q 2004 by the American Society of Ichthyologists and Herpetologists

Copeia, 2004(2), pp. 326–335

Bathyraja cousseauae sp.n.: A New Softnose Skate from the SouthwesternAtlantic (Rajiformes, Rajidae)

JUAN MARTıN DıAZ DE ASTARLOA AND EZEQUIEL MABRAGANA

A new species of softnose skate, Bathyraja cousseauae, is described from two adultand 19 juvenile specimens collected from different localities of the southwesternAtlantic. Bathyraja cousseauae is distinguished from all other southwestern Atlanticsoftnose skate species by its color pattern, squamation pattern of dorsal surface,lack of interspace between dorsal fins, and clasper morphology. The new specieshas a round pale area ocellus-like, margined with dark brown on posterior part ofeach pectoral base of dorsal surface, paler and dark spots over the disk, a contin-uous row of 21–27 median thorns from nuchal region to first dorsal fin and upperside of disc densely covered with dermal denticles. Males have rod-shaped claspersnot expanded at proximal section of glans as it does in Bathyraja brachyurops. Be-sides the external morphological features, skeletal characteristics (neurocranium,scapulocoracoids, and claspers) are also described and illustrated here.

Una nueva especie de raya de hocico blando, Bathyraja cousseauae, es descriptasobre la base de dos ejemplares adultos y 19 ejemplares juveniles capturados endiferentes localidades del Atlantico suroccidental. Bathyraja cousseauae se distinguede otras especies de rayas de hocico blando del Atlantico suroccidental por su pa-tron de coloracion, patron de espinulacion de la superficie dorsal, ausencia de es-pacio entre las aletas dorsales y por la morfologıa del clasper. La nueva especiepresenta una mancha palida ocelar rodeada de un anillo marron oscuro en la parteposterior de la base de cada aleta pectoral en la superficie dorsal, manchas palidasy oscuras dispersas sobre el disco, una hilera continua de 21 a 27 espinas medialesdesde la region nucal a la primera aleta dorsal y el lado superior del disco densa-mente cubierto de dentıculos dermicos. Los machos presentan claspers en formade vara no expandidos en la region proximal como sı ocurre en Bathyaraja brach-yurops. Ademas de los aspectos morfologicos, tambien se describen e ilustran lascaracterısticas osteologicas (neurocraneo, escapulocoracoides y claspers).

THE rajid genus Bathyraja Ishiyama is circum-global and the most diverse among all gen-

era of skates (Ishihara and Ishiyama, 1986). Spe-cies of this genus occur mainly in shelf watersat temperate and cold temperate latitudes andon continental slopes down to the deep-seaplains. More than 40 nominal species areknown (Ishihara and Ishiyama, 1986; Steh-mann, 1986). Compagno (1999) recognized 41valid species and at least two undescribed spe-cies worldwide. However, he assigned some spe-cies, previously situated in the genus Bathyrajaby several authors (Menni et al., 1984; Steh-mann, 1986; McEachran and Dunn, 1998) toRhinoraja Ishiyama. The genus is represented inthe southwestern Atlantic by 10 species (Steh-mann, 1986), seven of which are endemic onthe Argentinean continental shelf (Menni andStehmann, 2000). Species of Bathyraja havebeen a dominant proportion of the by-catch intarget fisheries for bony fishes on the south-western Atlantic shelf and at present are in-creasingly exploited as target species, since tra-

ditional bony fish stocks for the commercialfisheries became drastically depleted. Herein wedescribe a new species of Bathyraja from thesouthwestern Atlantic.

MATERIALS AND METHODS

Specimens examined were collected by theR/Vs CAPITAN OCA BALDA and DR. EDUARDO

HOLMBERG of the Instituto Nacional de Investi-gacion y Desarrollo Pesquero (INIDEP), and bythe F/V FEIXE of Uruguay, from different local-ities of the southwest Atlantic. Methods for mak-ing measurements and counts follow Hubbs andIshiyama (1968) and Ishihara and Ishiyama(1985). Two specimens of the new species (amale and a female) were dissected to examinethe skeletal structures (neurocranium, scapulo-coracoids, and clasper). Measurements of theneurocranium and scapulocoracoids were madeaccording to Hubbs and Ishiyama (1968), andMcEachran and Compagno (1979). Clasper ter-minology follows Stehmann (1970), Hulley

327DIAZ DE ASTARLOA AND MABRAGANA—NEW BATHYRAJA

(1972) and Ishihara and Ishiyama (1985), andcranium terminology follows Hulley (1972).The holotype and one of the paratypes were ra-diographed to verify skeletal observations basedon dissections and to count vertebrae and bothpectoral and pelvic radials. The holotype and 19paratypes are deposited in the fish collection ofthe Instituto Nacional de Investigacion y Desa-rrollo Pesquero, Mar del Plata, Argentina (INI-DEP). One paratype is housed in the fish col-lection of the Facultad de Ciencias, Universidadde la Republica, Montevideo, Uruguay (ZVC-P).

Bathyraja cousseauae, sp. nov.Figure 1, Appendix 1

Holotype.—INIDEP 719, 617 mm TL, juvenilemale, off northern Patagonian continentalshelf, 418219S, 578259W, 141 m, 2. x. 2001, R/VDR. EDUARDO HOLMBERG, cruise H-08/01, sta.31.

Paratypes.—INIDEP 720, 550 mm TL, juvenilemale, 720 mm disc width (tail broken), maturefemale, 750 mm TL, immature female, southMalvinas Islands, 528529S, 608159W, 277 m, R/VCAPITAN OCA BALDA, cruise OB-11/00, sta. 4, 13.x. 2000; INIDEP 721, 695 mm disc width (tailbroken), mature male, 575 mm TL, 610 mmTL, 515 mm TL, 650 mm TL, 860 mm TL, ju-venile males, 563 mm TL, 748 mm TL, 622 mmTL, 647 mm TL, 598 mm TL, 575 mm TL, 612mm TL, 412 mm TL, juvenile females, southIsla de los Estados, 548569S, 648209W, 397 m, R/V CAPITAN OCA BALDA, cruise OB-04/02, sta. 4,19. x. 2002; INIDEP 722, 535 mm TL, 540 mmTL, juvenile males, off northern Patagoniancontinental shelf, 418309S, 578359W, 119 m, 10.x. 2002, R/V DR. EDUARDO HOLMBERG, cruiseH-06/02, sta. 30; ZVC-P 4796, 502 mm TL, ju-venile male, off Mar del Plata, 388119S, 558189W,284 m, 25.vi.2003, F/V FEIXE, sta. 171.

Diagnosis.—Bathyraja cousseauae is characterizedby the combination of the following characters:A large and more distinct round pale area ocel-lus-like, margined with dark brown on posteriorpart of each pectoral base of the upper side ofdisc. Upper surface of disc completly coveredwith numerous small spinules, also placed oninterorbital area, tail, dorsal fins and posteriorpelvic lobes. In larger specimens the spinulesare scattered distributed in central parts of disc.No ocular thorns. A continuous row of 21–27median thorns from nuchal region to first dor-sal fin. Males with elongated, slender rod-shaped claspers. Dorsal fins densely spinulose

close to the end of tail and with no space be-tween them.

Description.—Measurements and counts are giv-en in Appendix 1. Differing values of the pa-ratypes following those for the holotype are inparentheses. Disc broadly rhombic, 1.26 timesas broad as long (1.15–1.40); margin of discnearly concave at sides of tip of snout, slightlyconvex to level of spiracles, nearly concave toouter corners that are abruptly rounded. Pos-terior margins of disc weakly convex with arounded inner corner to level of pelvic fins. An-terior margins of disc 1.46 times longer than theposterior ones (1.21–1.54). Preorbital length2.52 times (2.28–3.04) interorbital width. Orbitsmoderately large, their lengths 0.90 times(0.62–1.00) of the interorbital width. Orbitlength 1.37 times spiracle length (0.94–1.91).Distance between spiracles 1.69 times that be-tween orbits (1.30–1.78). Snout short, 8.45times in total length (6.83–8.62). Pelvic finsdeeply incised. Anterior lobe 0.89 times poste-rior lobe (0.75–1.05). Tail slender and moder-ately depressed with narrow lateral folds alongventrolateral surface originating at level of axilof pelvic fins, widening in posterior two thirdsof tail and extending to near tail tip. Length oftail from center of cloaca to tip 0.96 times thatfrom tip of snout to center of cloaca (0.90–1.20). Postdorsal fin tail length approximatelyone-third length base of second dorsal fin. Nospace between dorsal fins, neither between sec-ond dorsal fin and caudal fin. Height of dorsalfins approximately one-half of their bases’lengths. Preoral length 1.33 times mouth width(1.34–1.97). Preorbital length 2.80 times orbitlength (2.81–4.15). Nostril flaps are short, thickand tube-like. Anterior nasal flap (nasal cur-tain) well developed and fringed along distalmargin. Posterior nasal poorly developed andsmooth. Mouth transversally straight. Upperand lower jaws slightly arched, with 31 and 29(30–33, and 30–32) tooth rows, respectively. Dis-tance between first gill slits 2.23 times distancebetween nares (2.00–2.70). Distance betweenfifth gill slits 1.69 times internarial distance(1.40–2.07). Length of first gill slits 0.88 timeslength of fifth gill slits (1.00–1.60). First dorsalfin about equal in size and shape to second dor-sal fin. Vertebrae 38–39 (trunk) 1 78–80 (pre-dorsal caudal). Pectoral radials 86, pelvic radials18. Dorsal side of disc and tail densely coveredwith dermal denticles, with erect crowns andstellate bases. Posterior lobe of pelvic fin sparse-ly covered with dermal denticles. Ventral side ofdisc, pelvics, claspers and tail without dermaldenticles. Alar thorns well developed in one ma-

328 COPEIA, 2004, NO. 2

Fig. 1. Bathyraja cousseauae new species, holotype, INIDEP 719, 617 mm TL, juvenile male, off northernPatagonian continental shelf, Argentina. Scale bar 5 5 mm.

ture male of 695 mm DW and arranged in 4rows. Alar thorns of other males not yet devel-oped. Midrow thorns are continuous and widelyspaced on mid disc, with two thorns on nape,two on shoulder girdle, and four to six widelythorns between shoulder girdle and axil of pec-toral fins. Midline of tail with 14–19 strongthorns, with oval bases and backwardly directedcrowns.

Color when fresh.—A conspicuous white or paleocellus margined with brown on the hinder partof the base of each pectoral. Indistinct palerand dark spots of various sizes scattered over adark brownish colored disc. Darker spots morenumerous and rounded, some margined withdark brown. Thorns marked off pale milky-white. Rostral area and orbits as brown asground color of upper side, except side areas ofrostrum that are somewhat lighter, and one palemark on anterior inner side of each orbit. An-terior pelvic lobe edged whitish. Claspers dark

brown with lightened second half outer edgecreamy-white. Lateral tail folds brown. Two tothree incomplete pale cross-bands on the tail.Dorsal fins uniformly brown. Lower surface ofthe disc largely creamy-white, with posteriormargins of pectoral fins, and edges of posteriorlobes of pelvic fins narrowly edged dusky brown.Underside of tail from level of tips of pelvic’sposterior lobes to the tip of tail varies from afew dark brown spots to uniformly brown pig-mented pattern.

Etymology.—The new species is dedicated toProf. Dr. Marıa Berta Cousseau (Departamentode Ciencias Marinas, Universidad Nacional deMar del Plata, Argentina) in recognition of hercontribution to the marine fishes of Argentina.

Common name.—New English name: Joined-finsSkate; new Spanish name: Raya Aletas Juntas.

Neurocranium.—Neurocrania measurements of ajuvenile male (550 mm TL) and a juvenile fe-

329DIAZ DE ASTARLOA AND MABRAGANA—NEW BATHYRAJA

←

Fig. 2. Neurocranium of Bathyraja cousseauae, n.sp. (paratype, INIDEP 720, juvenile male, 550 mmTL) in dorsal (A), ventral (B), and lateral (C) views.Figure abbreviations: ac 5 anterior cerebral vein fo-ramen; af 5 anterior fontanelle; es 5 efferent spirac-ular artery foramen; ic 5 internal carotid artery fo-ramen; into 5 interorbital vein foramen; nc 5 nasalcapsule; of 5 orbital fissure; onc 5 orbitonasal canalforamen; os 5 optic stalk; pf 5 posterior fontanelle;poc 5 preorbital canal foramen; rb 5 rostral base; rs5 rostral shaft; II 5 optic nerve foramen; III 5 ocu-lomotor nerve foramen; VII 5 hyomandibular branchof facial nerve foramen. Scale bar 5 10 mm.

male (750 mm TL) are given as percentage ofnasobasal length. Cranial length 172.9–177.3;cranial width 103.0–105.9; cranial height 22.7–23.5; rostral shaft uncalcified, its length 72.7–76.5; rostral base 34.8–35.3; nasal capsulesrhomboid shaped, with straight anterior mar-gins, set at about 108 angle to transverse axis ofneurocranium; foramen for profundus nerveon leading edge of nasal capsule; internasalwidth 20.0–21.2; interorbital width 34.8–38.8;anterior fontanelle heart-shaped with a little pe-duncle in its base, its length 27.3–32.9, its width6.1–14.1; width across otic capsules 53.0–56.5;least width of basal plate 27.3–29.4; greatestwidth of nasal aperture 37.9–41.2; foramen foranterior cerebral vein on a vertical with dorsalrim of optic nerve foramen, posterior to lineconnecting foramina of preorbital and orbito-nasal canals; foramen for oculomotor nervedorsal to optic stalk; foramen for interorbitalvein anterior to orbital fissure and posterior toefferent spiracular artery foramen; basal and in-ternasal plates relatively narrow (Fig. 2).

Scapulocoracoids.—Scapulocoracoid elongated,expanded between mesocondyle and metacondy-le; anterior fenestra divided by an anteriorbridge into an anterodorsal and anteroventral fe-nestra; four postdorsal foramina: first, secondand fourth of about the same size, and third mi-nute (males); six postdorsal foramina: first, sec-ond and sixth are big and the third, fourth andfifth minute and form an inverted triangle (fe-males); seven small postventral foramina(males), eight postventral foramina (females), allof which are of about equal in size and shape;dorsal margin relatively straight. Posterior mar-gin with a pronounced concavity in its dorsal side(males), rounded in females (Fig. 3).

Clasper.—Clasper fully developed, its length 37%of disc width. Large pseudosiphon (ps) presentnear outer lateral edge of dorsal lobe, its length

330 COPEIA, 2004, NO. 2

Fig. 3. Lateral view of left scapulocoracoids of a female (A) and a male (B) specimens of Bathyraja cousseauaen. sp. (paratypes, INIDEP 720, juvenile female, 750 mm TL, and juvenile male, 550 mm TL, respectively).Figure abbreviations: ab 5 anterior bridge; adf 5 anterodorsal fenestra; avf 5 anteroventral fenestra; msc 5mesocondyle; mtc 5 metacondyle; pdf 5 postdorsal foramina; prc 5 precondyle; pvf 5 postventral foramina;rc 5 rear corner; scp 5 scapular process. Scale bar 5 10 mm.

25% of clasper length; inner surface of dorsallobe with pseudorhipidion (pr) and cleft (cf);inner surface of ventral lobe with projection(pj), sentina (se) and spike (sp). Dorsal margin-al (dM) with pointed distal elongation, formingpseudorhipidion externally; ventral marginal(vM) with pointed distal elongation, extendingto the tip of the clasper and forming projectionexternally; dorsal terminal 1 (dT1) shield-like,with notch along proximal and distal margins,curved around the axial onto ventral side andconnected with ventral terminal forming pseu-dosiphon at its outer edge; dorsal terminal 2(dT2) rectangular, its length half of the lengthof dT3, dT2 and dT3 firmly connected; dorsalterminal 3 (dT3) long and slender, extendingto tip of clasper; accessory terminal 2 (aT2)small, j-shaped, lying beneath tip of ventral mar-ginal and forming spike externally; ventral ter-minal (vT) spoonlike overlying tip of ventralmarginal and accessory terminal; axial cartilage(Ax) spatulate distally (Fig. 4).

DISCUSSION

Comparisons.—Ten species of Bathyraja occur inthe southwestern Atlantic (Stehmann, 1986). Ofthese, seven are known from the Argentineanand Uruguayan continental shelves (Cousseauet al., 2000). Bathyraja cousseauae is most similarto the Broadnose Skate Bathyraja brachyurops(Fowler). Bathyraja brachyurops has light spotsand blotches on the light brown colored dorsal

surface, has midrow thorns same color as disc,and ventral side of tail is largely white. Bathyrajacousseauae has light and dark colored spots overa dark brownish colored disc, midrow thornsmilky white, and underside of tail partly or en-tirely brown. Bathyraja brachyurops has a discon-tinuous row of mid row thorns on disc. Thoseof the new species are continuous but smallerand more widely spaced along the mid disc.Bathyraja brachyurops has an interspace andthorn between dorsal fin bases and lacks den-ticles on dorsal fins. Bathyraja cousseauae lacks aspace between dorsal fins which are densely cov-ered with denticles. The claspers of B. brachyu-rops are greatly expanded laterally along proxi-mal aspect of glans. Those of the new speciesare rod-shaped and little expanded distally (Fig.5).

Ventral coloration distinguishes the new spe-cies from the Butterfly Skate Bathyraja papilio-nifera Stehmann, the Whitemouth Skate Bathy-raja schroederi Kreft and the Dark-belly SkateBathyraja meridionalis Stehmann. Bathyraja papi-lionifera has outer corners and posterior mar-gins of underside of disc broadly edged darkand a characteristically butterfly-shaped palebrown blotch around anterior anus. Bathyrajaschroederi and B. meridionalis have plain colora-tion on underside of disc. That of the new spe-cies is creamy-white. Lack of ocular and scapularthorns, absence of an interdorsal space distin-guishes the new species from the PatagonianSkate Bathyraja macloviana (Norman), the Ma-

331DIAZ DE ASTARLOA AND MABRAGANA—NEW BATHYRAJA

Fig. 4. Right clasper cartilages of Bathyraja cousseauae, n. sp. (paratype, INIDEP 720, adult male, 695 mmDW) in external (A), dorsal (B and C) and ventral views (D and E). Figure abbreviations: aT2 5 accesoryterminal 2; Ax 5 axial; dM 5 dorsal marginal; cf 5 cleft; dT1 5 dorsal terminal 1; dT2 5 dorsal terminal 2;dT3 5 dorsal terminal 3; pj 5 projection; pr 5 pseudorhipidion; ps 5 pseudosiphon; se 5 sentina; sp 5 spike;vM 5 ventral marginal; vT 5 ventral terminal.Scale bar 5 10 mm.

gellan Skate Bathyraja magellanica (Philippi) andthe Multispine Skate Bathyraja multispinis (Nor-man).

Snout length distinguishes the CupheadSkate Bathyraja scaphiops (Norman) from thenew species. The former has a longer (25% ofwidth disc) translucid pointed snout, whereas

the latter has a shorter (17 to 18% width disc)obtuse, triangular-shaped and not translucidsnout. Presence of thorns along the midline ofthe disc distinguishes the new species from theGraytail Skate Bathyraja griseocauda (Norman). Acontinuous midrow of thorns and wider inter-orbital width distinguishes the new species from

332 COPEIA, 2004, NO. 2

Fig. 5. Dorsal view of left claspers of Bathyraja cous-seauae, n. sp. (A) (holotype, INIDEP 719, 617 mm TL,juvenile male) and Bathyraja brachyurops (B). Arrowindicates lateral process. Scale bar 5 10 mm.

Fig. 6. Geographic distribution of Bathyraja cous-seauae based on material collected. Symbols representmore than one capture. Square indicates original lo-cality of holotype.

the White-Dotted Skate Bathyraja albomaculata(Norman).

Distribution.—Type specimens of B. cousseauaewere collected off Mar del Plata at 284 m depth,off northern Patagonian continental shelf at119–141 m depth, southern Malvinas Islands at277 m depth and in the southernmost tip of thesouthwest Atlantic, off southern Isla de los Es-tados, at 397 m depth (Fig. 6). According to thecapture localities of the type specimens, the newspecies appears to have a wide distribution, atleast on the upper slope of the southwest Atlan-tic. Similar distributional pattern was observedin other Bathyraja species such as B. griseocauda,B. multispinis, and B. scaphiops (Cousseau et al.,2000). These species have a restricted range dis-tributions and show a strict-stenothermous-

stenohaline behavior, in relation to water mas-ses (Figueroa et al., 1999). Bathyraja cousseauaeseems to occur sympatrically with those softnoseskates primarily inhabiting the upper slope ofthe southwest Atlantic. In contrast, B. magella-nica is more eurythermic as well as euryhaline,feature also shared with B. albomaculata, B. brach-yurops, and B. macloviana (Figueroa et al., 1999),although the former is more restricted to south-ern Patagonia between latitudes 478 and 558S,whereas the others are widely distributedthroughout the southwestern Atlantic continen-tal shelf (Cousseau et al., 2000). Bathyraja papi-lionifera and B. schroederi are regarded deep-wa-ter skates occurring in the midcontinental slopeoff northern Argentina (Stehmann, 1987; Steh-mann and Shulze, 1996). Bathyraja meridionalis,although described as a new deep-water skatefrom the eastern slope of subantarctic SouthGeorgias (Stehmann, 1987) was also reported asa common rajid species in the deep-water, long-line fishery for the Patagonian toothfish Dissos-tichus eleginoides, from the southern part of theFalklands Outer Conservation Zone (Agnew etal., 1999). Bathyraja cousseauae can be includedwithin the morphotype group of transitional

333DIAZ DE ASTARLOA AND MABRAGANA—NEW BATHYRAJA

species defined by Stehmann (1986) and char-acterized by small to medium-sized skates thatreach a total length to about 100 cm, with areduced dorsal thorn pattern lacking orbitaland scapular thorns.

Comparative material examined.—Bathyraja albo-maculata: INIDEP 303, 625 mm TL, male, 452mm TL, female, 388509S, 558339W, 16. vii. 1981,R/V CAPITAN CANEPA, cruise C-09/81. INIDEP439, 500 mm DW, male, 488209S, 618059W, 13.ii. 1992, R/V DR. EDUARDO HOLMBERG, cruiseH-01/92, sta. 75. INIDEP 491, 350 mm TL,male, 548019S, 628549W, 20. xi. 1995, R/V CA-PITAN OCA BALDA, cruise OB-13/95, sta. 14.Bathyraja brachyurops: INIDEP 302, 750 mm TL,male, 388509S, 558339W, 16.vii. 1981, R/V CA-PITAN CANEPA, cruise C-09/81, sta. 15. INIDEP

552, 530 mm TL, female, 508439S, 658429W, 113m, 20. iii. 1997, R/V CAPITAN OCA BALDA, cruiseOB-04/97, sta. 16. Bathyraja griseocauda: INIDEP437, 348 mm DW, female, 488209S, 618059W,13.ii. 1992, R/V DR. EDUARDO HOLMBERG,cruise H-01/92, sta. 75. INIDEP 613, 401 mmTL, male, 378349S, 548489W, 400 m, 14. ii. 1998,R/V CAPITAN OCA BALDA, cruise OB-01/98.Bathyraja macloviana: INIDEP 304, 540 mm TL,female, 388509S, 558339W, 16. vii. 1981, R/V CA-PITAN CANEPA, cruise C-09/81, sta. 15. INIDEP438, 417 mm DW, female, 488209S, 618059W, 13.ii. 1992, R/V DR. EDUARDO HOLMBERG, cruiseH-01/92, sta. 75. INIDEP 493, 305 mm TL, fe-male, 528269S, 638329W, 244 m, 26. xi. 1995, R/V CAPITAN OCA BALDA, cruise OB-13/95, sta. 37.INIDEP 497, 570 mm TL, male, 548309S,618249W, 140 m, 19.xi. 1995, R/V CAPITAN OCA

BALDA, cruise OB-13/95, sta. 11. INIDEP 554,285 mm TL, female, 430 mm TL, male, 508079S,668279W, 92 m, 20. iii. 1997, R/V CAPITAN OCA

BALDA, cruise OB-04/97, sta. 12. Bathyraja ma-gellanica: INIDEP 553, 510 mm TL, female,498549S, 648409W, 115 m, 19. iii. 1997, R/V CA-PITAN OCA BALDA, cruise OB-04/97, sta. 7. INI-DEP 556, 265, 333 mm TL, females, 508039S,668019W, 98 m, 19. iii. 1997, R/V CAPITAN OCA

BALDA, cruise OB-04/97, sta. 9. Bathyraja multis-pinis: INIDEP 495, 480 mm TL, female, 548039S,618289W, 150 m, 19. xi. 1995, R/V CAPITAN OCA

BALDA, cruise OB-13/95, sta. 12. INIDEP 498,640 mm TL, female, 538159S, 628049W, 485 m,23. xi. 1995, R/V CAPITAN OCA BALDA, cruiseOB-13/95, sta. 31. Bathyraja scaphiops: INIDEP588, 718 mm TL, male, 398049S, 558399W, 187m, 22. x. 1997, R/V DR. EDUARDO HOLMBERG,cruise H-10/97, sta. 105. INIDEP 589, 624 mmTL, male, 408429S, 568479W, 135 m, 21. x. 1997,R/V DR. EDUARDO HOLMBERG, cruise H-10/97,sta. 66.

ACKNOWLEDGMENTS

We express our sincere thanks to P. Ibanez forcollecting numerous paratype specimens, A.Loureiro for providing one of paratypes fromoff Mar del Plata, L. Paesch for providing de-tailed information on one of the paratypes andtaking photographs, V. Segura for drawing thefigures, M. Tobıo for taking the photographs,D. Figueroa and L. Lucifora for suggestions thatimproved the manuscript, and the captains,crew, and colleagues of the R/Vs DR. EDUARDO

HOLMBERG and CAPITAN OCA BALDA of the INI-DEP for logistical and field assistance. Helpfulcomments of anonymous reviewers that im-proved earlier versions of this manuscript aregreatfully acknowledged. This project was sup-ported from grants of the Universidad Nacionalde Mar del Plata.

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( JMDA) DEPARTAMENTO DE CIENCIAS MARINAS,FACULTAD DE CIENCIAS EXACTAS Y NATURALES,UNIVERSIDAD NACIONAL DE MAR DEL PLATA.FUNES 3350, B7602AYL MAR DEL PLATA, AR-GENTINA. CONSEJO NACIONAL DE INVESTIGACIO-NES CIENTıFICAS Y TECNICAS (CONICET); AND

(EM) MUSEO DEL MAR, AV. COLON 1114, MAR

DEL PLATA, ARGENTINA. E-mail: ( JMDA) astar-loa@mdp.edu.ar. Send reprint requests toJMDA. Submitted: 11 May 2003. Accepted: 22Dec. 2003. Section editor: D. G. Buth.

335DIAZ DE ASTARLOA AND MABRAGANA—NEW BATHYRAJA

APPENDIX 1. MORPHOMETRICS (IN MILLIMETERS) AND MERISTICS OF THE HOLOTYPE (INIDEP 719) AND 20 PARA-TYPES OF Bathyraja cousseauae. Range and mean values expressed in % of disc width (DW), except total length

and DW in mm. SD 5 standard deviation.

Holotype

Paratypes

Range Mean SD

Total lengthDisc widthDisc lengthSnout length (preorbital)Snout length (preoral)Snout to maximum widthPrenasal lengthOrbit diameter

6174173307373

1945926

412–860282–72071.1–86.717.1–20.816.9–22.141.6–49.111.8–16.74.4–6.3

77.618.619.645.314.25.5

3.01.11.51.81.30.5

Distance between orbitsOrbit and spiracle lengthSpiracle lengthDistance between spiraclesMouth widthDistance: nare to mouthDistance between nostrilsWidth: first gill openingsWidth: third gill openings

293319495520487

10

5.3–7.76.7–7.82.8–5.16.8–12.2

10.9–12.82.9–6.8

10.5–12.61.6–2.51.8–2.5

6.87.34.1

11.011.84.5

11.61.92.1

0.60.40.61.10.50.70.60.30.2

Width: fifth gill openingsDistance: first gill openingsDistance: fifth gill openingsLength: anterior pelvic lobeLength: posterior pelvic lobe

8107815359

1.3–2.122.5–31.416.5–22.011.5–14.912.9–17.6

1.624.919.412.913.9

0.21.81.20.91.0

Tail width at axil of pelvic finDistance: snout to cloacaDistance: cloaca to caudal tipDistance: snout to 1st dorsal finDistance: snout to 2nd dorsal finLength: anterior margin of pectoral fin

29315302567589290

6.1–8.567.0–75.767.2–80.7

123.1–138.1128.2–144.160.5–72.4

7.171.973.9

131.2137.066.6

0.72.74.13.23.63.1

Length: posterior margin of pectoral finLength: 1st dorsal fin baseLength: 2nd dorsal fin baseLength: caudal fin baseClasper lengthNumber of trunk vertebrae

19824227

3239

44.2–56.25.1–7.34.7–7.31.9–4.16.3–37.138–39

49.56.25.82.98.4

2.40.60.60.79.5

Number of predorsal caudal vertebraeNumber of pectoral raysNumber of pelvic raysUpper jaw tooth rowsLower jaw tooth rows

78

3129

78–808618

30–3330–32

861831.730.7

1.11.5