A redescription of the type species of Oedicerina Stephensen, 1931 (Crustacea, Amphipoda, Oedicerotidae) and the description of two new species

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1 Zoosyst. Evol. 90 (2) 2014, DOI /zse museum für nturkunde A redescription of the type species of Oedicerin Stephensen, 1931 (Crustce, Amphipod, Oedicerotide) nd the description of two new species Chrles Oliver Colemn 1, Michel H. Thurston 2 1 Museum für Nturkunde, Leiniz-Institut für Evolutions- und Biodiversitätsforschung, Invlidenstrße 43, Berlin, Germny 2 Ntionl Ocenogrphy Centre, Europen Wy, Southmpton SO14 3ZH, UK Corresponding uthor: Chrles Oliver Colemn (oliver.colemn@mfn-erlin.de) Astrct Received 6 Septemer 2014 Accepted 8 Octoer 2014 Pulished 31 Octoer 2014 Acdemic editor: Crsten Lüter Key Words The poorly known species Oedicerin ingolfi Stephensen, 1931 (Crustce: Amphipod: Oedicerotide) is redescried, sed on new mteril from the Norwegin Se. Oedicerin vderi sp. n. from the northest Atlntic Ocen nd Oedicerin loerze sp. n. from New Zelnd wters re descried rising the numer of species in the genus to five. The three species treted here together with Oedicerin meglopod Ledoyer, 1986 nd Oedicerin denticult Hendrycks & Conln, 2003 re seprted y chrcters of the rostrum, mxilliped, gnthopods, epimer, nd y the dorsl rmture of pleonites nd urosomites. The genus is recorded from the Atlntic, Indin nd Pcific Ocens, minly t thyl depths. Txonomy Amphipod Oedicerotide Oedicerin new species Norwegin Se North Atlntic Ocen New Zelnd Pcific Ocen Introduction Prticipnts in workshop orgnised y Professor Wim Vder t the University of Tromsø field sttion t Skiotn in 2009 sorted nd identified mphipods from the Nturl History Collections of the University Museum of Bergen collected from the Norwegin Se. Among the extensive mteril in these collections were specimens of the poorly known oedicerotid genus Oedicerin Stephensen, The genus ws sed on single individul collected from the Norwegin Se y The Dnish-Ingolf Expedition (Stephensen 1931). Although the specimen consisted of just the hed nd pereonites 1 4 nd ws lcking gnthopods, the new genus ws creted for O. ingolfi Stephensen, 1931 to recognise the unique nture of the huge posterior loe of cox 4. The genus remined monotypic nd unreported until Ledoyer (1986) descried?oedicerin meglopod from the Mozmique Chnnel, western Indin Ocen, lso sed on single nterior frgment. This species ws defined y the chrcteristic shpe of cox 4 nd y the strongly developed crpl rticles of gnthopods 1 nd 2. The description of third species, O. denticult Hendrycks & Conln, 2003 from the northest Pcific Ocen, ws the first to record complete specimens nd ws ccompnied y more detiled pprisl of the genus. The new mteril from the Bergen Museum hs mde possile re-description of O. ingolfi. Specimens elonging to Oedicerin from the northest Atlntic Ocen nd Copyright Chrles Oliver Colemn, Michel H. Thurston. This is n open ccess rticle distriuted under the terms of the Cretive Commons Attriution License (CC BY 4.0), which permits unrestricted use, distriution, nd reproduction in ny medium, provided the originl uthor nd source re credited.

2 226 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... from New Zelnd wters demonstrte further diversity within the genus nd require the recognition of two new species which re descried elow. This pper is the third to utilize mteril sorted t Skiotn in 2009, following d Udekem d Acoz (2010) nd Krpp-Schickel nd Vder (2013). Mteril nd methods Norwegin Se mteril ssigned to O. ingolfi from the Nturl History Collection of the University Museum of Bergen (ZMBN) ws collected with n RP sledge (Rothliserg nd Percy 1977) y Torleiv Brttegrd (Brttegrd nd Fosså 1991) in the period Additionl mteril from the Norwegin nd Greenlnd Ses from the Museum of Zoology, Lund University ws collected during the NORBI expedition (Dhl et l. 1976) using n epienthic sledge (drgue Snders) (Guennegn nd Mrtin 1985). Specimens from the Discovery Collections t the Ntionl Ocenogrphy Centre, Southmpton were otined in the Est Icelnd Bsin on n RRS Discovery cruise tht contriuted to the Institute of Ocenogrphic Sciences investigtions of mid-wter nd enthic funs in the estern North Atlntic Ocen ( ). The mteril, from n epienthid sledge, ws fixed in 4% formldehyde nd lter trnsferred to 70% Industril Methylted Spirits. These specimens hve een deposited in the Amphipod collections t The Nturl History Museum, London. The New Zelnd mteril ws collected during the Ocen Survey 2020 expeditions with RV Tngro to the Chthm Rise nd the Chllenger Plteu (Knox et l. 2012) y mens of Brenke epienthic sledge (Brenke 2005). The mteril ws sorted on ord, fixed in 96% ethnol nd lter trnsferred into 70% ethnol. It hs een deposited in the Ntionl Institute of Wter nd Atmosphere Reserch (NIWA) Mrine Inverterte Collection in Wellington, New Zelnd. For hitus drwings the specimens were trnsferred into glycerol on cvity slide. Specimens were then dissected under stereomicroscope (Leic M205 or Wild M5) using dissecting needles. Mouthprts nd ppendges were mounted temporrily in glycerol on slides for microscopic exmintion nd drwing. Appendges were lter mounted s permnent slides with glycerol jelly, or trnsferred into smll glss microvils. Microvils were stoppered with cotton ll wrpped in Jpn pper to void the ppendges eing entngled in the cotton fires. After dissection, mouthprts nd ppendges of Discovery Collection mteril were mde directly into permnent mounts using Polyvinyl-lctophenol stined with lignin-pink. Drwings of hitus nd ppendges were mde using cmer lucid ttched to compound microscope (Leic DMLB or Wild M20). Pencil drwings were scnned, inked digitlly nd rrnged to pltes using the methods descried in Colemn (2003, 2009). Body lengths were mesured long the dorsl outline from the tip of the rostrum to the end of the telson. Lengths of individul rticles of gnthopods nd pereopods mesured long nterior or posterior mrgins cn vry depending on the degree of flexure of the ppendge. All rticultions except those etween coxe nd tergites nd etween merus nd crpus of gnthopods re icondylr. Mesurements mde etween condyles gives length tht is not ffected y lim flexure. Length rtios herein hve een derived using this principle. Systemtics Amphipod Ltreille, 1816 Oedicerotide Lilljeorg, 1865 Oedicerin Stephensen, 1931 Oedicerin Stephensen, 1931: 250. Brnrd nd Krmn 1991: 561, Ledoyer 1986: 832. Hendrycks nd Conln 2003: Dignosis (key chrcters emolded). Rostrum well-developed, modertely to strongly deflexed. Antenne sexully dimorphic or not, length medium. Antenn 1 out s long s hed nd pereonites 1 4 comined, peduncle rticle 1 longer thn rticles 2 nd 3. Antenn 2 suequl to or wekly longer thn ntenn 1; peduncle rticle 4 longer thn rticle 5. Lower lip, inner loes prominent, seprte. Mndile, molr triturtive; incisor 5-dentte. Mxill 1, outer plte 9-dentte; plp slender, rticle 2 suequl to or longer thn rticle 1. Mxill 2, pltes short, inner roder thn outer. Mxilliped, plp rticle 2 su-tringulr, redth gretest t hlf-length, inner mrgin strongly convex; rticle 3 produced mediodistlly; rticle 4 longer thn rticle 3. Coxl pltes 1 4 deep, s long or longer thn height of corresponding pereonite. Gnthopod 1, cox expnded distlly; crpus nd propodus suequl in length, strongly expnded posterodistlly. Gnthopod 2, crpus longer thn propodus, oth strongly expnded posterodistlly. Pereopods 3 nd 4 fossoril (setose); cox 4 deeply excvte posteriorly, posterodistl loe strong, surectngulr. Pereopod 5, cox ilote, posterior loe s long s cox 4. Pereopod 6, cox ilote, posterior loe strong. Pereopod 7, sis expnded. Pleonites, some or ll crinte or toothed. Epimer 1 3, 1 nd 3 rounded, 2 otusely rounded, posterior mrgin convex or sinuous. Uropods 1 2, outer rmus suequl to or shorter thn inner rmus. Uropod 3, peduncle short; rmi suequl, not extending s fr s pices of uropods 1 2. Telson notched 30 40%, pices cute. Type species. Oedicerin ingolfi Stephensen, 1931 Species composition. Oedicerin denticult Hendrycks & Conln, 2003; Oedicerin ingolfi Stephensen, 1931; Oedicerin loerze sp. n.; Oedicerin meglopod Ledoyer, 1986; Oedicerin vderi sp. n.

3 Zoosyst. Evol. 90 (2) 2014, Key to the species of Oedicerin 1 Rostrum mssive, sptulte, longer thn peduncle rticle 1 of ntenn 1... O. meglopod Ledoyer, 1986 Rostrum t most s long s rticle 1 of ntenn Pleonites 1 2 crinte, pleonite 3 with smll tooth...o. ingolfi Stephensen, 1931 Some pleonites smooth dorslly Pleonite 1 smooth... 4 Pleonite 1 2 crinte, pleonite 3 smooth...o. loerze sp. n. 4 Pleonite 1 2 smooth... O. vderi sp. n. Pleonite 2 crinte, pleonite 3 with short process...o. denticult Hendrycks & Conln, 2003 Oedicerin ingolfi Stephensen, 1931 Figs 1 5 Oedicerin ingolfi Stephensen, 1931, p. 250, fig. 72. Oedicerotide gen. et sp. n. Dhl 1979, p. 60 (ecology). Mteril exmined. 3 ovig. femles, 6 femles, 3 mles, 2 unknown sex, 4 juveniles, ZMBN 95143, St , N, W, 2630 m, F/F Håkon Mosy, RPsledge, T. Brttegrd, 14 August femle, 1 mle, 1 juvenile, ZMBN 95144, St , N E, 2908 m, F/F Håkon Mosy, RP-sledge, T. Brttegrd, 14 August ovig. femle, 4 femles, 7 mles, 3 unknown sex, 12 juveniles, ZMBN 95145, St N W, 3000 m, F/F Håkon Mosy, RP-sledge, T. Brttegrd, 24 Novemer mle, 1 juvenile, ZMBN 95146, St , N E, 2025 m, F/F Håkon Mosy, RPsledge, T. Brttegrd, 3 June femle, ZMBN 95147, St , N W; 2212 m, F/F Håkon Mosy, RP-sledge, T. Brttegrd, 26 July dult femles, 1 femle, 2 juveniles, NORBI St. 2, DS05, N E N E; 2970 m, N.O. Jen Chrcot, 21 July ovig. femle, 1 dult mle, NORBI St. 6, DS12, N E N E; 3200 m, N.O. Jen Chrcot, 2 August Description. Bsed on ovigerous femle, 10.3 mm, St Hed (Fig. 1): longer thn high, longer thn pereonites 1 2 comined; no eyes or oculr pigment visile; rostrum strongly deflexed, ventrl mrgin wekly convex. Antenn 1 (Fig. 1): out s long s ntenn 2; length rtios of peduncle rticles 1 3 1:0.9:0.6; flgellum 10-rticulte; ccessory flgellum 1-rticulte, minute, slender, less thn hlf length of first flgellum rticle. Antenn 2 (Fig. 1c): peduncle setose; length of rticle rticle 5; flgellum shorter thn peduncle rticle 5, 7-rticulte. Upper lip (lrum) (Fig. 1d): wider thn long, rounded piclly. Mndile: molr triturtive, with one ssocited set; incisors nd lcinie moiles 5-dentte; plp (Fig. 1i) 3-rticulte, rticle 2 swollen proximlly, rticle 3 tpered, length rtios of rticles 1 3 1:4.1:4.5. Lower lip: inner loes prominent nd rod, hypophryngel gp wide, outer loe mndiulr processes short nd rounded. Mxill 1 (Fig. 1e, f, g): inner plte ovl, with two distl sete; outer plte with nine cute setl-teeth; plp 2-rticulte, rticle length of rticle 1. Mxill 2 (Fig. 2): inner plte 1.2 wider thn outer plte; oth pltes with reltively sprse picl settion. Mxilliped (Fig. 2): inner plte short, extending just eyond se of plp rticle 1; outer plte extending 50% long plp rticle 2, concve medilly; plp 4-rticulte; rticle 1 tpered; rticle 2 rod, strongly expnded medilly, loe sutringulr; rticle 3 nrrow proximlly,expnded mediodistlly; rticle 4 cute, wekly flcte: length rtios of rticles 1 4 1:1.7:0.7:1.3. Pereon. Pereonite 1 (Fig. 1): longer thn 2; pereonites 3 5 successively longer; pereonites 6 nd 7 suequl in length to 5. Gnthopod 1 (Fig. 2c): cox sutringulr, posterior mrgin stright, nterodistl corner rounded, posterodistl corner rectngulr, distl mrgin stright, strongly setose; sis stright, wekly expnded, posterior mrgin with row of plumose sete; merus, posterodistl loe rounded, setose; crpus strongly expnded, sucute posterior loe with posterior nd distl mrgins setose; propodus strongly expnded, s long nd wide s crpus, nterior mrgin convex, plm trnsverse, convex, crenellte nd setose; dctylus curved, just longer thn plm. Gnthopod 2 (Fig. 3): cox s long s cox 1, wekly tpering distlly, pex rounded, setose; sis surectngulr, with row of plumose sete ner posterior mrgin; merus, posterodistl loe nrrow, setose; crpus strongly expnded, wider thn propodus, posterodistl loe sucute, distl mrgin olique; propodus shorter thn crpus, expnded distlly, plm strongly convex, crenellte; dctylus slender, flcte, s long s plm. Pereopod 3 (Fig. 3): cox suequl to cox 2; sis shorter thn cox, with very long slender sete on posterior mrgin nd plumose sete close to nterior mrgin; merus wekly expnded distlly; crpus 1.3 length nd out s wide s merus, posterior mrgin setose; propodus ovl, setose; dctylus 1.2 length of propodus. Pereopod 4 (Fig. 4): cox wider thn long, distl mrgin rounded, posterodistl loe very strong, surectngulr; sis shorter thn cox; merus wekly expnded; crpus shorter thn merus, ut suequl in width, setose posteriorly nd nterodistlly; propodus with nteromrginl rows of slender sete; dctylus rther stout, short nd stright. Pereopod 5 (Fig. 4): cox out s deep s cox 4, iloed, posterior loe

4 228 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... c d e f g h i Figure 1. Oedicerin ingolfi, ovig. femle, 10.3 mm; Norwegin Se, ZMBN 95143, St ) hitus (uropods rolled up, not illustrted); ) ntenn 1; c) ntenn 2; d) upper lip; e) mxill 1; f) mxill 1, setl teeth of outer plte; g) mxill 1, distl settion of plp rticle 2; h) telson; i) mndiulr plp. Scle rs: ; 2 mm:, c, e, h; 200 µm: d, i; 100 µm.

5 Zoosyst. Evol. 90 (2) 2014, c Figure 2. Oedicerin ingolfi, ovig. femle, 10.3 mm; Norwegin Se, ZMBN 95143, St ) mxill 2; ) mxillipeds; c) gnthopod 1. Scle rs: ; 100 µm:, c; 200 µm.

6 230 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... Figure 3. Oedicerin ingolfi, ovig. femle, 10.3 mm; Norwegin Se, ZMBN 95143, St ) gnthopod 2; ) pereopod 3. Scle rs:, ; 200 µm.

7 Zoosyst. Evol. 90 (2) 2014, Figure 4. Oedicerin ingolfi, ovig. femle, 10.3 mm; Norwegin Se, ZMBN 95143, St ) pereopod 4; ) pereopod 5. Scle rs:, ; 200 µm.

8 232 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... Figure 5. Oedicerin ingolfi, ovig. femle, 10.3 mm; Norwegin Se, ZMBN 95143, St , d) pereopod 6; ) pereopod 7; c) pleopod 1; e) uropod 1; f) uropod 2; g) uropod 3. Scle rs:, ; 1 mm: c, e, f, g; 200 µm.

9 Zoosyst. Evol. 90 (2) 2014, expnded distlly, distl mrgin stright, nterior loe 0.7 length of posterior loe, rounded distlly; sis shorter thn cox; merus s long s sis, crpus 0.5 length of merus; propodus slender, surectngulr, 0.8 length of merus, out s long s stright lnceolte dctylus; rticles 2 6 vriously setose. Pereopod 6 (Fig. 5): cox lmost s deep s cox 5, iloed, posterior loe long, distl mrgin stright, olique, nterior loe short, rounded distlly; sis surectngulr; merus posterior mrgin wekly convex; crpus tpering wekly, 0.5 length of merus; propodus with severl rows of sete long nterior mrgin, 0.9 length of merus; dctylus lnceolte; rticles 2 6 vriously setose. Pereopod 7 (Fig. 5): long, exceeding pices of uropods; cox wider thn long, posterodistl corner surectngulr; sis, posterior mrgin wekly sinuous, nterior mrgin strongly convex; merus nd crpus with groups of short slender sete on nterior nd posterior mrgins; crpus 1.2 length of merus; propodus nrrow, surectngulr, 0.6 length of merus; [dctylus unknown]. Pleon. Pleonites 1 2 (Fig. 1) with mid-dorsl, reltively long posteriorly directed crinte teeth; pleonite 3 with short, slender, upright tooth. Epimer:1 nd 3 evenly rounded; epimeron 2 surectngulr. Pleopod 1 (Fig. 5c): peduncle stout, 0.8 length of rmi. Urosome. Urosomite 1 (Fig. 1) longest, with n inconspicuous oss close to the posterior mrgin; urosomite 3 longer thn 2, with short, cute mid-dorsl projection. Uropod 1 (Fig. 5e): peduncle out s long s outer rmus, mrgins with short sete; inner rmus 1.3 length of outer rmus, with smll sete on oth mrgins; outer rmus with sete on lterl mrgin only. Uropod 2 (Fig. 5f): peduncle slightly tpering, with short sete on oth mrgins; inner rmus 1.7 length of outer rmus, with short sete on oth mrgins; outer rmus with sete on lterl mrgin only. Uropod 3 (Fig. 5g) peduncle short, out s long s telson, with ventrl sucute projection; rmi suequl, plumose sete on lterl mrgins. Telson (Fig. 1h) tpered, notched 30%. Sexul dimorphism. Mle ntenn 1 with shorter peduncle rticles in the rtio 1:0.7:0.3 nd more numerous flgellum rticles compred to femle. Article 1 of the flgellum is elongte, out s long s peduncle rticle 3. Susequent proximl rticles re shorter thn wide. The 1-rticulte slender ccessory flgellum is out 1/3 s long s rticle 1 of the primry flgellum. Distriution. Between the Froes nd Jn Myen (Stephensen 1931); Norwegin Se, Greenlnd Se (this study), m. 233 Remrks. Stephensen s (1931) specimen ws dmged nd incomplete. Only the hed nd pereonites 1 4, pereopods 3 4 nd coxe nd ses of pereonites 1 2 on one side were ville for study. The mteril from the Bergen Museum used for this description consists of numerous specimens of ll sizes, oth femle nd mle, nd ws collected reltively close to the type loclity of O. ingolfi, ut nevertheless we cnnot e solutely sure tht our mteril represents Stephensen s species (see discussion elow). Oedicerin vderi sp. n. Figs 6 10 Type mteril. Mle holotype, 7.3 mm; NHMUK , Discovery Stn 7709#73. Type loclity. North Atlntic, Est Icelnd Bsin: N W N W, 5 My 1971, BN 2.4, m. Prtypes. 1 femle, 6.3 mm; 2 specimens of unknown sex, 4.2 mm nd 5 mm; NHMUK , Discovery Stn 7709#73, from the type loclity. Etymology. The specific nme vderi recognises the importnt contriutions to mphipod studies mde y Professor Wim Vder. Description. Holotype mle, 7.3 mm. Hed (Fig. 6c): longer thn high, longer thn pereonites 1 2 comined; no eyes or oculr pigment visile; rostrum (Fig. 6c) strongly deflexed, the ventrl mrgin concve. Antenn 1 (Fig. 6f): length rtios of peduncle rticles 1 3 1:0.5:0.3; flgellum [roken], proximl flgellum rticles wider thn long; ccessory flgellum 1-rticulte, slender, out hlf the length of peduncle rticle 3. Antenn 2 (Fig. 6g): peduncle wekly setose; length of rticle rticle 5; flgellum shorter thn peduncle rticle 5, 9-rticulte. Upper lip (lrum) (Fig. 6d): wider thn long, piclly rounded. Mndile (Fig. 7, e): incisors nd lcinie moiles 5-dentte; plp 3-rticulte, rticle 2 swollen proximlly, rticle 3 tpered, length rtios of rticles 1 3 1:4.3:6.1. Lower lip: inner loes prominent nd rod, hypophryngel gp wide, outer loe mndiulr processes short nd rounded. Mxill 1 (Fig. 7d): inner plte tpered, with two distl sete; outer plte with nine cute setl-teeth; plp 2-rticulte, rticle 2 5 length of rticle 1. Mxill 2 (Fig. 7): inner plte 1.3 wider thn outer plte; oth pltes with reltively sprse picl settion. Mxilliped (Fig. 7c): inner plte short, extending just eyond se of plp rticle 1; outer plte extending 30% long plp rticle 2; concve medilly; plp 4-rticulte; rticle 1 tpered; rticle 2 rod, strongly expnded medilly, loe rodly rounded; rticle 3 nrrow proximlly nd expnded mediodistlly; rticle 4 curved, cute; length rtios of rticles 1 4 1:2.2:0.8:1.3. Pereon. Pereonite 1 (Fig. 6): longer thn 2; pereonite 2 shortest. Gnthopod 1 (Fig. 8): cox sutringulr, posterior mrgin stright, nterodistl corner rodly rounded, posterodistl corner rectngulr, distl mrgin stright, wekly setose; sis curved, posterior mrgin with row of plumose sete; merus, posterodistl loe rounded, setose; crpus strongly expnded, surectngulr posterior

10 234 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... c II III I d e f g Figure 6., c g) Oedicerin vderi sp. n., mle holotype, 7.3 mm; northest Atlntic Ocen, NHMUK , Discovery Stn 7709#73. ) hitus; c) hed; d) upper lip; e) epimerl pltes 1-3; f) ntenn 1; g) ntenn 2. Femle prtype, 6.3 mm; sme loclity. ) ntenn 1. Scle rs: ; 1 mm: g; 200 μm.

11 Zoosyst. Evol. 90 (2) 2014, c e d Figure 7. Oedicerin vderi sp. n., mle holotype, 7.3 mm; northest Atlntic Ocen, NHMUK , Discovery Stn 7709#73. ) right mndile; ) mxill 2; c) mxilliped; d) mxill 1; e) left mndile. Scle rs:, c; 200 μm:, d, e; 100 μm. loe with posterior nd distl mrgins setose; propodus strongly expnded, s long nd s wide s crpus, nterior mrgin convex, plm convex, trnsverse, crenellte, setose; dctylus flcte, s long s plm. Gnthopod 2 (Fig. 8): cox s long s cox 1, tpering distlly, pex truncte with few sete t the distl mrgin; sis surectngulr, with posterodistl group of plumose sete nd n nterodistl group of simple sete; merus, posterodistl loe nrrow, setose; crpus strongly expnded, wider thn propodus, posterodistl loe sucute, posterior nd distl mrgins setose; propodus s long s crpus, expnded distlly, plm convex, crenellte; dctylus slender, curved, s

12 236 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... Figure 8. Oedicerin vderi sp. n., mle holotype, 7.3 mm; northest Atlntic Ocen, NHMUK , Discovery Stn 7709#73. ) gnthopod 1; ) gnthopod 2. Scle rs:, ; 200 μm.

13 Zoosyst. Evol. 90 (2) 2014, c Figure 9. Oedicerin vderi sp. n., mle holotype, 7.3 mm; northest Atlntic Ocen, NHMUK , Discovery Stn 7709#73. ) pereopod 3; ) pereopod 4; c) pereopod 5. Scle rs: c; 200 μm. long s plm. Pereopod 3 (Fig. 9): cox suequl to cox 2, pex rounded, wekly setose; sis shorter thn cox, long plumose sete distlly long posterior nd nterior mrgins; merus expnded nterodistlly, setose; crpus 1.3 length nd out s wide s merus, posterior mrgin densely setose; propodus ovl, nterodistl nd posterior mrgins setose; dctylus lnceolte, suequl to propodus. Pereopod 4 (Fig. 9): cox wider thn long, nterior mr-

14 238 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... c d f e g Figure 10. d, f) Oedicerin vderi sp. n., mle holotype, 7.3 mm; northest Atlntic Ocen, NHMUK , Discovery Stn 7709#73. ) pereopod 6; ) pereopod 7; c) uropod 3; d) uropod 2, peduncle; f) telson; g) uropod 1. Femle prtype, 6.3 mm; sme loclity. e) pleonite 3 nd urosome with telson, uropods not shown, Scle rs: d; 200 μm: e; 1 mm.

15 Zoosyst. Evol. 90 (2) 2014, gin rodly rounded, distl mrgin stright, posterodistl loe very strong, surectngulr; sis shorter thn cox, nterior nd posterior mrgins setose distlly; merus expnded nterodistlly, setose; crpus 0.8 merus, ut suequl in width, posterior mrgin strongly setose, long sete nterodistlly; propodus, nterior mrgin setose; dctylus, stright, lnceolte, 1.5 propodus. Pereopod 5 (Fig. 9c): cox out s deep s cox 4, iloed, posterior loe expnded distlly, distl mrgin stright, nterior loe 0.7 length of posterior loe, rounded distlly; sis shorter thn cox; merus s long s sis; crpus 0.3 length of merus; propodus slender, surectngulr, 0.9 merus, out s long s stright lnceolte dctylus; rticles 2 6 vriously setose. Pereopod 6 (Fig. 10): cox 0.8 length of cox 5, iloed, posterior loe long, distl mrgin rounded, nterior loe surectngulr, 0.4 length of posterior loe; sis surectngulr; merus, posterior mrgin wekly convex; crpus surectngulr, 0.4 length of merus; propodus nd stright dctylus s long s merus; rticles 2 6 vriously setose. Pereopod 7 (Fig. 10): long; cox wider thn long, ovl, posterodistl corner rounded; sis posterior mrgin wekly convex, nterior mrgin strongly convex; merus elongte; crpus, 0.9 merus; merus nd crpus with groups of short sete on nterior nd posterior mrgins; [propodus nd dctylus unknown]. Pleon. Pleonites 1 3 (Fig. 6): smooth, lcking crine or teeth. Epimer (Fig. 6e): 1 nd 3 evenly rounded; epimeron 2, posterodistl ngle produced, rounded, posterior mrgin sinuous. Pleopod 1: peduncle stout, 0.8 length of rmi; rmi suequl in length. Urosome. Urosomite 1 (Fig. 10e): longest, with low oss close to the posterior mrgin; urosomite 3 longer thn 2, with short, cute mid-dorsl projection. Uropod 1: peduncle elongte, lterl mrgin with roust sete, mesil mrgin setose; inner rmus, [roken], oth mrgins setose; outer rmus 0.7 length of peduncle, sete on lterl mrgin only. Uropod 2 (Fig. 10d): peduncle not tpering, oth mrgins with short sete; [rmi dmged], inner rmus, oth mrgins setose; outer rmus, lterl mrgin setose. Uropod 3 (Fig. 10c): peduncle short, out s long s telson, with ventrl sucute projection; outer rmus just longer thn inner rmus. Telson (Fig. 10f) tpered, notched 40%. Vriility. The prtypes er smll posteriorly directed tooth on pleonite 3. It my e tht this process hs een present nd is worn down in the holotype. Antenn 1 of the femle (Fig. 6) hs longer nd more slender peduncle nd fewer nd more elongte flgellum rticles compred to the mle. Distriution. North Atlntic, south of Icelnd, m. Oedicerin loerze sp. n. Figs Holotype. Mle, 8.5 mm; NIWA 84727, TAN , Chthm Rise, S E, m, 5 April Prtypes. Mle?, 7.7 mm; NIWA 89970, TAN , Chthm Rise, S E, m, 24 April 2007; ovig. femle, 7 mm; NIWA 84740, TAN , Chthm Rise, S E, m; 9 April Etymology. The species is nmed for Dr. Anne-Nin Lörz to cknowledge her significnt contriutions to mphipod systemtics. Description. Holotype mle, 8.5 mm. Hed (Fig. 11): longer thn high, somewht longer thn pereonites 1 2 comined; no eyes or oculr pigment visile; rostrum (Fig. 11c) strongly deflexed, the ventrl mrgin wekly convex. Antenn 1 (Fig. 12): shorter thn ntenn 2; length rtios of peduncle rticles 1 3 1:0.5:0.3; flgellum 19-rticulte, first rticle s long s peduncle rticle 3, proximl rticles wider thn long; ccessory flgellum 1-rticulte, minute, 0.3 length of primry flgellum rticle 1. Antenn 2 (Fig. 12c): peduncle setose; length of rticle rticle 5; flgellum 23-rticulte, 1.9 length of peduncle rticle 5. Upper lip (lrum) (Fig. 11): wider thn long, truncte piclly. Mndile (Fig. 12, d): incisor 5-dentte; left lcini moilis wide nd multidentte, right nrrower; plp 3-rticulte, rticle 3 tpered, length rtios rticles 1 3 1:3.9:4.9. Lower lip (Fig. 11d): inner loes short nd rod, hypophryngel gp wide, outer loe mndiulr processes cute. Mxill 1 (Fig. 11e): inner plte tpered, with one distl set; outer plte with nine cute setl-teeth; plp 2-rticulte, rticle length of rticle 1. Mxill 2 (Fig. 12f): inner plte 1.1 wider thn outer plte; oth pltes with reltively sprse picl settion. Mxilliped (Fig. 13): inner plte (Fig. 12g) short, extending just eyond se of plp rticle 1; outer plte (Fig. 12e) extending 30% long plp rticle 2; concve medilly; plp (Fig. 13) 4-rticulte; rticle 1 tpered; rticle 2 rod, strongly expnded medilly, loe rodly rounded; rticle 3 nrrow proximlly, expnded mediodistlly; rticle 4 cute, wekly curved: length rtios of rticles 1 4 1:1.7:0.7:1.2. Pereon. Pereonite 1 (Fig. 11) longer thn 2; pereonite 2 suequl to 3; pereonite 7 longest. Gnthopod 1 (Fig. 13c): cox sutringulr, posterior mrgin stright, nterodistl corner rounded, posterodistl corner surectngulr, distl mrgin stright, setose; sis expnded distlly, posterior mrgin with scttered sete, nterior mrgin distl hlf with row of long sete; merus, posterodistl loe rounded, setose; crpus sutringulr, strongly expnded distlly, nterior, distl nd posterior mrgins setose; propodus strongly expnded distlly, s wide nd s long s crpus, nterior nd posterior mrgin convex, plm trnsverse, convex, crenellte, setose; dctylus slightly curved, just longer thn plm. Gnthopod 2 (Fig. 14, ): cox s long s cox 1, nterior nd posterior mrgins suprllel, pex truncte, wekly setose; sis little expnded, with some plumose sete ner posterior nd distl mrgins nd n

16 240 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... c d e Figure 11. Oedicerin loerze sp. n., mle holotype, 8.5 mm; southwest Pcific Ocen, NIWA 84727, TAN ) hitus; ) upper lip; c) rostrum; d) lower lip; e) mxill 1. Scle rs: ; 1 mm: e; 100 µm.

17 Zoosyst. Evol. 90 (2) 2014, c d e f g Figure 12. Oedicerin loerze sp. n., mle holotype, 8.5 mm; southwest Pcific Ocen, NIWA 84727, TAN ) ntenn 1; ) left mndile; c) ntenn 2; d) right mndile; e) mxilliped, outer plte; f) mxill 2; g) mxilliped, inner plte. Scle rs:,, d g; 100 µm: c; 200 µm.

18 242 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... c Figure 13. Oedicerin loerze sp. n., mle holotype, 8.5 mm; southwest Pcific Ocen, NIWA 84727, TAN ) outline of mxilliped; ) mxilliped plp; c) gnthopod 1. Scle rs: c; 200 µm.

19 Zoosyst. Evol. 90 (2) 2014, c d Figure 14. Oedicerin loerze sp. n., mle holotype, 8.5 mm; southwest Pcific Ocen, NIWA 84727, TAN ) gnthopod 2; ) gnthopod 2, detil of plm; c) pereopod 3, settion of crpus nd propodus omitted; d) pereopod 4. Scle rs:, c, d; 200 µm.

20 244 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... c Figure 15. Oedicerin loerze sp. n., mle holotype, 8.5 mm; southwest Pcific Ocen, NIWA 84727, TAN ) pereopod 5, ) pereopod 7; c) pereopod 6. Scle rs: c; 200 µm.

21 Zoosyst. Evol. 90 (2) 2014, Figure 16. Oedicerin loerze sp. n., mle holotype, 8.5 mm; southwest Pcific Ocen, NIWA 84727, TAN ) pleopod 1; ) telson; c) uropod 1; d) uropod 2; e) uropod 3. Scle rs: ; 200 µm; e; 100 µm. nterodistl group of long simple sete; merus, with ngulr posterodistl loe short, nrrow, setose; crpus strongly expnded, wider thn propodus, posterodistl loe sucute, distl mrgin olique; propodus shorter thn crpus, expnded distlly, plm stright, crenellte; dctylus curved, just longer thn plm. Pereopod 3 (Fig. 14c): cox suequl to cox 2, pex truncte; sis shorter thn cox, long plumose sete on posterior mrgin nd close to nterior mrgin; merus wekly expnded nterodistlly; crpus, length nd redth suequl to merus; propodus surectngulr, nterodistl nd posterior mrgins setose; dctylus 1.4 length of propodus. Pereopod 4 (Fig. 14d): cox wider thn long, nterior mrgin wekly convex, distl mrgin rodly rounded, nterodistl ngle surectngulr, posterodistl loe very strong, posterodistl ngle rounded; sis shorter thn cox, nterior nd posterior mrgins setose; merus wekly expnded nterodistlly, setose; crpus 0.9 merus, posterior mrgin convex, strongly setose, long sete nterodistlly; propodus, nterodistl mrgin strongly setose, posterior mrgin setose; dctylus stout, stright, 1.8 propodus. Pereopod 5 (Fig. 15): cox 0.9 length of cox 4, iloed, posterior loe expnded distlly, distl mrgin stright, nterior loe 0.6 length of posterior loe, rounded distlly; sis shorter thn cox, few plumose sete on ech mrgin; merus s long s sis, crpus 0.4 length of merus; propodus slender, 0.7 length of merus, shorter thn the stright lnceolte dctylus; rticles 2 6 vriously setose. Pereopod 6 (Fig. 15c): cox 0.7 length of cox 5, iloed, posterior loe sutringulr, nterior loe 0.5 length of posterior loe; sis surectngulr, long plumose sete nterodistlly; merus posterior mrgin convex; crpus surectngulr, 0.5 length of merus; propodus 0.9 length of merus; dctylus stright, lnceolte, s long s merus; rticles 3 6 vriously setose. Pereopod 7 (Fig. 15): long; cox wider thn long, rounded posterodistlly; sis, mrgins convex, posterodistl loe nerly

22 246 Colemn, C. O. & Thurston, M. H.: A redescription of the type species of Oedicerin... s long s ischium; merus with groups of short sete on nterior nd posterior mrgins [distl rticles unknown]. Pleon. Pleonites: 1 2 (Fig. 11) with mid-dorsl, posteriorly directed crinte teeth; pleonite 3 lcking crin nd tooth. Epimer: 1 nd 3 rodly rounded posterodistlly, epimeron 2 wekly ngulr. Pleopod 1 (Fig. 16): peduncle nd rmi suequl. Urosome. Urosomite 1 (Fig. 11): longest, with inconspicuous short oss close to the posterior mrgin; urosomites 2 nd 3 suequl in length, lcking dorsl projections. Uropod 1 (Fig. 16c): peduncle elongte, lterl mrgin with dense row of short sete, inner mrgin with fewer nd longer sete; inner rmus 0.7 length of peduncle, oth mrgins setose; outer rmus 0.9 inner rmus, lterl mrgin setose. Uropod 2 (Fig. 16d): peduncle not tpering, oth mrgins with short sete; inner rmus 0.9 length of peduncle, oth mrgins setose; outer rmus 0.9 length of inner rmus, lterl mrgin setose. Uropod 3 (Fig. 16e): peduncle short, out s long s telson; rmi suequl, 2.4 length of peduncle, outer rmus with plumose sete on the lterl mrgin. Telson (Fig. 16) tpered, notched 34%. Distriution. Chthm Rise, est of New Zelnd. Remrks. The femle specimen hs the sme ntenn 1 morphology s the mle: short peduncle rticles nd numerous flgellum rticles. The proximl rticles of the flgellum re shorter thn wide. Oedicerin sp. indet. Mteril exmined. 1 incomplete femle; NHMUK , Discovery Stn 7845: north-estern Atlntic, off the cost of Western Shr: N W N W, 24 Mrch 1972, BN 2.4, m. Remrks. Only the hed nd pereonites 1 2 re present. Coxe 1 2 er long sete long the distl mrgins. The niml ppers similr to O. ingolfi, ut s result of incompleteness it is impossile to ttriute it to ny species. Discussion The three species descried herein re morphologiclly very similr. Mouthprts nd ppendges show only minor nd sutle differences nd the species re est discriminted y hitus chrcters. Two of the species, O. ingolfi nd O. loerze sp. n. hve mid-dorsl crine on pleonites 1 2. Oedicerin ingolfi differs from O. loerze sp. n. in hving smll, slender, cute, upright tooth on pleonite 3 nd smll pointed process on the posterior mrgin of urosomite 3 oth of which re sent in O. loerze sp. n. Oedicerin vderi sp. n. hs smll pointed process on the posterior mrgin of urosomite 3, s found in O. ingolfi, ut pleonites 1 2 re evenly vulted lcking ny trce of crin. Pleonite 3 of the holotype of O. vderi sp. n. ppers to e dorslly unrmed, ut the prtypes hve smll cute process (see femle prtype, 6.3 mm in Fig. 10e). Cox 5 of O. ingolfi is longer thn wide, tht of O. vderi sp. n. is out s wide s long, nd tht of O. loerze sp. n. much wider thn long. Oedicerin sp. indet. ws collected from the wrm-temperte est Atlntic Ocen off Western Shr. The unique specimen is incomplete, only the hed nd pereonites 1 nd 2 re present, preventing full identifiction. Aprt from the type species O. ingolfi, two other species hd een descried in the genus prior to this study: Oedicerin meglopod Ledoyer, 1986, collected close to Myotte, Mozmique Chnnel, western Indin Ocen ( m) nd Oedicerin denticult Hendrycks & Conln, 2003 from the northest Pcific Ocen off Cliforni (4050 m). Knowledge of O. meglopod is limited s the unique specimen is incomplete, ut the mssive rostrum of this species differs mrkedly from ll other species of the genus. The plm of gnthopod 2 of O. meglopod is stright, similr to tht of O. loerze sp. n., nd thus different from the convex pttern seen in oth North Atlntic species. In Oedicerin denticult pleonite 1 is smooth nd pleonites 2 nd 3 hve posteromrginl process. The process on pleonite 3 is directed posteriorly nd is reminiscent of tht seen in prtype mteril of O. vderi sp. n. thus contrsting with the upright condition found in O. ingolfi. The posterior mrgin of urosomite 3 ers smll process (s do ll species except for O. loerze sp. n.) nd urosomite 1 hs smll upright process in the mle ut not the femle, chrcter unique within the genus. Cox 5 ppers to e longer thn wide, s in O. ingolfi. The mouthprts nd ppendges of ll species of this genus re remrkly similr to ech other. Exmintion of the extensive Norwegin Se mteril which we ttriute to O. ingolfi indictes tht intrspecific vriility is miniml, except for sexul dimorphism in ntenn 1. In femles of oth Atlntic species peduncle rticles of ntenn 1 re longer nd more slender thn in mles. Flgellum rticles in femles re uniformly much longer thn wide nd reltively few in numer, wheres in mles they re more numerous nd proximlly wider thn long, forming n incipient cllynophore. This sexul dimorphism is not pprent in O. loerze sp. n. where the structure of ntenn 1 is very similr in mles nd the one ovigerous femle prtype. Becuse of miniml differences mong ppendges nd mouthprts in Oedicerin species, differentition within the genus relies significntly on ptterns of ornmenttion of pleonites nd urosomites. As the posterior segments of the type mteril of O. ingolfi re missing, the question remins s to which of the two Atlntic species represents the species tht Stephensen descried. We llocte the mteril from the museums in Bergen nd Lund studied herein to O. ingolfi on geogrphicl grounds in tht it ws collected much closer to the type loclity of tht species, nd on the morphologicl grounds of the dense fringe of sete on the distl mrgins of coxe 1 nd 2 nd the shpe of the rostrum tht our mteril shres with Stephensen s originl description.

23 Zoosyst. Evol. 90 (2) 2014, Acknowledgments We re very grteful to Wim Vder for inviting us to the highly successful identifiction workshop t Skiotn which provided the mteril tht stimulted the present pper. We cknowledge with thnks Mr. Jon Anders Kongsrud (University of Bergen), Dr Lrs Lunqvist (Museum of Zoology, Lund University) nd Ms Sdie Mills (Ntionl Institute of Wter nd Atmosphere Reserch, Wellington, New Zelnd) for the lon of mteril. We thnk Ms Cmill Norrg, who drew the ppendges of O. ingolfi nd mde the line drwings of O. vderi sp. n. References Brnrd JL, Krmn GS (1991) The fmilies nd gener of mrine gmmriden Amphipod (except mrine gmmroids). Records of the Austrlin Museum 13(1/2): Brttegrd T, Fosså JH (1991) Replicility of n epienthic smpler. Journl of the Mrine Biologicl Assocition of the United Kingdom 71(1): doi: /S Brenke N (2005) An epienthic sledge for opertions on mrine soft ottom nd edrock. Mrine Technology Society Journl 39(2): doi: / Colemn CO (2003) Digitl inking : How to mke perfect line drwings on computers. Orgnism, Diversity nd Evolution, Electronic Supplement 14: Colemn CO (2009) Drwing sete the digitl wy. Zoosystemtics nd Evolution 85(2): doi: /zoos Dhl E (1979) Amphipod Gmmride from the deep Norwegin Se. A preliminry report. Srsi 64: Dhl E, Luier L, Siuet M, Strömerg JO (1976) Some quntittive results on enthic communities of the deep Norwegin Se. Astrte 9: Guennegn Y, Mrtin V (1985) Techniques de prélèvement. In: Luier L, Monniot C (Eds) Peuplements profonds de Golfe de Gscogne. IFREMER, Brest, Hendrycks EA, Conln KE (2003) New nd unusul yssl gmmriden Amphipod from the north-est Pcific. Journl of Nturl History 37(19): doi: / Knox MA, Hogg ID, Pilditch, CA, Lörz AN, Nodder SD (2012) Aundnce nd diversity of epienthic mphipods (Crustce) from contrsting thyl hitts. Deep-Se Reserch I 62: 1 9. doi: /j.dsr Krpp-Schickel T, Vder W (2013) Leucothoid nd merid mphipods (Crustce) from deep regions of the North Atlntic. Helgolnd Mrine Reserch 67: doi: /s Ledoyer M (1986) Crustcés mphipodes gmmriens. Fmilles des Hustoriide á Vitjzinide. Fune de Mdgscr 59(2). OR- STOM Institut Frnçis de Recherche Scientifique pour le Dévelopement en Coopértion, Pris, Rothliserg PC, Percy WG (1977) An epienthic smpler used to study the ontogeny of verticl migrtion of Pndlus jordni (Decpod, Cride). Fishery Bulletin. Ntionl Ocenogrphic nd Atmospheric Administrtion of the United Sttes 74(4): Stephensen K (1931) Crustce Mlcostrc VII (Amphipod III). The Dnish Ingolf-Expedition 3(11): d Udekem d Acoz C (2010) Contriution to the knowledge of Europen Liljeorgiide (Crustce, Amphipod), with considertions on the fmily nd its ffinities. Bulletin de l Insititut Royl des Sciences Nturelles de Belgique, Biologie 80: