104, NUMBER 2 263

is fully developed and lobate as in Iphi- duced medially. Coxae 2-4 scarcely acu- mediidae; the maxillipedal palp has more minate, ventral margins fitting normal ven- than 2 articles; coxa 1 is not significantly tral parabolic curve of anterior coxae; coxae reduced in size. Differing from the Ochlesi- 1 and 4 not shortened, coxa 1 expanded, dae also in the truncate coxae 2-3 and pres- wider than coxa 2, truncate or rounded be- ence of cusps on article 2 of pereopods 5-7. low, coxa 4 with posteroventral lobe, coxa Differing from the Acanthonotozomati- 5 shorter than 4. Gnathopods feeble, sim- dae, Acanthonotozomellidae, and Iphime- ple, merus not lobate, with slightly elongate, diidae (except Stegopanoploea) in the pres- weakly lobate or non-lobate carpus, prop- ence of a molar, the produced merus and odus shorter and narrower than carpus, nip- carpus of gnathopod 2, and the absence of ple-like, gnathopod 2 usually slightly longer

dorsal teeth on the pereon; instead, the pere- than gnathopod 1 . Article 2 of pereopods on has a flush dorsal keel or margin. 5-7 lacking posterior cusps or teeth. Epi- The Dikwidae are characterized by re- meron 3 lacking 2 large cusps. Urosomites duced head, well developed D-E setae on free. Uropods 1-3 biramous. Rami of uro- the mandibular palp, and flagellar gnatho- pod 3 longer than peduncle, flattened, lan-

pod 2. ceolate, usually 1 -articulate. Telson cleft. See Epimeriidae for mention of differ- Relationship.— Di^Qving from Stilipedi- ences from families outside of the families dae in the narrowness of the incisors, the included herein. presence of a setose molar and the reduction

List of genera. — Odius Liljeborg, 1865 of spines on the outer plate of maxilla 1 (=Otus Bate, 1862, homonym, Lepidop- Remarks. —We keep Eclysis distinct from tera) (=type genus); Postodius Hirayama, Astrya based on the expanded palp of max-

1983. illa 1 and the need to describe many un- known characters. Epimeriella is trans- ferred to Epimeriidae because of the

Astryidae ordinary outer plate of maxilla 1 and poorly dominant carpus of the gnathopods. Epi- Astryidae Pirlot, 1934:175. meriella victoria Hurley (1957) is trans- Diagnosis.— 'Qo&y compressed, with or ferred to Epimeria because of tapering

without dorsal teeth (sometimes confined to coxa 1. metasome). Rostrum of medium size or very List of genera.— Boeck, 1871 small. Antennae weakly elongate, flagella {^Chagosia Walker, 1909, =Parastyra Pir- with 5 + articles; accessory flagellum absent. lot, 1934); Eclysis K. H. Barnard, 1932. Mouthpart part field quadrately developed (box-like). Epistome and labrum broad, short, weakly incised. Incisor of mandible Stilipedidae ordinary, toothed; raker row strong; molar Stilipedidae Holmes, 1908:535. reduced, conical and pubescent; palp always present, 3-articulate. Lower lip without in- Diagnosis.— ^ody compressed, dorsal ner lobes (except tiny inner basal lobes), teeth present or confined to urosome. Ros- outer lobes widely spaced, without distinct trum very small. Antennae moderately

inner notches. Inner plate of maxilla 1 or- elongate, flagella with 5 + articles; accessory dinary, somewhat medially setose; outer flagellum 0-1 articulate. Mouthpart part

plate oblique, spines reduced to 7 (? or 8) field quadrately developed (box-like). Ep- in Eclysis; palp large, 2-articulate. Inner plate istome and labrum broad, short, incised or of maxilla 2 without facial setae. Palp of lobate. Incisor of mandible extremely wide, maxilliped 4-articulate, article 2 not pro- smooth or toothed; raker row present or .

264 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON absent; molar absent; palp always present, ^Pseudandaniexis Nicholls, 1938); Asty- 3 -articulate. Lower lip with or without weak roides Birstein & Vinogradova, 1960. inner lobes, outer lobes widely separated, without distinct inner notches. Inner plate of maxilla 1 ordinary, apically and some- Lafystiidae what medially setose; outer plate oblique, Lafystiidae Sars, 1895:382. normally spinose in 2 genera, very wide and superspinose in many species; palp large, Diagnosis. —Body broadened, without 2-articulate, greatly widened apically. Inner dorsal teeth. Rostrum large, flattened. An- plate of maxilla 2 without facial setae. Palp tennae scarcely elongate, flagella with 5 + of maxilliped 4-articulate, article 2 not pro- articles; accessory flagellum absent. Mouth- duced medially. Coxae 2-4 more or less acu- part part field conically developed (box-like).

minate, ventral margins flush; coxae 1 and Epistome and labrum of ordinary width and

4 not shortened, coxa 1 widely expanded, length, entire. Incisor of mandible ordinary, wider than narrowed coxa 2, ventral margin toothed; raker row absent; molar absent; rounded or quadrate, coxa 4 weakly acu- palp present, 3 -articulate. Lower lip without minate, with posteroventral lobe, coxa 5 inner lobes, without distinct inner notches.

shorter than 4. Gnathopods feeble, simple Inner plate of maxilla 1 small, apically se- or very poorly subchelate, merus not lobate, tose; outer plate oblique, normally spinose;

with slightly elongate, weakly lobate carpus, palp tiny, 1 -articulate. Inner plate ofmaxilla propodus irregularly ovate, shorter and nar- 2 without facial setae. Palp of maxilliped rower than dominant carpus, gnathopod 2 reduced to 2 articles. Only coxae 4-6 acu-

usually slightly longer than gnathopod 1. minate, coxa 4 longer than coxae 1-3, latter Article 2 ofpereopods 5-7 without posterior ordinary and quadrate, their ventral mar- cusps or teeth. Epimeron 3 lacking 2 large gins ffush; coxa 4 with weak posteroventral cusps. Urosomites free. Uropods 1-3 bira- lobe, with strong anteroventral lobe, coxae mous. Rami of uropod 3 longer than pe- 5-6 with strong, sharp posteroventral lobe,

duncle, flattened, lanceolate, 1 -articulate. scarcely shorter than or subequal to 4.

Telson weakly incised. Gnathopod 1 feeble and with scarcely elon- Remarks.— Astryoides and Bathypano- gate articles 5-6, gnathopod 2 slightly larger ploea retain the plesiomorphic characters of than gnathopod 1; gnathopod 1 simple,

rakers and normal outer plate of maxilla 1 gnathopod 2 weakly propodochelate, merus Relationship. —Sqq Astryidae and Acan- and carpus not produced. Article 2 of pereo- thonotozomellidae Relationships. Differing pods 5-7 without posterior cusps or teeth. from the Iphimediidae and Acanthonoto- Epimeron 3 lacking 2 large cusps. Uroso- zomatidae in the broadened and non-acu- mites free. Uropods 1-3 biramous. Rami of

minate coxa 1 . Differing from Epimeriidae uropod 3 elongate but peduncle also slightly

in the lack of molar, broad, ffat mandible, elongate, rami flattened, lanceolate, 1 -artic- and dominant carpus of the gnathopods. ulate. Telson entire, not longer than pedun- List of genera.— Stilipes Holmes, 1908 cle of uropod 3. (type genus); Bathypanoploea Schellenberg, Relationship.— T>\f^QT\ng from the La- 1939 (=Iphimediopsis Schellenberg, 1931, phystiopsidae in the normal outer plate of homonym, =Epimeriopsis K. H. Barnard, maxilla 1, the reduction of the maxillipedal 1931, void ab initio, misidentified type spe- palp, the sub(para)chelate gnathopod 2, and

cies, =Pseudiphimediopsis Ruffb, 1 949); Al- the lack of molar. exandrella Chevreux, 1911, \9\2h {=Par- List of genera.— Lafystius Kroyer, 1842 andaniexis Nicholls, 1938, =homonym. (type genus) {^Darwinia Bate, 1857, =Der- VOLUME 104, NUMBER 2 265

mophilus Beneden & Bessels, 1870, =Ich- and reduction of spines on outer plate of

thymozocus HessQ, 1873). maxilla 1 ; its unique apomorphic characters are elongate telson, and loss of rostrum. K. Laphystiopsidae H. Barnard (1930:342) thought that Lafys- tiidae and Laphystiopsidae should be syn- Laphystiopsidae Stebbing, 1899:211. onymized. We believe Prolafystius should

Diagnosis based only on Laphystiopsis; be reevaluated and place it here only pro- see Remarks for Prolaphystius. visionally. Diagnosis. —Body slightly broadened, Relationship.— T>i^Qv'\ng from Lafysti- with weak or strong dorsal teeth on few idae in the presence of a molar, reduction

pleonites. Rostrum large and flat or absent. of spines on outer plate of maxilla 1, simple Antennae elongate, first longer than second, gnathopod 2, and 4-articulate palp of the flagella with 5 + articles; accessory flagellum maxillipeds. absent. Mouthpart part field quadrately de- List ofgenera. —Laphystiopsis Sars, 1895 veloped (box-like). Epistome and labrum (type genus); 1Prolaphystius K. H. Barnard, broad, short, minutely incised. Incisor of 1930; Prolaphystiopsis Schellenberg, 1931.

mandible ordinary, toothed; rakers absent; (Note: We have determined that Sars (1 895) molar present, well developed but smooth overlooked the biarticulate condition of the

or triturative (Prolaphystius); palp present, palp on maxilla 1 in Laphystiopsis; this ne- 3-articulate. Lower lip with or without small gates the only published distinction of Pro- fleshy inner lobes, without distinct inner laphystiopsis Schellenberg; specimens of the

notches. Inner plate of maxilla 1 small, with latter will be examined to seek any other one apical seta; outer plate oblique, spines possible generic difference).

reduced to 5; palp large, 1-2 -articulate. In- ner plate of maxilla 2 without facial setae. Acknowledgments Palp of maxilliped 4-articulate, article 2 not The authors are grateful to Dr. Les Wat- produced medially. No member of coxae 1- ling and five referees for valuable improve- 4 acuminate downward, all short, broader ments. than long, ventral margins ofcoxae 1-4 flush; coxa 4 without posteroventral lobe; coxa 5 Literature Cited longer or shorter than 4. Gnathopods 1-2 feeble, simple, carpi not lobate, propodi Andres, H. G. 1985. Die Gammaridea (Crustacea: narrow and rectangular. Article 2 of pereo- Amphipoda) der deutschen Antarktis-Expedi- tionen 1975/1976 und 1977/1978. 4. Acan- pods 5-7 lacking posterior cusps or teeth. thonotozomatidae, Paramphithoidae und Epimeron 3 lacking 2 large cusps. Uroso- Stegocephalidae. — MiUeilungen aus den Ham- mites free. Uropods 1-3 biramous. Rami of burgischen Zoologischen Museum und Zoolo- uropod 3 elongate, peduncle ordinary, rami gisches Institut 82:1 19-153.

flattened, lanceolate, 1 -articulate. Telson Barnard, J. L. 1969. The families and genera of ma- rine gammaridean Amphipoda. — United States entire, not longer or greatly longer than pe- National Museum, Bulletin 271:1-535. duncle of uropod 3. , & G. S. Karaman. 1987. Revisions in clas- Remarks. —Prolaphystius combines char- sification of gammaridean Amphipoda (Crus- acters of Lafystiidae and Laphystiopsidae. tacea) Part 3. — Proceedings of the Biological So- Its lafystiid characters are larger coxae with ciety of Washington 100:856-875. Barnard, K. H. 1930. Amphipoda. — British Antarc- some degree ofacumination, a posterior lobe tic ("Terra Nova") Expedition, 1910. Natural on coxa 4, and shortened coxa its la- 5; History Reports, Zoology 8:307-454. phystiopsid are of (trit- characters presence . 1931. Diagnosis of new genera and species urative) molar, simple feeble gnathopod 2, of amphipod Crustacea collected during the 266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

'Discovery' investigations, 1925-1927.— An- ceedings ofthe Biological Society ofWashington nals and Magazine of Natural History (10)7:425- 104:269-278.

430. . 1991b. Amatiguakiusforsberghi, a nev/ genus

. 1932. Amphipoda. — Discovery Reports 5:1- and species from Alaska (marine Amphipoda: 326. Epimeriidae).— Proceedings of the Biological

. 1955. Additions to the fauna-list of South Society of Washington 104:279-287.

African Curstacea and Pycnogonida.— Annals Costa, A. 1851. Pp. 44^7, fig. 2 in Gugl. Hope's of the South African Museum 43:1-107. Catalogo dei Crostacei Italiani e di Molti Altri Bate, C. S. 1862. Catalogue of the specimens of am- del Mediterraneo. Azzolino, Napoli.

phipodous Crustacea in the collection ofthe Brit- . 1851-1853. Fauna del Regno di Napoli [and] ish Museum. British Museum (Natural Histo- Catalogo de' Crostacei del Regno di Napoli.

ry), London, iv + 399 pp., pis. 1, la, 2-58. [Miscellaneous pages ofincomplete editions seen; Bellan-Santini, D. 1972. Invertebres marine des see Stebbing 1888:247-250.] Xlleme et XVeme Expeditions antarctiques Della-Valle, A. 1893. Gammarini del Golfo di Na- francaises en Terre Adelie 10. Amphipodes poli.— Fauna und Flora des Golfes von Neapel Gammariens.— Tethys Supplement 4:57-238. und der angrenzenden Meeres-Abschnitte, Beneden, E. van, & E. Bessels. 1870. Memoire sur la Monographic 20:xi + 948 pp., atlas (Atlante) of formation du blastoderme chez les amphipodes, 61 pis. les lemeens et les copepodes.— Memoires Sa- Griffiths, C. L. 1974. The Amphipoda of southern vants Etrangers, Academic Royal Science Lettres Africa part 4. The Gammaridea and Caprellidea Beaux-arts Belgique 34(4): 1-59. [See footnote of the Cape Province east of Cape Agulhas. — in this paper.] Annals of the South African Museum 65:251-

Birstein, J. A., & N. G. Vinogradova. 1960. Donnye 336.

ultraabissal'nye gammaridy severo-zapadnoi . 1977. The South African Museum's Meiring

chasti Tixogo Okeana. I. Semeistva Liljeborgi- Naude Cruises Part 6 Amphipoda.— Annals of idae, Astryidae, Lepechinellidae, Gammari- the South African Museum 74:105-123. dae.— Akademiia Nauk SSSR, Instituta Okea- Gurjanova, E. 1955. Novye vidy bokoplavov (Am- nologii, Trudy 34:147-164. phipoda, Gammaridea) iz sevemoi chasti Tix- Boeck, A. 1871. Crustacea Amphipoda Borealia et ogo Okeana.— Zoologicheskogo Instituta Aka-

Arctica. — Forhandslinger i Videnskabs-Selska- demii Nauk SSSR, Trudy 18:166-218.

bet i Christiania 1870:3-280. Heller, C. 1875. Die Crustaceen, Pycnogoniden und

. 1876. De Skandinaviske og Artiske Amphi- Tunicaten der K. K. Osterr.-Ungar. Nordpol-

poder. A. W. Brogger, Christiania, iv + 7 1 2 pp., Expedition [sic]. — Denkschriften der Kaiser- 32 pis. liche Akademie der Wissenschaften Mathema-

Bruzelius, R. M. 1859. Bidrag till kannedomen om tisch-Naturwissenschaftliche Classe 25:25^6, skandinaviens Amphipoda Gammaridea.— 5 pis. Kongliga Svenska Vetenskaps-Akademiens Hesse, M. 1873. Memoire sur des crustaces rares ou Handlingar, new series 3:1-104. nouveaux des cotes de France.— Annales des Chevreux, E. 1906. Crustaces amphipodes.— Expe- Sciences Naturelles, 5 serie, Zoologie et Paleon- dition Antarctique Francaise (1903-1905) com- tologie 17(7): 1-1 6. mandee par le Dr. Jean Charcot. Sciences Na- Hirayama, A. 1983. Taxonomic studies on the shal- turelles. Documents Scientifiques 1-100, 56 figs. low water gammaridean Amphipoda of west

. 1911. Sur les amphipodes des Expeditions Kyushu, Japan. I. Acanthonotozomatidae, Am- Antarctiques Francaises.— Academic des Sci- pehscidae, Ampithoidae, Amphilochidae, An- ences, Paris, Comptes Rendus 153:1 166-1 168. amixidae, Argissidae, Atylidae and Colomastig-

. 1912a. Diagnoses d'amphipodes nouveaux. idae. — Publications on the Seto Marine Deuxieme Expedition dans I'Antarctique, diri- Biological Laboratory 28:75-150.

gee par le Dr. Charcot, 1908-1 9 10. -Bulletin Holmes, S. J. 1 908. The Amphipoda collected by the du Museum d'Histoire Naturelle, Paris 18:208- United States Bureau of Fisheries Steamer, "Al- 218. batross", off the west coast of North America,

. 1912b. Amphipodes.— Deuxieme Expedi- in 1903 and 1904, with descriptions of a new tion Antarctique Francaise (1908-1910) com- family and several new genera and species.— mandee par le Dr. Jean Characot, Sciences Na- Proceedings of the United States National Mu-

turelles. Documents Scientifiques 79: 1 86, 62 figs. seum 35:489-543.

Coleman, C. O., & J. L. Barnard. 1991a. Curidia mag- Hurley, D. E. 1954. Studies on the New Zealand ellanica, new species, from Magellan Strait amphipodan fauna no. 9. The families Acan- (Crustacea: Amphipoda: Ochlesidae). — Pro- thonotozomatidae, Pardaliscidae and Liljebor- VOLUME 104, NUMBER 2 267

giidae.— Transactions of the Royal Society of K.C.S., ifec.&c. Captain in the Royal Navy. In- New Zealand 82:763-802. cluding the reports of Commander, now Cap-

. 1957 Some Amphipoda, Isopoda and Tan- tain, James Clark Ross, R.N., F.R.S., F.L.S., &c. aidacea from Cook Strait.— Zoology Publica- and the discovery of the northern magnetic pole. tions Victoria University Collections 21:1-20. A. W. Webster, London. Karaman, G. S. 1980. Revision of the genus Iphi- Ruffo, S. 1949. Amphipodes (II). — Resultats du

media Rathke 1 843 with description of two new voyage de la Belgica en 1897-99. ... Rapports

genera, Anisoiphimedia and Stegopanoploea, n. Scientifiques, Zoologie: 1-58, 18 figs. gen. (Fam. Acanthonotozomatidae). — Poljopri- Sars, G. O. 1895. Amphipoda. An account of the

vreda I Sumarstvo, Titograd 26:47-72. Crustacea of Norway with short descriptions and

Krapp-Schickel, G. 1974. Camill Hellers Sammlung figures of all the species l:viii and 71 1 pp., 240 adriatischer Amphipoden— 1866 und Heute.— pis., 8 suppl. pis. Alb. Cammermeyers, Chris- Annalen der Naturhistorisches Museum, Wien tiania, Norway. 78:319-379. Schellenberg, A. 1926. Die Gammariden der Kroyer, H. 1842. Une nordiske Slaegter og Arter af deutschen Sudpolar-Expedition 1901-1903.— Amfipodemes Orden, henhorende til Familien Deutsche Sudpolar-Expedition 18:235^14. Gammarina. (Forelobigt Uddrag af et storre Ar- . 1931. Gammariden und Caprelliden des Ma- bejde).— Naturhistorisk Tidsskrift 4:141-166. gellangebietes, Sudgeorgiens und der Westant- . 1846. Karcinologiske Bidrag (Fortsaet- arktis. — Further Zoological Results ofthe Swed- telse).— Naturhistorisk Tidsskrift 2:1-211, pis. ish Antarctic Expedition 1901-1903 2(6):l-290. 1-2, 1-88, 115-123 [for amphipods only]. . 1939. Amphipoden des Kongo-Mundungs- Kunkel, B. W. 1910. The Amphipoda of Bermuda. — gebietes. — Revue de Zoologie et de Botanique Transactions ofthe Connecticut Arts Academy of Africaines 32:122-138. and Sciences 16:1-116. Stebbing, T. R. R. 1888. Report on the Amphipoda Ledoyer, M. 1982. Crustaces amphipodes gamma- collected by H. M.S. Challenger during the years riens families des Acanthonotozomatidae a 1873-76. — Report on the Scientific Results of Gammaridae.— Faune de Madagascar 59(1): 1- the Voyage of H.M.S. Challenger during the 598. Years 1873-1876, Zoology 2:xxiv + 1737 pp.. Liljeborg, W. 1865. Bidrag till kannedomen om un- Eyre and Spottiswoodie, London. derfamiljen Lysianassina inom underordningen . 1899. Revision of Amphipoda (contin- Amphipoda bland kraftdjuren. — ["Nova Acta ued).— Annals and Magazine of Natural History Regiae Societatis Scientiarum Upsaliensis III (7)4:205-211. Serie"] [journal title from Stebbing 1888, only . 1906. Amphipoda I. Gammaridea. — Das an unattributed reprint seen]. Tierreich 21:1-806. Moore, H. B. 1985. A new deep water species of . 1910. Crustacea. Part V. Amphipoda. Sci- Amphipoda (Crustacea) discovered off Otago, entific Results Trawling Expedition H.M.C.S. New Zealand and a note on another little known "Thetis".— Australian Museum, Memoir 4 2: species.— Zoological Journal of the Linnean So- 565-658, pis. 47*-60*. [Asterisks denote special ciety 83:229-240. plates.] Nicholls, G. E. 1938. Amphipoda Gammaridea.— Australasian Antarctic Expedition 1911-14, Stock, J. H. 1974. The systematics of certain Ponto- Caspian Gammaridae (Crustacea, Amphipo- Scientific Reports, C.-Zoology and Botany 2(4): 1-145. da). — Mitteilungen Hamburg Zoologisches Mu- seum Institut 70:75-95. Pirlot, J. M. 1934. Les amphipodes de I'expedition nev/ du Siboga. Deuxieme partie. ... II. -Les am- Thomas, J. D. 1983. Curidiadebrogania, a genus phipodes de la mer profonde. 2. Hyperiopsidae and species of amphipod (Crustacea: Ochlesi- dae) from the barrier reefs of Belize, Central ... Jassidae. — Siboga-Expeditie 33d:167-235. Rathke, H. 1843. Beitrage zur Fauna Norwegens.— America. — Proceedings of the Biological Soci- Verhandlungen Kaiserlichen Leopoldinisch- ety of Washington 96:127-133. Carolinischen Akademie Naturforscher, Bres- Thomson, G. M. 1880. A new species of Crustacea lau 20(1): 1-264, 264b, 264c, 12 pis. [Amphip- from New Zealand.— Annals and Magazine of

oda:63-98, pis. 3, 4.] Natural History (5)6:1-6.

Ross, J. C. 1835. Zoology. Pp. vii-c, pis. A-C, Ap- Walker, A. O. 1906. Preliminary descriptions of new pendix to the narrative of a second voyage in species of Amphipoda from the 'Discovery' search ofa north west passage, and ofa residence Antarctic Expedition, 1902-1 904. — Annals and

in the Arctic regions during the years 1 829, 1 830, Magazine of Natural History (7) 18:13-18.

1831, 1832, 1833. By Sir John Ross, C.B.,K.S.A., . 1907. Crustacea. 111. Amphipoda.— National 268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Antarctic Expedition, British Museum (Natural Iphimediidae. In J. A. Crane, ed., Origins and History) 3:1-39. evolution of the antarctic biota.— Geological

. 1 909. Amphipoda Gammaridea from the In- Society of America Special Publication 47:297- dian Ocean, British East Africa, and the Red 313. Sea.— Transactions of the Linnean Society of

London (2) Zoology 12:323-344. Watling, L., & H. Holman. 1980. New Amphipoda (COC) Fachbereich 7, Zoomorphologie, from the Southern Ocean, with partial revisions Universitat Oldenburg, Postfach 2503, of the Acanthonotozomatidae and Parampith- D-2900, Oldenburg, F.R.G.; (JLB) Depart- oidae.— Proceedings of the Biological Society of ment of Invertebrate Zoology, National Washington 93:609-654. Museum of Natural History, Smithsonian , & M. H. Thurston. 1989. Antarctica as an evolutionary incubator: evidence from the cla- Institution, Washington, D.C. 20560. distic biogeography of the amphipod family