PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 707-720 TWO NEW SPECIES OF PODOCERUS LEACH (CRUSTACEA: AMPHIPODA: PODOCERIDAE) FROM BERMUDA Adam J. Baldinger and Michael F. Gable Abstract.—Twonewspecies,PodocerustachyrheoandPodoceruslazowasemi, are described from Bermuda. Podocerus tachyrheo is a non-carinate, highly pigmented species lackinguropodalinterramal spines; P. lazowasemipossesses distinct uropodal interramal spines and dorsal carinations, but usually lacks pigmentation. In Kunkel's (1910) monograph on the 2withdorsalspinegroupsonly.Adultmales amphipodsofBermudanospecimensofthe and females with varying degrees of pig- genus Podocerus were recorded. Johnson mentation. Coxal plates 1-5 with a strong (1986) and Gable et al. (1988), however, distal spine. Male antennae 1-2, gnatho- bothrefertotheexistenceofasinglespecies pods 1-2, maxillae 1-2, mandibular palp, ofPodocerus in Bermuda. Examination of and maxilliped with plumose setae. Male collections at the Yale Peabody Museum antenna 2, flagellum 3-articulate and or- (YPM)andattheNationalMuseumofNat- namentedwith submarginal spines. Female uralHistory(USNM)clearlydocumentsthe antennae 1-2, gnathopod 1, maxillae 1-2, presenceoftwo speciesofPodocerusin Ber- mandibular palp, and maxilliped with plu- muda, both of which are newly described mose setae. Male gnathopod 2, article 5 in this paper. masked by articles 4 and 6, palm with ir- In the figures, body parts are marked by regular margin bearing a proximal conical abbreviations as follows: A, antenna; Gn, tooth and a distal rectangular process. Ar- gnathopod; LL, lower lip; Md, mandible; ticle 6 ofallpereopodswith strongbifurcate Mx, maxilla; Mxpd, maxilliped; P, pereo- spines. Uropods 1-2 with bifurcate spines pod; T, telson; U, uropod; UR, urosome. but lacking peduncular interramal spines. Description.—Male: Body and append- Podocerus tachyrheo, new species ageswithpigmentation,bodylackingdorsal Figs. 1-5 carinae,pereonsegments5-7andpleonseg- ments 1-2 with dorsal distoposterior spine Podocerus sp.—Johnson, 1986:378, fig. groups, pereon segments 5-7 also with lat- 125.-Gableetal., 1988:148-149. eral distoposterior spine groups. Coxae re- Etymology.—Namedfortheabilityofin- duced with serial discontiguity, coxae 1-5 dividuals ofthis species to live in areas of each with a large marginal spine, located extremely swift (G. tachy-) currents (G. anteriorly on coxae 1—4. Head less than pe- rheo), suchasthoseatTheFlatts, Bermuda, reonites 1 and 2 in length, cuboidal with whereall tidal flow from Harrington Sound ocular bulges. Eyes pigmented. moves through one narrow channel. Antenna 1, 33% oftotal body length, pe- Diagnosis.—hereon without dorsal cari- duncularratio 1:3:2.7,flagellum 5-articulate; nations. Pereon segments 5-7 with dorsal accessory flagellum 1-articulate and prom- andlateral spinegroups, pleon segments 1- inent. Antenna 2 greater than antenna 1 in ' 708 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON MMk •iMm^fm: M$ ^^^k r?5^!i^SmS^ mmmm ws .^^ .-^^j*»:s CBfe is^^i^ v^i^ \\^ ''iv bin' vV Fig. L Podocerustachyrheo, male, 4.0 mm, YPM No. 9239. length, articles 4 and 5 subequal, flagellum er plate reaching 50% length ofpalp, with 3-articulate with distinct submarginal an inner marginal spine row and submar- spines. ginal setae; palp 4-articulate, terminal ar- Mandible with 3-segmented palp, pen- ticle triangular. Lower lip, normal. ultimatearticletriangularwith marginal se- Gnathopod 1, coxal plate rhomboidal tae, terminal article clavate with facial and withadistinctdistoanteriorspine; article 6, apical setae; incisorand lacinia both with 4 palmar margin longer than hind margin; teeth; spine row of 3 short, broad spines; dactyl with 3 marginal setae. Gnathopod 2 molar normal. Maxilla 1, inner plate ves- heavilypigmented, robust, and muchlarger tigial;outerplatewith9strongapicalspines; than gnathopod 1; article 5 hidden behind palp 2-articulate, terminal article with 3 articles 4 and 6; palm ofarticle 6 with ir- apical, medially bulging spines and 1 apical regular margin, bearing a proximal conical seta,4submarginalsetae,and4facialspines. tooth and adistal rectangular process, mar- Maxilla 2, outer plate with 10 long apical gin heavily setose with 2 proximal spines. setae; innerplate with 1 apical setae, inner Basesofpereopods3-7withposteriorplate- margin with fine setae distally. Maxilliped, like extensions; article 6 of all pereopods inner plate with marginal and submarginal with distally bifurcate spines. setaeandasingleoutermarginal spine; out- Uropods 1 and 2 elongate, biramous. VOLUME 107, NUMBER4 709 Fig. 2. Podocerus tachyrheo. male, 4.0 mm, YPM No. 9239: Al, Gnl, Gn2, UR (with left Ul, right U2, rightand leftU3, T). Male, 2.9 mm, YPM No. 9242: A2. 710 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON Fig. 3. Podocenistachyrheo, male, 4.1 mm, YPM No. 9241. VOLUME NUMBER 107, 4 711 Fig. 4. Podocerustachyrheo, male, 2.9 mm, YPMNo. 9242: P3-P4. Male, 4.0 mm, YPMNo. 9239: P5-7. lacking interramal spines; peduncles and body segment, even on the most distal an- rami with marginal bifurcate spines, rami tennal segment. In females, however, pig- with distinct apical spines. Uropod 3 leaf- mentation appears to become much denser like with 3 apical setules. Telson dorsally on the ventral margin of the pereon, and produced and armed with 4 long setae and mayextendtothebasalportionsoftheoos- 6 short setules. tegites. Male/femaledifferencesinbodyspi- Female: All features same as those for nation, in the flagella ofboth antennae, in male except as noted. Body lacking dorsal both gnathopods, and in pigmentation ob- spinegroups. Antenna 1, 40% oftotal body viously make P. tachyrheo sexually dimor- length; flagellum 4-articulate. Flagellum of phic. antenna 2 lacking distinct submarginal Podocerus tachyrheo is morphologically spines. Gnathopod 1, article 6, proximal most closely related to Podocerus muhis- comer of palm demarcated by a spine. pinis K. H. Barnard, 1925 and Podocerus Gnathopod 2 resembling gnathopod 1 but muhispinis var. levis K. H. Barnard, 1925. twice the size and less ornamented. The only major difference between P. mul- Remarks.—The, degree and patterns of tispinis and P. muhispinis var. levis is the pigmentation are variable among individ- almost complete absence ofthe dorsal spi- uals of P. tachyrheo; juveniles, smaller niform tubercles in the variety (Barnard males, and females may completely lack 1925). Based upon the description and one pigment. Largermalesappeartohaveauni- illustrationofgnathopod2bybothBarnard form pigment band along the musculature (1925) and Griffiths (1976), the most sig- of the pereon segments. Pigment has also nificantdifferencesbetweenP. tachyrheoand been observed in males on almost every P. muhispinis are: the number of flagellar 712 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON Fig. 5. Podocemstachyrheo, female, 3.0 mm, YPM No. 9240. VOLUME 107, NUMBER4 713 YPM YPM articles ofantenna 1, P. tachyrheowith five paratype, 9242 (same data as and P. multispinis with eight; the lateral 9241), 2.9 mm. 1 female paratype, oviger- YPM YPM spinesonthepereon,presentinP. tachyrheo ous, 9244 (same data as 9243), and absent in P. multispinis; the number of 3.5 mm, highly pigmented. 5 paratypes, 2 setaepresentonthetelson,fouronP. tachy- males, 2 females, ovigerous, and 1 juvenile, rheoandtwoonP. multispinis; theirregular deposited by YPM in Bermuda Museum of YPM marginofthepalmofarticle6ofgnathopod Natural History (same data as 9243). 2, P. tachyrheowith onetoothandone rect- angular process and P. multispinis, as de- Podocerus lazowasemi, new species scribed by Barnard (1925), with two teeth Figs. 6-9 and one rectangular process. In addition, Etymology.—'Nz.v[\Qd. in appreciation for Barnard (1925) does not mention P. mul- the technical assistance, sound advice, and tispinis or the variety as having pigmenta- encouragement received for this and many tion. Finally, although close morphologi- other projects from our colleague and good cally to P. tachyrheo, P. multispinis is friend, E. A. Lazo-Wasem (YPM). endemicfromNataltoSaldanhaBay, South Diagnosis.—Pereon segments 6-7 and Africa (Griffiths 1975). Podocerus tachy- pleon segments 1-2 with dorsal carinations rheo, therefore, maybeconsideredaspecies andspinegroups. Coxalplates 1-7 eachwith endemic to Bermuda. 1 large marginal spine. Antennae 1-2, max- Otherthanitsmorphology,littleisknown illae 1-2, maxilliped, and male gnathopods aboutP. tachyrheo. In Bermuda most spec- 1-2 with plumose setae. Maxilla 1, outer imens collected were associated with hy- plate with 8 apical spines. Male gnathopod droids; one collection yielded several hun- 1, article 6 with anterior marginal setae; dred specimens taken from a small portion gnathopod 2, palm at dactyl hinge with a ofalargecolonialhydroid, Halocordyledis- pronounced rectangular process. Dactyls of ticha (Goldfuss), found attached in an area all pereopods with a proximoanterior plu- ofvery strong current. mose seta. Peduncle of uropods 1-2 with Material examined. —Male holotype, interramal spine. Dorsal lobe oftelson with YPM 9254, The Flatts, Harrington Sound, 2 long and 2 short setae. Bermuda, M. F. Gable, 23 May 1989, 5.0 Description.—Male: Body unpigmented mm, onHalocordyledisticha (Goldfuss) at- with conspicuous dorsal carinae and dorsal tached to a subtidal pipe. Female allotype, spine groups on pereon segments 6-7 and YPM 9245, The Flatts, Harrington Sound, pleon segments 1-2. Coxae reduced with Bermuda, M. F. Gable, 23 May 1989, 2.9 serial discontiguity, each with a large mar- mm, mature, on Halocordyle disticha ginal spine. Head equal to pereonites 1-2 (Goldfuss)attachedtoasubtidalpipe. 1 male in length, cuboidal with lateral ocular bulg- YPM paratype, 9239, Harrington Sound, es. Eyes pigmented. behind Bermuda Aquarium, Bermuda, M. Antenna 1, 40% oftotal body length, ge- F. Gable, 2 Jun 1987, 4.0 mm, within hy- niculate, article 2 slightlylongerthanarticle droids and algae. 1 female paratype, YPM 3, flagellum 5-articulate; accessory flagel- 9240, (same data as YPM 9239), 3.0 mm. lum, 1-articulate and prominent. Antenna 20 paratypes, males, females andjuveniles, 2, bigeniculate, article 5 nearlytwiceaslong YPM 9243,LongBirdCauseway, Bermuda, as article 4, flagellum 4-articulate and or- Ssideandundercauseway,E.A. Lazo-Was- namented with spines and setal groups. em, 20 Jun 1988, washing ofrocks. 1 male Mandible with 3-segmented palp, pen- YPM paratype, 9241, TheFlatts, Bermuda, ultimate segment with submarginal and M. F. Gable, 23 May 1989, 4.1 mm. 1 male marginal setae, terminalarticleclavatewith 714 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON Fig.6. Podoceruslazowasemi, male, 3.0mm,YPMNo.9260: body,Gnl,Gn2.Female,2.4mm,YPMNo. 9261:Gnl,Gn2. facial and apical setae; incisorwith 4 teeth; 9 apical setae. Maxilliped, inner plate cla- lacinia with 5 teeth; spine row of 3 short, vatewithapicalandfacialsetae,distalinner broad spines; molar normal. Maxilla 1, in- comer with a large marginal spine; outer ner plate vestigial; outerplate with 8 strong plate <50% length of palp, with 4 inner apical spines, 2 of them bifurcate; palp marginal spines and with submarginal and 2-articulate, terminal article with 4 strong marginal setae; palp 4-articulate with sub- apical spines and 3 apical setae. Maxilla marginal and marginal setae. Lower lip, 2, inner plate with 10 apical setae, inner normal. marginlinedwithfinesetae;outerplatewith Gnathopod 1, coxal plate rhomboidal VOLUME 107, NUMBER4 715 Fig. 7. Podoceruslazowasemi, male, 3.3 mm,YPMNo. 9262: Al,A2. Male, 3.1 mm,YPMNo. 9259: UR (with right Ul, left U2, rightand leftU3, T). 716 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON Fig. 8. Podoceruslazowasemi, male, 3.3 mm, YPM No. 8357. with 1 large distoanterior spine; article 6, pod 2 robust and much larger than gnatho- palm much longer than hind margin and pod 1; basis triangulate, formed by a dis- with marginal setae, anterior margin with 4 toanterior lobed projection; article 6, palm setal groups; dactyl with 4 marginal setules setose with a pronounced distal rectangular and a subterminal bifurcate spine. Gnatho- processand4distinctsubmarginalbifurcate