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FISCHER, P.-H., DuvAL, M. & RarFy, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs

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Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.

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Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.

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THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 72 ~~ # Band

May 1977 Mei

Part 12 #£Deel

THE VALIDITY OF MALACORAJA STEHMANN, 1970

(CHONDRICHTHYES, BATOIDEI, RAJIDAE)

AND ITS PHYLOGENETIC SIGNIFICANCE

PAE eAND ER HUI yi

MATTHIAS STEHMANN

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg. Kaap

THE VALIDITY OF MALACORAJA STEHMANN, 1970

(CHONDRICHTHYES, BATOIDEI, RAJIDAE) AND ITS

PHYLOGENETIC SIGNIFICANCE

P. ALEXANDER HULLEY

South African Museum, Cape Town

MATTHIAS STEHMANN

Tnstitut fiir Seefischerei, Hamburg

(With 5 figures and | table)

LMS. accepted 9 December 1976]

ABSTRACT

The clasper structure, neurocranial form and proportional dimensions of the first adult

male of Raja spinacidermis Barnard, 1923 are described. On the evidence presented the recog-

nition of the subgenus Malacoraja Stehmann, 1970 is substantiated. Ma/acoraja appears to

be the linking subgenus between Breviraja and rajids of the Dipturus/Rajella/ Amblyraja/

Leucoraja-line of evolution.

CONTENTS

PAGE

Introduction. DD

Material . ; 5 PAPAS

Morphology . 230

Clasper Structure . 230

Neurocranium . a 234:

Discussion . : + 236

Acknowledgements . 237

References . § 5, sil

INTRODUCTION

During the course of investigations on the systematics of the Rajidae of

the eastern North Atlantic, Stehmann (1970) defined a new monotypic sub-

genus, Malacoraja, for Raja mollis Bigelow & Schroeder, 1950 on the basis of

characters other than the clasper structure, since no adult male specimen had

been taken. Malacoraja was distinguished because of the extraordinary squama-

tion and tail coloration of the type species. Because of similarities in propor-

tional dimensions, tooth count and spination pattern, Hulley (1970) considered

Raja mollis to be synonymous with Raja spinacidermis Barnard, 1923, and later

followed Stehmann in the recognition of the subgenus Malacoraja to include

this species, although no claspers were examined (Hulley 1972a).

227

Ann. S. Afr. Mus. 72 (12), 1977: 227-237, 5 figs, 1 table.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

The Division of Sea Fisheries, Cape Town, has recently obtained the first

adult male specimen of this rare species, during the course of its 1973 Hake

Survey Programme, and has donated the specimen to the South African

Museum. The specimen not only calls for a description of the anatomy of the

clasper and the relevant taxonomic morphology, in an effort to validate the

subgenus, but also calls for comment on the phylogenetic position of Malacoraja.

MATERIAL

One specimen, an adult male (634,0 mm total length) trawled by R/V

Africana IT at station A 6139 (5 March 1973; 33°43’S 17°21’E; 914 metres;

bottom temp. 3,30°C; salinity 34,47%,; O, concentration 4,61 ml/litre); in the

collection of the South African Museum (SAM-—26879). Proportional dimensions

according to Hulley (1970) are given in Table 1.

TABLE 1

Raja spinacidermis (SAM-—26879). Measurements expressed in mm and as per mileage of

the total length.

Range

mm Yea (Hulley 1970)

Total length . E ; 5 : : ‘ 4 : 634,0 — —

Disc width . : : : j : ; : ; i 450,0 709 660-701

Disc length . B : : ‘ : : 5 335,0 528 515-542

Snout to greatest disc width : ; : : : : 198,6 313 307-335

Snout to middle of vent : ; 5 2 : ‘ : 306,2 483 466-487

Snout to axils of pelvics 5 ‘ ; : : : 272,4 430 —

Middle of vent to Ist dorsal origin , : : f Z 246,3 388 379-417

Snout length . : : : 3 : : : : : 92,6 146 145-166

Preoral length : ; : F F : : : 2 86,0 136 126-166

Prenasal length : : : : : é . : 66,5 105 106-136

Eye — horizontal diameter . 5 : : ' : } 17,9 28 33-37

Eye + spiracle . : ; : ; : , . ; 23,0 36 42-47

Spiracle . , ; : : ; 5 : : : 10,8 iN7 20

Interorbital distance 4 : : : ; : ; ; 2550 39 36-45

Interspiracular distance : : : : : : 41,5 65 64-70

Internasal distance : : ; “ ; : : 49,1 Wah 79-89

Mouth width . : ; : ; : : : ‘ 5 a2, 90 83-90

Gill slit lengths: Ist ; : ; : ; , ; : 9,3 15 13-15

3rd): : : : : : : : 12,0 19 16-17

Sth ; : 2 . ; : E 9,0 14 10-13

Distance between gill slits: Ist. : : ; ; ; 88,2 139 130-147

Sth. : : ; : : 49,7 78 84-91

Ist dorsal fin: height . i : 2 ; : - : 14,0 22 22-29

base length . : ; : : : X 28,0 44 47-50

2nd dorsal fin: height . j 4 : : : ‘ 17,6 28 19-27

base length . : : : : f ; 34,3 54 42-55

Interdorsal space . ; : ; : : é : : 0 0 0

Teeth (rows in upper jaw) . dee ee ee 56 54-60

Wertisormll @ouimes Wie gg | 24 28

Vprd . ; . , : : : : 63 60-65

Me ae ‘ i : : ‘ 87 88-93

THE VALIDITY OF MALACORAJA STEHMANN, 1970 229

Fig. 1. Raja (Malacoraja) spinacidermis. SAM -26879. A. Dorsal view. B. Ventral view.

Scale in cm and in.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

MORPHOLOGY (Fig. 1A-B)

The specimen fits the published descriptions of the species and gives

some additional information concerning general morphology:

1. Its squamation confirms the original definition of the subgenus, since

the upper surface of the disc and tail is completely devoid of thorns and is

entirely covered with spinules. A small patch of larger spinules (not typical

thorns) is situated in front of the left eye. Malar and alar spines are well

developed (Fig. 1A). The ventral surface is naked, except for a small patch of

spinules on the tip of the snout, small bands of spinules along the anterior

margins of the disc to about half the distance from tip of snout to level of

nostrils and the distal half of the tail, which is almost completely covered with

spinules.

2. The ventral surface of the disc and tail is uniformly dark brown (Fig. 1B),

except for small white areas at the corners of the mouth and between the gill

slits, at the axils of the pelvics and at the base of the tail. This coloration does

not correspond with the subgeneric and specific diagnostic character of dark

ventral tail colour, distinctly marked off from a predominantly white disc, a

character based only on juvenile specimens.

3. The teeth of the adult male are in close-set parallel rows, with those in

the middle part of the jaws having long, slender, sharply-pointed tips. The

teeth of the outer parts of the jaws have low, conical tips.

As far as the ventral coloration is concerned, Raja spinacidermis follows

a line of development which is well known for most deep-water rays. Juveniles

and adolescent specimens bear a few small thorns in the orbital, nuchal and

scapular regions and sometimes some enlarged spinules along the midline of

the back and tail; the ventral surface of the disc is usually white, with grey or

brown markings of varying extent. Thorns on the dorsal surface are almost

completely lost in adults, which furthermore show a change in the ventral

coloration of the disc to predominantly dark with small white markings in

certain areas.

CLASPER STRUCTURE (Figs 2-4)

Claspers moderately long, reaching to about 40 per cent of tail length

from axils of pelvics; rather slender, with terminal region barely broadened,

distal end pointed and somewhat fimbriate; dorsal and ventral surfaces

without dermal denticles; pseudosiphon absent; inner dorsal lobe with longi-

tudinal proximal cleft, upper end of which is covered by a transverse slit; inner

ventral lobe with well-developed shield, extending from above level of hypopyle

to about four-fifths the length of the glans, with pleated epithelia over most

of its surface and with cutting outer edge; insertion of the long rhipidion at

level of proximal tip of shield, its distal third fan-shaped and with porous

surface; sentinel well developed and slightly S-shaped, covered with fleshy

THE VALIDITY OF MALACORAJA STEHMANN, 1970 23)

integument; spike blunt and hardly projecting from midline of clasper, placed

below tip of sentinel; dike well developed along midline of distal half of glans,

rising in 90° angle from inner edge of shield and covered with thin integument;

medium-sized funnel below distal end of dike, formed as obtuse tip covered

10mm

Fig. 2. Raja (Malacoraja) spinacidermis. Lateral view of left clasper, opened to show structural

features of the glans.

cf—cleft; dk —dike; fn—funnel; hp —hypopyle; rh —rhipidion; sh—shield; sl—slit; sp—spike;

st —sentinel

DEB ANNALS OF THE SOUTH AFRICAN MUSEUM

TAREE Suber sguw set

Fig. 3. Raja (Malacoraja) spinacidermis. Cartilages of the terminal group of the right clasper.

A. Dorsal terminal 1 (dorsal view). B. Accessory terminal 1 (dorsal view). B’. Accessory

terminal 1 (lateral view). C. Ventral terminal (dorsal view). D. Dorsal terminal 2 and dorsal

terminal 3 (dorsal view). E. Accessory terminal 2 (dorsal view).

dT,—dorsal terminal 2; dT,—dorsal terminal 3; tb—terminal bridge.

THE VALIDITY OF MALACORAJA STEHMANN, 1970 233

with fleshy integument. Entire inner edge of shield darkly pigmented to level

of proximal end of dike, becoming diffuse laterally and distally on shield and

distally on dike.

Axial cartilage slender distally, but with slightly spatulate tip; dorsal

marginal with short distal extension, ventral marginal arched; dorsal terminal |

cartilage large, twisted and connected distally with distal tip of vental terminal

cartilage, without shelf for insertion of M. dilatator, but with 3 longitudinal

10mm

Fig. 4. Raja (Malacoraja) spinacidermis. Cartilages of right clasper (exploded).

Ax—axial; aT,—accessory terminal 1; aT,—accessory terminal 2; dM-—dorsal marginal;

dT,—dorsal terminal 1; dT,—dorsal terminal 2; dT,—dorsal terminal 3; vM—ventral

marginal; vI —ventral terminal.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

ridges; dorsal terminal 2 and 3 cartilages simple, forming framework of dorsal

lobe, with dT, and dT; making contact with a well-developed terminal bridge,

which is an offshoot of the Ax; dT, and Ax bordering proximal cleft; ventral

terminal cartilage large, with outer lateral margin forming the shield, with

dorsal crest the dike, with distal tip the funnel, and with anterior notch linked

with aT,; accessory terminal | cartilage somewhat U-shaped proximally and

with well-developed lateral projection forming the sentinel; accessory terminal

2 simple, with spatulate distal extremity, closely attached to the Ax proximally

and distally, but free medially.

With regard to the external components of ne glans, Malacoraja is charac-

terized by:

(i) very few structures on the inner dorsal lobe, but particularly a single, deep,

proximal cleft combined with a transverse slit,

(ii) a very long, prominent shield, covered with laminate epithelia and com-

bined with a dike and funnel at its distal end,

(111) a sentinel and spike located rather far proximally in the glans,

(iv) a medium-sized rhipidion and the absence of a pseudosiphon.

Diagnostic characters of the clasper skeleton include:

(1) the extraordinary shape of the aT, cartilage, which is unparalleled among

investigated Raja species,

(ii) the terminal bridge is neither formed by separate cartilages nor by dT

cartilages attached directly to the Ax, but is a massive offshoot of the Ax

itself.

NEUROCRANIUM (Fig. 5)

Neurocranium typically guitar-shaped and markedly constricted across

the orbital region, with well-developed post-orbital processes, short otic region,

and moderately-developed jugal arches; nasal capsules massive and directed

obliquely forward to about 55° to median axis, with ethmoidal nerve foramen

at leading edge; maximum width 60,6 per cent of total length of skull; rostral

cartilage projecting from cranium to tip of snout as strong but tapering rod,

without a segment; length of rostrum 53,8 per cent of total length of skull;

rostral appendices fused throughout their entire length to rostral bar and

extending posteriorly slightly more than two-thirds the distance from tip of

snout to level of anterior fontanelle or extend backward 60,6 per cent of length

of rostrum; radial cartilages of pectoral fin extending anteriorly, but falling

well short of tip of snout; anterior fontanelle extending forward to 15,5 per cent

of the length of rostrum, without anterior grooving and separated from posterior

fontanelle by narrow epiphysial bridge; orbito-nasal canal foramen compara-

tively small, optic foramen situated well forward; anterior cerebral vein foramen

well above level of optic foramen and situated close to internal foramen of

ophthalmic nerve; external foramen of ophthalmic nerve large and positioned

comparatively more posteriorly, at about level of antorbital processes.

THE VALIDITY OF MALACORAJA STEHMANN, 1970 235

ha.

F.n.eth.

F sup. oph.

: p.f.

oe ostor pr.

OE p p

Fend.

. S. par.dep.

ve, ‘

F. peri A ye:

Eoph.

eS Fa.cerv. !V_ Ill op.st.

Or-nas. can. F pro-ot.(V, VII)

ORO

Fin-orv. hy. fac.

Fig. 5. Raja (Malacoraja) spinacidermis. Neurocranium. A. dorsal view. B. lateral view.

C. posterior view.

a.f.—anterior fontanelle; an.pr.—antorbital process; F.a.cer.v.—anterior cerebral vein

foramen; F.aff.ps.a.—afferent pseudobranchial artery foramen; F.end.—endolymphatic

foramen; F.in-or.v.—interorbital vein foramen; F.].X—foramen of lateralis branch (X);

F.mag.—foramen magnum; F.n.eth.—ethmoidal nerve foramen; F.oph.—ophthalmic fora-

men; F.p.cer.v.— posterior cerebral vein foramen; F.peri.—perilymphatic foramen; F.pro-ot.

—pro-otic foramen; F.sup.oph.—superficial ophthalmic foramen; hy.fac.—hyomandibular

facet; j.a.—jugal arch; oc.con.—occipital condyle; or-nas.can.—oro-nasal canal; op.st.—

optic stalk; p.f.—posterior fontanelle; par.dep.—parietal depression; post-or.pr.—postorbital

process; pt.pr.—pterotic process; r.a.—rostral appendix; r.c.—rostral cartilage; I1—optic

nerve foramen; IIJ—oculomotor nerve foramen; IV—pathetic (trochlear) nerve foramen;

Vii—foramen of hyomandibular branch (VII); [X—glossopharyngeal nerve foramen;

X—vagus nerve foramen.

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnostic characters are:

(i) an anterior fontanelle clearly marked off on all sides,

(ii) a rostral cartilage longer than the neurocranium,

(iii) the maximum width of the skull more than SO per cent of its total length,

(iv) the rostral appendices more than half the length of the rostral bar, fused

with the latter over their entire length.

DISCUSSION

There is no doubt that the spination pattern of the disc and tail and the

complete absence of large thorns along the midline of the back and tail is unique

in Raja spinacidermis. The coverage of the dorsal surface by close-set spinules is

a character shared by species of Breviraja (B. stehmanni off South Africa), of

Bathyraja (B. smithii off South Africa) (Hulley 1972a, 19726), and rarely of

Raja (e.g. R. senta in the north-western Atlantic), but these species either have

specialized snout conditions and/or possess at least midline thorns on the dorsal

surface. While the enormous size of the nasal capsules appears to be correlated

mainly with depth distribution, e.g. Raja radiata and R. robertsi (Hulley 1972a),

the length of the rostral cartilage and especially its appendices, is rather extra-

ordinary among short-snouted Raja species. Both the rostral cartilage and the

appendices are typically rajid in form, but their characteristics can be interpreted

as an ancestral condition, which approximates Dipturus and single species of

other subgenera, e.g. Raja fullonica (Stehmann 1970: 146, pl. 22).

The anatomy of the clasper of Raja spinacidermis shows a number of simi-

larities to that of Breviraja-species (Hulley 19726; Stehmann 1976): the position

and general form of the sentinel and spike (aT, and aT,); the arrangement of

the dT, and dT, and their connection with a well-developed terminal bridge;

and the form of the vT, which distally is firmly bonded to the tip of the dT,.

The form of the aT, cartilage in Raja spinacidermis is unique among Raja-

species that have thus far been investigated. The form of the dT,, the association

of dT,, dT; and the terminal bridge, the form of the vT (resulting among others

in the formation of a prominent shield), and the lack of an external pseudo-

siphon, point to a closer association with species of the subgenus Dipturus

(Hulley 1972a). Malacoraja also shows a relation to Amblyraja, Leucoraja and

Rajella in the general form of the dT, and vT, especially in the position of the

anterior notch of the latter (Hulley 1972a: 38), and furthermore in the bonding

of the distal extremities of both these cartilages (cf. Stehmann 1970, pls 11-12).

In summary then, Malacoraja is also confirmed as a separate and valid

subgenus of Raja by conditions of clasper and skull, which are the most

important characters in modern rajid taxonomy. It possesses a true rayjid

condition of the clasper and skull, but shows characteristics in both, which are

intermediate between Dipturus and Breviraja-species. It may therefore be inter-

preted as the subgenus linking the genera Breviraja and Raja. With regard to

Raja, Malacoraja is closely associated to the evolutionary line of the subgenera

THE VALIDITY OF MALACORAJA STEHMANN, 1970 237)

Dipturus/Rajella/Amblyraja/Leucoraja. A strict application of Vprd values

would indicate a common ancestry for Malacoraja and Breviraja rather than a

direct lineage and would point to an origin from some ancestral Dipturus-species.

Further, the depth distribution pattern in Malacoraja would support the

hypothesis that Breviraja-species represent an early split from the rajid con-

dition, which penetrated abyssal regions, but which retained the neotenous

condition of the snout as an increased advantage in grubbing.

ACKNOWLEDGEMENTS

The authors would like to express their thanks to the Director and Staff

of the Sea Fisheries Branch, Department of Industry, Cape Town for the

donation of the rajid material taken during their Hake Surveys and especially to

Mr L. Botha of that institute. They would also like to thank Mr S. X. Kanne-

meyer, Department of Marine Biology, South African Museum and Mr V.

Branco for their assistance.

REFERENCES

BARNARD, K. H. 1923. Diagnoses of new species of marine fishes from South African waters.

Ann. S. Afr. Mus. 13: 439-445.

BIGELOW, H. B. & SCHROEDER, W. C. 1950. New and little known cartilaginous fishes from the

Atlantic. Bull. Mus. comp. Zool. Hary. 103: 383-408.

HuL Ley, P. A. 1970. An investigation of the Rajidae of the west and south coasts of southern

Africa. Ann. S. Afr. Mus. 55: 151-220.

HuLiey, P. A. 1972a. The origin, interrelationships and distribution of southern African

Rajidae (Chondrichthyes, Batoidei). Ann. S. Afr. Mus. 60: 1-103.

Huttey, P. A. 19725. A new species of southern African brevirajid skate (Chondrichthyes,

Batoidei, Rajidae). Ann. S. Afr. Mus. 60: 253-263.

STEHMANN, M. 1970. Vergleichend morphologische und anatomische Untersuchungen zur

Neuordnung der Systematik der nordostatlantischen Rajidae (Chondrichthyes, Batoidei).

Arch. FischWiss. 21: 73-164.

STEHMANN, M. 1976. Breviraja caerulea spec. nov. (Elasmobranchii, Batoidea, Rajidae); eine

neue archibenthale Rochenart und zugleich ein Erstnachweis ihrer Gattung im Nordost-

atlantik. Arch. FischWiss. 27: 97-114.

6. SYSTEMATIC papers must conform with the International code of zoological nomenclature

(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be

followed by the appropriate Latin (not English) abbreviation, e.g. gen. NOV., sp. nov., comb.

Nnov., syn. nov., etc.

‘An author’s name when cited must follow the name of the taxon without intervening

punctuation and not be abbreviated; if the year is added, a comma must separate author’s

name and year. The author’s name ‘(and date, if cited) must be placed in parentheses if a

species or subspecies is transferred from its original genus. The name of a subsequent user of

a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published

scientific names by which the species previously has been designated are listed in chronological

order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

the year is placed in front of each entry, and the synonym repeated in full for each entry, is

not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-

mens mentioned in the original description are to be designated paratypes; additional material

not regarded as paratypes should be listed separately. The complete data (registration number,

depository, description of specimen, locality, collector, date) of the holotype and paratypes

must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,

Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

e.g. . the Figure depicting C. namacolus . . in C. namacolus (Fig. 10) .

(b) The WeiRES of prefixed surnames in all inneaanest iia used in the text, if not pe

by initials or full names

e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a

book or article, such as

“Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation

to initial capital letter, provided the same generic name is used consecutively.

Name of new genus or species is not to be included in the title: it should be included in the

abstract, counter to Recommendation 23 of the Code, to meet the requirements of

Biological Abstracts.