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BULLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

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FISCHER, P.-H., DuvAL, M. & RarFy, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

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

By

PAE eAND ER HUI yi
&
MATTHIAS STEHMANN

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
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Court Road, Wynberg, Cape Courtweg, Wynberg. Kaap

THE VALIDITY OF MALACORAJA STEHMANN, 1970
(CHONDRICHTHYES, BATOIDEI, RAJIDAE) AND ITS
PHYLOGENETIC SIGNIFICANCE

By
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

(c) Scientific names, but not their vernacular derivatives
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.

SMITHSONIAN INSTITUTION LIBRARIES
“PACITY ANIA
3 9088 01206 6551

P. ALEXANDER HULLEY & MATTHIAS STEHMANN

THE VALIDITY OF MALACORAJA STEHMANN, 1970
(CHONDRICHTHYES, BATOIDEI, RAJIDAE)
AND ITS PHYLOGENETIC SIGNIFICANCE