A new species of deep-water skate, Breviraja africana sp. n. (Pisces, Batoidea, Rajidae), from the Eastern Central Atlantic slope, and remarks on the taxonomic status of Breviraja Bigelow & Schroeder, 1948

zyxwvutsr zyxwv zyxwvu zyxwv zyxw Bull. Mus. natn. Hist. nut., Paris, 4e sér., 5, 1983, section A, n o 3 : 903-925. A new species of deep-water skate, Brevirajce ceficana sp. n. (Pisces, Batoidea, Wajidae), from the Eastern Central Atlantic slope, and remarks on the taxonomic status of Brwimja 18;igd~w& Schroeder, 31948 zyxwv by Mattias STEHMANN and Bernard SERET Abstract. - ‘Three specimens (1adult $, 2 99)of a deep-water dwarf species of skate were obtained by thc authors during the course o€their investigations for the batoid volume of ORSTOM’s ‘‘ Faune tropicale ”. These specimens from 900-1 030 m depth off Gabon are described as Breoii-aja africana sp. n., which in the Eastern Atlantic forms the geographic link between three species known from the NE-Atlantic and another o m from the SE-Atlantic off South ilfrica. MCEACHRAN & COMPAGNO(1982) revised the genus Brevira,ja Bigelow & Schroeder, 1948, mainly on anatomical characters such as neurocranium, scapulocoracoid, and skeleton of the clasper as well as its external morphology. The investigation by these authors resulted in ordering the species into newly defined taxa, i.e. Breviraja for the minority, and Neoraja gen. n. for the majority of species, the latter taxon being subdivided into the new subgenera Neoraja and Ferzestraja. Breviraja africana, as described in the present contribution based on the complete range of modern taxonomic characters, appears intermediate between Breviraja and Neoraja with regard t o the diagnostic features given by MCEACHRAN & COMPAGNO(1982). This mainly urged the present authors to briefly discuss the revision of the latter authors and t o propose a renewed consideration of the status of Breviraja Bigelow & Schroeder, 1948, t o which the new species is assigned preliminarily until, such a careful consideration of the generic situation can be undertaken a t another occasion. RB5umé. - Trois exemplaires (1mâle adulte et 2 femelles) d’une petite espèce de raie de profondeur ont été récoltés au cours des recherches effectuées dans le cadre de l’étude que nous menons’ sur les poissons batoïdes de l’Est-Atlantique tropical. La synthèse de cette étude fera l’objet d’un numéro spécial de la série (( Faune tropicale )) de I’ORSTOM. L’espèce nouvelle Breoiraja africana est décrite h partir de ces trois exemplaires, qui ont été chalutés par 900-1 030 m de profondeur au large des côtes du Gabon. Elle établit la liaison géographique entre les trois espèces connues de l’Atlantique-NE et celle de l’Atlantique-SE (Afrique du Sud). MCEACHRAN & COMPAGNO (1982) ont révisé le genre Breuiraja Bigelow & Schroeder, 1948, en se fondant principalement sur les caractères anatomiques tels que le neurocrâne, le complexe scapulocoracoïde, le squelette et la morphologie externe des ptérygopodes. Des recherches de ces auteurs, il résulte que les especes sont classées dans deux genres, i.e. Breuiraja redéfini par MCEACHRAN & COMPAGNO,qui comprend quelques-unes des espèces, et Neoraja gen. n. qui regroupe la plupart d’entre-elles j ce dernier taxon étant lui-même divisé en deux nouveaux sous-genres Neoraja et Fenestrqja. Breuiraja africana, dont la présente description est fondée sur l’intégralité des caractères taxonomiques modernes, apparaît intermédiaire entre Breviraja et Neoraja, selon les caractkristiques des diagnoses données par MCEACHRAN & COMPAGNO(1982). De cette situation, nous avons été amenés à débattre de la révision effectuée par MCEACHRAN & COMPAGNO,et à proposer une nouvelle réflexion sur c zy zy zyxwvutsr zyxwvutsrqpo zyxwvuts - 904 - le status du genre Breviraju Bigelow & Schroeder, 1948, auquel la nouvelle espèce est provisoirement rat.tach&e,en attendant qu’une telle réflexion puisse être entreprise. M. STETIIWANN, Au.ssenstelle Iclath,yologie, Institut f iir Seefischerei, Zool. &luseuna Universitüt fIambu,rg, ~l/lnrtin-Lrcther-Kin.g-Platz 3, 0-2000 Hamburg 13, Federal Repu.blic of Gernmny. B. SERET,ORSTOM, Direction générale, 34, me Bayard, 75008 Paris, France. INTRODUCTION I n the course of the authors’ cooperation in the ORSTOM research project. for the purpose of preparing the batoid volume of ‘‘ Faune tropicale ” one of us, B. SERET,worked in t h e ORSTOM Center in Dakar for several years in order t o study the local fauna and collection material in Gorée as well as Pointe Noire, and t o obtain study material from fishmarkets, fishermen and through participation in research cruises of various vessels. A number of species unknown for t h e Eastern Central At,lantic or even for science were discovered, partly in scientific collections, but mainly during deep- trawling surveys along t h e continental slope. It is the purpose of the present contribution t o introduce a new rajid species from t h e slope off Gabon prior t o t h e main inventory and revision publication. Other papers of this nature may follow. The three type specimens were collected b y B. SEREI during the survey Seraie with t h e RV ‘ Nizery ’ off Gabon in April 1980. The holotype and one paratype will be deposited in the collection of t h e Muséum national d’E-Iistoire naturelle (MNHN) in Paris, t,he other paratype in t h e collection of t h e Institut für Seefischerei in Hamburg (ISH). zyxwvu Breviraja africana sp. n. MATERIALEXAMINED : Holotype : MNHN 1983-1, adult 8 of 288 mm TL. RV ‘ Nizery ’ cruise Seraie, stat. 35, 18.IV.1980 j 03025’ S, 09633’ E, 900-1 030 m depth, Tb 4.356 C, 9.6 m otter trawl. - Two paratypes : ISH 129/80, Ç? 304 mm TL. Capture data as for the holotype. - MNHN 1983-2, Ç? 284 mm TL. RIr ‘ Nizery ’ cruise Seraie, stat. 32, 17.1V.1980 j 02041‘ S, 08051’ E, 900-930m depth, Tb 4.660 C, 9.6 m otter trawl. DIAGNOSIS A dwarf-species of rajid skate of the genus Bieviraja Bigelow & Schroeder, 1948, with a maximum total length of about 300 mm. The new species is characterized by the combination of the following characters : Disc almost heart-shaped, with broadly rounded outer corners. Snout very short, bluntly angled (about 1350), and with a short triangular integumental process a t its tip. Tail long and rather slender, its length just under 60 % of the TL. Anterior pelvic lobe slender and as long as about 75 yo of the length of the posterior lobe. The two small dorsal fins very posterior on tail and with confluent bases, caudal fin with a low ventral fold. Nasal flaps and rear margin of nasal curtain fringed. Anterior disc margins strongly undulated in adult males. Upper side of disc and tail entirely and densely set with coarse spinules, except for posterior disc margins and origin as well as center of pectorals in adult males. 3-4 small preorbital thorns and 1-3 in postorbital position. At most a single posterior nuchal thorn and one on each shoulder. zy zyx zyxw - 905 - No further thorns on disc, except for malar and alar thorns in adult males. A median row of 17-28 thorns from level of pelvic axils along anterior three fifths of tail length, the remaining section t o first dorsal fin a narrow, shallow groove without thorns or spinules. Lower side smooth, narrow strips of spinules may occur only along edges of tail. 45-49 tooth rows in upper jaws. Teeth flattened and in quincunx arrangement in juveniles and females, but pointed and in parallel rows in adult males. Colour aIter preservation plain greyish-brown above, darker to margins of disc and pelvics. Several indistinct dark brown crossbars over tail length. Lower side predominantly white, but a broad brown border along margins of disc and pelvics, and anterior two thirds of tail brown. Head also largely brown, and brown blotches may occur on belly, inner pectorals and around anus. Fresh specimens with a distinct bluish shade on head and margins of disc and pelvics above, and margins of disc and pelvics below blaclusli rather than brown in fresh state. Upper side of clasper, which is moderately elongate and slender, with dermal denticles and a very large pseudosiphon, formed and supported in its distal third by dT1-cartilage. Glans clasper with components as generally described for other members of the genus, but particularly with terminal bridge, pseudorhipidion, rhipidion, flag, dike, and funnel. Clasper skeleton consisting of 4 dT-, 2 aT-cartilages and a ventral terminal. Dorsal marginal with distal extension, dT1 with proximal process and fused distally with the vT. Dorsal terminal 3 not fused with axial or dT4, which joins tip of axial, and ventral terminal with anterior notch and medial process. Neurocranium with rostral cartilage very delicate in its distal two thirds and joined to rostral node. Rostral appendices very elongate and flattened. Nasal capsules very large, ovoid in shape, without basal fenestrae. Preorbital processes and jugular arches poorly developed. Anteriormost pectoral radials and propterygia almost in contact with rostral appendices at snout tip. Scapulocoracoid subquadrangular and hardly expanded anteroposteriorly. Foramina expanded, only one postventrally. Rear corner not elevated, and postdorsal margin abruptly sloping. Mesocondyle almost equidistant to both the other condyles. Vtr : 23-24, Vprd : 68-70, pectoral radials : 61-63. zyxwvutsrqpo ETYMOLOGY : Named after its type locality off Central West Africa, with which it geographically links the Northeastern and Southeastern Atlantic representatives of the genus. DESCRIPTION OF THE H O L O T Y P E For detailed moryhometrics and nieristics see table I. External nrorphology (figs. 1-6) Disc almost heart-shaped, 1.3 x as broad as long, axis of maximum width a t about 65 yo of disc length behind level of shoulder girdle. Anterior margins strongly undulated in this adult male, i.e. weakly concave immediately behind snout tip, strongly convex a t level of snout length and orbits, and deeply concave again a t level of spiracles and nape. Outer pectoral corners broadly rounded and continuous with t h e relatively short, evenly convex posterior margins. Inner pectoral corners narrowly rounded. Pectoral axils deeply incised t o origin of anterior pelvic lobe. Snout very short, its preorbital length only 2.2 x the interorbital width, roundish and bluntly angled (1350). Tip of snout marked off as a short triangular projection. Orbits very large, their horizontal diameter about 1.5 x t h e interorbital width and 70 o/: of the preorbital snout length. Spiracles only half as long as the orbits, interspace between them twice as wide as the interorbital distance. Eight pseudobranchial folds in each spiracle. Pelvics large, with a slender and pointed anterior lobe about two thirds as long as the posterior lobe. Both lobes separated b y a deep notch. Glaspers with pointed tips, fully developed and extended t o about 40 % of tail - zy zy 906 - zyxwvuts zyx zyxwv FIG.1. - Breoirnja africana sp. n. ; holotype 6 MNHN 1953-1 in dorsal and ventral view. zy zy zyxwvut zyxwvu zyx zyxwvutsrqpo - 907 - ength. The glans extremely dilatated in tlie left, less so in the right clasper. Tail long and slender, almost GO % of t h e specimen's total length, obviously depressed over its whole length, a low oval in cross-section. Lateral tail folds short, in the posterior third of tail length, only one fiith of t h e total length. Folds widening a t level of dorsal fins and terminating distinctly anterior t o tip of tail level with end of second dorsal. The small dorsal fiiis a t t h e very end of tail, their hases confluent a t level ol about hall their vertical higlit, which is about 50 % of their base length. Second dorsal somewhat smaller t h a n first, both similar in shape with a steep ascending anterior, and a broadly rounded upper margin, a fan-shaped apex, and a posterior margin angled in forward direction. Postdorsal section o1 tail extremely short, only about 40 yo of tlie D2 base length. Caudal fin a low, evenly high fold continued veiitrally as a very low lold, resp. ridge t o below D2. Preoral snout length almost equal t o width of mouth, the latter being about 40 "/o of the ventral head length, which is about 3 x the internasal width. Distance between fifth gill slits alsout GO "i, of t h a t between first gill slits, tlie latter distance being 2 x the internasal width. Nostrils with a low fleshy flap somewhat extended laterally as a trapezoid tip with short fringes a t its edge. Nasal curtain subrectangular, with undulated outer margins, rounded apices, and almost transverse rear margins, which are set with short Eorked lobelets nearly t o tlie isthmus. No oronasal pits. Jaws protruded in an unnatural position, straight normally apart from the median convexity in upper and lower jaw. 46 close set obliquely parallel toot11 rows in upper jaw. Individual tooth in median third o1 jaws with erect, long awl-shaped tip on subquadratic base. Shape of teeth gradually changing toward corners of mouth in showing low conical central cusp on a broadly rectangular base. Anteriormost pectoral radials and propterygia extended forward over lull rostral length and almost in contact with rostral tip. Upper surface more or less densely set with coarse spinules, including orbits, dorsal and caudal fins. Snout tip smooth as well as centers of pectorals and sides of trunk, a broad strip along posterior pectoral margins, a narrow median strip along tail t o D l , and both pelvic lobes, but upper surface of claspers prickly. h wedge-shaped area of hooked thornlets in malar region continued as a Iwoacl strip of very coarse spinules along remaining anterior pectoral margin onto the entire pectoral apex. Spinules along sides of tail also distinctly coarser. Lower side completely smooth. Four small hooked thorns in line in preorbital position on left, three on right side, and two postorbital ones respectively. A similar small thorn on each inner shoulder. A median row 01 17 (15th lost) larger, curved thorns along only the anterior three fifths of tail length from aliout level of pelvic axils t o shortly behind clasper tips, their size decreasing rearward. A short strip of pointed alar thorns on each inner pectoral apex consisting of three longitudinal rows, each row with 5-7 thorns. No thorns on snout, nape, back of trunk, and the posterior tail section in front of first dorsal. Colouration (in alcohol) brown without any patterning on upper side, lighter along median axis of disc and on pectorals, darker t o margins of disc and pelvics, on claspers and on tail. Dorsal and caudal fins blackish-brown, lateral tail folds marbled brown with some white. Side areas of rostrum not marked off, but anterior part of nasal capsules dark brown, and a transverse strip of ochre across posterior part of nasal capsules and interorbital space. Orbits blackish. Areas of alar thorns marked lighter brown. Anterior pelvic lobes blackish brown towards their tips, a broad transverse band of creamy white zy zy - 908 - across the middle o l left lobe only. Tips of posterior pelvic lobes with a creamy blotch a t outer margin, more distinct on right than on left lobe. Basal part of claspers somewhat lighter brown than terminal region. Several irregular indistinct dark brown cross-bars over tail length. Lower side of disc predominantly white, but a dark brown disc border very broad a t outer corucrs and posterior margins, much narrower a t anterior margins. The entire prenasal snout region and a strip from outer nostrils t o level of first gill slits over propterygia also brown. Distal third of anterior pelvic lobes as well as almost the entire posterior lobes dark brown, except for their distal white blotches also shown dorsally. Claspers brown except for white basal third, but lighter in terminal region. Base of tail as well as posterior third white, with a few brown spots and blotches in latter part. Remaining area of tail plain dark brown. When freshly caught, the specimen showed dorsally a distinct bluish shade on orhits, margins of disc and pelvics, and over gill area. Below t h e outer margins of disc and ~ ~ e l v i cwere s blackish, whereas tail aiid inner pelvic areas were marked off in brown. zyxwvu zyxwvutsrq FIG.3. - Bseviraja africana sp. n. ; holotype 8 MNHN 1983-1, mouth and nasal region. Claspers (figs. 3-5). External description based on right clasper due t o the extreme dilatation of the left clasper. zy zy - 909 - Claspers moderately elongate and slender, the terminal region hardly marlred off from proximal part. Outer surface of dorsal lobe with a very long and deep pseudosiphon (ps), the inner margin of which is supported by the dT1-cartilage in its distal third, whereas the proximal two thirds are bordered b y and imbedded in the dorsal dilatator muscle. Outer dorsal surface densely set with fine dermal denticles (dd), except l‘or the area o€ the dilatator muscle and the deeper pseudosiphon groove, which is as long as the terminal egion and is located wholly proximally to it. dd zy zyxw zyx zyxwvutsrq zyxwvu h d’i FIG.3. - B r e h a j a afrkana sp. n. ; holotype 8 MNHN 1983-1, naturally dilatated right glans clasper. cf : cleft ; dd : dermal denticles ; di : dike ; fg : flag ; fn : funnel ; hp : hypopyle ; pr : pseudorhipidion ; ps : pseudosiphon ; rh : rhipidion ; sh : shield ; SIC : spilie ; sl : slit ; st : sentinel ; t b : terminal bridge ; dT3 and dT4 : position of dorsal terminal cartilages 3 and 4. ’ On inner dorsal lobe a deep proximal cleft (cf), placed longitudinally between axial, dorsal terminal 2, and terminal bridge (tb). An oblique oriented slit (sl) overlying proximal end of cleft and terminating inward a t proximal midline of glans a t the low vertical wall - zy z 910 - of the pseudorhipidion (pr), the distal end o€ which is hidden in a continued fleshy fold. Its outer margin, however, supported proximally b y the dT3, and distally by the dT4, with an unsupported space between both cartilages. Dominant component on inner ventral lobe is the long shield (sh) extending over the proximal three quarters of the glans. Its outer cartilaginous edge free and sharp. Level with its distal end and along its inner margin a distinct dike (di), which is a thin, almost vertically oriented cartilaginous wall covered with thin integument and somewhat curved inward. Distal end of both shield and dike is the horizontal tongue-like funnel (fn). Large fingerlike sentinsel (st) arising a t inner proximal half of shield and entirely covered with integument. Below its tip a broadly triangular spike (sk),which has upward curved clawlike free cartilaginous tip. Proximally on base of seiltinel a small, fan-shaped rhipidion (rh). Its whole inner surface structured with longitudinal lamellae, outer surface with large pores distally, but with lamellae towards its insertion. About level with the dike, a rudimentary but distinct flag (fg) in midline on axial cartilage. Inner surfaces of glans creamy white, with the following areas, or components sootygrey and/or brown : pseudosiphon entirely so as is outer dorsal lobe, integument flap over pseudorhipidion dark grey. However, loose pigmentation in proximal half of shield, oiiter surface of dike, distal end of axial cartilage and its vicinity on dorsal lobe, base of sentinel, inner side of the dorsal lobe integument edge, and marginal parts proximally 011 dorsal lobe. The clasper skeleton (figs. 4-Ei), dissected from the left clasper, consists of four dorsal, two accessory terminal cartilages, and a ventral terminal element grouped around the axial in the glans part. Axial (Ax) with a pointed distal end. Ventral marginal (vM) almost spoon-shaped distally, while the dorsal marginal (dM) shows a truncate distal end with a plate-like extension, this externally iorming the pseudorhipidion. ßeta-cartilage a relatively long, slender, plate-like element inserting a t the dorsal part. of the double-headed Ax-end and extending proxinially to half the length of the hi-element. Dorsal terminal 1 (dT1) (fig. 5a) very large, curved around the axial onto ventral side and with a long, pointed proximal extension, which supports the inner edge of the pseudosiphon in its distal third, Distally the dT1 is firmly connected with the large ventral terminal (vT) (fig. 5 a) on the ventral side of the slreleton. The outer lamella of the vT forms the shield, the short distal extension the funnel, and the broadly triangular, plate-like extension in the distal third forming the dike in curving upwards from the dorsal surface of the vT. Proximally the vT shows a distinct notch and a short conical process, which links this element with the ventral surface of the accessory terminal 1 (aT1). The distal extension of the latter of the dorsal side forms the sentinel. Spike formed by the distal end of the accessory terminal 2 (aT2). Both the latter cartilages illustrated separately in figures 5 b c. Dorsal ter-' m i n d 2 (dT2) subquLdrangular, with long slender, obliquely 'oriented' distal extension, which is fused with the inner proximal edge of the dT3. Dorsal terminal 3 fused with o u t e r edge of dT2-extension, and ending distally within the integument. Inner corner of dT3 connected with proximal end of the small rod-like terminal bridge cartilage (tb), which links the dT3 and the axial. The small plate-like and almost uncalcified dorsal terminal 4 (dT4) so delicate, t h a t it could not be isolated from the firm tissue. It is distally fused with the axial, and ends freely within the integument proximally. NO further components in distal part of the dorsal lobe. zyxw zy zyxwvu - zyx zyxwv + , - zy zy 911 - zyxwvutsrq zyxwvut zyxwvutsrq zyxwv FIG.4. - Breuiraja africana sp. n. ; holotype 8 MNHN 1983-1, left clasper skeleton in dorsal'and ventral view. 13 : beta cartilage of basal group ; Ax : axial ; dM and vM : dorsal and ventral marginal ; aT1 and aT2 : accessory terminals 1 and 2 ; dT1-dT4 : dorsal terminals 1 through 4 ; vT : ventral terminal ; t b : terminal bridge. Pelvic girdle (fig. 6 a, after radiograph) Relatively small, its maximum width 75 % of t h a t of the pectoral girdle. Front edge almost straight, rear edge a deeply concave rounded arc. Prepelvic processes short a n d massive, bluntly pointed, and oriented obliquely outward. Long iliac processes greatly curved inward and forward. Two iliac foramina. i zy - 912 - ventral dorsal zyxw zyx zyxwvutsrqponmlk zyxwvu d. V. V. d. FIG.5. - Breviraja africana sp. n. ; holotype 3 MNHN 1983-1, cartilages of left clasper in dorsal and ventral view ; a, fused dorsal terminal 1 and ventral terminal ; b, accessory terminal 1 i c, accessory terminal 2. Pectoral girdle (after radiograph) zyxwvu Maximum width 1.3 x t h a t of the pelvic girdle and almost equal t o greatest cranial: width. Mesocondyle of scapulocoracoid equidistant from procondyle and metacondyle. Neurocranium (after radiograph) Rostral cartilage short, only 34.4 yo of length of the cranium. Its basal part a moderately wide triangle, basal width 16.7 % of greatest cranial width, but abruptly tapering zy - 913 - distally. Distal third and appendices not visible in radiograph. Nasal calxdes very large and broadly extended. Their front edge greatly bulging, the rear edge moderately concave. Maximum cranial width 73.3 % of cranial length. Nasal capsules slightly angled forward a t 730 t o longitudinal axis of cranium. Nasobasal fenestrae absent. Orbital region long, greatly constricted, with rounded edges, preorbital processes poorly developed. Least interorbital width dorsally 29.4 % of cranial width. Otic region long and wide, its maximum width 63.3 % of cranial width. Postorbital processes distinct as short, broad triangles clearly separated from the slender pterotic processes. Jugal arches moderately developed, neither laterally, nor to rear exceeding the contour of the o c c i p t . Anterior fontanelle a n elongate rather narrow triangle with concave rear edge. Posterior fontanelle narrowly club-shaped, moderately constricted medially and rounded a t both ends, the broader part to the rear. Its length somewhat greater than t h a t of the anterior fontanelle and 35.5 % of cranial width. Vtr : 24, Vprd : 69, pectoral radials : 63. zyxwvuts zyxwvutsr zyx FIG.6. - Breviraja africana sp. n. ; pelvic girdles of (a) holoLype 8 and (b) MNHN paratype schematized after radiographs. 2 X natural size. 9, somewhat. zyxwvutsrq n E S C R I P T I O N O F TILE PARATYPE F E M A L E S (figs. 7-10) For detailed morphometrics and mcristics see table I. External morphology In general the same as the holotype, except for typical female features and the few head measurements, which in the male are due to artificial distortion of the jaws. Disc clearly heart-shaped, 1.2 x as broad as long, axis of greatest width a t about 6370 yoof disc length behind level of shoulder girdle. Anterior margins very weakly undulated 3, 16 - LG. 914 - zy zyxwvutsr zyxwvutsrqpo zy zyxwvutsrqponml BI 7. view. 3'-2 in 7rentra zy zyxwvuts - 915 - t o almost evenly convex, resp. straight in the median third, as is usual and coiitrary t o males. Preorbital snout length 2.5 x as long as tlie interorbital width. Snout rounded and bluntly angled (1370), its tip marked as a short triangular integumental process. Orbit diameter 1.4-1.8 x t h e interorbital width and about 57-71 yo o l preorbital snout length. Orbits 1.9-2.5 x as long as spiracles, interspace Letween the latter 2.2 x wider t h a n interorbital width. 7-9 pseudobranchial folds in spiracle. Anterior pelvic lobe two thirds of the length o€ t h e posterior lobe. Length of tail and its lateral folds in relation t o TL as in holotype, b u t shape in cross-section a low trapezoid rather t h a n an oval as in the male. Tail Eolds ending distinctly in front of tail tip. Short postdorsal tail section 45-69 yo of D2 base length. Preoral snout length 1.3 x as long as width ol mouth, the latter being 3 4 % of the ventral head length, which is 3.3-3.4 X the internasal width. Distance between fifth gill slits 59-62 yo of t h a t between first gill slits, t h e latter distance being 2.1 x t h e internasal width. Nostrils and nasal curtain (fig. 8) as in holotype, i10 oronasal pits. Jaws straiglit apart from the mediali convexity. 45, resp. 49 close-set tooth rows in upper jaw in quincunx arrangement. Individual tooth, a t least in the median section o l jaws, showing clear sexual dimorphism in having rhomboid flattened Lase with a very low conical cusp a t center or toward t h e inner corner. Anteriormost pectoral radials and propterygia extending almost t o snout tip. zyxwvu FIG.8. zyx zyxwvutsrq - Breuimja ufricana sp. n. ; paratype $2 ISH 129180, mouth and nasal region. zy zy - 916 - Apart from usual sexual dimorphism, spinulation and thorn pattern as in the holotype male. Upper surface entirely prickly, except €or tip of snout and a strip along posterior disc margins, a narrow median strip along tail from end of the thorn row t o the D1, and anterior pelvic lobe, b u t a central patch o l spinules on posterior lobe. Smaller female with three preorbital and three postorbital thorns on each side, a posterior median nuchal thorn, and a scapular thorn on either side. A suprascapular thorn may be worn off. Larger female with four preorbital thorns on left side, three on the right, and two in left, one in right postorbital position. Also a median nuchal thorn posteriorly, but a thorn on leit shoulder only. Median row of thorns only along anterior three filths o€ tail from about level of pectoral axils, 27 (2 lost) in the smaller, 28 (1lost) in the larger female, the line continued in both as a shallow groove to first dorsal fin without spinules or thorns. Lower side smooth in both specimens, except for an irregular narrow marginal strip of spinules along the tail, the spinules encroaching from the sides o€ the tail. Colouration (in alcohol) plain dark greyish-brown above, darker t o margins of disc and posterior pelvic lobes. About five indistinct dark brown cross-bars over length of the tail. I n the smaller paratype dorsally a white spot on middle of right anterior pelvic lobe, also one a t upper margin of first dorsal fin, and furthermore a large milky blotch a t right snout margin and two whitish markings a t left pectoral apex. Lateral tail folds milky white with some darker marbling. Lower side (fig. 7) generally as in holotype, but a large brown blotch on belly and irregular brown blotching along each pectoral origin. Head brown t o level of first gill slits, only margins of nostrils and nasal curtain as well as jaws white. Except €or its rear edge, anus surrqunded with sooty brown. The larger paratype dorsally (fig. 7) with a white spot a t about middle of left pectoral fin, two further ones in oblique orientation over left side of pelvis area, one a t right side o l tail a t level of tip of pelvic, and one anteriorly a t left side of second dorsal fin base. Lateral tail folds marbled brown and white. Lower side as in smaller female, but white center of disc extended forward as wedge-shaped areas a t sides of head and also t o lower jaw as in the holotype. Tail brown in both paratypes t o below first dorsal fin, remaining part white marbled with brown. When freshly caught, both females showed the sanie bluish shade on margins of disc, pelvics, and head as the Iiolotype. Disc margins below were also blackish, the tail as well as blotching in the center of the disc were brown, this latter pattern lacking in the male. zyx I Pelvic girdle (fig. 6 13, zyxwv after radiograph) Maximum width 72 % and 68.5yo of the pectoral girdle width for the smaller and larger paratypes respectively. Front edge as well as the rear edge almost straight, the latter formed as a shallow broad trapezoid. Otherwise as in holotype. Figures 6 a b show the same distinct sexual dimorphism discovered already in the Northeastern Atlantic Breoiraja caerulea by STEEIMANN (1976 b). + Pectoral girdle (after radiograph) Maximum width 1.4 and 1.5 x ,that of the pelvic girdle, 1.3 and 1.3 x as wide as the maximum craniai width for the smaller and larger paratypes respectively. Mesocondyle I zy zy zyxwvuts zyx - 917 - of scapulocoracoid in both females almost equidistant from both the other condyles. As compared with the male, there is an obvious sexual dimorphism also in t h a t the pectoral girdle is distinctly wider in relation to the pelvic girdle as well as to the cranial width. This has also been stated for Rreviraja caerulea b y STEHMANN (1976 b). mtc zyxwv zyx pvf msc FIG.9. - Breviraja africana sp. n. ; paratype $? MNHN 1983-2, right scapulocoracoid. af : anterior fenestra ; msc : mesocondyle ; mtc : metacondyle ; pdf : postdorsal fenestra ; prc : procondyle ; pvf : postventral foramen ; rc : rear corner ; scp : scapular process. Scapulocoracoid (fig. 9 ) dissected from the smaller MNHN paratype. It is subquadrangular in shape, only a little higher t h a n long. Mesocondyle slightly anterior of midlength. Postdorsal and postventral foramina single and expanded, anterodorsal foramen distinctly so. Dorsal margin concave, but rear corner not elevated, and postdorsal margin abruptly sloping. Neurocraiziz~in (fig. 10) Illustration based on radiograph o l the smaller female, the rostral features added b y dissection of the same specimen. Neurocrania of both paratypes generally in accordance with t h a t of the holotype. For the comparison of proportions with the male, those of t h e smaller paratype come first. Length of rostral cartilage 37.6 % and 34.4 ?< of the cranial length. Width of rostral base 14.6 yo and 17.4 yo of the cranial width, the latter being 72.2 o/: and 71.2 % of the cranial length. Least interorbital width dorsally 26.3 yo and 27.9 % of the cranial width. of which latter the width of the otic region is 61.4 % and zyxwvutsrqp zyxw TABLEI. - Breuiraja africana sp. n. Actual measurements (in niin) for the three type specimens or extcrnnl morphology and anatomical structures, and meristics (columns 1-111). Range of proportions in per cent of total length (column IV). ... (( )) indicates measurements of distorted regions. I Holotype Total length Disc, width length Snout, preorbital length Orbit diameter Interorbital width Spiracle Interspiracular width Orbit spiracle D 1,height base length D 2, height base length Distance D 1-D 2 C, base length Tail, postdorsal length height a t V-tips width a t V-tips height a t D 1 origin width a t D 1 origin Lateral tail folds, length Snout, preoral length prenasal length Head length, ventrally Mouth width Internasal width Nasal curtain, length width of each lobe distance betweeh lohes + 285.0 170.0 133.0 22.0 15.3 10.0 8.0 20.4 18.0 9.4 17.0 7.5 15.2 O 5.9 8 II Paratype Q III Paratype MNHN ISH 284.0 157.0 135.0 24.8 14.1 10.0 7.5 21.5 16.5 7.0 15.0 7.9 15.9 O 7.1 7.1 7.7 40.0 2.8 5.4 63.5 28.5 20.2 65.0 22.0 19.5 13.2 7.6 o5 304.0 169.0 138.0 24.7 17.6 9.7 7.0 21.0 18.9 9.8 13.0 8.7 15.0 O 40.4 10.4 s.1 11.0 3.0 5.0 68.9 30.5 20.5 67.0 23.0 19.5 13.5 8.2 9 IV 100.0 55.3-59.0 45.4-47.5 7.6-5.7 5.0-5.5 3.2-3.5 2.3-2.8 6.9-7.6 5.8-6.3 2.5-3.3 4.3-5.9 2.6-2.9 4.9-5.6 O 2.0-3.4 2.0-3.4 2.5-2.7 3.5-3.6 0.9-1.0 1.6-1.9 20.8-22.7 8.6-10.0 6.4-7.9 22.0-23.3 7.6-9.0 6.4-7.0 3.5-4.6 2.5-2.8 zyxwvutsr zyxwvuts zyx zyxwvutsrqponmlkj zyxwvu 5.9 7.2 10.0 2.7 5.5 60.0 (( 24.7 )) 15.5 67.0 (( 26.0 )I 20.3 10.0 7.2 R 1 1 n \\ o nr zyxwvutsrqp zyxwvuts zyxwvut 5th Interbranchial width, 1st’ 5 th’ V, 1ength.anterior lobe Clasper, postanus length Snout-middle of anus Middle of anus - D 1 -D2 . - tip of tail Snout - max. disc width 3.0 37.0 21.0 38.0 64.0 1.21.0 131.0 149.0 169.0 87.0 2.9 40.9 25.4 40.0 3.4 41.8 24.7 41.0 1.0-1.1 12.8-14.4 7.3-8.9 13.2-14.1 116.0 129.0 141.5 166.5 85.0 127.0 136.0 151.0 175.0 96.0 40.8-42.0 44.7-45.5 49.7-51.7 57.6-58.7 63.0-69.6 of disc length 1370 45 ’717 23 1370 49 919 24 68 63/63 - , - -- Angle of snout Tooth rows upper jaw Pseudobranchial folds, left/right Vtr Vprd P-radials leftlright Neurocranium, TL Rostrum, length Max. width cranium Min. dorsal interorbital width Max. width otic region Max. width a t jugal arches Width rostral base Post. fontanelle, length Post. angle nasal capsules Pelvic girdle, max. width Pectoral girdle, max. width Scnpulocoracoid, length height pre - msc - length post - msc - length number pvf 1350 46 818 24 69 63/63 45.0 15.5 33.0 9.7 20.9 19.8 5.5 11.7 730 25.9 34.2 13.2 (x-ray) - 6.0 (x-ray) 6.2 (x-ray) - ïo 61+/62+ 47.4 17.S 34.2 9.0 21.0 19.0 5.0 13.0 ca. 760 30.0 41.8 13.8 16.3 6.7 7.1 1 48.0 16.5 34.4 9.6 20.5 19.0 6.0 14.0 710 31.7 46.3 15.5 (x-ray) - 7.8 (x-ray 7.3 (x-ray) - - zy zy zyxwvutsr zyxwvutsr - 920 - 59.6 yo. Anterior fontanelle somewhat shorter t h a n the posterior one, the length of which is 38 yo and 40.7 yo of the cranial width. . Nasal capsules a t an angle of 760 and 710 t o the longitudinal axis of the slrull. Rostral cartilage abruptly tapering after basal triangle t o form an uncalcified delicate bar, which undulates laterally as well as vertically and is joined t o the median notch of the rostral appendices. These are delicate plates, with a large distal foramen and a slender, vertically undulated extension reaching rearward over two thirds of rostral length, but these long ends free of the rostrum. Anterior fontanelle as well as the posterior are similar in the Lhree types, except for a slight modification in the larger IS13 paratype. The narrow anterior p a r t of the posterior fontanelle in this specimen is as long as the posterior part and shows almost straight edges, i.e. no real median constriction exists. Vtr : 23 and 24, Vprd : 70 and 68, pectoral radials : 61-63. \ . z FIG.10. - Breviraja africana sp. n. i paratype pl MNHN 1983-2 ; neurocranium and snout skeleton, somewhat schematized in combination of radiograph and dissection. 1.92 x natural size. zyxwvu INTERSPECIFIC COMPARISON The only congeners known t o also possess a n external clasper pseudosiphon are the Northwestern Atlantic B. colesi Bigelow & Schroeder, 1943, and B. spinosa Bigelow & & COMPAGNO,1982). However, both these species have Schroeder, 1950 (MCEACHRAN - f zy zy zy 921 - short and stout anterior pelvic lobes, a different and heavier thorn pattern (especially B. spinosa). Furthermore, B. colesi shows a constant pattern of dark and light spots and blotches on the upper side. Additionally, both species possess the components promontory and hook in glans clasper and show corresponding modifications of the dorsal terminal 2 and 3 cartilages. Also sentinel and spike in both are located obviously distally within the glans (MCEACIIRAN & COMPAGNO, 1982). Nothing is known with regard t o clasper characters for B. inainiZZidens (Alcock, 1889) and a Breviraja sp. from the Indian and Indopacific Oceans. However, the former species (the holotype and only specimen lost) described as “ uniform jet-black throughout ”, has separate dorsal fins and a continuous median row of about 30 thorns from nape t o first dorsal fin. The latter Indopacific species is known from three juvenile males only, which have no thorns on disc other t h a n a single preorbital one on each side, a median row of about 40 tail thorns from level of pelvic axils to first dorsal fin, and separate dorsal fins. Both these species have been redescribed, discussed, and assigned t o Breviraja by STEHMANN (1976 a ) . . Another new Breviraja (or Neoraja) species (Ms, MCEACHRAN & STEHMANN) froin the NW-Atlantic shows thorns also along midline of the body, furthermore (MCEACIIRAN & COMPAGNO,1982) a rostral shaft failing to reach the rostral node, and a scapulocoracoid with an elevated rear corner as well as a diagonally sloping posterodorsal margin. Yet another undescribed Breviraja species from Surinam waters, not conipletely investigated though (STEHMANN, unpubl. results), is similar to B. spinosa in shape and dorsal spinulation as well as the heavy thorn pattern, and possesses the clasper components pseudosiphon, hook and rudimentary promoritory. However, its dorsal colouration is plain lead-grey to blackish-brown, and ventrally even darker in being uniformly blackish-brown. Hence, it differs from B. africana a t least in colouration and several clasper characters. Among the Eastern Atlantic congeners B. stehirianni Hulley, 1972, and B. caericlea Stehmann, 1976, lack a n external clasper pseudosiphon as stated in their original descriptions. Their rostral shaft fails t o reach the rostral node, and their scapulocoracoid has an elevated rear corner and a diagonally sloping posterodorsal margin (MCEACHRAN & COMPAGNO, 1982). 1973, An unnamed Breoirqja species from the southern Bay of Biscay (STEIIMANN, 1979) is very different in shape of the disc (adult male with straight anterior disc margins), has widely separated dorsal fins, and a median row of about 50 thorns from shoulder girdle onto the anterior two thirds of the tail. The single known specimen is in a n advanced stage of decomposition and hence, presence or absence of a pseudosiphon in this adult male’s claspers cannot be stated with certainty, although its clasper skeleton (STEIIMANN, unpubl. results) is very similar to t h a t of B. africana. Recently a number of juveniles of one more unknown Breviraja species have been obtained from moderately deep slope waters in the Northeastern Atlantic off the Iberian Peninsula. Although not yet investigated in detail (STEIIMANN & BARODOMINGUEZ, unpubl. results), this form is clearly distinct from E. africana in being light brown dorsally with an almost constant pattern of black dots and spots, and in having a plain white lower surf ace. Finally, B. africana is distinct from B. yucatanensis Bigelow & Schroeder, 1950, B. nigerrima De Buen, 1960, and B. Zongicauda De Buen, 1959. See discussion below. - zy zy 922 - zyxwvuts DISCUSSION The detailed generic revision b y MCEACHRAN & COMPAGNO(1982), based mainly on skeletal anatomy and clasper characters but somewhat neglecting external morphology, has resulted in splitting Breviraja Bigelow & Schroeder, 1948. They restricted the taxon t o B. colesi and B. spinosa, and erected the new genus Neoraja €or nine further species and subdivided it into the new subgenera Neoraja and Fenestraja. These consist of the species stehmanni Hulley, 1972, caerulea Stehmann, 1976, and a new NW-Atlantic species (Ms in preparation b y MCEACHRAN & STEHMANN) for the former subgenus, and of plutonia Garman, 1881, sibogae Weber, 1913, atripinna, cubensis, sinusmexicanzcs, all three described b y BIGELOW & SCIIROEDER in 1950, and ishiyamai Bigelow & Schroeder, 1962, for the latter subgenus. Apart from the above 11 species originally described as, or later assigned t o Breoiraja, MCEACTIRAN & COMPAGNO(1982) have revised, or commented on other nominal or valid species of the genus. They reallocated from Breuiraja t o Raja the Caribbean B. yucatanensis Bigelow & Schroeder, 1950, and the Chilean B. nigerrinza De Buen, 1960. The Indian Ocean B. manidlidens (Alcock, 1889) was tentatively assigned t o Neoraja. However, these authors left open the generic status of three further species assigned t o Breoiraja. Of these, B. longicazcda De Buen, 1959, from Chile, the holotype and only known specimen is lost (MCEACHRAN & COMPAGNO, 1982)) was inadequately described and illustrated originally, so t h a t its afiliation t o any rajid genus could not be stated. Should this species one d a y after all prove t o be a Breuiraja or Neoraja, it would most probably not effect B. africana, because the occurrence of B. longicauda in the Eastern Central Atlantic is most unlikely. A single old museum specimen, a n adult male from the southern Bay of Biscay, had been confirmed as a Breuiraja b y STEHMANN (1973, 1979) and was recently described in its external appearance (STEHMANN & BÜRIEEL,in press), but not named because of the bad condition of this Paris Museum specimen. It was also precisely described externally and perfectly illustrated b y VAILLANT(1888) and available for skeletal anatomy investigations. The third case are the three juveniles originally reported by WEBER(1913) from the Indopacific and identified as R a j a mamillidens Alcock, 1889. STEHMANN (1976 a ) described these three specimens in the Amsterdam Museum and assigned them t o Breuiraja, but did not name the species, which proved not t o be identical with Breviraja mamillidens (Alcock, 1889), due t o the juvenile stage of the three males and damage of the largest one. Although i t is admitted here t h a t MCEACIIRAN & COMPAGNO(1982) were confronted with the circumstance of inadequate, or even missing material and partly insufficient original descriptions and illustrations, the present authors are nevertheless critical concerning the nomenclatorial consequences caused by the generic revision in the latter three cases, in t h a t three species taxa originally described as, or later assigned to Breoiraja have lost their generic assignment. In this connection it appears unimportant, whether or not these perhaps are valid species, or named specifically. Such a case should not happen in taxonomic work, the less so since the present authors believe t h a t MCEACHRAN & COMPAGNO - ,. zy zy zy 923 - (1982) could have expressed and demonstrated their systematic conclusions in another nomenclatorial way as well. They could have, for example, provisionally subdivided Breuiiwaja into three subgenera Breuiraja, Neoraja, and Penestraja, and thus could have kept the generic assignment for the critical species taxa, which could for any reason no\ be fully investigated a n d hence, could not be arranged in their actual classification concept. Exactly the same problem, intensified however through a much larger number of species of worldwide abundance and only hitherto partly investigated, has to date restrained rajid workers from raising the various subgenera of R a j a Linnaeus, 1758, t o generic rank. After the above discussion of a more general nature related to the revision by MCEACHRAN & COMPAGNO (1982), the present authors wish to explain briefly the evaluation of B. africana as intermediate between Breuiraja and Neoraja sensu MCEACIIRAN & COMPAGNO (1982). Reference should be made t o t h e revision of t h e latter authors for the full details of their generic and subgeneric diagnoses, which cannot be repeated here completely. Furthermore, a renewed consideration of the conclusions b y MCEACHRAN& COMPAGNO (1982) must await further investigation of the species concerned, including material of the newly discovered lorms mentioned above. However, the present authors wish t o explain with a few examples, why in their opinion B. africana indicates t h a t the distinction between Breuiraja and Neoraja appears somewhat weak and is perhaps to a certain degree artificial. The generic diagnoses given b y MCEACHRAN& COMPAGNO (1982) have a nuinber of characters, which are common t o both genera. This is mainly due t o t h e circumstance t h a t a number of Neoraja generic features are statcd with the alternative “present or absent ”, which distiiiction in fact mainly refers t o thc two subgenera of Neoroja, but is not specified as such. Breviraja was characterized by these authors, among other features, in having a tail length of a t most 60 yo of the TL, a clasper with distinct pseudosiphon formed only by the dorsal dilatator muscle, a rostral shaft reaching rostral node and appendices, the latter being elongate and flattened, and a scapulocoracoid little expanded anteroposteriorly and with only one postventral foramen. These generic characters, e.g., are shared b y B. africana, which laclrs others such as, e.g., the thorn triangle over the nuchal/scapular region, short anterior pelvic lobes, oronasal pits, the components hoolr and promontory in glans clasper, a broad rostral base, moderately large rhomboidal nasal capsules, and well developed preorbital processes. Neoraja was characterized by M C E A C I ~ R A&N COMPAGNO (1982), among other features, in having separate nuchal and scapular thorns not forming a triangle, a short and broad iiitegumental process a t tip of snout, long anterior pelvic lobes, claspers moderately to very long and slender, clasper components flag and funnel, dM-cartilage with distal extension entering glans, rostral base relatively narrow, large ovoid nasal capsules, and poorly developed preorbital processes. These generic characters, e.g., are also shared b y B. africana, which lacks again others such as, e.g., one or three distinct thorn rows along midline of disc, a tail length of generally more than 60 yoof the TL, and a rostral shaft not reaching rostral node. Within Neoraja, B. africana is more similar to the subgenus Neoraja t h a n t o the subgenus Fenestraja, which latter is characterized, e.g., by having oronasal pits and nasal capsules with basal fenestrae, and through the lack of dermal denticles on the clasper, the lack of the dT1-cartilage and the anterior notch in vT-cartilage of clasper skeleton. zy zy zyxwvutsr - 924 - Additionally, B. africana shows unique features among the species so far arranged in Breviraja and Areoraia, in t h a t i t possesses four dT-cartilages, of which the dT4 joins the tip of the axial but is separated from the distal tip of the dT3. Furthermore, in t h a t the distal extension of the dM-cartilage appears as the component pseudorhipidion typically in median proximal position of the glans clasper, and in t h a t the distal third of the pseudosiphon groove is bordered and supported ])y the proximal outer edge of the dT1-cartilage. Although the specific validity of B. africana is quite clear, its generic afiliation a1)liears problematic with regard to the diagnostic characters combined b y MCEACHRAN & COMPAGNO (1982) t o describe Brecirajn and Neoruja. Apart from the above mentioned intermediate position of B. africana, the diagnoses given b y the latter authors are themselves somewhat weak, contain partly unprecisely stated features (e.g. pattern of orbital thorns, patterns of dorsal and ventral colouration), and contain shared features. Furthermore, t h e y mention in part characters not indicated in the specific descriptions and illustrations, such as the clasper component dike stated for Breviraja, b u t neither mentioned in the descriptions, nor indicated in figure 1 for B. colesi and B. spinosa (MCEACHRAN & COMPAGNO,1982 : 421, 402-403 respectively). The problem is further complicated, in t h a t MCEACIIRAN (pers. comm., 1982) explained t h a t the diagnostic generic characters should be understood in the light of their phylogenetic significance and interpretation mainly, through which viewpoints he strongly considered the present new species as a member of Neoraja, suhgerius Neoraja. The present authors do not intend to go into the very detail of such a basic discussion here (see above), Ilut would like t o underline a t least two major objections against MCEACIIRAN’S statement cited above. Firstly, significant characters combined in a diagnosis t o describe a generic taxon is one thing, the interpretation and analysis of such characters under terms of phylogenetic systematics is another. To assign a species taxon t o a genus means primarily its comparison t o generic diagnoses with regard to presence or absence of relevant features, a t most perhaps with regard t o a relative development or reduction of such characters. Considering these points of view B. u.fricanu certainly appears intermediate between Breviraja and Neoraja of MCEACHRAN & COMPAGNO (1982). Secondly, should the new species africana prove to be a member of Neoraja after a renewed consideration of the entire problem, then such important characters as a clasper pseudosiphon and a rostral shaft, which continues to join the rostral node and appendices, phyletically must have been developed independently twice. Although the present authors admit, t h a t such a case is not unusual in evolutionary processes, they nevertheless feel unable t o decide on the actual problem. The knowledge, in our opinion, about intrafamilial relationships within the Rajidae and about the phyletic significance of certain characters is not suficiently advanced, and in particular the revisiona1 information given by MCEACHRAN & COMPAGNO(1982) appears as an insuficient basis for t h e present case. As a consequence, the present authors feel unable t o assign with certainty the new species africana t o either Breviraja, or Neoraja in the meaning of MCEACHRAN & COMP A G N O (1982) and hence, the new species is preliminarily assigned to Breviraja sensu BIGELOW & SCHROEDER, 1948, and sensu ISIIIYARIA & HUBBS(1968). zy zy zyxwv - 925 - zyxwvutsrqp zy zyxwv Acknowledgements We are grateful t o Dr. A. CROSNIER(ORSTOM, Paris) for his permanent support o€ the project, to Dr. John D. MCEACHRAN (Texas A & M University, USA) for his comments on our manuscript on the revision draft and for having made available to us his manuscript with L. J. V. COMPAGNO of Breviraja prior t o its publication, and to M. STEIIMANN’S assistant, Mrs. Gudrun SCHULZE, for the preparation o€ all radiographs and photographs. Dr. D. L. BÜRICEL (Zool. Museum Universität Hamburg) kindly improved the English text of our manuscript and assisted in the technical reproduction of the drawings. We are especially obliged to Captain H. RIOU,master of the RV ‘ Nizery ’, whose fishing pxpertise mainly made the respective deep bottom hauls possible. LITERATURE REFERENCES 1 BIGELOW H. B. G., & W. C. SCHROEDER, 1948. - New genera and species of Batoid Fishes. J. n z a ~ . Res., 7 : 543-566, figs. 1-9. HULLEY, P. A., 1972. - A new species of Southern African brevirajid skate (Chondrichthyes, Batoidei, Rajidae). A m . S. Afr. Mus., 60 (9) : 253-263, figs. 1-5. ISHIYAMA, R., & C. L. HUBBS,1968. - Bathyraja, a genus of Pacific skates (Rajidae) regarded as phyletically distinct from the Atlantic genus Breoiraja. Copeia, 1968 (2) : 407-410, figs. 1-2. MCEACRRAN, J. D., & L. J. V. COMPAGNO,1982. - Interrelationships of and within Breviraja based on anatomical structures (Pisces : Rajoidei). BUZZ.mar. Sci., 82 (2) : 399-425, figs. 1-18. STEHMANN, M., 1973. - Rajidae, I n : Check-list of the fishes of the north-eastern Atlantic and of the Mediterranean. Eds J. C. Hureau & T. Monod. UNESCO Paris, 1 : 58-69. - 1976 a. - Revision der Rajoiden-Arten des nördlichen Indischen Ozean und Indopazifilc (Elasmobrancliii, Batoidea, Rajiformes). Beaufortia, 24 (315) : 133-175, Abb. 1-21. - 1976 b. - Breviraja caerzdea spec. nov. (Elasmobranchii, Batoidea, Rajidae) j eine neue archibcnthale Rochenart und zugleich ein Erstnachweis ihrer Gattung im Nordostatlantik. Arch. FischWiss., 27 (2) : 97-114, Abb. 1-11. - 1979. - Rajidae. In : Supplement to Check-list of the fishes of the north-eastern Atlantic and of the Mediterranean. Eds E. Tortonese & J. C. Hureau. Cybium, 3e sér., 1979 (5) : 341-342. STEHMANN, M., & D. L. B~RICEL, (in press). - Rajidae. I n : Fishes of the Northeastern Atlantic and Mediterranean. Eds P. J. P. Whitehead, M.-L. Bauchot, J. C. Hureau & E. Tortonese. UNESCO Paris. VAILLANT, L., 1888. - Poissons. I n : Expéditions scientifiques du ‘ Travailleur ’ et du ‘ Talisman ’ pendant les années 1880-83. Masson, Paris : 406 p., 28 pls. WEBER, M., 1913. - Die Fische der Siboga-Expedition. Siboga Ezped., 57 : XII 710 p., 123 figs, 12 pls. +