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- ANNALS OF THE _

SOUTH AFRICAN MUSEUM |

VOLUME 60

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Pais OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 60

SEE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN KAAPSTAD

1972-1973

% PRINTED IN THE REPUBLIC OF SO

THE RUSTICA PRESS (PTY.) LTD., \

1). 0nA

LIST OF CONTENTS

Cooper, M. R.

The Cretaceous stratigraphy of San Nicolau and Salinas, Angola (published Novem-

ber 1972)

GriFFiTHs, C. L.

The Amphipoda of southern Africa. Part 1. The Gammaridea and i amen of

southern Mocambique (published April 1973)

Hesse, A. J.

New Mydaidae (Diptera) from the Namib Desert and south-western Africa (pub-

lished November 1972) ; me

Huey, P. A.

Mesopelagic fishes from Vema Seamount (IK Station 52) (published November

1972)

Hurtzy, P. A.

A new species of southern African brevirajid skate (Chondrichthyes, Batoidei,

Rajidae) (published November 1972)

Hu tey, P. A.

The origin, interrelationship and distribution of southern African Rajidae (Chon-

drichthyes, Batoidei) (published November 1972)

Huey, P. A.

The rare plectognath fish, Macrorhamphosodes uradoi (Kamohara) (Triacanthodidae) in

South African waters (published November 1972)

Huttey, P. A.

A report on the mesopelagic fishes collected during the deep-sea cruises of R.S.

‘Africana II’, 1961-1966 (published November 1972)

KENSLEY, B. F.

Pliocene marine invertebrates from Langebaanweg, Cape Province ee

November 1972) : 3 : a ye :

PenritH, M. J. & PENRITH, M.-L.

Redescription of Pandaka silvana (Barnard) (Pisces: Gobiidae) (published November

1972) as Ss Ss he ae ns 3s Ee e:

Page

109

1gI

173

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Page

Janice Griffiths, 1973... age oy ty 2 #2 ry Ye Ae site| BBO

Mimadelphus Hesse, 1972 (Mydaidae, Diptera) 155

Namibimydas Hesse, 1972 (Mydaidae, Diptera) a a i si Be x, 158

Parectyphus Hesse, 1972 (Mydaidae, Diptera) a sé if a y. als 165

| VOLUME 60 PART 1 NOVEMBER 1972

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aNNALS OF THE SOUTH AFRICAN ' MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60 Band

November 1972 November

Part I Deel

THE ORIGIN, INTERRELATIONSHIPS

AND DISTRIBUTION OF SOUTHERN AFRICAN

RAJIDAE (CHONDRICHTHYES, BATOIDEI)

P. A. HULLEY

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 ORIGIN, INTERRELATIONSHIP AND DISTRIBUTION OF

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI)

PAS UL LEN.

South African Museum, Cape Town

(With 59 figures and 4 tables)

[Ms. accepted 27 March 1972]

CONTENTS

PAGE

Introduction . ‘ é i ; : I

Materials and methods ‘ , : 2

Pelvic girdle and clasper structure : : 3

Vertebral count : 3 ; : : 45

Neurocranium and rostral cartilages. : 56

Discussion : : : ; : : 66

Phyletic interrelationships . : ‘ ; 79

Distribution . é ; , : : 86

Summary : : : : ’ ; 97

Acknowledgements. : s : 2 98

References : : : ; : 5 99

INTRODUCTION

Skates, which show the greatest species diversity among elasmobranch

fishes, are included in the family Rajidae, and are characterized by their

dorso-ventrally flattened, rhomboidal disc, moderately slender tail, usually

with two dorsal fins and a membraneous caudal fin, and lack of serrate tail

spines. Although they are regularly taken in bottom trawls, together with

stockfish (Merluccius capensis), kingklip (Genypterus capensis), jacopever (Helicolenus

dactylopterus) and monkfish (Lophius piscatorius), they form a small proportion of

the total catch (0,001-0,075°%) by South African commercial trawlers, and

their landed weight has shown a sharp decline from 131 814 lb in 1963 to

14. 199 lb in 1965 (Division of Sea Fisheries Report, 1968).

In order to clarify the position of the Rajidae within the southern African

ichthyofaunal complex, a research programme was instigated by Dr F. H.

Talbot, formerly of the South African Museum, to investigate the systematics,

distribution and phylogeny of the group. Hulley (1966, 1969, 1970) has

revised the Rajidae of the west and south coasts of southern Africa, at the

species level, while Wallace (1967) has investigated the east coast species.

From this, it appears that natural relationships between the species are evident,

suggesting a regrouping of the Raja species at the generic or subgeneric level.

Because of difficulties associated with individual variation, Ishiyama

(1952, 1958, 1968) adopted a new approach to rajid systematics, employing

characteristics of the clasper, structure of the neurocranium, number of

Ann. S. Afr. Mus. 60 (1), 1972: 1-103, 59 figs, 4 tables.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

intestinal valves and valves in the conus arteriosus, and the vertebral count, as

well as standard morphometric procedures, and has established several new

genera and subgenera. Stehmann (1970) has followed this method, supple-

mented by investigations on the ampullae of Lorenzini and number of pseudo-

branchial folds in the spiracle, and has proposed 6 subgenera within the genus

Raja in the eastern North Atlantic. It appears, however, that of these characters

only the anatomy of the clasper, shape of the skull and rostral cartilages, and

vertebral count prove to be definite key characters.

These methods have been adopted in this investigation, not only for

comparison of southern African species with their European counterparts,

but also for establishing interrelationships within the family and the possible

origin of the southern African rajid fauna.

The scheme of classification followed in this paper is that of Bigelow &

Schroeder (1953).

This work formed part of a thesis submitted for the degree of Ph.D. at the

University of Cape ‘Town in October 1971, and the author is indebted to the

University, and particularly the Zoology Department, for permission to

publish the results.

MATERIALS AND METHODS

489 specimens, including the types of the following species were examined

in detail: Cruriraja parcomaculata, C. triangularis, Raja doutrei, R. pullopunctata,

R. stenorhynchus, R. lanceorostrata, R. springeri, R. alba, R. spinacidermis, R. miraletus,

R. clavata, R. straeleni, R. robertsi, R. radiata, R. wallacei, R. caudaspinosa, R. ravidula,

R. leopardus, R. confundens, R. dissimilis, Bathyraja smithi. All material in the

collections of the J. L. B. Smith Institute of Ichthyology, Grahamstown, and the

Oceanographic Research Institute, Durban, has been re-examined.

Material collected in the South Atlantic by the Discovery, FFS Walther

Herwig and the Belgian South Atlantic Expedition was seen during visits to

London, Hamburg and Brussels, while specimens from the eastern North

Atlantic, housed in the British Museum (Natural History), Institut fir

Seefischerei, Institut Royal des Sciences Naturelles de Belgique, and Muséum

National d’Histoire Naturelle were examined, including specimens from

Sierra Leone and Senegal.

Specimens of Cruriraja rugosa and Anacanthobatis americanus were obtained

from the National Museum of Natural History, Washington, for comparative

anatomical studies, and this institution also supplied X-ray photographs of

Raja garmani, R. oregoni, Pseudoraja fishert, P. atlantica and Gurgesiella furvescens.

Further X-ray photographs of Raja flavirostris, Psammobatis extenta, P. microps,

P. lima and P. scobina were supplied by the British Museum (Natural History),

of Raja straeleni by the Institut fiir Seefischerei and of Raja muraletus and

Rk. straeleni by the Institut Royal des Sciences Naturelles de Belgique.

The pelvic girdles of southern African Rajoidea have been examined by

means of X-ray photography and the drawings made from the negatives have

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 3

been scaled to the same width for comparative purposes. X-ray photographs

of the pelvic girdle of Springeria ort did not prove to be completely satisfactory,

but it appears that this is not unlike the girdle of Anacanthobatis marmoratus.

Claspers from adult males of the following species have been examined in

detail by dissection: Crurtraja parcomaculata, C. triangularis, C. rugosa, Raja doutrei,

R. pullopunctata, R. lanceorostrata, R. alba, R. caudaspinosa, R. miraletus, R. clavata,

R. straelenit, R. dissimilis, R. confundens, R. leopardus, R. wallacei, Bathyraja smithit,

Anacanthobatis marmoratus, A. americanus, Rhinobatos annulatus, Myliobatis cervus. All

material had been preserved in either 70% ethyl alcohol or 5% formalin.

Where possible several dissections were made for the particular species. Brief

examinations of the external morphology of the claspers of other South African

Batoidei have been made for comparative purposes: the species include

Rhinobatos ocellatus, Gymnura javanica, Mobula diabolus, Dasyatis thetidis, Urotrygon

daviest, Heteronarce garmani and Torpedo nobiliana.

Claspers of European Raja clavata, R. radiata, R. miraletus and R. batis and

the South American species R. flavirostris have also been dissected for

comparative purposes.

Vertebral counts were facilitated by the use of X-ray photography. The

number of predorsal caudal vertebrae (Vprd) has been taken as the number of

caudal vertebrae up to the origin of the first dorsal fin (Ishiyama 1952; Krefft

1968a); the number of trunk vertebrae (Vtr) and total count (V2) are given

according to Krefft (1968a). Vprd counts were not possible in Anacanthobatis

marmoratus, A. americanus and Springeria ori, as these species lack dorsal fins.

Although Bigelow & Schroeder (1953) have employed X-ray photography

in examination of the snout, this method has not been used in this study,

because of the small extent of calcification of the rostral cartilages and appen-

dices (Ishiyama & Hubbs 1968). In all cases, the rostral cartilages and rostral

appendices were examined by dissection, preserved material being soaked in a

weak (2%) solution of NaOH, according to the method of Stehmann (1970).

Where possible, the neurocrania were also examined in this way, but in several

cases, Cruriraja triangularis, Raja lanceorostrata, R. springert, R. stenorhynchus,

R. straeleni, R. robertsi, R. dissimilis, R. ravidula, R. wallacet and R. spinacidermis,

dissection of neurocrania could not be made. These were examined by X-ray

photography. The neurocrania of Anacanthobatis marmoratus and Springeria ori

could not be examined by dissection, while X-ray photography gave poor

results. The cranial characters of these species are therefore not included in the

study. Nerve foramina were affirmed by dissection.

PELvic GIRDLE AND CLASPER STRUCTURE

PELVIC GIRDLE

The pelvic bar is made up of a pair of anlage, which arise together with the

basals and peripheral radials in a continuous procartilaginous rudiment

(Balfour 1881). While the anlage remain separate in the Holocephali, they fuse

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

to form a single element, situated in front of the cloaca in Elasmobranchii.

Subdivision of the anlage in Cladodus suggests that the pelvics arose from a fusion

of originally independent basal fin elements.

Contrary to Wiedersheim’s theory (1892), Braus (1902) has shown that the

pelvic girdle consists of a transverse, bow-like cartilage, from which plate-like

girdles, e.g. Chlamydoselachus (Daniel 1934: fig. 85), may be derived. The trans-

verse band of cartilage is divisible into a median ischiopubic region and lateral

iliac regions, which may be slightly expanded and which are penetrated by

obturatoria! nerve foramina for the passage of the diazonal nerves. The number

of foramina may vary considerably, from one in Squalus acanthias to eight in

Chlamydoselachus (Kalin 1939).

Transverse pelvic bars occur in all batoid families, except the Dasyatidae,

Myliobatidae and Mobulidae (Garman 1913: figs 53, 54), in which they are

anteriorly arched and bear a single, median, prepelvic process—processus

a b

Fig. 1. Pelvic girdles of Batoidei. a. Rhinobatos

(after Daniel 1934); b. Platyrhina (after

Garman 1913); c. Arhynchobatis (after Garrick

1954); d. Dasyatis (after Garman 1913);

e. Mbyliobatis (after Garman 1913); f. Mobula

(after Garman 1913).

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI!) 5

praepubicus. However, in the Rhinobatidae, Platyrhinidae, Rajidae, Anacantho-

batidae and Arhynchobatidae, a pair of lateral prepelvic processes arise from

the antero-lateral edge of the pelvic bar and are directed anteriorly.

Besides the brief references to the pelvic girdle by Garman (1913) and

Garrick (1954, 1957), no comparative investigation of the girdle in Rajoidea

has been made.

Typically, the girdle is short and bar-like and is expanded laterally at the

iliac regions. Arising from the antero-lateral edges of the iliac regions are a pair

of prepelvic processes. In the Anacanthobatidae, the iliac regions are more

developed, which together with an increase in length of the prepelvic processes,

gives the girdle a U-shape. The girdle appears to be of intermediate shape in

Cruriraja species and in Arhynchobatis asperrimus (Garrick 1954: fig. 3), so that the

trend in this mode of development is shown by the series Raja, Cruriraja,

Arhynchobatis and Anacanthobatis (Fig. 2).

The lengths of the prepelvic processes vary considerably; in Raa species,

their lengths vary from 0,1 to 0,3 of the girdle width; in Bathyraja smithiu

Fig. 2. Pelvic girdles of southern African

Rajidae, showing sexual dimorphism.

a. Raja springeri (male); b. R. springeri

(female); c. Raja confundens (male);

d. R. confundens (female); e. Cruriraja

parcomaculata (male); f. C. parcomaculata

(female); g. Anacanthobatis marmoratus

(male) ; h. A. marmoratus (female).

tonct

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

about 0,3; in Cruriraja species from 0,1 to 0,2; in Arhynchobatis asperrimus

about 0,3; and in Anacanthobatis species from 0,8 to 0,9.

A short, stout, iliac process arises postero-laterally on each side from the

dorsal surface of the bar, but this recurves inwards to point anteriorly and

terminates in a bluntly rounded point. The iliac region is usually penetrated

by two (sometimes more) obturatorial foramina in most species, but in Cruriraja

parcomaculata, C. triangularis and Anacanthobatis marmoratus and in the North

Atlantic species Cruriraja rugosa and Anacanthobatis americanus, there is a single

foramen on each side (Fig. 4). Garrick (1954: fig. 3) has shown a single foramen

in the monotypic Arhynchobatis asperrimus.

Sexual dimorphism is evident in the pelvic girdles (Fig. 2), and although

it is not particularly marked in Raa and Bathyrga species, it is noticeable in

Cruriraja, Anacanthobatis and Springeria. In these genera, the posterior edge of the

ischiopubic region is strongly arched in males, due partly to the increase in size

of the iliac regions, while in females the posterior edge is only slightly arched,

giving rise to a much more elongate bar between the prepelvic processes. This

feature is probably linked with the oviparous behaviour of the suborder.

The ‘black-bellied’ skates, Raja doutret, R. stenorhynchus, R. springeri,

R. pullopunctata and R. lanceorostrata (Figs 3 a—e) are unusua! in that they show

a graded variation in anterior arching of the bar, which is unlike the simple,

transverse bar of most other species of Raja. This type of arched bar, as has been

pointed out above, approximates the condition in the Dasyatidae, Myliobatidae

and Mobulidae (Fig. 1). However, in these families there is a single, median,

prepelvic process, while in black-bellied skates the paired, lateral processes

indicative of the rajid condition, are present.

In the clavata-group (Figs 3 g, i, j), which in the eastern South Atlantic

consists of the species Raja clavata, R. straeleni and R. miraletus, the prepelvic

processes show an increase in length from Raja straeleni to R. clavata. In the latter

species the length of the prepelvic processes approximates to that of Arhynchobatis

asperrimus. Furthermore, slight anterior arching of the ischiopubic bar is seen in

Raja miraletus and R. clavata.

The girdle of Bathyraja smithti (Fig. 3 t) resembles that of Raja clavata and

R. miraletus in the length of the prepelvic processes, but differs in the iliac

processes. These are only slightly curved in Bathyraja smithu, and so attain a

position intermediate between that of Rhinobatos (Fig. 1 a) and most typical

rajids. It should be noted that a similar condition of the iliac processes is found

in Raja robertsi (Fig. 3 s), but in this species the prepelvic processes are much

shorter than in Bathyraja smithii, and the girdle represents a case intermediate

between that of Raja radiata (Fig. 3 r) and Bathyraja smithit.

A straight, transverse bar is found in the species Raja dissimilis, R. ravidula,

R. confundens, R. wallacei, R. leopardus, R. caudaspinosa and R. spinacidermis

(Figs 3 k-q), in which cases there is some variation in the prepelvic length.

However, in Raya wallacei and R. caudaspinosa (Figs 3 n, p), the iliac processes

appear to be more strongly developed than in the other species, except Raja

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE!) 7

ee ae

i i bce

ie a a

Fig. 3. Pelvic girdles of southern African Rajidae. a. Raja doutrei; b. R. stenorhynchus; c. R. springert;

d. R. pullopunctata; e. R. lanceorostrata; f. R. flavirostris (South American) ; g. R. straeleni; h. R. alba;

i. R. miraletus; j. R. clavata; k. R. dissimilis; 1. R. ravidula; m. R. confundens; n. R. wallacei;

o. R. leopardus; p. R. caudaspinosa; q. R. spinacidermis; r. R. radiata; s. R. robertsi; t. Bathyraja smithit.

spinacidermis (Fig. 3 q), which represents the extreme case. In this species the

iliac processes extend forward to the anterior edge of the pelvic bar, and in one

specimen (the type) the processes extend beyond the edge.

A similar condition is seen in Cruriraja species (Figs 4 a—c), where the

length of the prepelvic processes approximates that of typical rajids, but where

the iliac processes are more massively developed. However, in Cruriraja parco-

maculata, C’. triangularis and C. rugosa, the iliac processes do not reach the anterior

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

edge of the bar, as they do in Raja spinacidermis.

In Anacanthobatis marmoratus (Fig. 4 d), A. americanus and Springeria ori, it is

the prepelvic processes which are strongly developed, being almost equal in

length to the width of the girdle. On the other hand, the iliac processes are

reduced to small, truncate protuberances.

d e

Fig. 4. Pelvic girdles of southern African Rajoidea. a. Cruriraja

parcomaculata; b. C. triangularis; c. C. rugosa (North Atlantic) ;

d. Anacanthobatis marmoratus; e. A. americanus (North Atlantic).

CLASPER STRUCTURE

Although there are major differences in the siphon and clasper gland

structure between Selachii and Batoidei (Leigh-Sharpe 1926; White 1937), it is

the distal end of the clasper which is important in the systematics of lower taxa.

Anatomical investigations on the claspers of European rajids have been made

by Petri (1878), Jungersen (1899), Huber (1901), Leigh-Sharpe (1920-6) and

Stehmann (1970), while Ishiyama (1958) and Ishiyama & Hubbs (1968) have

made detailed studies of the structure and systematic significance of the organ

in Japanese rajids. Hulley (1966, 1969) has described the clasper structure of

several South African species, and although he has figured the external structure

of the claspers in eastern South Atlantic species (1970), detailed comparative

anatomical studies of these organs have not been made.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 9

Terminology of external features

Leigh-Sharpe (1920-6) defined most of the terminology used in clasper

anatomy, but it can be seen that his use of a particular term is not always

constant, since the same cartilaginous element is not always involved. The

reason for this is that he dealt only with the external morphology of the organ,

without reference to the internal cartilages.

Furthermore, differences in terminology of clasper components as employed

by Leigh-Sharpe (1920-6), and by Ishiyama (1958) and Ishiyama & Hubbs

(1968), have added to the confusion. In order to standarize, Stehmann (1969)

evaluated the terms pseudosiphon, slit, rhipidion and spur, and gave precise

definitions for his nomenclature. While the terminology used here is mainly in

accordance with his findings, there appears to be confusion in several other

structures. A discussion of all problematic components, applicable to southern

African Rajidae, follows below:

(1) pseudosiphon

Leigh-Sharpe (1921: 361) introduced the term pseudosiphon for Galeus

vulgaris, defining it as ‘a small blindly ending sac, whose wide aperture points

in a posterior direction’. He shows that the pseudosiphon is situated on the

outer surface of the dorsal lobe of the clasper glans (1921: fig. 2). However, in

Raa species (R. clavata, R. blanda, R. marginata), he indicates that the pseudo-

siphon lies on the inner surface of the dorsal lobe. Ishiyama (1958) gives a

| o

Fig. 5. Raja lanceorostrata. A. external view of right clasper

from dorsal side; B. lateral view of right clasper opened to

show structural features of the glans. Scale 2,0 cm.

cf—cleft; hp—hypopyle; rh—rhipidion; sh—shield; sp—

spike; st—sentinel.

IO ANNALS OF THE SOUTH AFRICAN MUSEUM

similar definition and points out that the pseudosiphon is always related to

modifications of the dorsal terminal cartilages.

In 1968, Ishiyama & Hubbs defined the pseudosiphon more precisely,

relating it to the degree of development of a single cartilage, the dorsal

terminal 1. They point out that a pseudosiphon is indicative of the genus

Bathyraja.

According to Stehmann (1969), Ishiyama’s first definition of the pseudo-

siphon could be interpreted to mean that the structure could be found either on

the outer surface or the inner surface of the dorsal lobe. However, Ishiyama

(1958: fig. 3) has shown that the pseudosiphon is situated externally. Further,

in his discussion of the slit, he points out that this structure may be related to

dorsal terminal cartilages, but (1958: 202) ‘has no close relation to the dorsal

terminal 1’, inferring that the pseudosiphon does have a relationship with that

cartilage. Therefore, it appears that there is no distinction between the

Ishiyama (1958) definition and the Ishiyama & Hubbs (1968) definition.

In all species investigated in this paper, except those of the genus Cruriraja,

the dorsal terminal 1 cartilage overlies the other dorsal terminals, so that if a

pseudosiphon is related directly to the dorsal terminal 1, then it must be

situated externally, on the outer surface of the dorsal lobe of the glans. It would

appear, therefore, that the structure referred to as a pseudosiphon by Leigh-

Sharpe (1920-6) in the species Raja clavata, R. blanda and R. marginata, which

occurs on the inner surface of the dorsal lobe, should be given a new name. In

! a

Fig. 6. Raja radiata (North Atlantic). A. external view of right clasper from dorsal

side; B. lateral view of right clasper opened to show structural features of the glans.

Scale 2,0 cm.

ap—apopyle; cf—cleft; hp—hypopyle; pe—pent; rh—rhipidion; ps—pseudo-

siphon; sl—slit; sh—shield; sp—spike; sr—spur; st—sentinel.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) II

Raja clavata, R. blanda and others, Stehmann (1969, 1970) has referred to this

structure as the ‘inner pseudosiphon’, but because it houses the sentinel in these

species, Hulley (1970) has proposed the term pocket. This has been followed by

Stehmann (19714, b), who recognizes a pocket, rather than an inner pseudo-

siphon, in Raja herwigi and R. maderensis. However, it should be noted that the

pseudosiphon which Leigh-Sharpe (1924) described in Raa marginata is

entirely different to the pocket, and has been referred to the term cleft.

Stehmann (1969) expresses some concern as to the question of priority in

terminology. However, an accurate definition for the pseudosiphon was never

given, since Leigh-Sharpe (1920-6) made numerous misidentifications of this

structure, presumably discriminating it by its position in the clasper alone.

Ishiyama’s definition can therefore be used (see p. 19).

It should be noted that Stehmann (1970) has distinguished a pseudosiphon

in the species Raja fyllae. I have found that in this species, as in Raja wallacei

(Fig. 7 A) and R. caudaspinosa, the cavity on the outer dorsal wall is formed by

an indentation in the musculature which is attached to the dorsal terminal 1

(m. dilatator), and further that this cavity is not found in all specimens. I have

therefore not described this as a real and separate structure, comparable to the

pseudosiphon. Stehmann (personal communication) feels that this must be left

to the discretion of the author.

In Cruriraja species, the dorsal terminal 1 is differently situated (Figs 40, 41,

42), so that it does not overlie the other dorsal terminal cartilages to any great

Fig. 7. Raja wallacei. A. external view of right clasper from dorsal side; B. lateral

view of right clasper opened to show structural features of the glans. Scale 2,0 cm.

ap—apopyle; cf—cleft; dd—dermal denticles; hp—hypopyle; pt— promontory;

th—rhipidion; rl—roll; sh—shield; sl—slit; sp—spike; sr—spur; st—sentinel.

(9 ANNALS OF THE SOUTH AFRICAN MUSEUM

extent. However, a small cavity is formed between the distal end of the dorsal

terminal 2 and the concavity of the dorsal terminal 1. This cavity is situated on

the inner surface of the dorsal lobe of the glans. Because of its close association

with the dorsal terminal 1 cartilage, I have termed this structure the pseudo-

siphon, but it is obviously formed in a different manner and lies transversally to,

rather than in the longitudinal axis of, the clasper. It may therefore necessitate

the introduction of a new term.

—~1

era .

Fig. 8. Cruriraja triangularis. A. external view of right clasper from dorsal side.

B. ventral surface of right clasper; C. lateral view of right clasper opened to show

structural features of the glans. Scale 1,0 cm.

ap—apopyle; dd—dermal denticles; ep—eperon; hp—hypopyle; kb—knob;

ps— pseudosiphon; rh—rhipidion; sh—shield; sp—spike; st—sentinel; th—thorn.

In some Batoidei (Figs 12 A—H), there is a cavity situated on the outer

surface of the ventral lobe of the glans, which I have termed the ventral

pseudosiphon, again because of its close association with the dorsal terminal 1

cartilage. This corresponds to the lateral pocket of Jungersen (1899). Leigh-

Sharpe (1920-6) has termed this the slit (see below), sentina and crumena in

Torpedo marmorata, Rhinobatos productus and Cestracion philippi respectively, but in

others, Trygon pastinacea and Myliobatis aquila, has left it unlabelled. Similarly,

Ishiyama (1958) does not label this structure in Rhinobatos schlegeli and Platyrhina

sinensis.

(2) slit

Leigh-Sharpe (1920-6) does not clearly define the term slit, mentioning it

only as a cavity, which may be internal, e.g. Raja batis, R. lintea, or may be

situated on the outer surface of the dorsal lobe, e.g. Raja murrayi, R. eatont.

From its position in the latter species, and from the fact that Raja eatoni is

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) 13

considered to be synonymous with Bathyraja smithi (Hulley 1970), it may be

concluded that these cavities correspond to pseudosiphons, according to the

above proposed definition.

Ishiyama (1958: 202) defines the slit as a ‘small, shallow cavity formed by

fleshy skin on the inner surface of the dorsal lobe and has no relation to dorsal

terminal 1’.

Fig. 9. Cruriraja rugosa. A. external view of right clasper from dorsal side. B. lateral

view of right clasper opened to show structural features of the glans. Scale 2,0 cm.

ap—apopyle; ep—eperon; hp—hypopyle; kb—knob; ps—pseudosiphon; rh—

rhipidion; sh—shield; sp—spike; st—sentinel; th—thorn.

Stehmann (1969) points out that Ishiyama has not used the term con-

sistently, especially where there are two or more cavities on the inner surface of

the dorsal lobe. He recognizes that two structures (not including the pseudo-

siphon) may be involved: (1) a skinny flap, situated proximally in the glans,

which has no relation to any dorsal terminal cartilage, and is so orientated that

the sac lies at right angles to the longitudinal axis of the clasper; and (2) a cavity

which may be formed between any dorsal terminal cartilage (other than the

dorsal terminal 1) and the axial or dorsal marginal cartilage, i.e. it is a cavity

which is bounded by cartilaginous elements. The first cavity is regarded as the

slit, the second has been termed the cleft. I agree with this distinction, and have

subsequently used Stehmann’s definitions for these dorsal cavities.

It should be noted that the cavity formed in the ventral lobe of the clasper

in Dasyatidae and Mobulidae, and which has been referred to as a slit by Leigh-

Sharpe (1920: fig. 15; 1926: figs 12, 14, 15), is now referred to as a ventral

pseudosiphon. The slit is therefore always situated on the inner surface of the

dorsal lobe of the glans.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

(3) rhipidion: pseudorhipidion

Definitions for both these structures have been given by Ishiyama (1958),

who relates the smooth-surfaced, dorsally situated pseudorhipidion to the distal

end of the dorsal marginal cartilage, while pointing out that the pitted, ventrally

situated rhipidion is composed entirely of erectile tissue and has no relation to

cartilaginous elements. Stehmann (1969), however, points out that Ishiyama is

incorrect in his restriction of the term rhipidion to Raja species (1958) or Raya

and Breviraja species (Ishiyama & Hubbs 1968), and pseudorhipidion to

Bathyraja, since members of his clavata-group (subgenus Raja: Stehmann (1970))

possess a pseudorhipidion. The presence of a pseudorhipidion is therefore not a

suitable character for the separation of the genus Bathyraja from other rajids.

Fig. 10. Anacanthobatis marmoratus. A. external view of right clasper

from dorsal side; B. lateral view of right clasper opened to show

structural features of the glans. Scale 1,0 cm.

ap—apopyle; cf—cleft; ep—eperon; hp—hypopyle; rh—rhipidion;

sh—shield; sp—spike; sr—spur; st—sentinel.

(4) spur

Leigh-Sharpe (1920-6) described this structure in Raa radiata, referring to

it as a hard, cartilaginous point, which curves outward from the dorsal lobe of

the glans (Fig. 6 B, sr). Stehmann (1969, 1970) has shown that this structure is

directly related to the dorsal terminal 3 cartilage. On the other hand, Ishtyama

(1958) refers the spur to a structure on the inner surface of the ventral lobe,

which is formed by the accessory terminal, while Ishiyama & Hubbs (1968:

figs 2 A, B) refer the spur to the ventral terminal cartilage. It is felt that

Leigh-Sharpe and Stehmann should be followed, and that Ishiyama should

redefine his structures.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 15

(5) funnel

The term was introduced by Ishiyama (1958: 203) as ‘a component

located on the distal portion of the ventral lobe, having a soft projection like the

foot of a bivalve mussel’. He relates this structure directly to the ventral

terminal cartilage.

It can be seen that in those species in which the funnel is present (Ishiyama

1958: fig. 4) it always overlies the ‘spur’, where this is present, on the inner

ventral surface. Examination of the corresponding cartilages (Ishiyama 1958:

fig. 12) reveals that in these species the only cartilage which can possibly overlie

the accessory terminal (‘spur’) is the distal end of the ventral marginal.

Furthermore, Ishiyama (1958) introduces the term claw for the species

Rhinoraja odai and Rhinoraa longicauda, relating this to a modification of the

accessory terminal cartilage. However, he shows (1958: fig. 11) that the

accessory terminal in these species is a single cartilage, which is flat and sharp-

edged and quite unlike the external claw. Again, examination of the figures

(Ishiyama 1958: fig. 12) shows that the cartilage which overlies the accessory

terminal (knife), and is extended distally, is the ventral marginal cartilage.

Fig. 11. Anacanthobatis americanus. Lateral view of

right clasper opened to show structural features

of the glans. Scale 1,0 cm.

g—clasper groove; ps—pseudosiphon; rh—

rhipidion; sp—spike; st—sentinel.

This means therefore that neither the ventral terminal nor the accessory

terminal can be related to the term funnel or claw, and that clearly the ventral

marginal is the associated cartilage in both cases. Further confusion is added

because the claw in Rhinobatos schlegeli and Rhinobatos annulatus (Ishiyama 1958:

figs 13, 14) is formed by the accessory terminal cartilage.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 7

Stehmann (1969) correctly follows Ishiyama’s definition, relating the term

funnel to the ventral terminal cartilage, while introducing Leigh-Sharpe’s

term, projection, for the distally situated, elongate, cartilaginous structure

formed by the ventral marginal cartilage. Hence the term projection of

Stehmann (1969, 1970) is equivalent to the terms funnel/claw of Ishiyama,

while the term funnel of Stehmann is clearly a different structure.

It should be noted that although Leigh-Sharpe (1926) did not include the

term projection in his list of clasper components, and does not adequately define

this structure, he shows that in Raja lintea (1924: fig. 6) the projection is an

expansion on the inner lateral edge of the shield. This can therefore be related

directly to the ventral terminal cartilage, and as such, is a different structure

to the projection of Stehmann.

It is suggested that the term funnel be applied to the distal protrusion of

the ventral terminal cartilage, as in Raja clavata (Fig. 22), while the structure

which is related to the distal protrusion of the ventral marginal be known as the

projection. The adoption of this convention would avoid further confusion,

especially with regard to Stehmann’s definitions.

The structure which is regarded as the projection by Leigh-Sharpe must

therefore be referred to some other term, if in fact it is an identifiable and real

clasper component. 3

Furthermore, the projection as now defined would cover the terms spike in

Raja murrayi and the sentinel in Raja eatoni (Leigh-Sharpe 1924: figs 13, 14). The

term claw should be retained and reserved solely for the structure formed by the

accessory terminal cartilage in Rhinobatos, Platyrhina and Myliobatis.

(6) pecten

The structure was recognized as a component in Raja marginata (= Raja

alba) and was defined (Leigh-Sharpe 1924) as a long, hard structure, bearing

six or more hard projections or serrations, which is situated on the outer lateral

margin of the dorsal lobe of the clasper.

Dissection of the clasper of Raja alba (Fig. 33) has revealed that the pecten

is directly related to the sharp protuberances or serrations, which are developed

on the outer lateral margin of the dorsal terminal 3 cartilage. A pecten was also

described in the clasper of Raja miraletus (Hulley 1969), but in this species the

dorsal terminal 2 cartilage is involved. However, since the serrations are

developed in a similar position to those in Raja alba, and since the orientation

and probably the function is identical, I have termed both structures the

pecten. This has been followed by Stehmann (1971a), who distinguishes a

pecten, formed by the dorsal terminal 2 cartilage, in Raja herwigi.

(7) scale and dermal denticles

Ishiyama (1958) pointed out that in two Japanese subspecies, Bathyraja

smirnovi smirnovi and B. smirnovi ankasube, minute scales were present on the

outer border of the ridge. As such a phenomenon had not been reported for

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

European rajids, he recognized them as a separate clasper component, which

he defined as the scale.

Dermal denticles are present on the outer surface of the dorsal lobe of the

claspers of Raja confundens, R. leopardus and R. wallace: (Hulley 1970: figs 17-19).

Although the lateral area would correspond with the ridge, if this had been

developed, I have preferred to regard such denticles as a separate component

to Ishiyama’s scale.

This means, therefore, that scale is a term which is restricted to dermal

denticles on the ridge, and is probably only applicable in the genus Bathyraja,

in which the ridge is developed. Dermal denticles are found in other rajid

genera. However, both these components have a similar derivation and as such

could possibly be referred to the same term.

(8) terminal bridge

Ishiyama (1958) first described the terminal bridge as a short, bar-like

cartilage, which connects the axial cartilage with the distal end of the dorsal

terminal 2 cartilage. He derived the terminal bridge from an offshoot of the

accessory terminal 2. However, he does not recognize this as a separate clasper

component (1958: tables 3, 4). Hulley (1966, 1969) also recognized the

existence of this linking cartilage in Raja pullopunctata, where it separates the two

dorsal elefts, and in Raja miraletus, where it separates the cleft from the pocket.

In both species, the terminal bridge links the axial with the dorsal terminal 2

cartilage.

Stehmann (1969, 1970, 1971a, 6) regards the terminal bridge as a real

component, identifiable in the external anatomy of the claspers of Raja brachyura,

R. montagui, R. herwigi, R. maderensis and R. straeleni, and directly related to the

development of one or two small, cartilaginous rods. In these species, the rods

link the axial with the dorsal terminal 2 cartilage and separate the cleft from

the pocket.

I have found this linking cartilage in Raja doutrei, where it joins the axial

to the dorsal terminal 2 (Fig. 30), thereby separating the two clefts in the

dorsal lobe. It is absent in Raja lanceorostrata (Fig. 31), where the dorsal

terminal 2 itself makes contact with the axial. This is the case with Raja radiata,

R. dissimilis, R. caudaspinosa, R. confundens and R. leopardus, but in R. wallacei the

connection is formed between the axial and the dorsal terminal 3 cartilage.

Furthermore, a cartilaginous connection is present in Cruriraja parcomaculata,

C. triangularis and C. rugosa (Figs 40-42), but in these species the connection is

completely different from the above. It is made between the accessory terminal 2

and the axial. It would appear, therefore, that the terminal bridge is not con-

stant in orientation and it is suggested that this element helps bind the cartilages

of the terminal group to the axial, thereby acting antagonistically to the

contraction of the m. dilatator.

Because of this, and because of the different areas of insertion (dorsal

terminal 2, accessory terminal 2), it is felt that the terminal bridge should not

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 1g

be recognized as a separate and distinct component, even though it might be

evident in the external and internal morphology of the glans (cf. Stehmann

19710).

(g) pent

The term was originally used to describe a ventral, ridge-like fold in

Trygon pastinacea (Leigh-Sharpe 1922: fig. 17). This fold is situated at about the

level of the hypopyle and presumably acts in much the same way as the

rhipidion. Ishiyama (1958: fig. 4 A) recognizes a pent in Bathyraa tobitukai,

which is similarly positioned in the glans.

The pent described by Stehmann (1969, 1970) is an elongate ridge, which

is pleated and runs parallel to the midline of the organ, from the distal end of

the rhipidion. Although this does not appear to be equivalent to the above,

Stehmann’s definition has been followed to avoid further confusion.

(10) sentina

The sentina originally described by Leigh-Sharpe (1922) was an external

cavity in Rhinobatos productus. ‘Vhis cavity was subsequently labelled as the slit or

crumena in later papers by this author. As has been pointed out above, the

external cavity on the ventral surface of the clasper has now been termed the

ventral pseudosiphon, because of its association with the dorsal terminal 1

cartilage. The term sentina has been reserved for an internal structure of the

glans, and is defined below.

(11) promontory

The promontory recognized by Ishiyama & Hubbs (1968: figs 2 C, D) in

Breviraja colesi is clearly related to the proximal region of the dorsal terminal 3

cartilage. On this basis, the promontory was recognized in Raja wallacer

(Hulley 1970: fig. 19). However, Leigh-Sharpe (1920-6) and Stehmann (1969,

1970) regard the promontory as the distal projection of the dorsal terminal 2

cartilage, while the structure associated with the dorsal terminal 3 is termed the

roll. This interpretation is accepted and to avoid confusion the structures have

been reidentified and relabelled in Raja wallace: (Fig. 7).

Description

Definitions of the external components of the clasper glans of southern

African Rajoidea are given below, and their occurrence is described and

figured (Figs 5-14; Hulley 1970: figs 4-21). For definitions of the apopyle and

siphon gland, the reader should refer to Leigh-Sharpe (1920-6).

Components of the dorsal lobe

(1) pseudosiphon (Ishiyama)

A cavity situated on the outer surface of the dorsal lobe of the glans at about the level of

the hypopyle; it has an oval aperture, the long axis of which is orientated in the longi-

tudinal axis of the organ; its degree of development is directly dependent upon the shape

and location of the dorsal terminal 1 cartilage.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

A pseudosiphon is present in Bathyraja smith (Hulley 1970: fig. 21), Raja

robertsi (Hulley 1970: fig. 12) and R. radiata (Fig. 6) where it is situated on the

outer surface of the dorsal lobe. In these species, the outer lateral margin of the

pseudosiphon is formed by the dorsal terminal 1 cartilage, so that the cavity

can be easily seen and opened by spreading this cartilage. A groove, which is

similarly located, is found in some specimens of Raja caudaspinosa and R. wallacei

(Fig. 7), but since this is developed as a fold in the musculature, and is not

present in all specimens, it has not been identified as the pseudosiphon (cf. Raja

fyllae, R. fullonica: Stehmann 1969; 1970).

In Cruriraja species (Hulley 1970: fig. 4; Figs 8, 9) a pseudosiphon has been

distinguished on the inner surface of the dorsal lobe. Unlike the above, this

cavity has a transverse aperture, but since it has a relationship with the dorsal

terminal i cartilage, it has been termed the pseudosiphon.

The outer lateral edge of the dorsal terminal 1 also forms the dorsal lip of

the pseudosiphon in Anacanthobatis americanus (Fig. 11), which, unlike the

South African species A. marmoratus, possesses a pseudosiphon on the outer

dorsal wall of the glans.

Both Rhinobatos annulatus (Fig. 13) and Myliobatis cervus (Fig. 14) possess a

pseudosiphon in the outer dorsa! wall. However, it should be noted that in these

species, as in Rhinobatos schlegeli and Platyrhina sinensis (Ishiyama 1958), the

dorsal terminal 1 is situated on the ventral side of the glans.

A brief examination of the following southern African Batoidei has

confirmed the presence of a pseudosiphon in Rhinobatos ocellatus, Gymnura

Fig. 13. Rhinobatos annulatus. A. external view of right clasper from dorsal side; B. distal tip of

right clasper; C. lateral view of right clasper opened to show structural features of the glans.

Scale 1,0 cm.

ap—apopyle; cl—claw; dd—dermal denticles; g—clasper groove; hp—hypopyle; ps—pseudo-

siphon; v.ps— ventral pseudosiphon.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE!) P41

; ig

Fig. 14. Myliobatis cervus. A. external view of right clasper

from dorsal side; B. lateral view of right clasper opened to

show structural features of glans. Scale 2,0 cm.

ap—apopyle; cl—claw; g—clasper groove; hp—hypopyle;

ps— pseudosiphon; v.ps—ventral pseudosiphon.

natalensis, Mobula diabolus, Rhinoptera javanica, Heteronarce garmani and Torpedo

nobiliana (Fig. 12). ‘The pseudosiphon in these species is dorsally situated and

should not be confused with the cavity on the ventral surface, which has been

identified as the ventral pseudosiphon (Fig. 12).

(2) cleft (Stehmann)

A small cavity, situated on the inner surface of the dorsal lobe of the glans, which is

formed between the axial or dorsal marginal cartilages and any other dorsal terminal

cartilage except the dorsal terminal 1, i.e. a cavity bounded by cartilages other than the

dorsal terminal 1 cartilage.

Two clefts are present in Raja doutrei, R. pullopunctata, R. lanceorostrata,

R. caudaspinosa and R. dissimilis (Hulley 1970: figs 5, 6, 7, 15; Fig. 5). The

proximal cleft in these species is formed as a cavity between the axial cartilage

and the dorsal terminal 2. A similar gap between these cartilages is found in

Raja wallacei, R. confundens and R. leopardus (Figs 35-37), but a proximal cleft is

not present externally. In Raja confundens and R. leopardus the axial and dorsal

terminal 2 are in close proximity, while in Raja wallacei the gap 1s closed by the

protrusion of the promontory (dorsal terminal 2). A slit is developed in this

region in the species.

The cleft in the species Raja clavata, R. straeleni, R. alba and R. muiraletus

(Hulley 1970: figs 8-11) is differently formed, in that the cartilages bounding

the cavity are the dorsal marginal and the dorsal terminal 2.

A distal cleft occurs in all species except Raja alba, R. clavata, R. straeleni and

R. miraletus; in R. pullopunctata (Hulley 1966: fig. 5) the distal cleft is formed

between the axial cartilage and the dorsal terminal 2; in R. doutrer and

R. lanceorostrata (Figs 30, 31) between the axial and dorsal terminal 3; and in

R. radiata, R. caudaspinosa, R. dissimilis, R. wallacei, R. confundens and R. leopardus

between the axial and dorsal terminal 4 (Figs 34~39). It is interesting to note

2D ANNALS OF THE SOUTH AFRICAN MUSEUM

that when a distal cleft is present, the proximal border of this cleft is always

formed either by the terminal bridge or the dorsal terminal 2 cartilage.

While a proximally situated cleft is present in Anacanthobatis marmoratus

(Fig. 10), itis absent in A. americanus. A cleft is also absent in Bathyraja smithii and

Rhinobatos annulatus.

(3) slit (Leigh-Sharpe)

A small, blindly-ending sac, formed as a skinny flap on the inner dorsal surface of the

proximal region of the glans; it is not bordered by cartilaginous elements and is not

associated with cartilages. Two or more slits may be developed (Ishiyama 1958; 1967).

This structure is present in Raja radiata, R. leopardus, R. wallacei, R. confundens

and Bathyraja smithu (Figs 6, 7; Hulley 1970: figs 17-19, 21). In these, the gap

formed between the axial cartilage and the dorsal terminal 2 appears to be

obscured by the development of other cartilages in this area, i.e. in Raja radiata

by the dorsal marginal (Fig. 34), in R. wallace: by the dorsal terminal 2 (Fig. 35)

and in R. confundens and R. leopardus by the close proximity of the dorsal

terminal 2 cartilage to the axial (Figs 36, 37).

(4) pocket (new term)

A small, shallow cavity, which is situated distally on the inner dorsal wall of the clasper

glans; it is bordered medially by the axial cartilage, while its outer lateral wall consists

of connective tissue, which runs between the dorsal terminal 2 and the axial tip. It

serves as a covering for the sentinel, in those species in which the sentinel is massively

developed.

A pocket is present in Raja clavata, R. straelent and R. miraletus (Hulley 1970:

figs g-11) and has been recognized in R. herwigi and R. maderensis (Stehmann

19714, b).

(5) pseudorhipidion (Ishiyama)

A cartilaginous tongue, situated on the inner lateral margin of the dorsal lobe of the

glans, close to the midline and at about the level of the hypopyle; it is formed by the

distal projection of the dorsal marginal cartilage into the glans. Its function is to

spread the ejaculating spermatozoa (Leigh-Sharpe 1920).

A pseudorhipidion is present in Raja clavata, R. straeleni and R. miraletus

(Hulley 1970: figs 9-11), where it runs parallel to the midline of the organ for

about one-third the length of the glans. However, in Raa alba (Hulley 1970:

fig. 8) the pseudorhipidion is slightly twisted and extends for almost half the

length of the glans; it is divisible into two areas: proximally the dorsal marginal

is covered by loose, fleshy skin, but distally the integument becomes tightly

bonded to the cartilage, so that the median, expanded region of the pseudo-

rhipidion is exposed as a sharp cartilage.

Although Ishiyama (1958) distinguishes a pseudorhipidion in several

Japanese rajids and restricts its occurrence to the genus Bathyraja (Ishiyama &

Hubbs 1968), a pseudorhipidion has not been distinguished in the South African

Bathyragja smithii. It should be noted, however, that in this species the dorsal

marginal cartilage extends well into the clasper glans and has a raised lip on its

outer lateral margin (Fig. 29); this would correspond to the pseudorhipidion,

if it were developed externally.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 23

(6) pecten (Leigh-Sharpe)

A cartilaginous structure located on the outer lateral margin of the dorsal lobe of the

glans, at about the level of the hypopyle; it consists of six or more sharp, comb-like

processes and is associated with the degree of development of the dorsal terminal 2 or

dorsal terminal 3 cartilage.

The pecten is present in Raya alba and R. miraletus (Hulley 1970: figs 8,

9, cm). In Raja straelem, the dorsal terminal 2 (Fig. 18 g) bears a number o

small, blunt protuberances, corresponding in position to the pecten. These do

not penetrate the integument of the outer wall of the clasper, and therefore do

not manifest themselves as a pecten in this species. Stehmann (1971a) recognizes

a pecten in Raa herwigi, which is formed in the same way as in R. miraletus.

(7) dermal denticles (new term)

Dermal denticles may be present on the outer border of the dorsal lobe of the clasper,

where they may be close-set or widely separated.

Dermal denticles are present in Afinobatos annulatus, Raja confundens,

R. leopardus and R. wallace: (Figs 7, 13; Hulley 1970: figs 17-19) on the outer

surface of the dorsal lobe. In Crurirqa triangularis (Fig. 8) they are developed on

the ventral lobe of the glans. The term dermal denticles is discussed above in

relation to the term scale.

(8) ridge (Ishiyama)

The ridge is the thickened and somewhat raised outer lateral border of the dorsal lobe;

it extends from about the level of the hypopyle almost to the distal end of the clasper;

it is directly related to the degree of development of the outer lateral edge of the

dorsal terminal 2 cartilage.

The ridge is present in a single species only, Bathyraja smithu (Hulley 1970:

. 21), in which it appears to be somewhat pleated and does not develop scale.

(9) spur (Leigh-Sharpe)

A hard, cartilaginous element, arising on the outer lateral margin of the dorsal lobe at

about the level of the rhipidion, and curving outwards; it is formed by the dorsal

terminal 3 cartilage, which appears to be movable, so that when the clasper glans is

opened it protrudes from the outer lateral margin.

A spur is present in Raja radiata, R. wallacet and Anacanthobatis marmoratus

(Figs 6, 7, 10). Although the sharp point of the dorsal terminal 3 cartilage did

not protrude through the integument of the outer wall of the cleft in the type of

Raa wallace: (Hulley 1970: fig. 19), examination of the cartilages revealed the

hooked dorsal terminal 3 cartilage. Further material of this species, which has

only just come to hand, confirms the presence of the spur (Fig. 7).

(10) thorn (new term)

A structure consisting or one or more small, sharp points, situated on the outer lateral

margin of the dorsal lobe, at about the level of the hypopyle; it is formed directly from

processes developed on the proximo-lateral edge of the dorsal terminal 1 cartilage.

A thorn has been found in Cruriraja parcomaculata, C. triangularis and

C. rugosa (Figs 8, 9; Hulley 1970: fig. 4).

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

(11) promontory (Leigh-Sharpe)

A cartilaginous element situated at the proximal end of the cleft; it consists of a

cartilaginous tongue, which is more or less covered by a thick layer of integument; it is

formed by the distal projection of the dorsal terminal 2 cartilage.

A promontory has been found only in Raja wallace: (Fig. 7), where it arises

at the medial region of the dorsal lobe and projects into the cleft.

(12) roll (Stehmann)

A fleshy structure situated in the proximo-lateral region of the dorsal lobe and

orientated to run paraliel to the outer margin; it is associated with the promontory and

probably arises as a fleshy expansion of the proximal end of the dorsal terminal 3

cartilage.

A roll has been found only in Raja wallace: (Fig. 7).

Components of the ventral lobe

It should be noted that while certain structures are developed on the outer

surface of the dorsal lobe (pseudosiphon, pecten, dermal denticles), no compo-

nents are developed on the outer surface of the ventral lobe in Rajoidea, except

for dermal denticles in Crunirqa triangulans (Fig. 8 B). However, in some

Batoidei a blindly ending cavity, which has been termed the ventral pseudo-

siphon, because of its relation with the dorsal terminal 1 cartilage, is present

(Fig. 12). The following structures are therefore all found within the glans:

(1) shield (Leigh-Sharpe)

An elongate, plate-like structure, extending along the outer lateral edge of the ventral

lobe, from about the level of the hypopyle; it may be dorsally convex and covered by

pleated epithelia, except along its outer lateral margin, which is sharp-edged; it is

directly associated with the degree of development of the outer lateral margin of the

ventral terminal cartilage.

With the exception of Bathyraja smithit (Hulley 1970: fig. 21) and Anacantho-

batis americanus (Fig. 11), a shield is present in all the examined rajoids. However,

it was found to be absent in other Batoidei.

In the majority of rajids, the shield extends from about the level of the

hypopyle to the distal tip of the clasper, and may be covered entirely by pleated

epithelia, e.g. Raja doutret, R. lanceorostrata, R. alba, R. miraletus (Hulley 1970:

figs 5, 8, 9; Fig. 5) or covered only on its distal half with pleated epithelia,

e.g. Cruriraja parcomaculata, C. triangularis, C. rugosa, Raja caudaspinosa, R. confundens,

Anacanthobatis marmoratus (Figs 8, 9, 10; Hulley 1970: figs 4, 7, 17). In Raja

clavata (Hulley 1970: fig. 10), unlike in R. straeleni (Hulley 1970: fig. 11), the

shield is not well developed.

(2) rhipidion (Ishiyama)

An elongate, fan-shaped structure, consisting of porous, erectile tissue, which is

situated on the inner border of the ventral lobe, and extends in the longitudinal axis of

the clasper, from about the level of the hypopyle to about half the length of the glans;

it is not associated with any cartilaginous element.

The rhipidion is elongate and free in Raja doutrei, R. lanceorostrata, R. pullo-

punctata, R. caudaspinosa, R. radiata, R. dissimilis, R. confundens, R. leopardus,

CC SE

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) 25

R. wallacei, Anacanthobatis americanus and A. marmoratus (Figs 5, 6, 10, 11; Hulley

1970: figs 5—7, 15, 17-19), but is small, triangular and fixed in Cruriraja parcoma-

culata, C’. triangularis and C. rugosa (Figs 8, 9; Hulley 1970: fig. 4).

(3) boss (new term)

A small, flattened, cartilaginous process, which is situated just below the rhipidion,

and more medially than the knob; it is directly related to the development of a recurved

lateral process of the accessory terminal 2 cartilage.

The boss has been found in a single species only, Cruriraja parcomaculata

(Hulley 1970: fig. 4). Although a small process is developed on the accessory

terminal 2 cartilage in both Cruriraja rugosa and C. triangularis (Fig. 27), the

corresponding boss is not present as an external structure in the glans (Figs 8, 9).

(4) knob (new term)

A process, or processes, situated medially to the shield at about one-half the length of

the clasper glans; it consists of a number of flattened processes (usually 4), or a single,

plate-like cartilaginous element, from which a laterally directed, sharp, recurved

point arises; the knob is related to the development of processes on the medio-dorsal

side of the accessory terminal 1 cartilage.

The knob was so called to describe the blunt protuberances found in

Cruriraja parcomaculata (Hulley 1970: fig. 4), but is now defined so as to include

the plate and laterally projecting point in C. triangularis (Fig. 8) and the single,

flattened process in C. rugosa (Fig. 9).

(5) eperon (Leigh-Sharpe)

A sharp, anteriorly directed, spur-like structure, which is situated proximally to the

level of the hypopyle on the outer lateral margin of the clasper; it is an anterior

projection of the ventral terminal cartilage and may be interpreted as the sharp apex of

the shield.

An eperon is found in Cruriraja parcomaculata, C. rugosa and Anacanthobaiis

marmoratus (Figs 9, 10; Hulley 1970: fig. 4) and although Hulley (1970) pointed

out that it is absent in Crurtraja triangularis, examination of further material has

revealed that the eperon is present in this species.

In Cruriraqa triangularis, C. rugosa and Anacanthobatis marmoratus, the eperon

appears to be associated with the development of small processes or sharp

serrations on the outer lateral edge of the shield. However, in Cruriraja

parcomaculata the shield is smooth-edged.

The structure was first described by Leigh-Sharpe (1924) to cover a

process in two South American species, Raja cyclophora and R. platana.

(6) spike (Leigh-Sharpe)

A cartilaginous element of varying size and shape, which is covered by integument and

is located in the distal region of the glans; it is formed by the distal tip of the accessory

terminal 2 cartilage.

A spike was found to be present in all the species of Rajoidea examined,

except Raja alba and Bathyraja smithii, and was found to be absent in the

Rhinobatidae and Myliobatoidea.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

In Cruriraja parcomaculata, C. triangularis and C. rugosa (Figs 8, 9; Hulley

1970: fig. 4), the spike resembles a flattened, leaf-like tongue, which is situated

on the median axis of the clasper. In Raa doutre: and R. pullopunctata (Hulley

1970: figs 5, 6) the spike is also medially placed, but is sharp-pointed and

straight, while in R. lanceorostrata (Fig. 5) it is hooked. The sentinel obscures the

spike in R. miraletus and R. radiata. The spike in R. miraletus is somewhat sharp-

pointed, but in R. radiata it is more or less bilobed. A bulbous, bilobed spike is

found in R. confundens (Hulley 1970: fig. 17), while in R. wallace: (Fig. 7) the

spike is bilobed and elongate. A simple, bulbous spike, which in some cases

appears to arise from the dorsal lobe of the glans, is found in R. caudaspinosa,

R. dissimilis and R. leopardus (Hulley 1970: figs 7, 15, 18). A distally situated,

sharp-pointed spike, forming a border to the sentina is found in R. straelent and

R. clavata (Hulley 1970: figs 10, 11). Anacanthobatis marmoratus and A. americanus

(Figs 10, 11) possess a somewhat similarly shaped spike, which in A. americanus

is directed laterally outward.

(7) sentinel (Leigh-Sharpe)

A cartilaginous structure, varying considerably in shape and size and which may or

may not be covered by fleshy integument, located on the midline of the clasper; it is

formed by the distal region of the accessory terminal 1 cartilage.

This structure is present in all species of Rajoidea except Bathyraja smithu

and Raa pullopunctata and in Rhinobatidae and Myliobatoidea. It should be

noted that the terms knife and claw, which are also developed from the accessory

terminal, when only one accessory terminal cartilage is present, are used for

Bathyraja smith, Rhinobatidae and Myliobatoidea (cf. Stehmann, 1970).

The sentinel in the genus Cruriraja (Figs 8, 9; Hulley 1970: fig. 4) is a

flattened, spatulate, cartilaginous structure, which is situated medially to the

shield. An enlarged sentinel, with a sharp, knife-like, outer lateral edge is found

in Raja clavata and R. straeleni (Hulley 1970: figs 10, 11); an enlarged, but spade-

shape sentinel is found in R. miraletus (Hulley 1970: fig. 9); a curved, dorsally

convex sentinel is found in R. radiata (Fig. 6); and an enlarged, elongate

sentinel is found in R. alba (Hulley 1970: fig. 8), where it extends to the distal

tip of the glans.

Raja doutret and R. pullopunctata (Fig. 5; Hulley 1970: fig. 5) possess small,

knob-like sentinels; in R. dissimilis, R. confundens and R. leopardus (Hulley 1970:

figs 15, 17, 18) the sentinel is foot-like and may protrude laterally from the

median axis of the glans; the sentinel in R. wallacei (Fig. 7) is bulbous, while in

R. caudaspinosa (Hulley 1970: fig. 7) it is small and dorso-ventrally flattened.

The sentinel in Anacanthobatis marmoratus (Fig. 10) is tongue-like and quite

dissimilar to the spike, but in A. americanus (Fig. 11) the sentinel resembles the

spike, although it points laterally inward.

(8) signal (Leigh-Sharpe)

A fleshy pad, which is located posterior to the hypopyle at about the same level as the

pseudorhipidion, and which can rotate about the longitudinal axis of the clasper; it is

formed by the accessory terminal 3 and accessory terminal 4 cartilages.

a a,

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) 27

The structure is present in Raja clavata and R. straeleni (Hulley 1970:

figs 10, 11), where it is formed by both the accessory terminal 3 and 4 cartilages.

It can rotate so as to protrude from the opened glans. In Raja miraletus (Hulley

1970: fig. 9) the signal is formed from a single cartilage only, the accessory

terminal 3, and does not appear to be movable.

(9) knife (Ishiyama)

A movable, cartilaginous structure with an axe shape, which is covered by integument

at its base, but exposed along its ‘cutting’ edge; it is situated in the distal region of the

ventral lobe and is usually partly obscured by the projection; it is formed by the

accessory terminal 1 cartilage.

A knife is present in a single species only, Bathyraja smith (Hulley 1970:

fig. 21), in which it is almost entirely obscured from view by the well-developed

projection.

(10) funnel (Stehmann)

A bluntly-rounded, cartilaginous tongue, resembling the foot of a bivalve mollusc,

which is situated at the distal region of the ventral lobe of the clasper, so as to extend

beyond the cavity of the glans, when this is opened; it is formed by the elongate, distal

end of the ventral terminal cartilage.

The funnel is found in the species Raja clavata and R. straeleni (Hulley 1970:

figs 10, 11). Ishtyama (1958) is of the opinion that this structure is present in the

Japanese genera Bathyraja and Rhinoraja, but this is incorrect (see discussion on

terminology).

(11) Projection (Stehmann)

An elongate, finger-like, cartilaginous rod, which is covered by integument and which

projects from the ventral lobe; it is formed by the distal elongation of the ventral

marginal cartilage.

A projection has been recognized in Bathyraja smithit (Hulley 1970: fig. 21),

where it projects laterally inwards and obscures the underlying knife from view.

It is thickly covered by integument, so as to resemble the funnel, but may

easily be distinguished from that structure (see above).

(12) sentina (Leigh-Sharpe)

A blindly ending sac, whose aperture points posteriorly, and which is located at the

distal end of the clasper; it may be associated with the distal tip of the accessory

terminal 2 cartilage.

A sentina is present in Cruriraja parcomaculata, Raja caudaspinosa, R. miraletus,

R. clavata, R. straeleni, R. confundens and R. wallacei (Hulley 1970: figs 4, 7, 9-11,

17, 19). It is not easily identifiable, and should not be confused with the ventral

pseudosiphon.

(13) claw (Ishiyama)

A small, sharp, cartilaginous point, which is situated on the ventral lobe at about the

level of the hypopyle; it is formed by the accessory terminal 1 cartilage and is usually

associated with a ventral pseudosiphon.

Among the species which have been examined in detail, a claw has been

found only in Rhinobatos annulatus and Myliobatis cervus (Figs 13, 14), but it may

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

well be present in other Rhinobatoidea and Myliobatoidea, e.g. Rhinobatos

ocellatus (Fig. 12), Rhinobatos schlegeli, Platyrhina sinensis (Ishiyama 1958: fig. 20).

(14) pent (Leigh-Sharpe/Stehmann)

An elongate ridge, which is covered by pleated integument and runs from the distal end

of the rhipidion, parallel with the mid-line of the clasper; it is a fleshy structure, which

is associated with the inner lateral edge of the ventral terminal cartilage.

A pent has been recognized in Raja dissimilis, R. confundens, R. wallace: and

R. radiata (Figs 6, 7; Hulley 1970: figs 15, 17 (not labelled)), and may also be

present in R. caudaspinosa and R. leopardus.

The skeleton of the clasper is a continuation, in the median axis, of the

basipterygium, and is composed of three groups of cartilages, the basal group

(including the basipterygium), the axial group and the terminal group.

Basal group

The basal group consists of three types of cartilages (Fig. 15): the

basipterygium (B), the Beta-cartilage and a varying number of intermediate

proximal segments (b,—b,). Ishiyama (1958) has pointed out that the basal

group is not species specific, and although the number of proximal segments may

vary within large groups of elasmobranchs (Huber r1go1), the basal group may

only be used in the interpretation of phylogenetic relationships (White

1937).

Huber (1901) pointed out that in the Platosomeae (Batoidei) the number

of proximal segments varies between two and four. Ishiyama (1958) found that

there are four intermediate segments in Platyrhinidae (Platyrhina sinensis), three

in Rhinobatidae (Rhinobatos schlegeli), and only two in Raja, Bathyraja and

Rhinoraja among the Rajidae, and on this basis has postulated that the Rhino-

batidae are intermediate between the Platyrhinidae and the Rajidae. Some

criticism of this theory may be raised by the fact that Jungersen (1899) and

Huber (1go1) recorded four intermediate segments in Rhinobatos columnae, but

the present investigation supports Ishiyama, in that only three intermediate

cartilages are present in Rhinobatos annulatus (Fig. 15 a) and Rhinobatos

ocellatus.

This investigation of the basal group of cartilages, while including Raja and

Bathyraja, has been expanded to incorporate the genus Cruriraa of the family

Rajidae and Anacanthobatis of the family Anacanthobatidae, so that an overall

picture of the basal group in the Rajoidea may be obtained.

It appears that in the Rajoidea (Fig. 15), the basipterygium is connected

to the axial cartilage by two intermediate segments (b,—b,), which are partially

covered dorsally by a flat, plate-like Beta-cartilage, serving as an area of

attachment for the m. flexor internus (Jungersen 1899; Hulley 1966). Unlike the

Rhinobatidae, the Beta-cartilage in the Rajoidea does not extend on to the axial

cartilage, but terminates above the junction of the b,—and axial cartilages

(Fig. 75).

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) 29

Fig. 15. Intermediate proximal segments of the claspers. a. Rhinobatos annylatus;

b. Bathyraja smith; c. Raja clavata; d. Cruriraja parcomaculata; e. Anacanthobatis

americanus. Scale 2,0 cm.

B—basipterygium; b,, b., b; —intermediate segments; 8 — beta cartilage; ax—axial

cartilage.

Axial and terminal groups

The axial, dorsal marginal and ventral marginal cartilages are included in

the axial group, but as it is the distal regions of these cartilages which are

important, and since they extend into the terminal complex, they are described

under this heading. The terminal group consists of three different types of

cartilages: dorsal terminal cartilages, a series situated dorsally to the axial, to

which some are joined; the ventral terminal cartilage, a single element which is

ventrally situated and overlies the accessory terminals; the accessory terminal

cartilages (Hulley 1966: fig. 3 G). The number of cartilages in the terminal

group varies between seven and ten.

Huber (1901) and White (1937) suggest that the terminal (accessory)

structures are constant within large groups of elasmobranchs, and Ishiyama

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

(1958) has discussed the specificity and evolutionary significance of the terminal

group in Japanese rajids.

(1) axzal cartilage

The axial cartilage, which is usually much less calcified than the other

cartilages, forms the basic foundation of the clasper. At its junction with the

b,- and Beta-cartilages, it is almost cylindrical in cross-section, and it extends

as a bar-like cartilage to terminate at the distal end of the clasper. From its

proximal end to the level of the commencement of the glans, the axial is nearly

completely enclosed by dorsal and ventral marginal cartilages, so as to form a

tube on the outer lateral surface of the axial. This tube is the clasper groove.

The distal end of the axial cartilage is bluntly pointed in Rhinobatos

annulatus, Anacanthobatis americanus, Bathyraja smith, Raja straelent and R. clavata

(Figs 29, 32, 44, 45; Hulley 1966: fig. 3), but is expanded and spatulate in

Raja lanceorostrata, R. pullopunctata, R. doutrei, R. radiata, R. wallacer, R. confundens,

R. caudaspinosa, R. leopardus, R. miraletus, and Myliobatis cervus (Figs 30-31,

34-39, 46; Hulley 1966: fig. 5; 1969: fig. 3). In Crurtraja parcomaculata, C’. rugosa,

C. triangularis and Anacanthobatis marmoratus (Figs 40-43) the distal end is

expanded laterally and recurved to form a J-shaped cartilage. The well-

developed lateral expansion of the axial in Raa alba (Fig. 33) is not formed in a

similar manner, and should rather be seen as further development of the

spatulate type, through some intermediary like Raa miraletus.

(2) dorsal marginal cartilage

The dorsal marginal cartilage attaches tightly to the axial, starting where

the axial is connected to the b,/Beta joint, and runs along the outer lateral

margin of the axial. Proximally, the cartilage is somewhat broadly pointed, but

the distal region is characteristically expanded. Three different types may be

recognized: the dorsal marginal may be truncate at its junction with the dorsal

terminal 2 cartilage as in Raja caudaspinosa, R. dissimilis, R. confundens, R. leopardus ;

it may be asymmetrically bifurcate, with the larger of the two limbs developed

from the inner lateral edge as in Raja doutrei, R. pullopunctata, R. lanceorostrata,

R. radiata, R. wallacei, Anacanthobatis americanus and Mpyliobatis cervus; or may

possess a marked distal elongation as in Raja alba, R. clavata, R. straeleni,

R. miraletus and Bathyraja smithi (Figs 29, 32, 33; Hulley 1966: fig. 3; 1969:

fig. 3). The distal elongation of the dorsal marginal into the clasper forms the

dorsally situated pseudorhipidion. The extension is best developed in Raja alba

(Fig. 33), where it narrows proximally, but becomes expanded distally and

terminates in a blunt point at about half the length of the clasper glans. The

distal elongation is shorter in Raja miraletus and is stepped to accommodate the

dorsal terminal 2 cartilage (Hulley 1960: fig. 3). In Raja clavata and R. straelent,

it is only slightly stepped (Fig. 32; Hulley 1966: fig. 3).

A modified form of asymmetrically bifurcate type is found in the genus

Cruriraja (Figs 40-42) and in Anacanthobatis marmoratus. In these species, it is the

| i cs,

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 31

outer lateral edge which is elongate. This elongation is bluntly rounded in

Cruriraja parcomaculata and C’. triangularis, but is truncate in C. rugosa.

In Rhinobatos annulatus (Fig. 45) the dorsal marginal cartilage terminates in

a bluntly rounded point.

(3) ventral marginal cartilage

The ventral marginal cartilage, unlike the dorsal marginal, arises some

little distance posteriorly to the axial and b,/Beta junction. It is a flat, tongue-

like structure, whose outer lateral edge forms the ventral margin of the clasper

groove. It extends further along the length of the clasper than the dorsal

marginal cartilage, and distally is usually expanded and ventrally convex.

In the majority of the species examined, the terminal end of the ventral

marginal cartilage is evenly curved. However, in Bathyraja smith (Fig. 29) the

outer lateral edge of the ventral marginal is elongate, extending almost to the

distal end of the clasper. This distal prolongation of the cartilage forms the

projection in the clasper glans.

The genus Anacanthobatis is unusual in that the distal end of the ventral

marginal is truncate in A. americanus (Fig. 44), as itis in Myliobatis cervus (Fig. 46),

and is anteriorly arched in A. marmoratus.

(4) dorsal terminal 1 cartilage

The dorsal terminal 1 cartilage (‘cover-piece’ of Jungersen 1899) is

situated on the dorsal side of the clasper glans, usually partially overlying the

other dorsal terminal cartilages. Ishiyama (1958), besides pointing out the

species specificity of this cartilage, has shown that phylogenetic relationships

may be based on its degree of development and orientation, together with its

associated structure, the pseudosiphon. The pseudosiphon develops as a blindly

ending sac, underlying the dorsal terminal 1. Dorsally the dorsal terminal 1 is

flat and shield-like, but in some species it may be strongly curved medially to

wrap around the inner margin of the axial cartilage. The m. dilatator is attached

to the proximal edge of the dorsal terminal 1, but in Raja miraletus, R. straelent

and R. clavata, there is a proximal shelf (Figs 16 g—h), which serves as the area

of attachment of the muscle. A smaller shelf is found in Bathyraja smithu

(Fig. 16 a).

In the Rhinobatidae, e.g. Rhinobatos annulatus (Fig. 17 a) and Rhinobatos

schlegelt (Ishiyama 1958: fig. 20), the dorsal terminal 1 is wholly situated on the

ventral surface of the clasper, and is associated with the development of both

the dorsal pseudosiphon and the ventral pseudosiphon. Similarly, in Myliobatis

cervus, the dorsal terminal 1 is ventro-laterally positioned.

While the dorsal terminal 1 is dorsally situated in Anacanthobatis marmoratus

(Fig. 17 g), it is positioned laterally in A. americanus (Fig. 17 h), so that the

major portion of the cartilage is situated on the ventral surface of the clasper.

Amongst the Rajidae, seven types of dorsal terminal 1 cartilage may be

distinguished. In Bathyraja smithii (Fig. 16 a), the dorsal terminal 1 is dorsally

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

Y n

J |

Fig. 16. Dorsal view of dorsal terminal 1 cartilages. Scale

4,0 cm.

a. Bathyraja smithii; b. Raja doutrei; c. R. lanceorostrata;

d. R. pullopunctata; e. R. alba; f. R. miraletus; g. R. clavata;

h. R. straeleni; i. R. radiata; }. R. confundens; k. R. leopardus;

1. R. dissimilis; m. R. caudaspinosa; n. R. wallacet.

situated and shield-like, but it is markedly curved laterally and wraps around

on to the ventral side, cf. Bathyraja diplotaenia, B. parmifera (= B. simoterus),

B. smirnovi (Ishiyama 1958: fig. 8); in Raja doutrei, R. lanceorostrata and R. pullo-

punctata (Figs 16 b—d), the dorsal terminal 1 is shield-like dorsally, but is not so

markedly curved along the inner lateral margin, and does not have the proximal

shelf for the insertion of the m. dilatator; the dorsal terminal 1 in Raja alba

(Fig. 16 a) is unique amongst the species examined in that curvature is well

developed, while the dorsal area runs as a transverse band to the outer edge of

the clasper; the cartilage in the species Raja confundens, R. leopardus, R. dissimilis,

R. caudaspinosa and R. wallacei (Figs 16 j-n) have been classed together and

appear to be not too different from that of Raja alba, but in these species the

dorsal region of the cartilage is directed proximally; in Raja radiata (Fig. 16 1),

there is virtually no ventral curvature, but the dorsal region of the cartilage is

large and directed proximally, with an enlarged, hammer-shaped, outer lateral

margin. The dorsal terminal 1 in Raja miraletus, R. clavata and R. straelent

(Figs 16 f-h) is carried almost entirely on the dorsal surface as a well-developed,

shield-like structure with a proximal shelf. The similarity between the cartilages

of Raja clavata and R. straeleni is marked.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 33

Fig. 17. Dorsal view of dorsal terminal 1 cartilage.

Scale 1,0 cm.

a. Rhinobatos annulatus; b. R. annulatus (ventral view) ;

c. Cruriraja parcomaculata; d. C. triangularis; e. C. rugosa;

f. Mbyliobatis cervus; g. Anacanthobatis marmoratus;

h. A. americanus.

A different type of cartilage is possessed by the genus Cruriraja (Figs 17 c-e).

In this, the dorsal terminal 1 is situated within the short arm of the J-shaped

axial, and only partially overlaps the dorsal terminal 2 proximally, so that the

cartilage forms the dorsal wall of the clasper glans. Sharp points are developed

on the proximo-lateral edge of the dorsal terminal 1, which are directly related

to, and responsible for, the development of the thorn. A simple point is found in

Cruriraja parcomaculata and C. triangularis, while a series of points is found in the

North American species C. rugosa,

(5) dorsal terminal 2 cartilage

The dorsal terminal 2 is situated along the proximal dorsal border of the

glans and is joined to the distal region of the dorsal marginal cartilage, or fits

into a step in that cartilage, when the dorsal marginal has a well-developed

distal extension (Hulley 1969: figs 3 A, C). The dorsal terminal 2, together with

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

the dorsal terminals 3 and 4 (when present), extend distally as bar-like elements

and form the framework of the dorsal lobe. Short, sharp processes may be

developed on the outer lateral margin of the dorsal terminal 2, and these may

protrude through the thin integument as the pecten.

The dorsal terminal 2 cartilage in Rhinobatos annulatus (Fig. 19 a) is not

connected to the distal end of the dorsal marginal, but is tightly bonded along

the inner lateral margin of that cartilage (Fig. 45). It is only slightly convex

dorsally. A dorsally convex dorsal terminal 2 is also found in Myliobatis cervus

(Fig. 19 d), but in this species it develops from the distal end of the dorsal

marginal (Fig. 46).

As with the dorsal terminal 1, the dorsal terminal 2 cartilage can be divided

into six types, although the grouping does not correspond with that of the dorsal

terminal 1.

Fig. 18. Dorsal view of dorsal terminal 2 cartilage. Scale 2,0 cm.

a. Bathyraja smithii (dorsal terminal 2 and 3); b. Raja doutrei; c. R. lanceorostrata;

d. R. pullopunctata; e. R. clavata; f. R. miraletus; g. R. straeleni; h. R. alba; i. R. wallacet;

j. R. radiata; k. R. dissimilis; 1. R. caudaspinosa; m. R. confundens; n. R. leopardus.

i !”)U a i a,

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 35

In Bathyraja smith (Fig. 18 a) the dorsal terminal 2 arises at the distal end

of the dorsal marginal and runs distally for a short distance, before turning

laterally inward to become fused to the axial cartilage. Closely approximating

to this type is the dorsal terminal 2 in Raya pullopunctata, R. doutrei and R. lanceoro-

strata (Figs 18 b—d), which becomes narrower proximally, to form a bar-like

element. In Raja doutre: and R. pullopunctata, a short, cartilaginous bridge

(terminal bridge) is developed, which attaches the dorsal terminal 2 to the

axial. In Raja lanceorostrata (Fig. 31) the dorsal terminal 2 itself makes contact

with the axial. A dorsal terminal 2/axial junction is also shown by the species

Raja radiata, R. dissimilis, R. caudaspinosa, R. confundens and R. leopardus

(Figs 18 j—n, 34, 36-39), but in these species the proximal end of the dorsal

terminal is expanded, while the distal end is contracted. Although the dorsal

terminal 2 is not attached to the axial in Raa wallace: (Figs 18 i, 35), it is

similarly shaped. In this species the free end of the cartilage is responsible for

the formation of the promontory. In Raja miraletus and R. straelen the dorsal

terminal 2 is flat and shield-like, with a series of protuberances situated on its

outer lateral edge. An elongate cartilage is found in Raa alba (Fig. 18 h),

Fig. 19. Dorsal view of dorsal terminal 2

cartilage. Scale 1,0 cm.

a. Cruriraja triangularis; b. C. parcomaculata;

c. C. rugosa; d. Mpyliobatis cervus; e. Rhinobatos

annulatus; i: Anacanthobatis marmoratus ;

g. A. americanus.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

which is similar to the cartilage in R. miraletus and R. straeleni (Figs 18 f, g), but

does not form lateral processes. ‘The dorsal terminal 2 is strongly curved in

Raga clavata (Fig. 18 e) and is thickened along its side of greatest curvature, to

form a flat ledge, which articulates with the dorsal terminal 1 cartilage. Although

the dorsal terminal 2 appears to be species specific in Cruriraja species, they are

of the same general type and are quite different from the structure of the

cartilage in Raga species.

In the genus Cruriraja, the dorsal terminal 2 is a short and slightly curved

bar-like element, which arises at the distal end of the dorsal marginal and

terminates in a truncate tip, within the curvature offered by the dorsal terminal 1

(Figs 40-42).

As with the dorsal terminal 1, the dorsal terminal 2 cartilage in Anacantho-

batis marmoratus (Fig. 19 f) and A. americanus (Fig. 19 g) is very dissimilar. In

A. marmoratus, the dorsal terminal 2 is truncate and plate-like, while in

A. americanus the proximal end of the tongue-like cartilage is bifurcate and the

distal end is free and protrudes laterally from the axial.

(6) dorsal terminal 3 cartilage

This cartilage was found to be absent in Cruriraja parcomaculata, C. triangu-

laris, C- rugosa, Raja pullopunctata, R. miraletus, R. clavata, R. straeleni and Anacantho-

batis americanus.

In Rajidae, the dorsal terminal 3 extends distally from the dorsal terminal 2

cartilage and may either have a free, distal end, as in Raja caudaspinosa, R. con-

fundens, R. dissimilis, R. leopardus, R. wallacei and R. radiata (Figs 20 c—h, 34-39),

Fig. 20. Dorsal view of dorsal f

terminal 3 cartilage. Scale 2,0 cm.

. Raja doutrei; b. R. lanceorostrata;

. R. caudaspinosa; d. R. dissimilis; /

. R. confundens; f. R. leopardus;

R. wallacei; bh. R. radiaia; i

R. alba; j. Rhinobatos annulatus J

(scale 0,5 cm); k. Anacanthobatis

marmoratus (scale 0,5 cm).

“7 oA ®

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 37

or the distal end may be fused to the expanded terminal end of the axial

cartilage, as in Raja doutre: and R. lanceorostrata (Figs 20 a—b, 31, 32).

The dorsai terminal 3 cartilage in Bathyraja smithu (Fig. 18 a) is not firmly

attached to the bluntly pointed axial. However, it is tightly bonded to the

dorsal terminal 2 along its inner lateral margin, while its somewhat thickened,

outer lateral edge forms the ridge in the clasper glans. ‘The dorsal terminal 3 in

Raja lanceorostrata and R. doutre: (Figs 20 a, b) is flat and spatulate, and arises

either from the terminal bridge (RA. doutrez) or directly from the dorsal terminal 2

(R. lanceorostrata).

The ‘free-ended’ dorsal terminal 3 species consist of two groups: (1) the

distal region of the cartilage is curved laterally outward in a sharp point,

forming the spur in Raja radiata and R. wallacei (Figs 34, 35); (11) the cartilage is

straight, and the free end is bluntly pointed, as in R. caudaspinosa, R. confundens,

R. dissimilis, and R. leopardus (Figs 36-39). In the latter group the cartilage

forms the outer lateral border of the distal cleft.

In Rhinobatos annulatus (Fig. 20 }) the dorsal terminal 3 is simple, while in

Mylhobatis cervus (Fig. 19 d) the cartilage is flat and attached along its inner

lateral margin to the dorsal terminal 2 (Fig. 46), as in Bathyraja smithi. A spur is

formed by the dorsal terminal 3 in Anacanthobatis marmoratus (Fig. 20 k).

(7) dorsal terminal 4 cartilage

This cartilage is present only in Raya alba, R. radiata, R. dissimilis, R. cauda-

spinosa, R. confundens, R. leopardus and R. wallace: cf. R. fyllae, R. fullonica

(Stehmann 1970)—see discussion. In all these species, the dorsal terminal 4

attaches the dorsal terminal 3 to the axial, so completing the dorsal framework.

On the basis of the attachment of the dorsal terminal 4 to the dorsal terminal 3,

the cartilages may be divided into two groups. In the first, the dorsal terminal 4

attaches to the dorsal terminal 3 externally, some little distance behind the

) |

Fig. 21. Dorsal view of dorsal

terminal 4 cartilage. Scale 2,0 cm.

a. Raja radiata; b. R. wallacei; é f 2

c. R. caudaspinosa; d. R. confundens; =

e. R. leopardus; f. R. dissimilis; g

g. R. alba.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

dorsal terminal 3/dorsal terminal 2 junction, so that the distal end of the dorsal

terminal 3 is free, e.g. Raja radiata, R. caudaspinosa, R. confundens, R. dissimilis,

R. leopardus and R. wallace: (Figs 34-39). In the second group, e.g. Raja alba

(Fig. 33), the dorsal terminal 4 is attached to the distal end of the dorsal

terminal 3, thereby forming a link between this cartilage and the axial.

(8) ventral terminal cartilage

This cartilage was found to be absent only in Anacanthobatis americanus. The

ventral terminal cartilage is located on the ventral side of the clasper, and

extends from about the level of the distal end of the ventral marginal to the

distal tip of the clasper. It therefore forms the ventral lobe of the clasper glans.

Amongst the Rajidae, Bathyraja smithi (Fig. 22 a) has the simplest ventral

terminal cartilage. This is more or less spoon-shaped, with a somewhat pointed

proximal extension and thickened distal region, so that it resembles the type

usually associated with this genus (Ishiyama 1958: figs 10 B, E, F, H, I, L). The

ventral terminal cartilages of Raja doutrei, R. pullopunctata and R. lanceorostrata

have common characteristics and may be grouped together. In these species

(Figs 22 b—d) the cartilage may be divided into three areas: the outer lateral

margin is well developed and is dorsally convex, forming the shield of the glans;

there is an anterior notch, which attaches to the accessory terminal 1 cartilage;

the distal region is expanded on its inner lateral margin and this plate-like area

folds around the axial, so that the ventral terminal attains a J-shape. The outer

lateral margin is narrower and flat in Raja miraletus, R. straelem and R. clavata

(Figs 22 e-g), so that in these species the shield is not well developed. Although

the medio-distal inner margin of the cartilage is expanded to wrap around the

axial (Hulley 1966: fig. 3 G; Figs 22 e-g), the distal end of the cartilage is well

developed and forms a structure which resembles the foot of a bivalve mollusc.

This distal end forms the funnel of the glans. There is no anterior notch.

The ventral terminal in Raja alba (Fig. 22 h) appears to be unique in that,

while the inner median lateral edge of the cartilage is expanded to curve

around the axial cartilage, the outer lateral margin is well developed and

dorsally convex.

Unlike the above, the ventral terminals in the following rajids are different

in that neither the distal tip, nor the inner lateral margin are expanded to

encompass the axial. The anterior notch, for attachment to the accessory

terminal 1, is shifted posteriorly and is situated at about half the length of the

cartilage. Furthermore, it is the proximal region of the ventral terminal which

is expanded, and in the species Raja caudaspinosa, R. confundens, R. dissimilts,

R. leopardus and R. wallacei (Figs 22 j-n) is characterized by an inner lateral

expansion, cf. projection of Raja lintea (Leigh-Sharpe 1924: fig. 6). On the basis

of the above, Raja caudaspinosa, R. confundens, R. dissimilis, R. leopardus and

R. wallacei have been grouped together, and while R. radiata is somewhat similar,

it has been kept separate because of its unique shape and its median notch

(Fig. 22 i).

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 39

Fig. 22. Dorsal view of ventral terminal cartilage. Scale 2,0 cm.

a. Bathyraja smithii; b. Raja doutrei; c. R. lanceorostrata; d. R. pullo-

punctata; e. R. clavata; f. R. straeleni; g. R. miraletus; h. R. alba; i. R.

radiata; j. R. confundens; k. R. leopardus; 1. R. dissimilis; m. R. cauda-

spinosa; n. R. wallace.

Once again the ventral terminal in the genus Cruriraja is more or less

constant in shape, although it appears to be species specific. In these species

(Figs 23 a—c), a point is developed on the proximal outer lateral edge of the

narrow shield. This is the eperon. The notch is situated anteriorly in all, but in

C. triangularis the distal region is bifurcate and in C. rugosa it is expanded.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 23. Dorsal view of ventral terminal cartilage.

Scale 1,0 cm.

a. Cruriraja parcomaculata; b. C. triangularis; c. C. rugosa;

d. Anacanthobatis marmoratus; e. Rhinobatos annulatus;

f. Myliobatis cervus.

The ventral terminal in Anacanthobatis marmoratus (Fig. 23 d) is unusual.

It possesses a well-developed outer lateral edge, which forms the shield, and

which is serrate, as in some Cruriraja species. Medially the cartilage is expanded

and runs around the axial, as in Raja alba. However, in Anacanthobatis marmoratus

the inner lateral plate is windowed.

(9) accessory terminal 1 cartilage

In all Batoidei the accessory terminal 1 arises at the distal end of the

ventral marginal cartilage, but its form in the Rhinobatoidea and Mylio-

batoidea (Figs 45, 46) is quite different from that in the Rajoidea, although the

accessory terminal 1 in Anacanthobatis marmoratus and A. americanus resembles

that of Rhinobatos annulatus (Figs 25 c-e) and Myliobatis cervus (Fig. 46), particu-

larly in its sharp, pointed distal end. The accessory terminal 1 is closely asso-

ciated with the accessory terminal 2, although in Rhinobatos annulatus, Myliobatis

cervus and Bathyraja smithii only one accessory terminal cartilage is present. ‘The

distal end of the accessory terminal 1 has been termed the sentinel in all cases

in which this cartilage manifests itself within the glans, except in the above-

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) 4I

mentioned species, in which it is variously termed claw and knife, depending

on its degree of development, cf. Stehmann (1970). The distal end of the

accessory terminal 1 cartilage varies in shape from elongate, sharp-pointed to

expanded and flattened. In Cruriraja species, there is a lateral process, which

develops at about half the length of the cartilage, in the form of the knob. This

consists of a flattened protuberance in C. rugosa (Fig. 25 f), a series of four blunt

lobes in C. parcomaculata (Fig. 25 a, a!), or asharp, recurved spine in C. triangularis

(Fig. 25 b, b?).

The accessory terminal in Bathyraja smithi (Fig. 24 a) is unusual and quite

J } k / m n

Fig. 24. Ventral view of accessory terminal 1 cartilage.

Scale 2,0 cm.

a. Bathyraja smithii; b. Raja doutrei; c. R. lanceorostrata;

d. R. pullopunctata; e. R. clavata; f. R. straeleni; g. R. miraletus;

h. R. alba; i. R. radiata; j. R. confundens; k. R. leopardus;

l. R. dissimilis; m. R. caudaspinosa; n. R. wallacei.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

unlike the cartilage in other rajid species. ‘The cartilage is fan-shaped with a

blade-like posterior margin, similar in shape to the accessory terminal in

Rhinoraja odai and Rhinoraqja longicauda (Ishiyama 1958: fig. 11). In all other

rajids, the accessory terminal 1 cartilage is Y-shaped, although in Raa clavata,

R. straeleni, R. miraletus and R. alba (Figs 24 e-h) there appears to be a single

arm. Ishiyama (1958) has commented on the change from a symmetrical to an

asymmetrical shape.

Furthermore, Ishiyama (1958) has grouped the accessory terminal 1

according to the shape of the distal region of the cartilage. However, in this

study the accessory terminal 1 has been grouped on the basis of general overall

appearance. Excluding the species Bathyraja smithi, five types of cartilages can

be identified in southern African rajids.

In Raa pullopunctata, R. doutrei and R. lanceorostrata, the cartilage is of

asymmetrical Y-shape, with the posterior arm varying from elongate and

pointed (R. doutrei, Fig. 24 b) toa small blunt process (R. pullopunctata, Fig. 24 d).

Fig. 25. Ventral view of accessory terminal 1 cartilage.

Scale 1,0 cm.

a. Cruriraja parcomaculata; a. C. parcomaculata (dorsal view);

b. C. triangularis; b‘. C. triangularis (dorsal view) ; c. Rhinobatos

annulatus; d. Anacanthobatis marmoratus; e. A. americanus;

f. Cruriraja rugosa.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 43

The accessory terminal 1 in Raya clavata, R. straelen and R. miraletus (Figs 24 e-g)

is characterized by a single, outwardly directed proximal arm and a very well-

developed and dorso-ventrally flattened distal blade. Although the form of the

cartilage is somewhat similar in Raya alba (Fig. 24 h), it has not been included

with R. clavata because of its elongate- rather than Z-shape, and because of the

lack of a distinguishable proximal arm.

The U-shaped accessory terminal 1 in Raja radiata is unique (Fig. 24 1) and

quite unlike the Y-form of the cartilage in Raja confundens, R. leopardus,

R. dissimilis, R. caudaspinosa and R. wallace: (Figs 24 j-n), whose anterior limbs

resemble that of R. lanceorostrata, and yet whose general form is quite different.

The species specificity of the posterior extension should be noted.

(10) accessory terminal 2 cartilage

This cartilage, like the accessory terminal 1, arises at the distal end of the

ventral marginal cartilage. It is situated on the median side of the accessory

terminal 1. Although it may possess a median, lateral process, which can form

the boss (Cruriraja parcomaculata, Fig. 27 a, a‘), it is the distal end which is

Fig. 26. Ventral view of

accessory terminal 2 cartil-

age. Scale 2,0 cm.

b. Raja doutrei; c. R. lanceoro-

strata; d. R. pullopunctata;

e. R. clavata; f. R. straeleni;

g. R. miraletus; h. R. alba;

i. R. radiata; j. R. confundens;

k. R. leopardus; 1. R. dissi-

milis; m. R. caudaspinosa;

n. R. wallacei.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

important and which forms the spike. An accessory terminal 2 cartilage is

absent in Rhinobatos annulatus, Myliobatis cervus and Bathyraja smithit.

Groupings similar to those shown by the accessory terminal 1 cartilage can

be seen. In the species Raja doutrei, R. pullopunctata and R. lanceorostrata (Figs

26 b—d) the accessory terminal 2 projects distally as a simple rod, which may be

hooked distally, and may or may not possess an attachment process. The

cartilage is strongly united with the accessory terminal 1 in Raa clavata (Hulley

1966: fig. 3 E) and AR. straeleni, while in R. miraletus the association does not

appear to be that marked (Hulley 1969). Although not strongly united with the

accessory terminal 1, the accessory terminal 2 cartilage in Raja radiata is

somewhat curled (Fig. 26 1), so as to lie within the cavity of the accessory

terminal 1 (Fig. 34). The accessory terminal 2 in Raa alba (Fig. 26 h) is simple,

with a shortened distal end, so that a spike is not formed in the glans. It has a

process on its inner lateral margin, which butts against the axial cartilage. The

species Raja dissimilis, R. caudaspinosa, R. confundens, R. leopardus and R. wallacei

possess a characteristically shaped accessory terminal 2 (Figs 26 j—n).

Fig. 27. Ventral view of accessory terminal 2 cartilage.

Scale 1,0 cm.

a. Cruriraja parcomaculata; at. C. parcomaculata (dorsal view) ;

b. C. triangularis; c. C. rugosa; d. Anacanthobatis marmoratus;

e. A. americanus.

In the genus Cruriraja (Figs 27 a—c), the cartilage is elongate, and in the

three species examined a lateral process was found. Distally the cartilage is

rounded.

A similarly shaped accessory terminal 2 cartilage is found in Anacanthobatis

marmoratus (Fig. 27 d), except that there is no lateral process and the flattened,

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 45

distal region is curled to form a spoon-shaped end. In Anacanthobatis americanus

(Fig. 27 e), the accessory terminal 2 is Y-shaped, and similar in appearance to

the accessory terminal | cartilage in this species.

(11) accessory terminal 3 and 4 cartilages

These cartilages are present only in Raja clavata, R. straeleni and R. miraletus,

and form the signal of the clasper glans. As with other accessory terminal

cartilages, they are attached at the distal end of the ventral marginal, more

ventral than the attachment of the accessory terminal 1 and 2 cartilages. In

Raja miraletus (Fig. 28 c) there is a single cartilaginous element, the accessory

terminal 3, while in R. clavata and R. straeleni (Figs 28 a—b) two cartilages are

present. In the latter two species, the accessory terminal 4 is movable on the

accessory terminal 3.

Fig. 28. Ventral view of accessory terminals 3 and 4.

Scale 2,0 cm.

a. Raja clavata; b. R. straeleni; c. R. miraletus.

VERTEBRAL COUNT

Vertebral numbers have been employed as taxonomic characters in

teleosts (Bailey & Gosline 1955; Lagler et al. 1962). Their possible use in

elasmobranch taxonomy was reviewed by Springer & Garrick (1964), while

their species specificity in Japanese rajids has been investigated by Ishiyama

(1952, 1958) and in European rajid species by Stehmann (1970). Vertebral

counts for South African west and south coast Rajidae have been given by

Hulley (1970). Krefft (1968a) has analysed the terminology, and has proposed

a standard method for distinguishing the various groups of vertebrae.

In rajids, the vertebrae may be divided into two groups, trunk (Vtr) and

predorsal caudal (Vprd) vertebrae, of which the latter appears to be more

suitable for taxonomic purposes. The number of predorsal caudal vertebrae is

not species specific (Ishiyama, 1958), although the count may, in some cases,

be used to distinguish between closely related species (Hulley 1970; Stehmann

1970).

Interpreting the predorsal caudal count according to the method of Hubbs

& Hubbs (1953), Ishiyama (1958) recognizes two forms of rajids, the ‘northern’

form (Bathyraja and Rhinoraja) with more than 61-62 Vprd, and the ‘southern’

form (Raja) with less than 61-62 Vprd. Stehmann (1970) can find no clear-cut

distinction between the two genera, Bathyraja and Raja, in European species, but

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 29. Bathyraja smithii. Cartilages of right clasper (exploded)

with d.tr, removed. Scale 2,0 cm.

a.tr.— accessory terminal; ax—axial; d.mg—dorsal marginal;

d.tr—dorsal terminal; v.mg.— ventral marginal; v.tr—ventral

terminal.

Fig. 30. Raja doutrei. Cartilages

of right clasper (exploded) with

d.tr, removed. Scale 2,0 cm.

a.tr—accessory terminal; ax—axial;

dm.g—dorsal marginal; d.tr—dorsal

terminal; t.br—terminal bridge;

v.mg—ventral marginal; v.tr—ven-

tral terminal.

a tr.

Vv tr

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 47

vy tr

a tr

—SSSS,

Fig. 31. Raja lanceorostrata. Cartilages of right clasper

(exploded) with d.tr, removed. Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—dorsal marginal;

d.tr—dorsal terminal; v.mg—ventral marginal; v.tr—ventral

terminal.

—_——4{

Fig. 32. Raja straeleni. Cartilages of right clasper

(exploded) with d.tr, removed. Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—dorsal

marginal; d.tr—dorsal terminal; v.mg—ventral

marginal; v.tr—ventral terminal.

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

—————,

Fig. 33. Raja alba. Cartilages of right clasper (exploded)

with d.tr, removed. Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—dorsal

marginal; d.tr—dorsal terminal; v.mg—ventral

marginal; v.tr—ventral terminal.

dtr,

Fig. 34. Raja radiata. Cartilages of right clasper (exploded)

with d.tr, removed. Scale 2,0 cm.

a.tr— accessory terminal; ax—axial; d.mg—dorsal marginal;

d.tr—dorsal terminal; v.mg—ventral marginal; v.tr—ventral

terminal.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) 49

Fig. 35. Raja wallacei. Ventral view of cartilages

of right clasper. Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—

dorsal marginal; d.tr—dorsal terminal.

Fig. 36. Raja confundens. Ventral view of

cartilages of right clasper, with d.tr,, a.tr, and

v.tr removed. Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—

dorsal marginal; d.tr—dorsal terminal; v.mg—

ventral marginal.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 37. Raja leopardus. Ventral view of cartilages

of right clasper, with d.tr,, a.tr, and v.tr

removed. Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—

dorsal marginal; d.tr—dorsal terminal; v.mg—

ventral marginal.

Fig. 38. Raja dissimilis. Ventral view of cartilages

of right clasper, with d.tr,, a.tr, and v.tr

removed. Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—

dorsal marginal; d.tr—dorsal terminal; v.mg—

ventral marginal.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 51

v mg

Fig. 39. Raja caudaspinosa. Ventral view of

cartilages of right clasper, with d.tr,, a.tr,; and

v.tr removed. Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—

dorsal marginal; d.tr—dorsal terminal; v.mg—

ventral marginal.

v mg

d tr,

t br

d tr,

a tr,

ax v tr

Fig. 40. Cruriraja parcomaculata. Cartilages of right

clasper (exploded). Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—dorsal

marginal; d.tr—dorsal terminal; t.br—terminal

bridge; v.mg—ventral marginal; v.tr—ventral

terminal

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 41. Cruriraja triangularis. Cartilages of right clasper

(exploded). Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—dorsal marginal;

d.tr—dorsal terminal; t.br—terminal bridge; v.mg—ventral

marginal; v.tr—ventral terminal.

Fig. 42. Cruriraja rugosa. Cartilages of right clasper (exploded).

Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—dorsal marginal;

d.tr—dorsal terminal; t.br-—terminal bridge; v.mg—ventral

marginal; v.tr—ventral terminal.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE!)

Fig. 43. Anacanthobatis marmoratus. Cartilages of right

clasper (exploded) with d.tr, removed. Scale 1,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—dorsal

marginal; d.tr—dorsal terminal; v.mg—ventral

marginal; v.tr—ventral terminal.

a tr.

Fig. 44. Anacanthobatis americanus. Ventral view

of cartilages of right clasper with d.tr, removed.

Scale 1,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—

dorsal marginal; d.tr—dorsal terminal; v.mg—

ventral marginal.

ANNALS OF THE SOUTH AFRICAN MUSEUM

d tr, = i; a tr

d tr,

_ |

Fig. 45. Rhinobatos annulatus. Dorsal view of

cartilages of right clasper, with d.tr, removed.

Scale 0,5 cm.

a.tr—accessory terminal; ax—axial; d.mg—

dorsal marginal; d.tr—dorsal terminal; v.mg—

ventral marginal; v.tr—ventral terminal.

Fig. 46. Mpliobatis cervus. Lateral view of

cartilages of right clasper with d.tr, removed.

Scale 2,0 cm.

a.tr—accessory terminal; ax—axial; d.mg—

dorsal marginal; d.tr—dorsal terminal; v.mg—

ventral marginal; v.tr—ventral terminal.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 55

divides them into four groups. The group limits appear to be arbitrary and are

exceeded by some of the species (Stehmann 1970: fig. 14), so that this clustering

is artificial and bears no resemblance to the groupings suggested by clasper

structure and cranial anatomy. The picture is further complicated by the fact

that, as in teleosts (Tester 1938; Taning 1950; Post 1968), there appears to be

some correlation between water temperature and vertebral number in skates

(Ishiyama 1958). However, Stehmann (1970) states that an intra-specific

dependence of the number of predorsal caudal vertebrae on water temperature

is not noticeable in eastern North Atlantic species. In the case presented, i.e. Raja

miraletus, it may be that we are dealing here with isothermic distribution rather

than temperature differences based on geographic distribution.

Predorsal caudal vertebral counts can be used in the interpretation of the

phylogeny of rajids. Ishiyama (1958: 237) has pointed out that ‘the variation

in vertebral count in each group of species (as ascertained from clasper structure)

can be considered corresponding with phylogenetic progression along each

branch’ and has shown that with evolution there is a trend in reduction in

number of predorsal caudal vertebrae. It is within this context that the predorsal

caudal count of southern African rajids will be discussed.

TABLE I

Predorsal caudal vertebral counts (Vprd) for southern African Rajidae.

Species No. of Range Mean

specimens

[35 SUCHE oe a ne 6 68-71 69,2

i SECAGE ae 4 48-52 49,3

ICIQUGIM es 10 45-53 47,7

R. miraletus “Ge ae ee 17 44-52 48,5

2. GG co, i 6 62-67 64,0

Hespullopunciata =. 2. s 14 50-58 53.4

R. doutrei En eae ee ts 4 43-49 46,0

ESPHUROCTE . ww 3 53-56 55,0

Mea stenorhynchus . . . . I = 49,0

R. lanceorostrata 2 56-57 56,5

R. robertsi I aa 55,0

R. radiata oe eG eae tah 2 58-62 60,0

RR olig | 10 64-74 69,0

R. ravidula . 3 69-70 69,3

R. dissimilis 3 65-69 67,0

R. caudaspinosa 15 66-73 67,0

R. leopardus 7 55-58 56,9

R. confundens 5 55-63 59,2

R. spinacidermis 3 60-65 62,7

C. parcomaculata 6 66-69 67,8

C. triangularis 4 65-70 67,3

The predorsal caudal vertebral count for southern African species is given

in Table 1, and is represented graphically in Figure 47, in subgeneric groupings.

Vprd counts for Raja miraletus and R. straeleni given by Krefft (1968) are

incorporated in the results.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

CS Roc

WEB © parcomaculata

parcomacula

CRURIRAJA SS Cr fanigularis

(RAGS a) UL eos

40 50 60 70 80

Fig. 47. Ranges in number of predorsal caudal

vertebrae (Vprd) in southern African Rajidae.

NEUROCRANIUM AND ROSTRAL CARTILAGES

In order to establish the phylogenetic positions of suborders within the

Batoidei, comparative anatomical studies on various neurocrania have been

made (Gegenbaur 1872; Parker 1879; Garman 1913; Holmgren & Stensio

1936; Davies 1948; Melouk 1949; El-Tourbi & Hamdy 1959; Gohar &

Bayoumi 1959; Hamdy 1964) and these have been supplemented by develop-

mental studies (Parker 1879; De Beer 1926, 1932, 1937; Holmgren 1940, 1941;

Hamdy 1956).

From a taxonomic viewpoint at the generic level, Bigelow & Schroeder

(1948) studied the X-ray structure of Psammobatis, Sympterygia and some Raja

species, with particular reference to the rostral projection and the degree of

forward extension of the anterior radials of the pectoral fins. This resulted in the

distinction of a new genus Breviraja. Following on this, Ishiyama (1952, 1958,

1968) recognized this genus in Japanese rajids and was able to show that a

further new genus, Rhinoraja, could be established on rostral characters.

Subsequently, Ishiyama & Hubbs (1968) pointed out that the Pacific

brevirajid species were distinct from the Atlantic species, not only in terms of

clasper structure, but also in the shape and size of the rostral cartilage and

rostral appendices. They therefore defined a new genus, Bathyraja, confining it

solely to the Pacific region. However, this genus has now been recognized in the

ee ee ee ee

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) 57

North Atlantic (Stehmann 1970) and in the southern African region (Hulley

1970).

Further differences in the anatomy of the neurocrania of Japanese rajids

were found by Ishiyama (1958), who showed a certain degree of species specificity

in the shape and extension of the anterior and posterior fontanelles. However,

Heintz (1962) found that the cartilaginous neurocranium could be modified by

external factors, and noted differences only in the anterior fontanelle, orbital

and otic regions of the neurocrania of Raya batis, R. nidarosiensis and R. oxyrhynchus,

so that on the basis of structure alone it was impossible to draw any conclusions

of the relationship of the three species.

On the other hand, Stehmann (1970) found that while species specificity

may not necessarily be evident from cranial structure, there appeared to be

certain parts, which could be used as auxiliary characters in the grouping of the

species within the proposed subgenera. ‘These included:

1. The shape and orientation of the nasal capsules with regard to the median

axis of the skull.

2. The appearance and size of the processus praeorbitalis, including the crista

praeorbitalis.

3. ‘The form of the regio orbito-temporalis.

The size and position of the jugal arches.

5. The form and position of the anterior and posterior fontanelles.

Differences in cranial morphometry for taxonomic use were suggested by

Ishiyama (1958) and some 18 measurements have now been proposed as

standards (Hubbs & Ishiyama 1968). However, both Heintz (1962) and

Stehmann (1970) find that these possess no species specificity, although

Stehmann (1970) finds that the length of the rostrum in comparison with the

length of the cranium can be used for grouping the species. The course of the

hypothetical line, drawn from the tip of the rostrum and over the Foramen

ophthalmicus profundus V to the otic region (Stehmann 1970) is obviously some

complex function of the rostral length/cranial width, and is difficult to interpret

as a single character.

Description

Descriptions are based mainly on neurocrania which have been prepared.

However, in cases where this was not possible, some results, such as general

shape, orientation of the nasal capsules, form of the anterior and posterior

fontanelles and size of the jugal arches, have been obtained from X-ray photo-

graphs. Details of the cranial anatomy of Rhinobatos halavi have been given by

El-Tourbi & Hamdy (1959) and are used in this description.

The neurocranium is a violin-shaped structure, which is dorso-ventrally

compressed and is contricted at the orbital region. The nasal capsules are fused

to this anteriorly and the auditory capsules posteriorly. The rostral bar projects

from the mid-region as a hard bar, to which are attached the rostral appendices.

ANNALS OF THE SOUTH AFRICAN MUSEUM

F. oph.

prae. pr. : p os an. pr.

F. sup. oph.

p.f.

: AG haem Post - or. pr.

F. end. ; s

lake sont Par. dep.

F. oph. Wl

F. a. cer. v. op. st.

; os F. pro-ot. (V, Vil)

Or-Nnas. Can, yi A

j.a.

Vil

F. p.cer.v.

F.mag.

Fig. 48. Neurocranium of a typical rajid showing positions of

foramina and fontanelles.

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.—inter-

orbital vein foramen; F.]. X—foramen of lateralis branch (X);

F.mag.—foramen magnum; F.n.eth.—ethmoidal nerve foramen;

F.oph—ophthalmic foramen; 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; prae.pr.—praeorbital

process; post-or.pr.—postorbital process; pt.pr.—pterotic process;

r.a.—rostral appendix; r.c.—rostral cartilage; I]—optic nerve

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

nerve foramen; VII—foramen of hyomandibular branch (VII);

IX—glossopharyngeal nerve foramen; X—vagus nerve foramen.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 59

The nature of the bar and appendices have been described by Hulley (1970),

while the varying lengths of the rostra in relation to cranial length are given in

Table 2.

TABLE 2

Rostral length expressed as a percentage of the cranial length

in southern African Rajidae.

Species No. of Mean %

specimens

C. parcomaculata 3 39,6

C. triangularis . 2 47,6

R. doutrei P 63,5

R. pullopunctata 5 56,7

R. lanceorostrata I 65,7

R. springeri . 2 65,3

R. stenorhynchus I 70,7

R. miraletus 5 45,6

R. clavata 10 48,1

R. straeleni . 2 49,0

R. alba . 4 57,0

R. radiata 2 46,2

R. robertsi . I 46,5

R. wallacei . 2 4554

R. caudaspinosa 6 37,6

R. leopardus 10 49,6

R. confundens 3 50,1

R. dissimilis 3 46,4

R. ravidula . 3 52,4

R. spinacidermis I 41,3

B. smithii 3 4555

The course of the hypothetical line drawn from the tip of the rostrum and

through the ophthalmic foramen to the occipital region, cuts the otic region in

Raja alba, R. leopardus and Cruriraja parcomaculata, runs at a tangent to the otic

region in Raja doutrei and R. confundens, and passes the otic region without

touching in Bathyraja smithii, Raja pullopunctata, R. clavata, R. miraletus, R. radiata

and R. caudaspinosa.

The nasal capsules are relatively thin-walled structures, which are attached

to the side walls of the brain case. They are orientated to form a right angle

with the median axis of the cranium in Rhinobatos halavi (El-Tourbi & Hamdy

1959: pl. 1), Bathyraja smithit, Raja clavata, R. straeleni, R. caudaspinosa, R. confundens

and Cruriraja parcomaculata (Figs 49 A, E; 51 A, D; 53 A, E). The angle is slightly

acute in Raja stenorhynchus, R. springert, R. lanceorostrata, R. doutrei, R. pullopunctata,

R. miraletus, R. radiata, R. robertsi, R. dissimilis, R. ravidula and R. leopardus

(Figs 50 A, D, E, F, G; 51 E; 52 D, E; 53 D; 54 A, D), more so in R. wallacei

(Fig. 54 F), and extremely acute in R. alba and R. spinacidermis (Figs 52 A; 54 E).

In Raja spinacidermis, R. ravidula, R. dissimilis and R. roberisi (Figs 52 D; 53 D;

54 D, E), the nasal capsules are comparatively larger than in the other species.

There is a well-developed articular process for the attachment of the antorbital

process at the latero-posterior edge of each nasal capsule, while antero-dorsally

there is usually one small foramen (sometimes more, e.g. Raja clavata (Fig. 51 A))

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

al LL

\ "AZ SQ ()

©) B J)

-——

We a We Z

a S pe \

KS & : c \

ee Sen 7 Be, ‘\ aa

Cc N= G ae

i -——

Fig. 49. Dorsal, lateral and posterior views of the neurocrania of:

A. B. C. Bathyraja smithit; D. Cruriraja triangularis (X-ray reconstruction) ; E. F. G. C. parcomaculata.

for the passage of the ethmoidal nerve, a small, lateral branch of the superficial

ophthalmic nerve. The foramen appears to be absent in Bathyraja smithu, Raja

leopardus and R. confundens (Figs 49 A; 53 A; 54 A). However, the close proximity

of this foramen to the ophthalmic foramen in Raja caudaspinosa (Fig. 53 E)

suggests that the ethmoidal nerve and the superficial ophthalmic nerve may

have a common foramen in these species.

The crista supraorbitalis is continued anteriorly to expand on the postero-

dorsal region of the nasal capsule, where it develops to form the preorbital

process (crista praeorbitalis of Heintz (1962); preorbital cartilage of Holmgren

(1940)). The preorbital process is best developed in Rhinobatos halavi (El-Tourbi

& Hamdy 1959: pl. I A), Bathyraja smithii, Raja clavata, R. miraletus and R. alba

(Figs 49 A; 51 A, E; 52 A), not so marked in Raja doutrei, R. pullopunctata,

Rk. radiata, R. leopardus, R. confundens and R. caudaspinosa (Figs 50 A, G; 52 E;

53 A, E; 54 A), and least developed in Cruriraja parcomaculata (Fig. 49 E).

The crista supraorbitalis, which forms the dorsal border of the orbit, is

perforated by a series of small foramina for the superficial ophthalmic nerves.

The number of foramina varies in specimens of the same species and may even

vary on opposite sides of the same specimen. The orbital region is characterized

by the anterior position of the optic foramen, which is typical of the Rhino-

batidae (El-Tourbi & Hamdy 1959) as well as of rajids (Figs 49-54). The optic

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 61

Fig. 50. Dorsal, lateral and posterior views of the neurocrania of:

A. B. C. Raja pullopunctata; D. R. lanceorostrata (X-ray reconstruction) ; E. R. stenorhynchus (X-ray reconstruction) ;

F. R. springer (X-ray reconstruction); G. H. I. R. doutrei.

foramen appears to be somewhat more anteriorly positioned in Raja radiata,

R. confundens, R. leopardus and R. caudaspinosa (Figs 52 F; 53 B, F; 54 B) than it is

in the other species. Above the optic foramen are a series of two or three small

foramina for the passage of the pathetic (trochlear) nerve, the most anterior of

which is the largest. Although these branches of the pathetic all innervate the

superior oblique muscle, the single, large foramen, typical of Rhinobatos (Daniel

1934; El-Tourbi & Hamdy 1959) has not been found in rajids. The position of -

the pathetic foramen given by Heintz (1962) is erroneous and has obviously

been confused with the foramen for the anterior cerebral vein.

In the antero-ventral region of the orbit is the large orbito-nasal canal

foramen, which runs into the nasal capsule and accommodates the anterior

facial vein (Holmgren 1940; El-Tourbi & Hamdy 1959). In Cruriraja parco-

maculata (Fig. 49 F), the orbito-nasal canal is much narrower. Hyman (1942)

points out that in the skate, the deep ophthalmic nerve leaves the orbit via the

orbito-nasal canal, but dissection of several species has revealed that this nerve

joins the superficial ophthalmic nerve at the antero-dorsal region of the orbit,

the two nerves passing through a single foramen, the ophthalmic foramen. This

is also found in Rhinobatos halavi, Rhynchobatos djiddensis and Trygon kuhliu

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

(El-Tourbi & Hamdy 1959) and would therefore appear to be a constant feature

in batoids.

Between the ophthalmic foramen and the foramen for the orbito-nasal

canal is a small foramen for the passage of the anterior cerebral vein. In

Bathyraja smithit, Raja miraletus and R. clavata (Figs 49 B; 51 B, F), it is situated at

about the same level as the optic foramen, in Raja alba (Fig. 52 B) it is displaced

ventrally, and in Raja doutrei, R. pullopunctata, R. radiata, R. confundens, R. leopardus

and R. caudaspinosa (Figs 50 B, H; 52 F; 53 B, F; 54 B) it is displaced dorsally.

Posterior to, and at about the same level as, the optic foramen is the

cartilaginous optic stalk, above which is a single foramen for the oculomotor

(III) nerve. The most conspicuous foramen in the orbit is the large pro-otic

foramen, which is separated from the smaller foramen of the hyomandibular

branch (VII) by the prefacial commissure. Positioned anteriorly to the prefacial

commissure and almost at the ventral edge of the orbit is a very small foramen

for the afferent pseudobranchial artery. Between this and the pro-otic foramen

is another small aperture for the interorbital vein (pituitary vein of El-Tourbi &

Hamdy (1959)), which runs into the cranial cavity in the sella turcica, placing

the two orbits in communication with one another. This has been incorrectly

termed the abducent foramen by Heintz (1962), although it has been labelled

as IV (trochlear) in the figure (fig. 2 B). Dissection has revealed that, as in the

Fig. 51. Dorsal, lateral and posterior views of the neurocrania of:

A. B. C. Raja clavata; D. R. straeleni (X-ray reconstruction); E. F. G. R. miraletus.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 63

case of Rhinobatos halavi, Rhynchobatos djiddensis and Trygon kuhli (El-Tourbi &

Hamdy 1959), the abducent nerve enters the orbit through the pro-otic

foramen in rajids. :

The posterior border of the orbital region is marked dorsally by the small

postorbital process. It is most prominent in Bathyraja smithu (Fig. 49 A). On each

side of the neurocranium, at its posterior end, are the auditory capsules,

between which is the parietal depression (endolymphatic fossa of El-Tourbi &

Hamdy (1959)). This is similarly situated in all rajid species and contains two

pairs of foramina leading to the auditory capsules. The anterior pair are the

endolymphatic foramina and the posterior pair, which lead directly to the

perilymphatic cavity, are the perilymphatic foramina.

At the posterior edge of the lateral surface of the auditory capsule is the

articular facet of the hyomandibular cartilage, which is in the form of an

elongate, slightly concave groove, running antero-dorsally. It is bordered

dorsally by a shallow depression, the postorbital groove, which contains the

posterior region of the postorbital sinus. This vessel passes through the jugal

arch to become the anterior cardinal vein (jugular vein of Heintz (1962)) at the

junction with the posterior cerebral vein (O’Donoghue & Abbott 1928). The

jugal arch links the protuberance of the hyomandibular facet to the posterior

region of the auditory capsule. Its size and position (Figs 49-54) seem to bear

Fig. 52. Dorsal, lateral and posterior views of the neurocrania of:

A. B. C. Raja alba; D. R. robertsi (X-ray reconstruction); E. F. G. R. radiata.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

some relationship to the groupings suggested by the clasper structure.

The occipital region of the neurocranium is more or less similar in form

in all species (Figs 49-54), although the skull appears to be comparatively

higher and narrower in Raja confundens and R. caudaspinosa (Figs 53 C, G).

Foramina for the posterior cerebral vein are absent in Cruriraja parcomaculata

(Fig. 49 G), so that in this species the vein probably leaves the cranial cavity

together with the vagus nerve (X), as in the case of Squalus acanihias (O’ Donoghue

& Abbott 1928).

The ventral surface of the neurocranium runs forwards from the occipital

region as a flat plate, although it forms a shallow subethmoidal depression

between the bases of the nasal capsules. Medially, it is perforated by one

(sometimes two) small foramen, through which the internal carotid artery

enters the cranial cavity. Hyrtl (1872) has described the cross-over system of the

internal carotids in Raja clavata. Anterior to the subethmoidal depression, the

prolongation of the trabecular plates leads to the formation of the rostral

cartilage (El-Tourbi & Hamdy 1959), which runs in an even curve towards the

tip of the snout. As has been pointed out (Hulley 1970: fig. 20), some lateral

undulation of the rostral cartilage occurs in Bathyraja smithu, but dorsal/ventral

undulation of the rostral cartilage, especially at its anterior extremity, is found

———— [SS

Fig. 53. Dorsal, lateral and posterior views of the neurocrania of:

A. B. C. Raja confundens; D. R. dissimilis (X-ray reconstruction) ; E. F. G. R. caudaspinosa.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]) 65

only in Raja caudaspinosa (Fig. 53 F).

The dorsal surface of the neurocranium is perforated by two large fonta-

nelles, separated by the narrow epiphysial bridge. The most anterior of these,

usually termed the anterior fontanelle, is an amalgamation of the precerebral

fontanelle and the anterior supracranial fontanelle (Hamdy 1960). The

posterior fontanelle should therefore be correctly termed the posterior supra-

cranial fontanelle (El-Tourbi & Hamdy 1959; Hamdy 1960). In Rhinobatos

halavi and Rhinobatos productus (Daniel 1934: fig. 62), the anterior fontanelle is

moderately wide at its posterior end, and narrows anteriorly to run as a groove,

almost to the tip of the rostral cartilage, i.e. there is no marked anterior border

to the fontanelle in these species. Anterior grooving of the rostral cartilage is

found only in Raja doutrei, R. pullopunctata, R. lanceorostrata and R. stenorhynchus

(Fig. 50) and reaches its maximum development in R. springert (Fig. 50 F).

In these species the posterior region of the anterior fontanelle is moderately

wide, while the posterior margin is produced in Raja doutrei and R. lanceorostrata

(Figs 50 D, G), and evenly curved in Raja pullopunctata, R. stenorhynchus and

R. springert (Figs 50 A, E, F). In Bathyraja smithu (Fig. 49 A) the anterior

fontanelle is pear-shaped, with a broad posterior region and an evenly curved >

anterior margin, which is not elongated to form a furrow. In Raja clavata,

Fig. 54. Dorsal, lateral and posterior views of the neurocrania of:

A. B. C. Raja leopardus; D. R. ravidula (X-ray reconstruction) ; E. R. spinacidermis (X-ray recon-

struction); F. R. wallacei (X-ray reconstruction).

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

R. straeleni, R. miraletus, R. alba and R. wallace: (Figs 51 A, D, E; 52 A; 54 F) the

fontanelle is moderately obovate, while in Raja dissimilis, R. ravidula, R. confundens

and R. leopardus (Figs 53 A, D; 54 A, D) it is narrowly obovate, with a convex

posterior margin. In Raja radiata (Fig. 52 E) it is broadly obovate, and in

R. robertsi (Fig. 52 D) ellipsoid with a truncate anterior margin. The fontanelle

in Raja spinacidermis (Fig. 54 E) is reduced and extends only for about one-third

the length of the rostral cartilage, but in Raja caudaspinosa, and Cruriraja parco-

maculata (Figs 49 E; 53 E) it is broadly obovate and reaches the level of the

rostral appendices. Crurirqa triangularis (Fig. 49 D) differs from C. parcomaculata

in that the anterior fontanelle does not reach the level of the rostral appendices.

The posterior fontanelle is difficult to follow, especially in X-ray photo-

graphs, but it appears to be more or less similar in all species, being constricted

to form a waist medially. This does not appear to be the case in Raja pullopunctata

and R. dissimilis (Figs 50 A; 53 D), while in R. ravidula (Fig. 54 D) two posterior

fontanelles are present. The extent of the posterior fontanelles, their anterior

and posterior margins, and the degree of constriction vary in the different

species (Figs 49-54).

Discussion

It is recognized (Bigelow & Schroeder 1962) that the suborder Rajoidea

may be divided into four families, Rajidae, Arhynchobatidae, Pseudorajidae

and Anacanthobatidae, on the basis of the number of dorsal fins and the nature

of the anterior lobe of the pelvic fin. De Buen (1959) recognized a further family

Gurgesiellidae, based on a single specimen (Gurgesiella furvescens) from the

abyssal region off Valparaiso, Chile. However, Bigelow & Schroeder (1962)

hold that the characters on which the family were based appear to fall within

the Pseudorajidae and have synonymized the two. It is now apparent that, not

only is the family Gurgesiellidae valid, but also that Pseudoraja atlantica, known

from the Atlantic coast of Nicaragua, falls within this family (Hulley 1972). It

appears that the Rajidae possess common, but distinguishable characters in the

structure of the pelvic bar and in the number of proximal basal segments in the

myxopterygia.

As has been pointed out, two intermediate segments connect the basiptery-

gium with the axial cartilage in Raja, Cruriraja, Bathyraja and Anacanthobatis.

This would support the view that the number of basal segments varies within

large groups of elasmobranchs (Huber 1901), and cannot be used for the

separation of families within the Rajoidea. In. terms of the pelvic girdle, the

basic plan of a transverse (or only slightly arched) pelvic bar, with a pair of

lateral prepelvic processes, is adhered to in the Rajoidea. However, the girdles

can be divided into four (six, with the inclusion of the Pseudorajidae and

Gurgesiellidae) distinctive groups.

Group 1

Pelvic bar simply transverse; lateral prepelvic processes of varying length;

iliac processes recurved; two or more obturatorial foramina.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE!) 67

The prepelvic processes appear to vary considerably in length between

closely related species, e.g. Raja clavata and R. straeleni (Figs 3 g, j), so that this

does not appear to be a suitable diagnostic character within the group. On the

basis of anterior arching of the bar, the black-bellied skates may be recognized

as a separate, identifiable subgroup. In the remaining species, there appears to

be a gradation in the recurvature of the iliac processes, with Bathyraja smithii and

Raja spinacidermis representing the two limits of the sequence (Fig. 3).

Group 2

Pelvic girdle transverse, with iliac region poorly developed; single

obturatorial foramen.

This group includes the monotypic genus Arhynchobatis, known only from

New Zealand.

Group 3

Pelvic girdle simply transverse, with iliac regions moderately developed;

one obturatorial foramen; prepelvic processes poorly developed; iliac processes

large and recurved.

This group includes all species of the genus Cruriraja from South Africa, as

well as C. rugosa from the Gulf of Mexico.

Group 4

Pelvic girdle transverse, with iliac regions greatly developed; single large

obturatorial foramen; iliac processes small; prepelvic processes large.

This group includes both the South African and North American represen-

tatives of the family Anacanthobatidae.

The present taxonomy recognizes that groups 1, 2 and 4 form separate

families within’ the Rajoidea, namely Rajidae, Arhynchobatidae and

Anacanthobatidae respectively, while group 3 (Cruriraja) is considered to be a

genus within the Rajidae. However, it is now proposed that, on the basis of the

pelvic girdle, a new family Crurirajidae be erected to contain the genus

Cruriraja. This is further supported by the structure of the clasper. The family

will be defined at a later stage in this paper. Similarly, on the basis of the

structure of the pelvic girdle, neurocranium and hyomandibular cartilage, the

Gurgesiellidae and Pseudorajidae are now recognized to be distinct families

(Hulley 1972). Examination of the X-ray photographs of Psammobatis lima,

P. scobina, P. extenta and P. microps has revealed that the girdles are not unlike

those of Raja clavata and R. miraletus (Figs 3 i, j}) and would therefore not

necessitate the recognition of a further family at this stage. However, detailed

studies of the structure of the claspers of these species is required.

The southern African Rajidae can be grouped into eight distinguishable

types, on the basis of the clasper structure:

Type A: Bathyraja smithii (Miller & Henle).

Claspers very slender and elongate, with club-shaped distal glans; well-developed

pseudosiphon on outer dorsal wall; internally, shield and spike absent, but sharp-edged

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

knife lying beneath projection; rhipidion absent, pseudorhipidion present; dorsal

terminal 3 situated laterally to dorsal terminal 2; ventral terminal simple.

Although the single species was referred to the genus Bathyraja because of

the structure of the rostral bar and appendices (Hulley 1970), there is no doubt

that this type of clasper is characteristic of the genus (Ishiyama 1958; Ishiyama

& Hubbs 1968; Stehmann 1970). However, Ishiyama & Hubbs (1968) have

given diagnostic characters, which are not altogether correct for the genus

(Hulley 1970). The pseudorhipidion and pseudosiphon are not wholly confined

to the genus: the former is found in Raja clavata, R. straelem, R. miraletus and

R. alba (in Bathyraja smithu (Hulley 1970: fig. 21), the pseudorhipidion is not

labelled, although the associated dorsal marginal cartilage is well developed

(Fig. 29)); the latter is present in Raja radiata, R. robertsi and possibly in R. wallacei

and R. caudaspinosa. Differences based on distribution for the separation of Bathy-

raja from Breviraja given by Ishiyama & Hubbs (1968) add to the confusion, and it

is now considered that Bathyraja is a bipolar, antitropical genus, while Breviraja

is confined to slope areas in the tropics (Hulley 1970).

In terms of clasper structure, the Japanese subgenus Rhinoraja can be

separated from Bathyraja by its distinct, external pseudosiphon, large dorsal

terminal 1 cartilage and elongate ventral marginal cartilage. However, these

characters do not seem to be that significant, for a distinct external pseudosiphon

is present in Bathyraja matsubarai, B. aleutica and B. diplotaenia (Ishiyama 1958:

fig. 3), the dorsal terminal 1 is not markedly reduced in some Bathyrqa species

(Ishiyama 1958: fig. 8), and the ventral marginal is particularly elongate in

Bathyraja trachouros and B. abasiriensis. The subgenus Rhinoraja can therefore be

distinguished only on the segmented nature of the rostral bar. “

It should be noted that the shape of the accessory terminal cartilage in

Bathyraa smith (Fig. 24 a) is very similar to that in Rhinoraja odai and Rhinoraja

longicauda (Ishiyama 1958: fig. 11). However, the nature of the rostral bar

precludes its identification within the genus Rhinorqa.

Type A corresponds to Stehmann’s spinicauda-type, which includes

Bathyraja spinicauda, B. pallida and B. richardsoni (with reservation), and which is

now held to be equivalent to the genus Bathyraja (Stehmann 1970).

Type B. Raja doutrei Cadenat; R. pullopunctata Smith; R. lanceorostrata Wallace;

(and probably includes R. springert Wallace and R. stenorhynchus

Wallace).

Clasper moderately long, naked, with spatulate distal tip more or less dorso-ventrally

flattened ; pseudosiphon absent; inner dorsal lobe of glans with two clefts, separated by

terminal bridge; rhipidion well developed; shield large and covered with pleated

epithelia; sentinel usually small and bluntly rounded or absent; spike always present,

sometimes hooked; ventral terminal with dorsally convex, outer lateral margin;

accessory terminal 1 with asymmetrical, proximal arms; accessory terminal 2 with or

without attachment process.

All southern African ‘black-bellied’ skates are included in this group, which is

comparable with the oxyrhynchus-group of the eastern North Atlantic (Stehmann

1969, 1970). The group apparently has a world-wide distribution with the

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 69

following species: Raja batis, R. oxyrhynchus and R. nidarosiensis in the eastern

North Atlantic; Raja laevis, R. oregoni, R. bullist, R. floridana and R. garricki in the

western North Atlantic; R. flavirostris in the western South Atlantic; Raja rhina

and R. binoculata in the eastern North Pacific; Raja australis, R. nasuta and

possibly <earaja gudgert around Australia and New Zealand; and the species of

the subgenus Tengwe: Ishiyama (Raja tengu, R. pulchra, R. macrocauda and

R. gigas) in the western North Pacific. ‘These species inhabit the continental

shelf area and penetrate the slope regions to between 350 m and 550 m, with a

maximum recorded depth of 740 m in the case of Raja springert (Wallace 1967).

The poor degree of development of the distal projection of the accessory

terminal 1 appears to be characteristic of all the species, so that the sentinel is

poorly developed in the glans. Of the species examined, Ishiyama (1958)

reports a massive development of the accessory terminal 1 cartilage in a single

species, Raja pulchra, where the condition of the cartilage (Ishiyama 1958: fig. 16;

1967: fig. 9) approximates to the condition found in Raja clavata and R. straelent.

However, there are marked dissimilarities in the detachment process and in the

accessory terminal 2 cartilage between Raja pulchra and R. clavata and

R. straeleni.

Raja pullopunctata does not possess a sentinel in the glans, as the distal

projection of the accessory terminal 1 does not develop (Fig. 24 d).

A well-developed and hooked spine is found in Raa lanceorostrata and in

the Japanese species R. gigas (Ishiyama 1958, 1967) and further similarities are

shown in the parallelogram-shaped dorsal terminal 1 and the distal projection

of the dorsal marginal cartilage (Fig. 31; Ishiyama 1967: fig. 11). However,

differences can be seen in the dorsal terminal 2 and 3 cartilages, the proximal

elongation of the outer edge of the dorsal terminal 1 cartilage, the expansion of

the distal projection of the accessory terminal 1 and the attachment process of

the accessory terminal 2. Furthermore, the outer lateral edge of the ventral

terminal appears to be comparatively broader in Raja gigas.

The spatulate terminal end-to the distal projection of the accessory

terminal 1 is found in Raja lanceorostrata (Fig. 24 c), R. batis (Hulley 1966: fig. 7)

and &. garricki, but there are marked differences in the structure of the dorsal

terminal 1, ventral terminal and accessory terminal 2 cartilages between the

species. The degree of hooking of the accessory terminal 2 in the geographically

separated species Raja batis, R. lanceorostrata, R. macrocauda, R. gigas and

R. garricki may be considered in terms of parallel evolution.

The possession of an anteriorly arched pelvic bar is peculiar to the southern

African species of ‘black-bellied’ skate (see above) and in the South American

species Raja flavirostris (Fig. 3 f) and has been confirmed in the case of R. garricki

and R. oregoni from the western North Atlantic. It would appear, therefore, that

this condition of the pelvic bar is characteristic of the group as a whole.

‘Black-bellied’ skates were recognized as a separate group, Gammaraia, by

Leigh-Sharpe (1925), who proposed the pseudogenus on the basis of external

clasper structure. However, this taxon has no nomenclatural standing (Jordan

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

1925; Bigelow & Schroeder 1953). On the other hand, Ishiyama (1958)

identified the Japanese representatives of the group as a separate subgenus,

Tengwei. Stehmann (1969) has reviewed the taxonomic validity of Ishiyama’s

subgenus, and proposed that the name Laeviraja Bonaparte, 1838 (type-species

Raja oxyrhynchus) be employed. It appears, however, and has been confirmed

(Stehmann 1970), that the name Dipturus Rafinesque, 1810 (type-species

Raa batis) has priority.

Type C: Raga clavata Linnaeus; R. straelent Poll; R. miraletus Linnaeus.

Claspers moderately long, with naked outer surface; some species with pecten on outer

dorsal lobe; pseudosiphon absent, but inner dorsal lobe with pocket and cleft; sentinel

massively developed, with knife-edged outer lateral margin, fitting into pocket; spike

situated distally in sentina; fleshy signal situated proximally at about level of well-

developed pseudorhipidion; ventral terminal characteristically J-shaped, with distal

protuberance, forming funnel.

Hulley (1970) has pointed out that the species Raja clavata, R. straeleni,

R. herwigi and R. maderensis form a complex of very closely related species within

the genus Raja. It would appear that the species Raja miraletus can now be

identified with the complex. Stehmann (1970) terms this complex the clavata-

group, and considers it to be a separate subgenus Raja Linnaeus (type-species

Raja miraletus).

Stehmann (1969: 137) is of the opinion that the species Raja picta and

R. alba are members of the subgenus, although they represent ‘slightly divergent

forms’. This may be true in the case of Raja picta, where, although there appear

to be marked differences in the structure of the ventral terminal and accessory

terminal 2 cartilages, some similarities are shown by the dorsal terminal 1 and 2

cartilages. Furthermore, Raja picta possesses a signal. However, Raa alba should

be considered as a separate subgenus. It will be defined at a later stage.

It has been pointed out (Hulley 1970) that Raja straeleni closely resembles

R. clavata, but that the two were held to be distinct on the basis of colour pattern

and of differences in the shield of the clasper glans. ‘There are further differences

in the clasper cartilages of the two species. In Raja straeleni, the dorsal terminal 1

is narrower and more blunt distally (Figs 16 g, h), the accessory terminal 2 is

comparatively broader and distally more pointed (Figs 26 e, f), and the outer

lateral margin of the ventral terminal is better developed (Figs 22 e, f). The

most marked difference occurs in the dorsal terminal 2 (Figs 18 e, f), which in

Raja straelent possesses numerous, small, blunt processes on its outer lateral wall.

While the clasper cartilages are species specific without exception (Ishiyama

1958), the differences between Raja clavata and R. straeleni appear to be minor,

suggesting that R. straeleni could be regarded as a subspecies of R. clavata.

Recently, Stehmann (1971) has re-examined this question, and concludes that

the species are distinct.

The subgenus Raja appears to be confined to shelf areas of the eastern

North Atlantic and eastern South Atlantic (Stehmann 1970), with a bipolar

distribution pattern shown by the species Raja clavata (Hulley 1966). Raa

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 71

miraletus is now known to extend into the south-western Indian Ocean (Hulley

1969) and has been reported from the Kerala coast in India (Samuel 1963).

Type D: Raja alba Lacépéde.

Clasper massive, with pecten on outer dorsal surface, but without pseudosiphon;

pseudorhipidion well developed; inner dorsal lobe with proximal cleft, but without

distal slit or pocket; shield prominent with laminate integument; sentinel massive and

situated far distally; spike and signal absent; four dorsal terminal cartilages, with pecten

developed from outer lateral margin of dorsal terminal 3; ventral terminal with

dorsally convex, wide, outer lateral margin.

While Leigh-Sharpe (1924) considered that Raja alba belonged to a

separate pseudogenus Jotaraia, Stehmann (1969: 136) pointed out that R. alba is

closely related to the clavata-group, although ‘in etwas abweichender Richtung

entwickelte Formen gelten’. It now appears obvious from the examination of

the number and arrangement of the internal cartilages that Leigh-Sharpe’s

supposition was correct, but his pseudogenus has no taxonomic standing

(Jordan 1925; Bigelow & Schroeder 1953).

Firstly, there are four dorsal terminals in Raa alba, which are arranged so

that they extend distally from the dorsal marginal to the axial tip, so forming

the framework of the dorsal lobe (Fig. 33). In this arrangement, Raa alba

approximates to Raja radiata, R. caudaspinosa, R. dissimilis, R. confundens,

R. leopardus and R. wallacei. In the subgenus Raja, there are only two dorsal

terminal cartilages and the connection between these and the axial tip is

formed from connective tissue only. Secondly, Raja alba lacks the accessory

terminal 3 and 4 cartilages (forming the signal) which are characteristic of the

subgenus Raja.

It would appear that parallel evolution has occurred in the formation of

the pecten, which presumably acts as a holdfast structure. However, as has

been pointed out above, the pecten in Raja miraletus (subgenus Raja) is formed

from the dorsal terminal 2, while in Raja alba, it is formed by the dorsal terminal 3

cartilage.

Leigh-Sharpe’s (1924) classification of Raja alba (= Raja marginata

Lacépéde) in the pseudogenus Jotaraia has no taxonomic standing. A new sub-

genus, Kostroraja, is therefore proposed to include the species Raja alba, and is

defined on page 77. The subgenus shows bipolar distribution patterns in the

eastern Atlantic (Hulley 1966: fig. 8) and is reported to extend as far north as

off Barra Falsa, Mocgambique, in the Indian Ocean (Wallace 1967). It inhabits

shelf areas from 50 m to 360 m.

Although the following groups are held to be equivalent to their European

counterparts, there are differences in the interpretation of the number of dorsal

terminal cartilages and in the shape of the axial cartilage.

Stehmann (1969, 1970) considers that in the fullonica-type (subgenus

Leucoraja), radiata-type (subgenus Amblyraja) and fyllae-type (subgenus Rajella),

the axial is completely recurved on itself, so that it attains a marked J-shape.

In Amblyraja and Rajella, the recurved tip of the axial then connects with a small

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

dorsal terminal 4 cartilage, to form the framework of the dorsal lobe, while in

Leucoraja, which has a similarly recurved axial, this cartilage connects directly

with the external surface of the dorsal terminal 3. In this subgenus, a dorsal

terminal 4 is absent.

These interpretations are now held to be incorrect. It has been found that

in the species Raja dissimilis, R. confundens, R. caudaspinosa and R. leopardus

(Figs 36-39) a dorsal terminal 4 cartilage is present, and consists of a flattened,

slightly dorsally convex plate, which runs from the external surface of the dorsal

terminal 3 to the axial tip. In these species the cartilage is heavily calcified and

well demarcated, both from the axial and from the dorsal terminal 3 cartilages.

The axial in these cases is therefore spatulate or only very slightly recurved, and

is somewhat similar to the condition of the cartilage in Japanese rajids of the

southern form (Ishiyama 1958). In ‘Type E (Raa radiata: subgenus Amblyraja of

Stehmann) the dorsal terminal 4 is not heavily calcified, so that its texture

resembles that of the axial. It is held that Stehmann (1969, 1970) has incorrectly

termed the calcified proximal end as the dorsal terminal 4 and has referred the

distal end to the axial. In Raya fullonica, R. circularis and R. naevus (subgenus

Leucorqa), Stehmann (1969, 1970) finds no dorsal terminal 4 cartilage, but in

the southern African species Raja wallace: (Fig. 35), which is obviously of the

same subgenus, a dorsal terminal 4 cartilage, which is usually well calcified, is

easily identifiable. Therefore, the axial cartilage is spatulate or only slightly

recurved. Furthermore, the subgenera Amblyraja, Leucoraja and Raella show an

obvious sequence, especially in regard to the dorsal terminal 3 cartilage, when

considered in the light of these findings.

‘Type E: Raga radiata Donovan; R. robertsi Hulley.

Claspers short and club-like; pseudosiphon in outer dorsal lobe; spur well developed,

forming outer border of distal cleft; sentinel partly covering spike; pent extending

distally from rhipidion; distal end of prominent shield with laminate integument; four

dorsal terminal cartilages; ventral terminal typically J-shaped; accessory terminal I

characteristic, with concave dorsal surface, enveloping the accessory terminal 2.

The pseudogenus Deltaraia was proposed by Leigh-Sharpe (1924) to

include the species Raja radiata and R. naevus. However, Stehmann (1970) has

grouped the species Raja radiata and R. hypoborea as a separate subgenus,

Amblyraja Malm, 1877, and, mainly on the basis of clasper structure, has shown

that Raja naevus should be referred to the subgenus Leucoraja Malm, 1877.

From its morphometry and the presence of a pseudosiphon in the claspers

of the juvenile male type, Hulley (1970) has suggested that Raja robertst should

be included with R. radiata, so forming a group which encompasses the species

Raa duellojuradoi, R. frerischi and R. georgiana in the western South Atlantic,

R. jenseni from the western North Atlantic and R. badia from the Gulf of Panama.

The bipolar distributional pattern of the twin species has already been pointed

out (Krefft 1968). All species of this subgenus inhabit slope areas of the archi-

benthal and some penetrate to abyssal depths. It should be pointed out that Raja

badia is at present the only species of the subgenus not found in the Atlantic.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 73

Type F: Raa wallace: Hulley.

Claspers short and spatulate; dermal denticles usually present externally; cavity in

musculature of outer dorsal lobe usually present (? pseudosiphon); proximal slit

bordered by roll; distal cleft bordered by small spur, usually covered by integument;

promontory present; spike and sentinel small and situated far distally; rhipidion fan-

shaped; four dorsal terminal cartilages, with dorsal terminal 3 forming small spur.

This type of clasper structure is comparable with that of the fullonica-type

of Stehmann (1969), the species of which have been included in the subgenus

Leucorgqa Malm, 1877. While the subgenus is represented by three species in the

eastern North Atlantic, only one species is known from southern African waters,

where it has been recorded at depths varying from 73 m to 445 m from west of

Cape Point north to the Limpopo River mouth (Wallace 1967). Therefore,

Raja wallace: like R. fullonica, shows a depth distribution into the upper limits

of the archibenthal. Recently, Stehmann (1971c) has recognized a further

species of the subgenus, Raja (Leucoraja) leucosticta, from the tropical waters of

West Africa. This species had previously been confused with Raja ackleyi,

Garman.

It has been pointed out that the cavity found on the outer surface of the

dorsal lobe of the glans is formed in a different manner to the pseudosiphon and

is related only to the musculature. However, the two cavities may prove to be

related. As can be seen (Figs 34, 35) in both Raa radiata and R. wallacei, the

dorsal terminal 3 cartilage is developed into a laterally projecting spur, which

is large and S-shaped in Raja radiata, but smaller and less well developed in

Raa wallace. However, in the form and orientation of the accessory terminal 1

and 2 cartilages and in the ventral terminal, Raja wallace: shows a closer

similarity to Raja caudaspinosa, R. confundens, R. leopardus and R. dissimilis

(Figs 24, 26). Stehmann (1970) indicates an accessory terminal 3 cartilage in

Raja fullonica, but not in R. circularis and R. naevus. Only two accessory terminals

are found in Raja wallace.

Type G: Raa caudaspinosa Von Bonde & Swart; R. leopardus Von Bonde &

Swart; R. confundens Hulley; R. dissimilis Hulley; (and probably

R. ravidula Hulley).

Claspers small and short, with or without dermal denticles on outer dorsal lobe; some

specimens with a small cavity in musculature; two clefts or a cleft and a slit present;

spike and sentinel small; rhipidion fan-shaped and usually associated with pent; shield

not well developed; axial cartilage spatulate; four dorsal terminal cartilages; dorsal

terminal 3 pointed, but not developed into lateral spur.

Species with this type of clasper structure have been grouped into the

subgenus Rajella Stehmann (type-species Raja fyllae), which, in the North

Atlantic, may also include the species Raja garmani, R. erinacea, R. ocellata and

R. lentiginosa. On the basis of Raja fyllae, Stehmann (1970) confines the subgenus

to the archibenthal, but depth distribution records for the South African species

(Hulley 1970) indicate that some of the species may occupy the shelf areas

(Raja leopardus), while others are known only from abyssal depths (Raa

dissimilis, R. ravidula).

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

The presence of four dorsal terminal cartilages and the shape of the axial

have already been discussed above in relation to Stehmann’s ideas. Similarities

in the accessory terminal 1 and 2, dorsal terminal 1 and ventral terminal

cartilages, and the framework arrangement of dorsal terminals 2-4, can be

seen between this group and ‘Type F. However, the dorsal terminal 3 is

differently developed.

Type H: Cruriraja parcomaculata (Von Bonde & Swart); C. triangularis Smith;

C. rugosa Bigelow & Schroeder; (and probably includes C. durbanensis

(Von Bonde & Swart)).

Claspers small and spatulate; dermal denticles usually absent, but one species with

denticles on ventral border; thorn present on lateral edge of dorsal lobe, eperon on

ventral lobe; inner dorsal lobe with pseudosiphon; rhipidion small and closely attached;

knob and sentinel present; spike tongue-shaped and situated medially; axial cartilage

characteristically J-shaped; dorsal terminal 1 enclosed by axial limb; terminal

bridge between axial and accessory terminal 2.

This particular type of clasper structure is readily distinguished from all

other types and is associated with the genus Cruriraja. In fact, it is further

removed from the typical rajid type than Bathyraja, and, as in the case of the

Anacanthobatidae, suggests that a further family within the Rajoidea be

distinguished. Examination of the pelvic girdles and neurocrania seems to

agree with this. A new family will therefore be defined (see page 78).

The genus is, at present, known only from two areas, the southern African

coast (Cruriraja durbanensis, C. parcomaculata and C. triangularis), where the species

inhabit shelf areas and the upper regions of the archibenthal, and the Gulf of

Mexico (Cruriraja atlantis, C. poeyt, C. rugosa and C. cadenati), where they are

recorded from archibenthal regions.

As has been pointed out, the predorsal caudal vertebral count (Vprd) is

not species specific, but it would seem that in certain cases it may be used to

distinguish between closely related species (Stehmann 1970; Hulley 1970) and

may even be used for taxonomic discrimination at the subgeneric level.

Stehmann (1970) suggests that Raja alba may be separated from the subgenus

Raga by both Vtr and Vprd counts. While there appears to be some overlap in

Vprd values for Raja alba and R. clavata in the eastern North Atlantic (Stehmann

1970: fig. 14), the counts for the southern African specimens appear to be well

separated (Fig. 47), substantiating the recognition of Rostroraja, which was based

on clasper structure.

The true significance of this method of application is open to question,

since the separation of species on the basis of Vprd counts is not always supported

by clasper distinctions. For example, in the southern African subgenus Rayella,

two distinct species groups can be recognized (Fig. 47), but the clasper structure

of the species Raja leopardus, R. caudaspinosa, R. confundens and R. dissimilis are

almost identical. It is probable that with increasing knowledge, species of this

particular subgenus, with intermediate values in Vprd count, may be found, as

with Bathyraja and Dipturus.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 75

Apparent differences seem to exist between Bathyraja pallida (Vprd 66-67)

and B. spinicauda (Vprd 77-82) (Stehmann 1970). However, within this genus

intermediate values have been recorded (Ishiyama 1958; Hulley 1970).

Similarly, the high Vprd values for Raja (Dipturus) nidarosiensis (Stehmann 1970)

are approximated by the southern African Dipturus species, Raja pullopunctata and

R. lanceorostrata (Table 1), and are exceeded by the Japanese species Raja

macrocauda and R. gigas.

While the shape of the anterior and posterior fontanelles may be species

specific in bathyrajid species (Ishiyama 1958), marked differences are not

evident in other rajids. It would appear therefore that neurocranial characters

are mainly applicable above the species level. There can be no doubt that

characters such as the structures of the rostral cartilage and appendices are of

significant value in taxonomy (Bigelow & Schroeder 1948; Ishiyama 1952,

1958; Stehmann 1970). These have already been employed in the recognition

of genera in southern African Rajidae (Hulley 1970). However, it appears that

certain cranial characters can be used to separate the proposed subgenera of

Raja, while the majority can best be interpreted in terms of the phylogenetic

interrelationship between these subgenera.

Stehmann (1970) suggests that the length of the rostrum in relation to the

cranial length may be used for the grouping of species and for the interpretation

of phylogenetic relationships. He has pointed out that the rostral length is

greater than the ‘cranial length’ (not cranial length of Hubbs & Ishiyama

(1968: fig. 1) but equal to total cranial length minus rostral length) in Dzpturus,

but less than the ‘cranial length’ in all other subgenera of Raja. While high

values for the rostral length/cranial length proportion are found in Dipturus

species (Table 2), equally high values have been found for Rostroraja, and values

above 50% have been found in Raya confundens and R. ravidula. Furthermore, the

course of the hypothetical line drawn from the tip of the rostrum through the

ophthalmic foramen appears to be a complex function of the rostral length and

cranial width, and bears little relation to the suggested subgeneric groupings.

However, Dipturus is characterized by the forward extension of the anterior

fontanelle, so that in all species of this subgenus the rostrum is grooved, there

being no definite anterior limit to the fontanelle. The extent of the rostral

grooving varies in the different species (Fig. 50) and its presence can be regarded

as primitive, since Dipturus shares this character with the Rhinobatidae.

The size of the nasal capsules varies to some extent. Enlarged capsules are

found in Raja robertsi, R. dissimilis, R. ravidula and R. spinacidermis (Figs 52 D;

53 D; 54 D, E), a fact which seems to be correlated with depth distribution

rather than proposed groupings, since these species (of different subgenera) are

all found to occur at depths of 1 000 metres or more (Hulley 1970). Although no

comparison was made of the number of Schneiderian and secondary folds or

the number of sensory cells per unit area between these species and the conti-

nental shelf species, the massive development of the nasal capsules would

substantiate the theory that olfaction is probably of greater importance in a

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

deep-water habitat (Weichart 1951). As with the rostral length, the orientation

of the nasal capsules bears little relationship to the groupings, but their shape,

which is altered by the degree of development of the preorbital processes, may

be of importance.

The preorbital process is well developed only in Raa and Rostroraja

(Figs 51; 52 A), a character which these subgenera share with both the Rhino-

batidae and with Bathyrga (Fig. 49 A; Ishiyama 1958: fig. 28; Stehmann 1970:

fig. 27). This character, together with the orientation of the nasal capsules, may

be employed to distinguish Raa and Rostroraa from the other subgenera.

Similarly, the presence of a comparatively large postorbital process in Bathyraja

smithi (Fig. 49 A) and other bathyrajids (Ishiyama 1958: fig. 28; Stehmann

1970: fig. 27) may be used as a taxonomic character for distinguishing this genus.

A marked degree of constriction of the neurocranium across the orbital

region is characteristic of Rajella (Figs 53; 54 A, D), but the noticeable anterior

displacement of the optic foramen and the position of the anterior cerebral vein

foramen in this subgenus and in Amblyraja and Dipturus are best interpreted in

terms of their phyletic relationship. The possession of a small orbito-nasal canal

foramen in Crurtraja parcomaculata (Fig. 49 F) could be taxonomically employed,

but further evidence is required from other species of this genus.

The jugal arches are angular and prominent in Amblyrqa (Figs 52 D, E), in

which they project as wing-like extensions from the laterally expanded auditory

capsules. Angular arches are also found in Rajella (Figs 53; 54 A, D), but are

directed postero-laterally from moderately expanded auditory capsules. In

Raa, Leucoraja and Rostroraa (Figs 51; 52 A; 54 F), the jugal arches are more or

less evenly curved and project posteriorly, while in Dzpturus (Fig. 50) they are

comparatively smaller and do not disrupt the contour of the auditory capsule

region. In Raja spinacidermis (subgenus Malacoraja) (Fig. 54 E) the arches are

poorly developed and thin, while in Crurirqja (Figs 49 D, E) they are particularly

thin at their junction with the process of the hyomandibular facet.

The taxonomic significance of the shape and position of the posterior

fontanelle is open to question, since these characters and even the differentiation

into two posterior fontanelles vary within the same species (Stehmann 1970:

figs 17, 19, 20, 24). As has been pointed out above, the shape of the anterior

fontanelle is species specific in Bathyraja, and its shape and degree of develop-

ment in other rajids are only suggestive of the subgeneric groupings, which are

based on clasper structure.

In conclusion, it should be pointed out that while certain neurocranial

characters, such as the constriction of the orbital region, the degree of develop-

ment of the preorbital and postorbital processes, the orientation of the nasal

capsules and jugal arches, and the shape and extension of the anterior fontanelle,

may be used in combination as taxonomic characters at the subgeneric and

generic level, their significance is more difficult to interpret than that of the

clasper structure. However, they may be suitably employed in elucidating

phyletic interrelationships.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) Ti.

To summarize, the southern African Rajidae can be identified within the

framework of the genera and subgenera proposed by Stehmann (1970), which

is based mainly on clasper structure, as follows:

Family Rajidae

Genus Bathyraja Ishiyama, 1968

type-species Raja isotrachys Gunther, 1877

species : Bathyraja smithu (Muller & Henle, 1841)

Genus Raja Linnaeus, 1758

type-species Raja miraletus Linnaeus, 1758

I. subgenus Aaa Linnaeus, 1758

type-species Raja miraletus Linnaeus, 1758

species: Raja clavata Linnaeus, 1758; Raja straeleni Poll, 1951

2. subgenus Dzupturus Rafinesque, 1810

type-species Raja batis Linnaeus, 1758

species: Raja doutrer Cadenat, 1960; Raa pullopunctata Smith,

1964; Raja springeri Wallace, 1967; Raja lanceorostrata

Wallace, 1967; Raja stenorhynchus Wallace, 1967.

3. subgenus ROSTRORAJA subgen. nov.

type-species Raja alba Lacépéde, 1803

Definition

Clasper massive, with pecten on outer dorsal surface, but without pseudo-

siphon; pseudorhipidion well developed; inner dorsal lobe with proximal cleft,

but without distal slit or pocket; shield prominent with laminate integument;

sentinel massive and situated far distally; spike and signal absent; four dorsal

terminal cartilages, with pecten developed from outer lateral margin of dorsal

terminal 3; ventral terminal with dorsally convex, wide, outer lateral margin.

Rostrum produced and elongate, its length more than 50% of cranial length;

nasal capsules moderately developed and orientated anteriorly; anterior

cerebral vein foramen ventral.

4. subgenus Amblyraja Malm, 1877

type-species Raja radiata Donovan, 1808

species: Raja robertst Hulley, 1970

5. subgenus Leucoraja Malm, 1877

type-species Raja fullonica Linnaeus, 1758

species: Raja wallace: Hulley, 1970

6. subgenus Rajella Stehmann, 1970

type-species Raya fyllae Liitken, 1888

species : Raja caudaspinosa Von Bonde & Swart, 1923; Raga

leopardus Von Bonde & Swart, 1923; Raja confundens

Hulley, 1970; Raja dissimilis Hulley, 1970; Raa

ravidula Hulley, 1970

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

Stehmann (1970) has defined a further subgenus, Malacoraja, based on

Raja mollis. It is now considered that Raja mollis is synonymous with Raja

spinacidermis Barnard (Hulley 1970). As no adult male specimen of this species

has yet been taken, this identification is based on characters other than the

clasper structure. Marked differences in the rostral length, the orientation and

massive development of the nasal capsules and the poorly developed jugal

arches, serve to distinguish Malacoraja from all other subgenera.

7. subgenus Malacoraja Stehmann, 1970

type-species Raja spinaciderms Barnard, 1923

The family Crurirajidae is now erected to include all species of the genus

Cruriraja, which were formerly identified with the Rajidae.

Family CRURIRAJIDAE fam. nov.

Definition

Pectorals with radials of ordinary form, without lateral processes; outer

margins of pelvics deeply notched, to form an anterior, limb-like structure,

consisting of three articulated segments, externally distinct from the posterior,

fin-like lobe of the pelvic; tips of anterior rays of pectorals falling short of tip of

rostral cartilage; two dorsal fins. Otherwise external characters as for suborder.

Claspers small and spatulate; dermal denticles usually absent on dorsal

border; thorn present on lateral edge of dorsal lobe, eperon on ventral lobe;

inner dorsal lobe with pseudosiphon; rhipidion small and closely attached;

knob and sentinel present; spike tongue-shaped and situated medially; axial

cartilage characteristically J-shaped; dorsal terminal 1 enclosed by axial limb;

terminal bridge between axial and accessory terminal 2.

Pelvic girdle simply transverse, with iliac regions moderately developed;

one obturatorial foramen; prepelvic processes poorly developed; iliac processes

large and recurved.

Neurocranium without preorbital and postorbital processes; orbito-nasal

canal foramen small; jugal arches poorly developed; posterior cerebral vein

foramen absent.

Genus Cruriraa Bigelow & Schroeder, 1948

type-species Cruriraja atlantis Bigelow & Schroeder, 1948

species: Cruriraja parcomaculata (Von Bonde & Swart, 1923);

Cruriraja durbanensis (Von Bonde & Swart, 1923); Cruriraja

triangularis Smith, 1964.

An interesting case is presented in the family Anacanthobatidae, which at

present contains two genera, Anacanthobatis and Springeria. The discussion is only

in relation to Anacanthobatis marmoratus and A. americanus, so that before definite

taxonomic changes are made, A. longirostris from the Gulf of Mexico and

A. borneensis from the South China Sea should be examined.

Basic major differences in clasper structure can be seen between Anacantho-

batis marmoratus and A. americanus (Figs 10, 11, 43, 44), the most important of

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 79

which is the absence in A. americanus of the ventral terminal cartilage. Hence

there is a lack of an associated shield and eperon in the clasper glans in this

species. Probably associated with this is the fact that the spike and sentinel in

Anacanthobatis americanus are positioned laterally and are capable of rotation

(Fig. 11). Furthermore, Anacanthobatis marmoratus lacks an external pseudo-

siphon, while in A. americanus there is an external pseudosiphon. Examination

has shown that the structure and orientation of the cartilages is different in the

two species.

It therefore appears that, on the basis of clasper structure, these two species

should not be referred to the same genus and that the family Anacanthobatidae,

as defined at present, may be diphyletic. In terms of priority, the South African

species Anacanthobatis marmoratus becomes the type-species for the genus

Anacanthobatis, which is then considered to be monotypic, while Anacanthobatis

americanus should be referred to a new genus. Although Bigelow & Schroeder

(1953) consider that a second South African species, Anacanthobatis dubius

Von Bonde & Swart, 1923, can be referred to the genus Springeria, they later

(1962) hold that A. dubius is probably an immature A. marmoratus, so that

Wallace (1967) has synonymized the two under Anacanthobatis marmoratus. As the

type specimens of these species are missing, this synonymy has been tentatively

accepted.

Family Anacanthobatidae

Genus Anacanthobatis Von Bonde & Swart, 1923

type-species Anacanthobatis marmoratus Von Bonde & Swart, 1923

Genus Springeria Bigelow & Schroeder, 1951

type-species Springeria foliorostris Bigelow & Schroeder, 1951

species: Springeria ort Wallace, 1967

PHYLETIC INTERRELATIONSHIPS

White (1937) and Saint-Sienne (1949) consider that recent elasmobranchs,

which do not include the archaic families Heterodontidae and Hexanchidae,

arose from a common ancestor, which was not unlike Palaeospinax (family

Heterodontidae). A monophyletic origin is also held by Disler (1966) on the

basis of the embryonic development of Trygon pastinacea. Examination of the

structure of the neurocranium led Holmgren (1941) to suggest a diphyletic

origin for the recent Elasmobranchii, a view supported by Hasse’s (1885)

observations, and by Melouk (1947), who considers that the benthic mode of

life is more primitive. However, it has now been shown that the features stressed

by Holmgren (1941) represent minor ontogenetic modifications, and Schaeffer

(1967) therefore considers separate origins for galeoids, squaloids and batoids,

the latter probably arising from some benthic hybodont stock.

Within the batoid line, it is generally considered (Regan 1906; Leigh-

Sharpe 1924; Holmgren 1941; Melouk 1947; Gregory 1951) that the Rajoidea

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

arose from the Rhinobatidae, since they show a strong resemblance in dentition

and in the possession of longitudinal folds on the tail. Ishiyama (1958), in

comparing skull, rostral and clasper structure, arrives at the following

conclusions:

(1) Rajids have arisen from a rhinobatid-like stock rather than a platyrhinid,

since there is a trend in the reduction of basal segments in the myxoptery-

gium and in denticulation, and a trend in the development of the clasper

siphon.

(2) The genus Bathyrga originated directly from the ancestral stock, with

subsequent neotonous modification of the rostral cartilage, but with

retention of primitive characters in the clasper.

(3) Both Raja and Rhinoraja species can be derived from the bathyrajid-type,

with the genus Raja representing an earlier split.

(4) Amongst the species of the genus Raja, those of the subgenus Tengujei

(= Dipturus) are considered to be more primitive, since they show

primitive characters in their rostral structure.

Stehmann (1970) has presented a phylogeny for the rajid species of the

eastern North Atlantic, which, as has been shown above, form subgeneric

groupings identical with those of the species of the southern African region. This

phylogeny is based on clasper anatomy, rostral structure and geographic

distribution. Stehmann (1970) selects the pseudosiphon as the definitive

character, so leading to a division of the rajids into Rajidae I (without a

pseudosiphon — Dipturus, Raja) and Rajidae II (with a pseudosiphon — Bathyrga,

Rhinoraja, Leucoraja, Amblyraja, Rajella) (Fig. 55).

There is no doubt that the pseudosiphon is a primitive structure, and that

DIPTURUS AMBLYRAJA LEUCORAJA

RAJA RAJELLA BATHYRAJA

rhipidion pseudorhipidion rhipidion rhipidion rhipidion pseudorhipidion

no pseudosiphon no pseudosiphon pseudosiphon ? pseudosiphon pseudosiphon Ppseudosiphon

RAJIDAE | RAJIDAE II

RAJIDAE

HYPOTHETICAL ANCESTOR

Fig. 55. Relationship within the Rajidae as suggested by Stehmann (1970), showing

the possession of rhipidion, pseudorhipidion and pseudosiphon.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 81

Ishiyama’s (1958) interpretation of the ancestral nature of the genus Bathyraja

is correct, for in this genus the ventral terminal cartilage is simple and not

expanded into a lateral shield, the accessory terminal is reminiscent of Rhinobatos,

and the axial cartilage is bluntly pointed rather than spatulate. Ishiyama (1958)

also considers that the primitiveness of the group, especially the species

Rhinoraja kujiensis, can be seen in the possession of scale on the ridge, and in the

simple construction of the dorsal terminal 1 cartilage. However, it should be

pointed out that the development of dermal denticles may be secondary,

e.g. Raja wallacer, R. leopardus, R. confundens, and that the position of the dorsal

terminal 1 in Rhinoraja kujiensis probably represents the case most removed from

Rhinobatos, if we accept Ishiyama’s theory of rotation of the dorsal terminal 1

from the ventral to the dorsal side of the clasper.

The genus Bathyraja represents a direct modification of the basic, ancestral

stock, in which, by the process of neotony, the rostral bar is reduced to a thin

rod. This may be related to an increased advantage in grubbing (Ishiyama

1958). The widespread, discontinuous, antitropical distribution of the genus

(Hulley 1970) supports its antiquity, as does the possession of comparatively

well-developed preorbital and postorbital processes in the neurocranium.

Rhinoraqja, with its segmented rostral bar, represents a further specialization of

the bathyrajid condition, which would allow for even greater flexibility of the

snout.

If the subgenus Raja is closely associated with the subgenus Duzpturus

(Stehmann 1970), then it must be accepted that the rhipidion was evolved

twice (Fig. 55). However, the persistence of the pseudorhipidion, i.e. retention

of a well-developed dorsal marginal cartilage, is evidence of an association of

the subgenus Raja with the bathyrajid condition. This is supported by the

simple form of the dorsal terminal 1 cartilage, with its proximal shelf for the

insertion of the m. dilatator, conditions which approximate those found in

Japanese bathyrajids (Ishiyama 1958: fig. 8). Furthermore, in the subgenus

Raja the axial cartilage retains its primitive, terminal point. The proximal

position and orientation of the dorsal terminal 2 can then be explained as a

continuation of the trend in development of the dorsal terminal 2 and 3 cartilages

from their position in Rhinobatos (Fig. 45) through some intermediate bathyrajid-

like form (Ishiyama 1958: figs 10 K, D). The subgenus Raja may therefore be

regarded as a side branch of the ancestral stock, close to the bathyrajid condition

(Fig. 57), in which the pseudosiphon was lost (possibly with complete dorsal

rotation of the dorsal terminal 1) and the ventral terminal more developed, but

in which the neotonous condition of the snout was never retained. Rather there

was a simple length reduction of the snout from its ancestral condition. In

support of this, the neurocranium of Raja shows a closer affinity with that of

Bathyraja, rather than with that of Dipturus. The preorbital process is well

developed in Raja and Bathyraja, while its size and position in Dipturus approxi-

mates the condition in Amblyraja, Rajella and Leucoraja. Furthermore, the position

of the foramen for the anterior cerebral vein in Raja approximates the bathyrajid

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

condition, while in Dzpturus, Amblyraja, Rajella and Leucoraja, it is displaced

dorsally.

The acceptance of this proposition would mean that the rhipidion was only

evolved once (Fig. 56), while the pseudosiphon, whose function is unknown,

was lost in two separate evolutionary lines. ‘This appears to be more acceptable

than Stehmann’s suggestion.

The subgenus Rostroraja appears to be associated with those groups in

which the ancestral pseudorhipidion is retained. As has been pointed out above,

it differs markedly from the subgenus Raa in axial shape, in the framework

arrangement of the dorsal terminal cartilages and in the heavy development of

the lateral edge of the ventral terminal cartilage. The possession of an elongate

snout (the anterior fontanelle of which is not primitive) and a discontinuous

distribution indicate a greater age for this group. This is supported by the Vprd

count, which for Raja alba is quite distinct from that of the subgenus Raja. The

higher count in Rostrorqa (Fig. 47) is indicative of its greater age, so that this

subgenus represents an early split from the Bathyraja/Raa line of evolution. The

extreme ventral position of the anterior cerebral vein foramen would indicate a

specialized condition, when compared with its median position in Raja and

Bathyraja.

Some criticism may be levelled against this theory in terms of the accessory

terminal cartilages, of which there are four in the subgenus Raja, two in

Rostroraqa, but only one in bathyrajids. However, a marked similarity can be

seen between the extension of the ventral marginal cartilage of bathyrajid

species and the tightly bonded accessory terminal 1 of Rostroraja and Raa.

DIPTURUS LEUCORAJA BATHYRAJA ROSTRORAJA

AMBLYRAJA RAJELLA RAJA

thipidion rhipidion thipidion rhipidion pseudorhipidion pseudorhipidion pseudorhipidion

No pseudosiphon _pseudosiphon pseudosiphon 7 pseudosiphon pseudosiphon no pseudosiphon _no pseudosiphon

HYPOTHETICAL ANCESTOR

pseudorhipidion

pseudosiphon

Fig. 56. Proposed relationship within the Rajidae, adapted from Fig. 57, showing

the possession of rhipidion, pseudorhipidion and pseudosiphon.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 83

Furthermore, the accessory terminals 3 and 4 might well be derived from

laterally positioned dorsal terminal elements.

On the evidence presented by their distribution (Hulley 1966: fig. 8), both

Raja and Rostroraja, which developed as shelf species from the boreal bathyrajid

ancestors, penetrated the Mediterranean Subregion at the beginning of the

Pleistocene glacial period (Klausewitz 1968). Possibly at this same time, a

complete north-south distribution on the continental shelf may have been

attained by Rostroraja, since equatorial cooling was pronounced along the

eastern shores of the Atlantic (Hubbs 1952). ‘The subsequent increase in sea

temperatures at the end of the last glacial would have given rise to (?) equatorial

submergence in the case of Raja alba (Hulley 1966), and would have induced a

southerly spread of the subgenus Raja. This would then support Crowson’s

(1970) and Parin’s (1970) ideas on the correlation between group age and

distribution.

All other rajid subgenera in the eastern Atlantic can be derived from an

ancestor, which possessed a pseudosiphon and long snout, and which had

developed a shield and fleshy rhipidion (Fig. 57). In all, the longitudinally

segmented pattern of the dorsal terminal cartilages, which form the framework

of the dorsal lobe, is retained. This ancestor probably inhabited the archibenthal

regions in boreal and antiboreal latitudes.

The subgenus Dzipturus represents an early split from this ancestor, as

indicated by the world-wide distribution of the group, in which the primitive,

rigid, rostral bar, with elongate anterior fontanelle and grooved rostrum was

retained, but in which the pseudosiphon was lost. The group apparently

colonized the edge of the shelf and upper regions of the archibenthal, a fact

which probably allowed for their subsequent spread to all oceans (Stehmann

1970). The low Vprd values in the southern African species of Dipturus seem to

CRURIRAJIDAE RAJI DAE ' ANACANTHOBATIDAE

Bathyraja

Cruriraja

Rostroraja

seo Sea er eo SO eS Soe SSS SoS eae Se6eccsss

Fig. 57. Phyletic interrelationships as suggested by southern African Rajidae.

The thickness of the lines corresponds to the relative number of species.

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

indicate that the group is not particularly ancient, but, as has been pointed out

above, in several Japanese species of this subgenus, the count is somewhat

higher (Ishiyama 1958). A strict interpretation of Vprd count to phylogeny

would therefore mean that the origin of the subgenus Dzpiturus must be sought in

areas other than the eastern North or eastern South Atlantic.

Examination of the clasper structure of Breviraja coles: (Ishiyama & Hubbs

1968) would seem to suggest that this genus can be closely associated with the

Dipturus group. The species of this side branch penetrated the abyssal regions of

the western central Atlantic, and, as in bathyrajids, retained the neotonous

condition of the snout as an increased advantage in grubbing.

Although Stehmann (1970) suggests an association of Amblyraja, Leucoraja

and Rajella with the bathyrajid condition, Rajidae II (Fig. 55), a closer affinity

with the subgenus Dzpturus is now evident. Not only are there similarities in the

structural arrangement of the cartilages and in the presence of a well-developed

rhipidion and strongly developed shield, but there are also marked resemblances

in the dorsal terminal 1, ventral terminal and accessory terminal 1 cartilages.

Similarly these subgenera share common neurocranial characters with Dzpiurus.

They do not have well-developed praeorbital processes and the position of the

anterior cerebral vein foramen is always dorsal to the optic foramen.

In terms of the pseudosiphon, it would appear that Amblyraja might be

considered to be ancestral to both Leucoraja and Rajella, and Stehmann (1970)

goes as far as to derive Rajella directly from Amblyraja. Counts of the number of

predorsal caudal vertebrae show a low value for Amblyraja (48-52) and a high

value for Rajella (55-73), suggesting that Rajella might be the more primitive.

However, the subgenus Amblyraja has a much wider distribution in the Atlantic

than either of the other two subgenera, and, in terms of the evolution of the

clasper, particularly the retention of the pseudosiphon, may be considered to be

an early split from the Leucoraja/Rayjella line.

The phylogenetic position of Raja spinacidermis, as based on Vprd count, is

not clear, although comparatively high values indicate an age equivalent to

that of the subgenus Amblyraja, with the possibility of an even greater age. It

must be emphasized that the validity of Malacoraja as a separate subgenus rests

solely on its peculiar spination pattern, rostral length, nasal capsule orientation

and depth distribution, the taxonomic values of which are uncertain. But until

examination of the clasper structure can be made, Stehmann’s (1970) proposals

must be accepted.

The claspers of the genus Cruriraja, which possess a shield and pitted

rhipidion, but lack an external pseudosiphon, bear a marked resemblance to

those proposed for the ancestral Dipturus stock. However, there are major

differences, especially in the arrangement of the dorsal terminals. The com-

paratively high values of the Vprd count in both Cruriraja parcomaculata and

C. triangularis would seem to indicate an early split from the ancestral to the

crurirajid condition (Fig. 57). The structure of the clasper of the mid-western

Atlantic species Cruriraja rugosa appears to be much simpler than that of the

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 85

southern African species, particularly as regards the accessory terminals. While

noting that a simpler structure may not necessarily mean more ancestral

(Mayr 1969), the structure of the shield in Cruriraja rugosa approximates more

closely to the ancestral condition. Furthermore, the Vprd count for Cruriraja

rugosa, which is based on a single specimen only, is slightly higher than that of

the southern African species. However, before theories are advanced as to the

possible method of evolution, i.e. west to east migration (cf. Clinidae: Springer

1970), the archibenthal and abyssal rajid fauna of the tropical western

Atlantic needs further examination.

The evolutionary position of the Anacanthobatidae is not evident at this

stage, for, as has been pointed out, on the basis of the clasper structure of

Anacanthobatis marmoratus and A. americanus, the group as defined as present

appears to be diphyletic. Anacanthobatis americanus, which is now considered to

belong to another genus, appears to be closer to the Dipturus ancestral stock and

its lack of a shield can be seen as the ultimate case in the general trend in

reduction of that structure, which is shown by Amblyraja, Leucoraja and Raella

species. The possession of similarly formed pelvic girdles but structurally different

clasper arrangements can therefore only be interpreted as an early origin for

the family (Fig. 57). This theory is tentatively advanced until examination of

further anacanthobatid material becomes possible.

In conclusion, it must be emphasized that, unlike the majority of South

African marine fishes, which appear to have their origin in the central Indo-

Pacific (Smith 1961), the Rajidae show a closer affinity with the eastern North

Atlantic. This is not surprising, since the tropical East African region

apparently forms a barrier zone, in which rajids have never been recorded

(Playfair & Giinther 1866; Peters 1868; Fourmanoir 1954; Smith & Smith 1963).

A warm-water barrier zone to temperate species has apparently been in

existence in this region since early Permian times, when a palaeo-equatorial

current impinged on the African—Arabian coastline (Frakes & Crowell 1970).

This is well before the proposed mid-Cretaceous origin of the Rajidae (White

1937). During the Pleistocene period, when major colonization of the southern

African shelf region was in progress (M. M. Smith 1970), this tropical barrier

was probably not broken down, as it was in the tropical eastern Atlantic

(Hubbs 1952). The following hypothesis is offered in support.

Biggs (1966), on the basis of endemism in fishes of oceanic islands, has

shown that Pleistocene temperature fluctuations have not been the same in all

oceans. Similar conclusions have been reached by Ericson et al. (1964) and

Emiliani (1970) on palaeontological evidence.

During the periods of glaciation, the Northern Hemisphere was charac-

terized by a southern spread of the polar ice. This had a marked effect in

lowering sea temperatures, especially where the thermal capacity of the ocean

was small and where the ice was in direct contact with the sea, e.g. North

Atlantic. This resulted in a southerly displacement of certain species. However,

in the Southern Hemisphere, glaciation was comparatively slight (Charles-

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

worth 1957). Coupled to this is the fact that the Indian Ocean is bounded to

the north by a continental mass, rather than an ice-cap, so that while extensive

glaciation may have occurred in the Himalayan region, this only had a

localized effect on the sea temperatures.

Therefore, because of the lack of a northern boundary ice-cap and because

the major portion of the Indian Ocean is situated south of the equator, it is

held that the tropical marine environment of the East African region did not

vary during the Pleistocene. This is supported by Cox (1927: 18), who states

for Mollusca that ‘there is no evidence for the invasion of the area by colder

water forms in Pleistocene times’. ‘The East African region has therefore always

formed an effective barrier zone to the migration of the Rajidae.

The southern African region has therefore been one of secondary coloniza-

tion rather than primary development, and the Rajidae can be envisaged as

arising from two sources:

(1) an older Gondwanaland distribution: these ancestral forms, i.e. Bathyraja,

Dipturus, probably had their origin in some other region. In the Southern

Hemisphere at least they were distributed in the area of the Cape sulcus

(Frakes & Crowell 1970), which was in existence until the end of the

Jurassic. This would mean that these forms arose from the rhinobatid

stock somewhat earlier than the proposed mid-Cretaceous origin for ‘Raja’

proposed by White (1937);

(2) a later north to south distribution, following the continental shelf and slope

regions: these subgenera (Raja, Amblyraja, Leucoraja, Rajella) probably had

their origin in the North Atlantic, and penetrated the southern African

region in a series of waves, the most recent dating from the end of the last

glacial period of the Pleistocene.

DIsTRIBUTION

Skates are widely distributed in all oceans, where they occur from polar

(Clark 1926; Nikolskii 1961; Andriashev 1966) to tropical latitudes (Fowler

1936; Bigelow & Schroeder 1962; Blache et al. 1970; Hulley 1972), but have

their greatest species diversity in warm temperate and boreal latitudes (Bigelow

& Schroeder 1953; Stehmann 1970). However, they have never been recorded

from Micronesia, Polynesia, Hawaii and the tropical region of East Africa,

around Zanzibar. Although found predominantly on the continental shelf and

upper regions of the slope, recent investigations have shown that skates are

present in archibenthal and abyssal regions (Garrick 1965; Forster 1965;

Hulley 1970). Besides brief descriptions given by Barnard (1925), Norman

(1935), Smith (1961) and Wallace (1967), no detailed account of the distri-

bution of southern African Rajidae has been given, although Hulley (1966,

1969) has commented on the distribution of Raja pullopunctata, R. clavata, R. alba

and R. miraletus.

The limits of the southern African region have been variously defined

(Barnard 1925; Knox 1960; Smith 1961; Day 1967; Penrith 1970) and appear

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 87

to depend largely on personal choice. For the purposes of this discussion, the

southern African region, as defined by Barnard (1925) has been used; that is,

the subcontinent south of the fifteenth parallel, extending from about

Mocamedes in southern Angola on the west coast, to north of the Zambezi

River mouth in Mocambique on the east, and taken down to a depth of

I 000 metres. The northern limits of this region on the west coast have been

fairly well covered by research cruises of the Mercator, Walther Herwig, Atlantide

and the Belgian South Atlantic Expedition, while the area north of Beira on the

east coast has been covered during Cruise 8 of R.V. Anton Bruun.

The distribution of the dominant elements of the intertidal flora and fauna

of South Africa has been summarized by Stephenson (1939, 1944, 1948), who,

on the basis of surveys of rocky intertidal shores, recognized three faunal

provinces with areas of overlap between them: the west coast, from the Orange

River mouth to Cape Point; the south coast, from Cape Agulhas to Port

Elizabeth; and the east coast, from Port St. Johns to northern Natal. In

discussing the distribution of polychaete worms, Day (1967) considers that the

west and south coasts are not distinct below the intertidal zone, and has

pointed out that bottom temperatures at 100 metres (12°—14°C) are uniform

from Port Elizabeth to Liideritzbucht. He recognizes four faunistic provinces

in southern Africa:

(1) the Mocgambique—Madagascar province, dominated by tropical species:

this reaches Lourengco Marques;

(2) the Natal province, with tropical species, but also with fair numbers of

endemics and Atlantic species; this reaches Bashee River;

(3) the Cape and South West African province, dominated by endemics, but

with a few tropical and several other components;

(4) the Angola province, dominated by tropical West African species: this

extends north of Cape Frio. Penrith & Kensley (1970, b) suggest that the

southern limit of this province may extend to between Liideritzbucht and

Walvis Bay, at least intertidally.

These faunal provinces allow for the greatest number of generalizations

and, since certain of the boundaries remain speculative, may be expanded or

contracted to suit the discussion of particular groups of organisms. Fishes,

however, are unsuitable subjects on which to base conclusions regarding faunal

distribution boundaries, owing to their mobility. In his discussion on the

zoogeography of the fishes of the Indian Ocean, Cohen (1971) postulates that

the 20°C isotherm is the most reliable parameter marking the boundary

between tropical and temperate waters, and limits the tropical region of the

east coast to Natal. Myers (1939) considers that the boundary of the tropical

Indo-Pacific is best defined by the distribution of the genus Scarus (Pisces,

Callyodontidae), which would mean that, as far as the ichthyofauna is con-

cerned, the boundary of the tropical region may be further south, in the region

of East London. Smith (1961) places the boundary of the Indo-West Pacific at

Great Kei River mouth.

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

On the basis of distribution beyond the limits of the southern African

region, the ichthyofauna has been divided into seven categories (Barnard 1925;

M. M. Smith 1970): deep and mid-water species; cosmopolitan and circum-

tropical species; Indo-Pacific species; Atlantic species, confined to the west

coast; austral species; eastern Atlantic and Mediterranean species, which pass

around Cape Point to various points on the east coast; and endemics. The

distribution of these categories is intimately connected with hydrographic

conditions (M. M. Smith 1970). |

The known latitudinal and depth distributions of the 22 species of Rajidae,

recorded from the southern African region, are shown in Figures 58, 59, while

the frequency of occurrence is given in Table 3. The sampling data does not

allow for comment on the relative abundance at various depths and hence the

variation in depth distribution with latitude.

Of the species, 13 are endemic, 8 are known from the eastern Atlantic, and

I species is also reported from Kerguelen. Of the endemics, Cruriraja durbanensis,

Raa dissimilis, R. ravidula and R. robertst are known only from the west coast;

Raa leopardus and R. caudaspinosa are probably confined to the west coast, but

have been recorded once from the east coast; Cruriraja triangularis, Raja steno-

rhynchus, R. springert and R. lanceorostrata are known only from the east coast;

while Cruriraja parcomaculata, Raja pullopunctata and R. wallacec have been

recorded both east and west of Cape Point. It is remarkable that no Indo-

Pacific species of Rajidae are known from the southern African region, and a

theory has been advanced to explain this phenomenon (see page 85).

Bottom temperatures, corrected to the nearest degree, at various depths

throughout the southern African region have been taken from Rand (1965),

Bang & Pearse (1970), Annual Reports of the Division of Sea Fisheries (1958,

1960, 1961, 1962, 1964) and unpublished station lists of R.V. Mezring Naude,

and are summarized in Table 4. For latitudes 17°S to 34°S on the west coast,

the range in bottom temperatures over a three-year period is given, while at

other positions, temperatures from a single cruise tract are shown. It should be

noted that the temperatures have mainly been taken from reversing bottles, so

that the temperature at sounding depth is not available; the discrepancy varies

from 5 metres inshore to 100 metres in depths over 300 metres. However, the

records are only used to show major trends in temperature variation.

TABLE 3

Frequency of occurrence of southern African Rajidae. Rare/little known = 2-6 records;

regularly present = 6-50 records; common = more than 50 records.

Single record only Rare/little known Regularly present Common

C’. durbanensis R. springeri R. pullopunctata R. alba

R. stenorhynchus R. spinacidermis R. confundens R. miraletus

R. lanceorostrata R. ravidula C. parcomaculata R. clavata

R. dissimilis R. doutrei C.. triangularis R. straelent

R. robertsi R. radiata R. caudaspinosa

B. smithii R. leopardus

R. wallacet

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI)

TABLE 4

Bottom temperatures, corrected to the nearest degree, in the southern African region.

WEST COAST (continued)

Latitude

g°S

17°S

19°S

21°S

23°S

25°S

26,5°S

29°S

30,5°S

32°S

WEST COAST

Depth Temperature

(metres)

150-199

200-299

300-800

I 000

50-99

100-149

150-199

200-299

300-800

I 000

FI 8

100-149

150-199

200-299

300-800

I 000

(°C)

Latitude Depth Temperature

(metres) (°C)

34S 50-99 g-10°

100-149 10°

150-199 —

200-299 8—10°

300-800 5-8°

I 000 3°

AGULHAS BANK

Latitude Depth ‘Temperature

(metres) (°C)

S 30 16°

Mossel Bay 80 10°

350 (a

I 000 5

S 50 10°

Knysna 100 10°

goo 4°

I 000 4°

S) 100 II-12°

Plettenberg Bay 200 12°

400 7

I 000 Aa

S 70 12°

Port Elizabeth 200 8°

800 5°

I 000 5°

50 Lo”

East London 400 Tels

600 hig

I 000 5°

EAST COAST

Latitude Depth Temperature

(metres) (°C)

33°59 70 TD

I 000 7

32°S 600 a

I 000 5-6°

1°S 50 Pah

: 800 oe

1 000 6°

29°S 150 16°

I 000 5

28°S 300 ie

400 g-12°

27°S 400 Ly

600 Q-

800 9°

I 000 4°

26°S 30 24°

400 10°

600 9°

2 000 2°

go ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

Classic zoogeography, which is concerned with the distribution of taxa in

relation to physical and chemical parameters, rests solely on two foundations:

the correct identification of the particular taxon on a world-wide basis, and

extensive sampling, especially towards the limits of ranges, where specimens

dwindle in numbers. Unfortunately, as far as rajids are concerned, both criteria

have not been fully met. The new approach to rajid systematics is still in its

infancy, and only the faunas of the western North Pacific (Ishiyama 1967),

eastern North Atlantic (Stehmann 1970) and eastern South Atlantic (Hulley

1970) have been extensively examined. Furthermore, in the southern African

region the majority of specimens have been taken by commercial trawlers,

fishing between 250 and goo metres, so that for the west coast at least there are

few inshore records.

In certain instances, confusion of specimens and their localities rather than

misidentification has taken place. This is obviously the case with Von Bonde &

Swart’s paper (1923), in which the type locality for Cruriraja durbanensis is given

as 30°10.00'S, 14°38.00’E in 859 metres, i.e. west coast, whereas the specific

name indicates east coast, while the type localities for Raja leopardus and

R. caudaspinosa are given as off Natal in 512 metres. Since the paratype of Raja

albalinea (= Raja caudaspinosa) was also taken at 512 metres, but at 32°3.00'S,

16°2.00’E, and since these two species have not been recorded from Natal by

Wallace (1967), it is considered that they are confined to the west coast.

While other factors such as salinity and bottom topography may play an

important role in determining the distribution patterns of fishes (Gilchrist

1905; M. M. Smith 1970; Zoutendyk, personal communication), temperature

has been considered the most determinate parameter (Ekman 1953; Hedgepeth

1957; Cohen 1971), and it is within this context that the distribution of southern

African Rajidae will be considered. The effect of temperature can be twofold

(Penrith 1970): it can act directly on the species concerned, or act in an indirect

manner in limiting other organisms, which make a particular habitat more

suitable. The ecology of the southern African Rajidae is still too little known to

estimate the relative importance of direct and indirect influence of temperature.

Although no fish family is endemic to the southern African region, the

degree of endemism at the species level in this region appears to be somewhat

higher than in the Mediterranean or Galapagos Islands (Tortonese 1963;

Rosenblatt & Walker 1963), and has been estimated at 31,5°% (Barnard 1925).

A lower value of 25,4°% has been estimated by M. M. Smith (1970: 5), but she

considers that ‘true endemics’ cannot be considered to come from deeper than

200 metres, as ‘it is merely a matter of time and opportunity before they are

found in other parts of the oceans’. There is no doubt, at least among elasmo-

branchs (Hulley 1971) that this is true, where the number of endemics is being

constantly reduced with further systematic work. However, the estimated values

approximate to that given by Day (1967) for polychaetes (36%). The percentage

endemism varies from family to family; in some the value may be as high as

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) gI

70% (Scyliorhinidae: Barnard 1925) and 100% (Clinidae: Penrith 1970).

Stehmann (1970) indicates that of 22 species of Rajidae represented in the

eastern North Atlantic 11 (50%) are endemic, and it would seem, therefore,

that the value of 59,1% for the southern African rajid fauna is reasonable.

There are no endemic genera or subgenera in the southern African region. All are

represented in the eastern North Atlantic. Not only this, but the specific

composition of the southern African rajid fauna shows a closer affinity with this

region (Hulley 1970).

Day (1967) and Penrith (1970) have pointed out that the Cape/South West

African province is dominated by endemics, and this is true of rajids, where

46,2% are confined to the west coast, while only 32,2°% are entirely east coast in

distribution. However, Cruriraja durbanensis, Raja stenorhynchus, R. lanceorostrata,

R. dissimilis and R. robertst are only known from their type localities. I consider

their distribution too imperfectly known to be discussed further here.

Only one species, Raja miraletus (Fig. 59), shows discontinuous distribution

in the southern African region, as defined. ‘This species has been recorded from

the Mediterranean (Clark 1926) and extends southwards along the north-west

African and West African coasts, being recorded from Morocco (Clark 1926),

Cape Bojador (Murray & Hjort 1912), Rio de Oro, Cape Blanc to Cape Verde

(Pellegrin 1914), Senegal (Cadenat 1950) and from Cape Lopez in Gabon to

Baie les Tigres in Angola (Fowler 1936; Poll 1951; Krefft 19682). The southern-

most record for the species at 18°30’S, 11°27’E was taken by FFS Walther

Herwig (Hulley 1970). It is absent from regions further south, but is recorded in

False Bay and extends northwards along the east coast to Richard’s Bay

(Wallace 1967).

While depth distribution records (Fig. 58) indicate that the species may

occur down to 440 metres, Poll (1951) points out that Raya miraletus is essentially

a shallow-water species, most abundant between 50 and 150 metres, where

temperatures vary between 24,1°C and 12,7°C. From Table 4 it can be seen

that for depths up to 200 metres on the west coast, a bottom temperature of

13°C or above is maintained as far south as about 19°S, but that from about 21°S

to 34°S, the minimum temperature at 200 metres is lower, although at shallower

depths may be between 12°C and 14°C as far south as about 23°S. This would

suggest that this dominant member of the tropical West African fauna has a

lower limiting temperature of about 13°C, and that its most southerly record

(18°30’S) might mark the boundary zone from tropical West African to cold

west coast fauna. .

From False Bay to Durban, Raja miraletus is commonly taken in depths less

than 100 metres, where temperatures vary seasonally (Bang, personal communi-

cation) between 9°C and 14°C. Although the present data allows for no

prediction of onshore and offshore seasonal migrations of this species over the

Agulhas Bank, it appears that the limiting temperature for Raja miraletus in this

region may be slightly lower than on the west coast, i.e. there may be physio-

logical differences between West African and South African specimens. It

Q2 ANNALS OF THE SOUTH AFRICAN MUSEUM

' '

| nes C triangularis

' '

’ i @ C.durbanensis

: '

mene € )rComaculata

‘ '

es Fullopunctata

as R. doutrei

R. lanceorostrata @

mms 59110921

: @ 8. stenorhynchus

een renee errr ereeeeee een eeeeeee eee eee cece eee ea R.clavata

ST ES SE R. straeleni

Seer Coeeee eer TU

inn |

R. robertsi@

' ‘

@ R.radiata

! eee FB dissinilis

! @ F.ravidula

annem R.confundens

eee R. caudaspinosa

Ea eee § leopards

R. Spinacid e777 S quam

Ro S10) (11) Sameer a TE

100 200 306 4 600 700 800 900 1000 1100 1200 1300 1400

DEPTH (metres)

een ee eo Or----- --

i—)

‘continental shelf break

Fig. 58. Depth distribution ranges of southern African Rajidae.

should be pointed out that there are some morphological! details (Hulley 1969)

which would substantiate the distinction of a South African subspecies.

Although the most northerly record on the east coast is at Richard’s Bay,

bottom temperatures indicate that the species could inhabit waters further to

the north (Table 4).

Two other species found in tropical West African waters also penetrate the

southern African region. Raja straeleni is common between 20°N and 15°S

(Blache et al. 1970). Poll (1951) points out that this species, which inhabits

colder waters than Raja miraletus, has regularly been taken in depths greater

than 200 metres in this region, but may be found in shallower depths in more

southerly latitudes. Temperatures at 200 metres in the tropical region vary

from 11,6°C to 15,75°C (Poll 1951). From Table 4 it would appear that for

depths greater than 200 metres, a lower limiting temperature of about 12°C

occurs in the region between 21°S and 23°S. The most southerly record of this

species at 22°03'S, 13°12’E (Fig. 59) would substantiate this temperature limit.

Again it can be seen that inshore temperatures remain higher from 23°S to

about 29°S, which would mean that the species may be expected to be found at

shallower depths in this region.

Raja doutrei, the other tropical West African species, has been taken off the

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) 93

coast of Senegal (Cadenat 1960) at a depth of 450-600 metres, where bottom

temperatures are in the order of 6°—10°C (Ingham 1970). This would mean that

Raja doutrei would be capable of extending as far south as Cape Point, although

its most southerly record to date is only 27°S.

Therefore, on the basis of rajid distribution alone, the southern boundary

of the tropical West African fauna cannot be accurately fixed, since the

temperature dependency of each species varies considerably. However, the

distribution pattern of the only inshore species, Raja miraletus, would support

the recognition of a boundary zone between 18°S and 20°S, as proposed by

Penrith & Kensley (1970a, 5) for rocky intertidal shores.

The phenomenon of bipolar (antitropical) distribution is well reported in

the Rajidae (Hubbs 1952; Bigelow & Schroeder 1953; Hulley 1966; Krefft

1968); Stehmann 1970), where certain species exhibit equatorial discontinuity

in latitudinal distribution. Four species in the southern African region show this

type of distribution pattern, which in this instance should be termed anti-

tropical rather than true bipolarity. Raja clavata and R. alba are known from the

eastern North Atlantic and Mediterranean (Clark 1926; Stehmann 1970), and

extend southwards to about 20°N (Cape Blanc). They are not recorded over

the tropical West African region, but reappear in the waters off South West

Africa, from where they extend around Cape Point to Barra Falsa in

Mocambique (Wallace 1967), southern Madagascar and Mauritius (Hulley

1966: fig. 8). In order to allow for gene flow from north to south and vice versa,

equatorial submergence of these species probably takes place, so that it is

possible that they will be discovered in deeper waters in tropical latitudes along

the West African coast. The records of both species east and west of Cape Point

(Fig. 59) support the idea of suitable, uniform temperatures for the west, south

and east coasts at depths greater than 100 metres. It should be noted that the

temperature range at 100 metres, 12°-14°C, given by Day (1967) appears to be

somewhat high, for although there are seasonal variations, the temperatures

east and west of Cape Point at 100 metres vary from 9°C to 10°C (Table 4).

It is interesting to note (Fig. 59) that, in the South Atlantic, Raja clavata

appears to have a northern limit at about 23°S, and is replaced by the closely

allied species, R. straeleni, over the area 22°S to 15°S, in the southern African

region. Direct temperature dependence seems an insufficient explanation for

this, and it is probable that competition between the species plays an important

role in limiting the spread of Raja clavata towards the end of its possible tempera-

ture range at that depth, since both species have a similar depth distribution

range (Fig. 58).

The other species exhibiting antitropical distribution patterns, Raja

Spinacidermis and R. radiata, also show an east/west distribution in the North

Atlantic (Stehmann 1970). In the southern African region, Raja radiata is

known from a single record (Fig. 59) and Raja spinacidermis only from deep

water off Cape Point (Hulley 1970), so that it is impossible to comment further

on their distribution.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

ee eee eee eee ee ee a

v R. straeleni

(ARE @ C. parcomaculata

See. Mocamedes ¥V C. triangularis @ R-ravidula

Ey O C.durbanensis

Vv R.dissimilis ;

Cape Frio 4

Ra Aro

! {

[\ Walvis Bay :

N \

Lt \\ y, L

on Lourengog a A |

ft! Luderitzbucht 1 \

alae — i t

| ;

A Pt Nolloth Durban JA 7 lcastoat \ / alannah T14

Hondeklip Bay Pt St f ! 5 | a Spst pasate 2

Lambe: i i y | [ [ ae \\ 7 + —t- Tt a

~ + ——

Bi: “ ast a | 14) ‘\s Aaa 1

: a 7 | | ea bed - | i ilk q

Pn ae ui | | 1 y | |

| il | i H = ti

| | | calm nae e.

| 0 (a 1 J setae m 30 | Chast i t an

oO » & jo 15 a Las

a areas ee er a

mm @ R. clavata

@ R.miraletus

Lett

‘ifaa 1

RD,

lea aly @.R.alba @ R. radiata

a Vv R.robertsi A i

(= E

)

aaa ct

SSE Ha

me) \ ia

LE EN eae LSI

Lae MOBS s —t+ aS

ECE | Ht

mane n f Saaes

see ee t—t—}-_}

i. FEY sanan

al + Hit ode + \ -

poi et He

Fig. 59. Distribution of Rajidae and 'Crurirajidae in the southern African region. Open symbols

indicate doubtful records.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 95

as oe ee es ee es I = D

@ R.pullopunctata

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B R.springeri ” a DE [Th

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ee | ET i

_SERRE =

JER

V B.smithii Ge

@ R. spinacidermis wee

@ R.confundens

Fig. 59. (continued)

LS OF THE SOUTH AFRICAN MUSEUM

@ R.ravidula

VY R.dissimilis

SOUTHERN AFRICAN RAJIDAE (cHonpRicuTHyEs, BATOIDE!)

@ R.clavata

y R. straeleni

@.R.alba

94 ANNA

WC. parcomaculata

¥C.triangularis

aos OC.durbanensis

Cn toe a

prea By

LI bras

Tele Leder yUraa

oe rina Seer’

Horde .

co arses Ene

CEL eee 9

L | eS

rl ppt ct

@ R.miraletus

fale

(OG

| ml

ig (ala olBt

1 30) | 35

@ R. radiata

WR.robertsi

AM © R. pullopunctata

~ ‘eR. wallacei -stenorhynchus

ry +H

SHOO0)

rH + Iniatslaes

[ 5 f Tite

: ibeee ce

ma 5 1s a

mere Se

a @ R.caudaspinosa A @ R. leopardus

ne LHe iE

| Sereeeey ie +t

aa | | it

+t 1 |

te: 4 te

Tet L i i

a a”) RGR at

HEH a ff Ci a ET

7 [ > | 7 = im

10s) wat

TT | i 4 H aa tf PH

moe ets H Ne ea

@ R.confundens

Fig. 59. Distribution of Rajidae and Crurirajidae in the southern African region. Open ®

indicate doubtful records.

ymbols

Y B smithii

@ R. spinacidermis

Fig. 59. (continued)

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

While the west/south/east coast distribution of the endemic species,

Cruriraja parcomaculata, Raja pullopunctata and R. wallace: (Fig. 59), may be

explained in terms of uniformity of temperatures at depths greater than 100

metres, the distribution of the east coast endemics, Cruriraja triangularis and

Raja springert (Fig. 59), and the west coast endemics, Raja caudaspinosa and

R. leopardus (Fig. 59), are more difficult to account for in terms of temperature

dependence. However, both Cruriraja triangularis and Raa springert are known

only from the northern limits of the east coast (Durban to Barra Falsa) in

230-420 metres and 400—740 metres respectively, where minimum temperatures

are higher than further south. Raja leopardus and R. caudaspinosa are apparently

confined to the west coast, west of Cape Point, despite the fact that their type

localities have been given as off Durban (Von Bonde & Swart 1923).

M. M. Smith (1970) reports an entirely west coast distribution for certain

South African fishes, although it appears that this is inexplicable in terms

of temperature dependence.

Raja confundens (Fig. 59) is not now regarded as a southern African endemic

species, and has been reported at 2°09’N, 9°27’E in 260-650 metres (Krefft

1968a; Hulley 1970). Because of this range, it is probable that the species will

be taken east of Cape Point in the future.

Bathyraqa smithi (Fig. 59) has been recorded from Kerguelen and from

the slope regions of the western edge of the Agulhas Bank, where its depth

distribution (Fig. 58) indicates an association with the Antarctic Intermediate

Water mass. This water mass is characterized by a salinity minimum core

(34,33%,) at 600-1 500 metres and temperatures of 4°-5°CQ (Orren 1963;

Shannon 1966), and is sandwiched between the warm Deep Water and Central

Water, which forms the source for the Benguela Current. Since the Antarctic

Intermediate Water extends northwards in the Indian Ocean at 200-300 metres

deeper than in the southern Atlantic (Orren 1963), it is probable that Bathyraja

smith is not confined to the western slope regions, and will be taken in depths of

800-1 800 metres off the Natal coast.

In conclusion, it must be pointed out that as far as the southern African

rajid fauna is concerned, distribution patterns are complex and do not follow

precisely the faunistic provinces as limited by Day (1967), although these

limits were based on distribution data only to 200 metres. There is no doubt

that tropical West African species extend into this region, but the southern

limit of this fauna is indistinct, although inshore species indicate some boundary

zone in the region between 18°S and 22°S. Furthermore, while several species

seem to be confined to the northern regions of the east coast and indicate

another boundary zone in the region south of Durban, i.e. a Natal province, the

majority of the species are widely distributed throughout the whole southern

African region. There is also some evidence that the direct influence of

temperature may not be the only limiting factor.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 97

SUMMARY

The structure of the pelvic girdle, neurocranium and the predorsal caudal

vertebral count are described for southern African Rajoidea, as are the

structures of the claspers of 17 species of Rajidae, Rhinobatos annulatus, Myliobatis

cervus and Anacanthobatis marmoratus, and the North American species Cruriraja

rugosa and Anacanthobatis americanus. A discussion of clasper terminology is

included.

On the basis of the above, the southern African Rajoidea have been

grouped into the following families, genera and subgenera:

Family Rajidae

Genus Bathyraja Ishiyama, 1968

species: Bathyraja smithu (Miller & Henle, 1841)

Genus Aaa Linnaeus, 1758

subgenus Aaja Linnaeus, 1758

species: R. miraletus Linnaeus, 1758; R. clavata Linnaeus,

1758; R. straelent Poll, 1951

subgenus Dzpturus Rafinesque, 1810

species: R. doutrei Cadenat, 1960; R. pullopunctata Smith,

1964; R. lanceorostrata Wallace, 1967; R. springer: Wallace,

1967; R. stenorhynchus Wallace, 1967

subgenus Rostroraja subgen. nov.

species: R. alba Lacépéde, 1803

subgenus Amblyraja Malm, 1877

species: R. radiata Donovan, 1808; R. roberts: Hulley, 1970

subgenus Leucoraja Malm, 1877

species: R. wallace: Hulley, 1970

subgenus Rajella Stehmann, 1970

species: R. caudaspinossa Von Bonde & Swart, 1923;

R. leopardus Von Bonde & Swart, 1923; R. confundens Hulley,

1970; R. dissimilis Hulley, 1970; R. ravidula Hulley, 1970

Family Crurirajidae fam. nov.

Genus Cruriraja Bigelow & Schroeder, 1948

species: C. durbanensis (Von Bonde & Swart, 1923); C. parcomaculata

(Von Bonde & Swart, 1923); C. triangularis Smith, 1964.

Family Anacanthobatidae

Genus Anacanthobatis Von Bonde & Swart, 1923

species: A. marmoratus Von Bonde & Swart, 1923

Genus Springeria Bigelow & Schroeder, 1951

species: S. ort Wallace, 1967

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

The subgenus Rostroraja is described as new and includes the single species,

Raja alba, while a new family, Crurirajidae, is defined to include all species of

the genus Cruriraja. The family Anacanthobatidae is diphyletic and, for the

moment, it is considered that the genus Anacanthobatis is monotypic. The North

American species Anacanthobatis americanus should be referred to a new genus.

The phyletic interrelationships of the suborder Rajoidea, based on these

morphological details, are discussed and an evolutionary pattern, different

from that of Stehmann (1970), is proposed. On this evidence, and on the

evidence presented by world-wide zoogeographical distribution, a theory as to

the origin of the southern African rajid fauna is advanced. ‘The effectiveness of a

warm-water barrier zone off the East African coast and its prevention of either

a northward migration of temperate rajid species or a southern invasion by

Indo-Pacific species appears to be significant and is discussed in relation to the

origin of the rajid fauna.

The distribution of the southern African Rajidae is discussed with particular

reference to temperature parameters, so far as the limited data will allow.

ACKNOWLEDGEMENTS

I am deeply indebted to Dr N. A. H. Millard, formerly of the Department

of Zoology, University of Cape Town, for helpful criticism and guidance

throughout this work, and to Mr S. X. Kannemeyer, of the South African

Museum, for his assistance during all phases; to Dr G. Krefft, Institut fiir

Seefischerei, Hamburg, for many suggestions and valuable criticism and for

much useful material; to Dr M. Stehmann, Institut fiir Seefischerei, for many

inspiring discussions and recommendations; and to Prof. R. Ishiyama, Tokyo

University of Fisheries, for his advice.

I am grateful to the late Prof. J. L. B. Smith and Mrs M. M. Smith,

J. L. B. Smith Institute of Ichthyology, Grahamstown; to Mr J. H. Wallace,

Oceanographic Research Institute, Durban; and to the Directors of the

British Museum (Natural History), the Institut fiir Seefischerei, the Institut

Royal des Sciences Naturelles de Belgique and the Muséum National d’Histoire

Naturelle, for access to their collections.

I wish to express my thanks to Mr Stewart S. Springer, formerly of the

National Museum of Natural History, Washington, not only for many X-ray

photographs, but also for the donation of specimens of Anacanthobatis americanus

and Cruriraqja rugosa; to Dr F. M. Mombeck, Institut fiir Seefischerei, for the

collection of the Walther Herwig specimens; and especially to Miss Christa

Liibben, Institut fiir Seefischerei, for photographs and X-ray plates of Raja

straelem and R. spinacidermis, and for her kind hospitality during my stay in

Hamburg.

My thanks are also due to the following individuals, either for placing

specimens in existing collections at my disposal or for help with X-ray photo-

graphy: Miss P. Verity, Nuffield Institute for Comparative Medicine; Drs P. H.

SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDE]I) 99

Greenwood, N. B. Marshall and Mr A. Wheeler, British Museum (Natural

History); Dr W. Templeman, Fisheries Research Board of Canada, St. Johns;

Dr J. P. Gosse, Institut Royal des Sciences Naturelles de Belgique, Brussels;

Dr M. Poll, Musée Royal d’Afrique Centrale, Tervuren; Dr M. L. Bauchot,

Muséum National d’Histoire Naturelle, Paris; Messrs M. J. Holden and

C. N. Humphries, Fisheries Laboratory, Lowestoft; and Drs H. Wormald and

R. Hindle, Durban.

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Notolepis rissot (Bonaparte, 1814) (Osteichthyes, Iniomi, Paralepididae). Arch. FischWiss.

1g: 103-113.

Ranp, R. W. 1965. Cruise 251 of R/S Africana IT during June/July, 1961. Station list. Investl Rep.

Div. Sea Fish. S. Afr. 51: 50-67.

Recan, C. T. 1906. A classification of the selachian fishes. Proc. zool. Soc. Lond. 1906: 722-758.

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Sea Fish. S. Afr. 58: 1-22.

Smiru, J. L. B. 1961. The sea fishes of southern Africa. 4th ed. Cape Town: C.N.A.

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with an account of the genus Cruriraja Bigelow & Schroeder, 1954 in South Africa. Vidensk.

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Ichthyology, Rhodes University.

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SOUTHERN AFRICAN RAJIDAE (CHONDRICHTHYES, BATOIDEI) 103

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Part III. Ann. Natal Mus. 11: 207-324.

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Nature, Lond. 165: 28-30.

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temperature. 7. Cons. perm. int. Explor. Mer 13: 71-75.

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mare Mediterraneo. Annali Mus. civ. Stor. nat. Giacomo Doria 74: 156-185.

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S.S. ‘Pickle’. Rep. Fish. mar. biol. Surv. Un. S. Afr. 3 (Spec. Rep. 5): 1-22.

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Wuite, E. G. 1937. Interrelationships of the elasmobranchs with a key to the order Galea.

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INSTRUCTIONS TO AUTHORS

Based on

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Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins, arranged in the following order:

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Figure captions and tables to be on separate sheets.

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Examples (note capitalization and punctuation)

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

FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

FiscHer, P.-H., Duva, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.

Archs ool. 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. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

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

Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature issued

by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).

The Harvard system of reference to be used in the synonymy lists, with the full references

incorporated in the list at the end of the article, and not given in contracted form in the synonymy

list.

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Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

P. A. Hulley

THE ORIGIN, INTERRELATIONSHIPS

AND DISTRIBUTION OF SOUTHERN

AFRICAN RAJIDAE (CHONDRICHTHYES,

BATOIDEI)

ANNALS

E SOUTH AFRICAN

OF THE S MUSEUM

CAPE “TOWN

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60 Band

November 1972 November

Part: 2 Deel

REDESCRIPTION OF PANDAKA SILVANA

(BARNARD) (PISCES: GOBIIDAE)

M. J. PENRITH & MARY-LOUISE PENRITH

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

REDESCRIPTION OF PANDAKA SILVANA (BARNARD)

(PISCES: GOBIIDAE)

M. J. PenrirH & Mary-Louise PENRITH

State Museum, Windhoek

(With 1 figure)

[MS. accepted 1 Fune 1972]

CONTENTS

PAGE

Introduction : : : a EOS

Description . : : : : 106

Discussion . : ; “ : 107

Summary . : : : ate

Acknowledgements : : : 108

References . : : : 2y; “tos

INTRODUCTION

During a check of the type collections of the South African Museum, the

syntypes of Gobius siluanus Barnard, described in 1943 from the upper reaches

of the Knysna estuary, were found. We discovered that this species had been

omitted from both Smith’s revisions of southern African gobies (1959, 1960)

and from Jubb’s (1967) account of the freshwater fishes of southern Africa.

On examination the specimens were found to agree closely with the genus

Pandaka Herre, 1927, described originally for two species from the Philippine

Islands. The species included in Pandaka are minute gobies with a low number of

dorsal and anal rays; about 20—22 rows of large ctenoid scales on the body;

the nape and head naked; a feeble pelvic fraenum; two rows of teeth in each

jaw, the inner row of the upper jaw minute, the inner row of the lower jaw

equal to or only very slightly smaller than the outer row; and the tongue

rounded. Gobius silvuanus agrees with Pandaka in all these respects as well as in

the general colour pattern, with the one exception that it has two predorsal

scales. Smith (1959) described Pandaka minuta from Ibo and Wamizi Islands in

Mocambique. This southern African species also differs from the Philippine

species of Pandaka in having predorsal scales, given as 3-4 in Smith’s descrip-

tion, although in the three paratypes of Pandaka minuta we have examined we

could find only two predorsal scale pockets.

The species Gobius silvuanus Barnard is therefore placed in the genus Pandaka

Herre. We have designated the female specimen illustrated (Fig. 1) as the

lectotype, the remainder of the syntype series being paralectotypes. Pandaka

stlvana is redescribed below. ‘The dorsal and anal ray counts differ from those

105

Ann. S. Afr. Mus. 60 (2), 1972: 105-108, 1 fig.

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

given in the original description because the last ray in each fin, which is split

to the base, is here counted as one element, while Barnard apparently counted

it as two. Unless otherwise stated, all lengths referred to are standard length.

DESCRIPTION

Pandaka silvana (Barnard, 1943)

Gobius silvanus Barnard, 1943: 258

Material

I specimen, female, 18 mm, lectotype; 22 specimens (14 males, 8 females),

13,3-18,0 mm, paralectotypes; all from Knysna Lagoon, Republic of South

Africa. All specimens in the South African Museum (S.A.M. 26208, lectotype,

S.A.M. 19355, paralectotypes), with the exception of a single stained para-

lectotype retained by the authors and in the collection of the State Museum,

Windhoek (S.M. P. 561).

Fic. 1. Pandaka silvana (Barnard), lectotype, female, 18,0 mm.

Description

D VI+I1 6-7. A I 6. P16. Dorsal fins well separated, first dorsal not

elevated in either sex. Ventral fins united, fraenum at base very poorly

developed. Caudal fin truncate. Branchiostegal rays: 6.

Maximum body depth 22-31 per cent of standard length. Body covered

with large ctenoid scales, 20-21 longitudinally, seven transversely at vent.

Nape, head, and breast naked.

Head 29-35 per cent of standard length, slightly depressed, naked; nape

naked except for two large predorsal scales. Snout rounded; lower jaw slightly

projecting. Mouth moderate, jaw reaching first third of eye. Eye large rounded,

8-11 per cent of standard length. Anterior and posterior nostrils tubular. The

arrangement on the head of pores and sensory papillae as far as could be

determined from the minute, rather poorly preserved specimens is shown in

Figure 1. A dentary pore, not able to be shown, is present at the lower angle

of the jaw. The main rows of papillae are: a row along lower edge of preopercle

to tip and continued along under lower jaw to chin; two horizontal rows on

REDESCRIPTION OF PANDAKA SILVANA (BARNARD) (PISCES: GOBIIDAE) 107

cheek from subocular pore, upper row of fewer and larger papillae than lower

row, which consists of numerous small papillae; a vertical row down opercle

behind preopercular flange, and a horizontal row running back across opercle

from lower third of vertical row; three short horizontal series between hind

margin of eye and upper posterior corner of opercle.

Teeth in two rows in each jaw, those of lower jaw almost equal in inner

and outer rows, the inner row of upper jaw minute, very much smaller than

outer row. Tongue rounded to subtruncate. Gill openings restricted, not

extending beyond pectoral base. No vomerine teeth.

Males with genital papilla narrow, acutely conical, and elongate; female

genital papilla short and broadly conical; several females ovigerous.

Colour

Faint darker marks along middle of flank; very faded owing to long

preservation.

DiIscussiION

Pandaka siluana was compared with three paratypes of Pandaka minuta

from Mogambique. P. silvana is apparently a larger species (all the ovigerous

females were over 16 mm, and most of the specimens were 16-18 mm standard

length, while none of the ten specimens of P. minuta exceeded 16 mm total

length). P. minuta has 22 scales in the lateral series (20-21 in P. silvana), and a

very dark mark at the pectoral base, apparently not present in P. svlvana,

although the latter species has a faint mark at the pectoral base which may

merely have faded. P. siluana has the teeth slightly smaller than P. minuta, and

the scales are more markedly striated in P. silvana. The genital papilla is

situated relatively slightly further back in P. minuta; in P. minuta the distance

genital papilla to caudal base is markedly less than the distance genital papilla

to eye, while in P. silvana these distances are about equal. P. silvana differs from

the Philippine species P. pusilla Herre and P. pygmaea Herre in the possession

of predorsal scales and further from P. pusilla in the lack of an elevated first

dorsal fin in the male.

Pandaka silvana is so far known only from the type locality. It appears

to be unique in the genus Pandaka as a temperate estuarine species, the other

species being tropical and marine. As the fresh waters of the Cape Province are

well known it does not seem likely that Pandaka silvana extends into the fresh-

water headwaters of the Knysna estuary.

SUMMARY

Gobius silvuanus Barnard, described in 1943 from the Knysna Lagoon,

and subsequently ignored in literature on South African gobies, is placed in

the genus Pandaka Herre, redescribed, and a lectotype selected from the syntypic

series and figured.

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

ACKNOWLEDGEMENTS

We are indebted to Dr T. H. Barry, Director of the South African Museum,

to Dr P. A. Hulley, South African Museum, and to Mrs M. M. Smith, Director

of the J. L. B. Smith Institute of Ichthyology, for lending us specimens for

examination, and to Dr D. Cohen of the Ichthyological Laboratory, Washing-

ton, for generous assistance with literature.

We acknowledge the permission of the Secretary for National Education

to publish this paper.

REFERENCES

BARNARD, K. H. 1943. Revision of the indigenous freshwater fishes of the S.W. Cape region.

Ann. S. Afr. Mus. 36: 101-262.

HeErrE, A. W. 1927. Gobies of the Philippines and the China Sea. Monogr. Bur. Sci. Manila 23:

I — 352.

Juss, R. A. 1967. Freshwater fishes of southern Africa. Cape ‘Town: Balkema.

Situ, J. L. B. 1959. Gobioid fishes of the families Gobiidae, Periophthalmidae, Trypauchenidae,

Taenioididae and Kraemeriidae of the western Indian ocean. Ichthyol. Bull. Rhodes. Univ

13: 185-225.

SmitH, J. L. B. 1960. Fishes of the family Gobiidae in South Africa. Ichthyol. Bull. Rhodes Univ.

18: 299-314.

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins, arranged in the following order:

(1) Heading, consisting of informative but brief title, name(s) of author(s), address(es) of

author(s), number of illustrations (figures, enumerated maps and tables) in the article.

(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-

headings to be used sparingly and enumeration of headings to be avoided. (4) Summary

(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

ILLUSTRATIONS

To be reducible to 12 cm X 18 cm (19 cm including caption). A metric scale to appear

with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their

proper place in the text), with arabic numbering; items of composite figures to be designated

by capital letters (A, B, C etc.).

REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

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

FiscHer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

~ 88: 100-140.

FiscHEr, P.-H., Duvat, M. & Rarry, 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. 1960. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

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

Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature issued

by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).

The Harvard system of reference to be used in the synonymy lists, with the full references

incorporated in the list at the end of the article, and not given in contracted form in the synonymy

list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, 6; Liste: 11. Turton, 1932: 80.

M. J. Penrith & Mary-Louise Penrith

REDESCRIPTION OF PANDAKA SILVANA

(BARNARD)

(PISCES: GOBIIDAE)

_” Se

- VOLUME 60 PART 3 NOVEMBER 1972

i...

{

ANNALS

i J]

t 4

OF THE SOUTH AFRICAN

CAPE TOWN

_~uyAiSs OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60 Band

November 1972 November

Part 3 Deel

mery MYDAIDAE (DIPTERA) FROM THE NAMIB

DESERT AND SOUTH-WESTERN AFRICA

A. J. HESSE

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na beskikbaarheid

van stof

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad

OUT OF PRINT/UIT DRUK

I, 2, 35 5s 7-8), Be. 5> tt Pale)5 5(1-2, 5> iS):

6(1, ESDuils)p 7(1-3); 8, g(1-2), 10(1),

ini (=P, 5, 7s Dela) 24(2), 27) SiN (hess) 33

Price of this part/Prys van hierdie deel

R6,30

Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum

1972

ISBN 0) 949940" 17 ‘3

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The Rustica Press, Pty., Ltd, Die Rustica-pers, Edms., Bpk.

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

NEW MYDAIDAE (DIPTERA) FROM THE NAMIB DESERT AND

SOUTH-WESTERN AFRICA

A. J. HEssE

South African Museum, Cape Town

(With 8 figures)

[MS. accepted 12 Fune 1972]

CONTENTS

PAGE

Introduction s : : , 109

Descriptions : : : 2 LEO

Summary . ‘ : : $2 70

Acknowledgements : ‘ ze 7O

iNeferences; >. 2 : : eT REG

INTRODUCTION

Since my revision (Hesse 1969) of the Mydaidae of southern Africa some

interesting new genera and species of this family, from various parts of South

West Africa and the Namib Desert, have been submitted to me for identification

and others have been very kindly presented to the South African Museum.

As South West Africa is faunistically one of the most interesting territories

in southern Africa, with its southern arid parts, its semi-desert east passing into

the Kalahari, its western coastal Namib Desert, its central grassland savannah

and broken thorn bush, and its north and north-eastern subtropical and tropical

enclave, the description of new and strange species from such different environ-

ments is a necessary obligation in a faunistic survey of the territory.

The mydaid fauna is comparatively rare anywhere in southern Africa,

if not in Africa, and the acquisition and descriptions of these various new forms

from this part of the subcontinent, as a sort of addendum to my revision of the

family, become necessary.

Up to date 12 species of Mydaidae have been described from South West

Africa. These, listed more or less in taxonomic order and with the dates of

description in brackets, are:

Eremohaplomydas desertorum Bequaert (1959)

Lachnocorynus kochi Hesse (1969)

Afroleptomydas pseudolanipes Bequaert (1963)

Afroleptomydas pulverulentus Hesse (1969)

Afroleptomydas pseudo-opacus Bequaert (1963)

Afroleptomydas (Crossoprosopus) kaokoensis Hesse (1969)

109

Ann. S. Afr. Mus. 60 (3), 1972: 109-171, 8 figs.

IIo ANNALS OF THE SOUTH AFRICAN MUSEUM

Afroleptomydas (Crossoprosopus) rudebecki (Bequaert) (1959 and 1969)

Namadytes vanson Hesse (1969)

Namadytes prozeskyt Hesse (1969)

Notosyllegomydas brincki (Bequaert) (1959 and 1969)

Nothomydas gariepinus Hesse (1969)

Cephalocerodes eremobius Hesse (1969)

In this paper 16 new species, including 3 new genera, are described. The

total number of known species from the territory is now 28, divided among

10 genera.

More organized and intensive collecting in this vast territory is bound

to increase this number of genera and species considerably in future.

DESCRIPTIONS

Subfamily Syllegomydainae

Tribe Syllegomydaini

Genus AFROLEPTOMYDAS Bequaert

Afroleptomydas lindneri n.sp.

A single ¢ specimen, among the Mydaidae from South West Africa sub-

mitted by Professor E. Lindner of the Staatliches Museum ftir Naturkunde in

Stuttgart, and labelled as “Gen. nov. pr. ? Nomoneura Bezzi’ is obviously a

new species of Afroleptomydas, which I have the pleasure in naming for Pro-

fessor Lindner. It has no relationship with Nomoneura, and is characterized as

follows:

Body with the vertex of head on each side infused with dark reddish brown;

antennae dark blackish brown, joint 1 also slightly infused reddish brown;

proboscis black, its base more brownish; the following parts pale reddish

brown: humeral tubercles, notopleural part, an abbreviated submedial streak

on each side of mesonotum ending before middle, postalar calli to an obscure

extent, anterior and posterior parts of mesopleuron, the pteropleuron, an

infusion on metapleural parts, area below and behind halteres, sides of tergite 1,

base laterally of 2 (the last three more yellowish), extreme sides of 2—5 to an

obscure extent, apical parts of lobes of tergite 9, sternites 8 and 9, hypopygial

structures (especially apical parts of processes of sternite 9, and the aedeagal

apparatus); venter also mainly reddish brown, paler near base; hind margins

of tergites 2-6 and very narrowly of 7 whitish, narrower across sides; bullae

reddish brown; hind margins of sternites on extreme sides also pallid; legs

mainly reddish brown, the femora darkened above.

Integument of central part of frons shining; that of antennae dull; proboscis

mainly shining; thorax above dull; pleurae subshining, with microscopic

areolar microsculpture; metanotum mainly dull, posteriorly subshining, also

NEW MYDAIDAE (DIPTERA) Loa

with indications of fine areolar microsculpture; abdomen above and below

more or less shining, more so towards apex, finely setiferously punctured, the

genital parts, including sternite 8 duller, finely microscopically microsculptured ;

integument of legs also dull, with very fine areolar microsculpture.

Vestiture with the hairs on head moderately long and dense, snow white,

those on each side just behind antennae and on clypeus the densest, directed

downwards, those on upper part of frons laterally, on vertex and upper part of

occiput directed more upwards, the occipital part fairly densely snow-white-

haired all round, the upper occipital part also with distinct and conspicuous

brownish postvertical spines; sides of frons and sides of occipital part behind

eye-margins with dense white tomentum; hairs on the following parts snow

white: humeral tubercles, dense ones along notopleural part, anteriorly on the

two submedial mesonotal streaks, across base of mesonctum, the finer shorter

and sparser ones on the submedial pale mesonotal streaks, very short and sparse

ones along the central mesonotal streak, longish ones across hind part of

mesopleuron, those on pteropleuron, metapleural part, on sides of metanotum,

dense ones on sides of tergite 1, as well as less dense longish ones on tergite I

discally, longish ones on sides in basal half of 2, much shorter and sparser ones

on extreme sides of 3, fine and short ones on genital segments, sparse and

longish ones medially on sternite 2, a tuft of longish hairs on each side of pro-

sternum, shorter ones basally and anteriorly on front coxae, sparse ones laterally

on middle coxae, sparse ones on metasternum, and a tuft of longish hairs

laterally and basally on hind coxae; short decumbent hairs in setiferous punc-

tures on tergites 2-5 dark or blackened, those discally and laterally on 6 and 7

gleaming more sericeous in certain lights; the short ones on extreme sides of

tergites 4-7 paler, more sericeous yellowish; short backwardly-directed hairs in

setiferous punctures on sides of venter also dark, those along middle appearing

paler, and those on sternites 7 and 8 distinctly more sericeous yellowish; hairs

on legs very short, seta-like, denser on femora above, sparser on sides and below,

appearing dark in certain lights and silvery gleaming in others, those on tibiae

even shorter, gleaming pale sericeous yellowish and whitish.

Head broader than thorax; eyes large, very convex; interocular space on

vertex comparatively narrow, only a little less than a third width of head

broader than space below head (as 16:11), the inner margins of eyes widest

apart opposite antennae where the distance is wider than interocular space,

the margins thus slightly converging to vertex and more so towards head below;

vertex itself distinctly sunk in; clypeus only slightly prominent; proboscis

(cf. Fig. 1, top left) about 2,48 mm long, the stem slightly broadened and

compressed to labella in side view, the labellar lobes ploughshare- or hoof-

shaped, sharply pointed apically, the heel-part rounded, striated transversely

and covered with fine spinules, the rest of stem very sparsely spinulated; palps

subequal in length to first antennal joint; antennae (cf. Fig. 1, top left) close

together, long, nearly or quite as long as mesonotum plus scutellum, joint 1

thickened, subspindle-shaped, about twice length of joint 2, the latter a little

le ANNALS OF THE SOUTH AFRICAN MUSEUM

broader than long, with a crown of short blackish brown hairs, joint 3 rod-like,

a little less than 4 times length of 1 and 2 combined, longer than club, not

demarcately thickened apically, obscurely transversely and shallowly ringed,

except in apical part, the club elongate, broadest at about middle, from there

to base gradually narrowing and with only a slight neck-like base, apicalwards

at first slightly constricted then again widening subapically, the apical sensory

part sloping up conically to tubercle.

Wings relatively broad, not projecting beyond abdomen, infuscated smoky

brownish in more than anterior half, more or less to end of second submarginal

cell, to basal part of first posterior cell at level of apex of discoidal cell, to

faintly along posterior vein of third posterior cell, to apical part of latter and

posteriorly and across to apex of anal cell, being darkest in middle part of

wings in greater parts of marginal, first submarginal and first basal cells and

along veins in hinder half; greater part of first posterior cell, the apical part,

broad hind border, broad axillary lobe and middle parts of anal, third posterior

and discoidal cells being clearer; veins brownish; second submarginal cell with

a well-developed appendix; first posterior cell much or sharply narrowed

apically, shortly stalked on costal margin; discoidal cell shortly stalked apically;

hind border of wings comparatively broad; axillary cell markedly broad, with

distinct flattened cilia along hind margin at broadest part and minute ones

from there to its apex; alula well developed, broad, lobe-like; halteres pale

yellowish brown, but more than posterior half of broadened apical part black.

Legs moderately long; hind femora much thickened, subspindle-shaped,

thickest a little beyond middle, armed below with a double row of strong,

backwardly-directed, blackish brown spines on tubercles, beginning a little

before middle, with about 7 in outer row and 7 in inner one (one of the hind

femora in the specimen is damaged and tibiae of both missing) ; front and middle

tibiae, apart from their apical spicules, with about 3-5 dark spicules along

posterior lower aspect and a few much shorter ones along anterior upper part,

the middle ones also with 3—5 longer ones along anterior ventral part; anterior

tibiae below with a ventral streak of dense, brush-like, short hairs beginning

near base and more evident in apical half; front and middle tarsi about, or a

little less than, half length of corresponding tibiae; claws well developed,

yellowish brown, darker at bent-down apices; pulvilli well developed.

Hypopygium (cf. Fig. 1, bottom left) with the lateral lobes of tergite 9

angularly pointed; sternite 9 roundly prominent posteriorly, slightly laterally

compressed apically, its processes in side view gradually narrowed apically

and the apices directed upwards, in dorsal view strap-like, slightly narrowed to

blunt and rounded apex; aedeagal apparatus rather elongate, the anterior

or dorsal epimere long, extending much beyond the two phallic tubes which

are also comparatively long.

From the ¢ holotype in the Stuttgart Museum.

Length of body: about 14,5 mm

Length of wing: about 11,5 mm

NEW MYDAIDAE (DIPTERA) 113

Distribution

South West Africa: Guinas (? 29/9/54) (No regional locality is given and

no collector’s name). From the handwriting the collector probably was Herr

I’. Gaerdes of Okahandja.

This species is to be placed in the westermanni-section of Afroleptomydas

in which the hind femora are distinctly thickened or incrassate and the wings

in a number of $¢ are infuscated. Characters such as the presence of delimited

bare streaks on mesonotum, white hairs on mesonotum, the short dark hairs on

the abdomen above, the type of wing-infuscation, the apically-narrowed and

stalked first posterior cell, the relatively dark legs, etc., however exclude it from

any of the subsidiary sections of the westermanni-group of species.

CROSSOPROSOPUS Hesse subgenus of AFROLEPTOMYDAS

All the other species of Afroleptomydas, collected in South West Africa and

submitted to me for identification and described in this paper, belong to the

subgenus Crossoprosopus, defined by me in 1969. This subgenus is apparently

well represented in the territory.

Afroleptomydas (Crossoprosopus) angolensis n.sp.

Three $4 from Angola, collected by Dr Brown, though not from South

West Africa, are described below because the locality from which they come

is just north of the Kunene River and similar environmentally to that of

north-western South West Africa in which this species might also occur. ‘This

new species is very near tulzensis Hesse from Rhodesia and is characterized as

follows:

Body on the whole much darkened above, dark brown to blackish brown;

antennal joints 1 and 2, about basal 3 of joint 3, and greater part of club, except

black base and sometimes the slightly darkened subapical part of latter, orange

yellowish; clypeus yellowish; head below pale yellowish brown; proboscis

black, pale yellowish brown below; humeral tubercles, sides of thorax above,

postalar calli, medial basal part of mesonotum in front of scutellum, scutellum,

sides of metanotum to a variable extent, pteropleural and metapleural parts

in front of halteres to a variable extent, and hypopleural part pale yellowish

brown; sides and also broad hind margin of tergite 1 also pale yellowish brown,

the middle of the hind margin inclining to be more yellowish; abdomen brown

to dark brown or even blackish brown; hind margins of tergites 2-7 ivory

yellowish or yellowish white, becoming progressively narrower, the last 2

being much narrower than the very broad ones across tergites 2-4, the sides

of tergites sometimes more yellowish brown or brown to a variable extent,

especially on 2-5, contrasting with the darker discal dorsum which latter tends

to appear darker, more patch-like basally dorsally; bullae rather large, bean-

shaped, the broad anterior border mainly shining black, the median anterior

discal part just behind the black anterior border tending to be orange brownish

Ii4. ANNALS OF THE SOUTH AFRICAN MUSEUM

to a variable extent, the space between bullae much broader than transverse

length of bullae; venter either mainly dark or to a variable extent yellowish

brown, especially on sternites 1-5 (or 6), the hind margins of sternites 1-6

however broadly whitish, especially on sides; genitalia mainly yellowish brown,

sometimes dark reddish brown; legs yellowish brown to reddish brown, the

femora above darker brownish, the bases of hind ones more yellowish, the

knees and bases of tibiae also paler, more yellowish, more than basal halves

of hind tibiae above, especially when viewed from behind, also more yellowish,

only the apical part appearing darker, the front and middle coxae dark

brownish, the hind ones paler, more yellowish, the tarsi more yellowish, the

apices of the joints below darkened, and claw joint also blackish towards apex.

Integument of central raised part of frons, clypeus, and head below smooth

and shining; that of thorax above dull, leathery; metanotum dull, leathery;

greater part of pleurae dull, the outer part of anterior spiracle, and sutural parts,

anterior part of pteropleuron, and sclerites below wing-bases however shining;

abdomen above mainly dull, somewhat shining across extreme bases on sides

of tergites 3—5, with fine, separated, not very dense, setiferous puncturation,

slightly denser on sides of tergites 2 and 3 and absent from ring-like hind margins

of tergites, those on 6 and 7 more ‘nadelrissig’; venter also with separated

setiferous puncturation, absent from broad transverse basal parts of sternites

and from broadish shining hind margins, and also from extreme sides of ster-

nites, the last two sternites with indications of transverse striae or rugae,

especially on sides; legs mainly dull.

Vestiture not very long; that on vertex and head in front dense and white,

shorter and less dense on occiput, with dense, greyish white tomentum on sides

of frons and on occiput; hairs on thorax above mainly white, those discally

sometimes with a slight sericeous yellowish tint, dense and longer on sides and

medially basally in front of scutellum, the short decumbent ones on disc

arranged in more or less 2 broadish submedial streaks and a narrow, linear,

central one along the middle, all these longitudinal streaks, as well as hairy

sides, separated by more or less bare streaks, the hairy streaks being on streaks

of tomentum; hairs on metanotum also white, the integument of metanotum

itself with slight greyish tomentum; pleurae with fine greyish, not very dense,

tomentum, the posterior margin of mesopleuron, the pteropleuron, metapleural

part in front of spiracle, and metasternal part with white hairs, not much

longer than those on sides of thorax above; tergite 1 also with dense, longish,

white hairs on sides; base of tergite 2 especially sides, with similar longish white

hairs; extreme base laterally of tergite 3 also with some long white hairs;

fine, short, backwardly-directed hairs in punctures on dorsum of abdomen

gleaming slightly sericeous yellowish, the integument of tergites however covered

with greyish white to faintly yellowish white tomentum; venter with some

longish white hairs basally on sternite 2; the backwardly decumbent ones on

greater part of venter slightly longer than short ones on dorsum and also with a

slight sericeous yellowish tint; hairs on hypopygium not very dense, sericeous

NEW MYDAIDAE (DIPTERA) 115

white to faintly sericeous yellowish; hairs on legs comparatively short and not

dense, mainly gleaming sericeous white, denser and longer on coxae, the

femora below bare; spines on hind femora below and spicules on tibiae whitish,

those on tarsi tending to be more faintly yellowish white to yellowish.

Head broader than thorax; vertex sunk in; interocular space on vertex

about as wide as distance of central frontal ridge between vertex and base of

antennae, wider than space below head; yellowish white postvertical spines

present; clypeus convex; antennae longer than mesonotum, about 3,9—4,1 mm,

joint 1 slightly thickened, about twice as long as 2; joint 3 elongate, about

3,5-3,7 times combined length of 1 and 2, longer than club, with a little less

than its apical fourth thickened; club itself (excluding articulation between it

and joint 3) distinctly shorter than joint 3, elongate pyriform, broadest at about

apical fourth, beyond that sloping up or narrowing to the transverse crater-like

terminal tubercle, the base of club constricted neck-like; proboscis about 2,8 mm

long, only a little longer than vertical length of eye; palps small, but distinct.

Wings greyish hyaline, with a very faint indication of faint yellowish

borders to the veins in apical part beyond apex of first basal cell; veins near

base and costal vein yellowish, in rest of wings brown or dark brown; first

posterior cell broadly open on costal vein; second submarginal cell with or

without a short appendix at base; halteres yellowish white, the knobs infused

with brown above to a variable extent.

Legs moderately long; hind femora thickened, spindle-shaped, with a

double row of pallid spines (7-10 in both rows) on tubercles below, beginning

a little distance away from base; front and middle tibiae slightly curved,

without any distinct spicules (excepting apical spurs) on front ones, but with

2 rows of widely separated pallid spicules on middle tibiae; hind ones with

granules below and with an outer row of pallid spicules on granules in apical

half; basal joint of hind tarsi subequal to or only a little longer than claw-joint;

pulvilli just falling short of apices of claws.

Aypopygium (cf. Fig. 2, top left) with the lobes of tergite g slightly angularly

produced apically; sternite 9 conical, medially incised apically, its processes in

side view slightly S-curved, in dorsal and ventral views hollowed, narrowed

towards apex and bluntly pointed; aedeagal apparatus with the dorsal epimere

longer than the phallic tubes.

From a ¢ holotype and 2 ¢ paratypes in the South African Museum.

Length of body: about 19-20,5 mm

Length of wing: about 12,5-14 mm

Distribution

Angola: Mocamedes District: 42 km south of Sao Nicolau (H. D. Brown,

16/4/1971).

From the Rhodesian species tuliensis Hesse, nearest to it, it differs in being

slightly larger, with comparatively longer, more slender abdomen, distinctly

longer antennae, joint 3 and club being proportionally longer, longer labella

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

of proboscis, less dense, sparser and slightly shorter vestiture, relatively broader

anal cell, its posterior vein more sinuous, distinctly longer slender part (side

view) of processes of sternite 9, their apices blunter in dorsal view, and larger

sternite Q.

Afroleptomydas (Crossoprosopus) ovamboensis n.sp.

A somewhat damaged rather small 3 specimen from Ovamboland in the

collections before me, and belonging to a new species, is characterized as

follows:

Body mainly dark; head, including clypeus, black; proboscis very dark

blackish brown; antennae dark yellowish brown, the first joint, thickened

apical part of joint 3, base and apex of club darker, more black, the broad

middle part of club orange yellowish; humeral tubercles yellow; thorax above,

scutellum, pleurae and metanotum black; sclerites below wing-bases brownish

yellow; abdomen mainly brownish, the dorsal discal basal parts of the tergites

darker, more blackish brown, more so on tergites 1-6, the sides basally of

tergites 1-3 distinctly more yellow and to a lesser extent also those of 4-7, the

hind margins of 2-8 broadly pale yellowish white, those of 7 and 8 narrower,

and that of tergite 1 the narrowest; venter more yellowish brown, the hind

margins of sternites 1-7 also broadly pale yellowish white; bullae shining black,

widely separated; hypopygium pale yellowish brown, the anal lobes more

yellowish; legs mainly yellowish, the coxae dark or black, more or less apical

half of hind femora darker above, more brown, and claw-joint of tarsi also

more brown.

Integument of medial part of frons, clypeus, head below and proboscis

shining; that of thorax above dull, leathery; scutellum subshining; metanotum

dull, with indications of some transverse grooves on sides; pleurae mainly dull,

the propleural part and sutural part of sternopleuron somewhat shining;

abdomen discally on dark discal patches dull, the sides of basal segments and

sides of rest of tergites up to 6 more or less shining, the tergites with setiferous

puncturation and tending to be striated on sides towards hinder parts; processes

of sternite g and aedeagus shining; venter shining, with sternites 7 and 8

somewhat transversely striate; legs subshining, with fine setiferous puncturation

lodging the hairs, the outer bases of front and middle femora, and to a finer

and lesser extent the tibiae transversely striate, the granulation on hind tibiae

below rather feebly developed and not dense.

Vestiture fairly dense, but not very long, mainly snow or sericeous white;

that on head in front dense, snow white; that on head below sparser; that on

sides of thorax above, postalar calli, and basally in front of scutellum dense

and snow white; the backwardly-decumbent, shorter ones on disc gleaming

more sericeous yellowish, arranged more or less in streaks, a narrow central

streak and 2 broader submedial ones, with a bare sublateral streak on each side

which becomes entirely bare in hinder half; hairs on metanotum denser on

sides; pleurae mainly bare, with fine greyish tomentum, with very sparse white

NEW MYDAIDAE (DIPTERA) 117

hairs across hind margin of mesopleuron, sparse on pteropleuron, and denser

in front of halteres; abdomen with dense, longish, white hairs on tergites 1 and 2,

especially on sides, some longish ones also on sides basally of 3 and 4, with the

other short, decumbent hairs in fine punctures on tergites gleaming slightly

sericeous yellowish in certain lights, sparser on tergite 7, also comparatively

sparse on hypopygium; hairs on venter slightly longer than on dorsum, mainly

white, longer on sternites 1-4; hairs on legs gleaming mainly sericeous white,

not very long, except on coxae where they are also denser, those on front and

middle femora above also longer and denser than on rest of legs, slightly sparser

and shorter on hind femora above, the shorter ones on all the tibiae gleaming

slightly more sericeous yellowish; spines on hind femora below and spicules on

tibiae and tarsi pale yellowish white; tomentum on sides of head in front greyish

white, rather narrowly confined along eye-margins, that on occiput also greyish

white, more extensive; fine greyish tomentum on disc of thorax more or less

confined to hair streaks; that on metanotum slightly denser than that on entire

pleurae; the fine greyish tomentum on tergites 1-6 visible in certain lights,

even on black basal discal patches on 3-6.

Head much broader than thorax; vertex sunk in; no distinct postvertical

spines detectable; interocular space on vertex only a little broader than space

on head below; antennae subequal in length to mesonotum plus scutellum,

joint 1 only slightly thickened, about 24 times length of 2, and with only a few

hairs, joint 3 nearly or about 3 times length of 1 and 2 combined, thickened in a

little less than apical 4, the club elongate pyriform, slightly longer than joint 3,

narrowed neck-like at base, broadest at about or at a little less than apical 4,

its apical sensory area more rapidly narrowed on inner than outer part to

crater-like prominence; proboscis short, only a little longer than club of

antennae, very much shorter than vertical length of eyes, only about 1,24 mm

long; palps small, but distinct, with yellowish hairs apically; buccal cavity

below projecting rather prominently ledge-like.

Wings comparatively short, clear hyaline, with a faint whitish tint in certain

lights; veins mainly yellowish, becoming more yellowish brown in apical and

hinder parts of wings; first posterior cell narrowly opening (or sessile) on costal

margin; second submarginal cell with a short appendix; discoidal cell stalked

apically; halteres whitish.

Legs with the front and middle femora unarmed below; hind femora

moderately thickened, spindle-shaped, armed below with a double row of

spines on tubercles, about 8 in outer row and g in inner one, the latter beginning

before the outer one from near base; front and middle tibiae slightly curved,

the pale spicules on middle ones more developed, longer; hind tibiae with

distinct spicules on granules, only present along a little more than outer apical

third, the rest in the row minute or small, the granules below the hind tibiae

on the whole poorly developed; basal joint of hind tarsi subequal in length to

claw-joint.

Hypopygium (cf. Fig. 2 top middle) with the lateral lobes of tergite g rather

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

angularly pointed; sternite g bluntly rounded apically, medially dorsally and

apically with the usual incision, the dorsal processes of the sternite rather long,

in side view rather slender, slightly upcurved, in dorsal view slightly curved

outwards in apical part and sharply pointed apically; aedeagal apparatus with

the central and dorsal shoe-horn-shaped epimere rather long, projecting much

beyond the two phallic tubes.

From a single ¢ holotype in the South African Museum.

Length of body: about 13,5 mm

Length of wing: about 8 mm

Distribution

South West Africa: Ovamboland: 35 km north-west of Oshakati (H. D.

Brown, 21/4/1970).

Superficially this species resembles the 3 of vansoni Hesse in the nature of

the vestiture, short proboscis and yellow legs, but differs in its entirely dark

clypeus, longer and less thickened antennal clubs, more slender legs, a first

posterior cell which opens more narrowly on costal margin, comparatively

longer processes of sternite 9g, and distinctly longer epimere of aedeagal

apparatus.

From 36 of certain forms or varieties of mauricezx Beq. 1t may at once be

distinguished by its longer hairs, denser hairs on head in front, slightly longer

hairs on rest of body and legs, comparatively much shorter antennae, very

much shorter proboscis, darker clypeus, paler legs, comparatively longer and

more slender processes of sternite 9, and comparatively longer and more

slender epimere.

Afroleptomydas (Crossoprosopus) nigrescens n.sp.

An almost entirely black-bodied ¢ and a 9, which I take to be the same

species, in the collections from South West Africa before me, represents a new

species which is near maurice: Beq.

The 3 specimen is chacterized as follows:

Body almost entirely dark blackish brown to black; clypeus reddish brown;

antennae brown, joints 1 and 2 dark blackish brown, greater part of clubs,

excepting brownish, neck-like, basal part and apical sensory area, orange

yellow; proboscis reddish brown, its labella slightly darker; humeral tubercles

pale yellowish and area around them more reddish brown; rest of thorax,

excepting dark brownish postalar calli and sclerites below wing-bases, entirely

dark; abdomen very dark blackish brown to black, the lateral part of tergite 1

reddish brown and its hind margin more dark reddish; broad, ring-like hind

margins of tergites 2-8 ivory whitish (those of 6-8 narrower); hind margins of

sternites 2—5 also broadly ivory whitish, that of sternite 6 much narrower and

those of 7 and 8 very narrow and scarcely differently coloured from the dark

colour of those sternites; apical part of lateral lobes of tergite 9, apical part of

sternite 9, processes of latter, and aedeagal apparatus slightly paler reddish

NEW MYDAIDAE (DIPTERA) 119

brown; legs with the coxae dark, the base of middle ones yellowish brown, the

membranous articulations between front and hind coxae and their respective

sternal parts more yellowish, the legs themselves brownish, the hind femora

in more than apical half laterally and below darkened blackish brown, and

front and middle ones along outer or hinder aspect also darkened; all the

tibiae appearing paler yellowish brown, the last 3 joints of front and middle

tarsi, and last 2 of hind tarsi tending to be dark blackish brown, and apical

curved part of claws black.

Integument of head in front, first and second antennal joints, proboscis,

and rest of body mainly shining; propleural and prosternal parts, hinder half

of mesopleuron, the pteropleuron, metapleural part however dulled by greyish

white tomentum; a narrow central streak, a broader submedial streak on each

side of mesonotum, a lateral notopleural streak, and a medial patch in front

of scutellum also dulled by greyish tomentum; abdomen with tergite 1 markedly

transversely convex, the rest with saddle-shaped dull patches discally above on

more than basal halves of tergites 2-6, appearing dull black in certain lights,

but covered with fine greyish white tomentum in other lights; integument of

tergites with comparatively widely-spaced setiferous puncturation lodging the

hairs, denser on tergites 1 and 2; sides of tergites 2-7 also showing indications of

transverse striation in certain lights; bullae large, bean-shaped, shining black,

separated by a space slightly narrower than transverse length of a bulla, their

anterior margin marked off as a transverse row of coarse punctures, and

anterior part of the disc of the bullae behind the anterior ridge with a row of

pit-like punctures; legs subshining, with fine, slightly raised, setiferous punc-

turation lodging the hairs, the femora, especially hind ones and near the apex,

with indications of transverse striae, and the tibiae too, especially hind ones,

with indications of transverse striation.

Vestiture mainly snow white or silvery white, not very long, the longest and

densest hairs in tufts on head in front, on vertex, upper occipital region, on

humeral tubercles, sides of thorax above, in a patch in front of scutellum, on

metanotum, especially sides, as sparse ones across hind margin of mesopleuron,

and slightly denser ones on pteropleuron and metapleuron; hairs on sides of

tergites 1 and 2 also long and dense, and also with a few sparse longish ones

basally on sides of 3; hairs on disc of mesonotum arranged in the usual narrow

central streak and 2 broader submedial ones, these hairs short and backwardly-

decumbent and, excepting the longer white ones anteriorly, with a very faint

sericeous yellowish tint; short decumbent hairs on abdomen above rather

widely spaced, slightly denser discally on tergite 2 and to a lesser extent on 3,

all with a very slight sericeous yellowish tint; hairs on venter more or less

confined to middle part of sternites 2-6, also widely spaced and slightly gleam-

ing sericeous yellowish, but slightly longer than on dorsum; hairs on sternites 7

and 8 like those dorsally; hairs on hypopygium equally widely spaced and also

tinted sericeous yellowish; hairs on coxae dense and silvery white, those on

front and middle femora above and behind dull white, longer and denser than

I20 ANNALS OF THE SOUTH AFRICAN MUSEUM

those on hind femora, and those on tibiae short, dense and gleaming very

slightly sericeous yellowish; tomentum behind eyes greyish white, that on

thorax and abdomen as stated above, but that on tergites 6,7 and sides of 8

not confined to saddle-shaped discal patches, but also faintly evident laterally;

tomentum on hypopygial parts also faintly indicated.

Head much broader than thorax; interocular space on vertex subequal

in width to the interocular space on head below, the inner margins of eyes

converging above and below; some distinct postvertical spines detectable;

clypeus comparatively short centrally, tending to be slightly longitudinally

convex; proboscis comparatively long, about 2,8 mm long fully extended,

much longer than vertical length of eyes; palps distinct, slender; antennae

comparatively long, much longer than mesonotum, with joint 1 not much

thickened, about 2} times length of 2, joint 3 the longest, slightly more than 3

times length of 1 and 2 combined, a little less than its apical $ slightly thickened;

clubs elongate pyriform, shorter than joint 3, broadest at about apical third,

their bases (a little less than basal +) markedly and sharply demarcated neck-like,

with the conical apical sensory area also sharply demarcated by an almost

encircling ridge, interrupted dorsally or laterodorsally where club passes

apically into the vertical, slightly elongated, crater-like, apical prominence.

Wings greyish hyaline, with a very faint whitish tint in certain lights;

veins yellowish brown; second submarginal cell with a short appendix; first

posterior cell broadly opening on costal margin; hind border of wings much

narrower than either first and third posterior cells; halteres pallid, the knobs

darkened above.

Legs with the front and middle femora unarmed below; hind femora

markedly thickened, subspindle-shaped, armed below with a double row of

pallid spines on tubercles, about 5—7 in the inner row and 6 in the outer one

which begins a little before the inner one; front and middle tibiae slightly

curved, the spicules on front ones shorter than those on middle ones; hind

tibiae also slightly curved beyond middle, the granules below distinct, those

in outer row below with stoutish spine-like spicules in apical half, the rest of

spicules on hind tibiae finer and shorter; basal joint of hind tarsi subequal to

claw-joint.

Hypopygium (cf. Fig. 2, top right) with the lateral lobes of tergite g slightly

produced apically; sternite g conical, slightly grooved obliquely on sides

basally, its process on each side, in side view, not very long, slightly sinuous,

the apex, in dorsal view, sharply pointed; aedeagal apparatus with the dorsal

shoe-horn-shaped epimere rather narrow and much longer, quite 2 times,

length of phallic tubes.

The 9 specimen which I take to be the @ of this species is characterized as

follows:

Body also mainly very dark blackish brown, the apical part of abdomen

becoming more dark reddish brown; clypeus also reddish brown; antennae

coloured as in $; proboscis however darker above; thorax with the humeral

NEW MYDAIDAE (DIPTERA) 121

tubercles also pale yellowish as in 3, the area around them, especially anteriorly

on inner side of tubercles, the sides of thorax, postalar calli, base of mesonotum,

and scutellum however also yellowish; metanotum dark reddish brown antero-

laterally, its central part black; pleurae as in §; abdomen with tergite 1 also

markedly inflated and convex as in J, the sides of this tergite and its broad hind

margin reddish brown; the very broad, ring-like, hind margins of tergites 2-6

ivory yellow, and a broad, discal spot occupying apical half of tergite 7 and

extending on each side narrowly and obliquely towards hind margin also ivory

yellowish; bullae shining black; venter also with broad ivory yellow hind

margins across sternites 2-6, but 7 and 8 however without yellow hind margins;

coxae more dark reddish brown than black; legs coloured as in ¢.

Integument of head and thorax as in 3; abdomen however mainly smooth

and shining, polished in appearance, with widely-spaced, fine, setiferous

puncturation lodging the forwardly-directed hairs only on tergites 6-8 and

with fine, scarcely detectable, setiferous puncturation on disc of tergite 2 and

base of 3; venter mainly smooth and shining, with widely-spaced, fine punctura-

tion only on sternites 7 and 8; tergites and sternites 7 and 8 also with transverse

striation; integument of legs as in ¢.

Vestiture also mainly snow white; that on head, thorax, basal part of

abdomen, and coxae similar to that of 4, only slightly shorter; that on sides of

tergites 1 and 2, metasternum and hind coxae also slightly shorter; fine and

short, separated hairs on tergites 2 and 3 gleaming slightly sericeous yellowish

in certain lights; middle parts of abdomen above mainly smooth; fine reversed

hairs on tergite 6 and longer stiffer ones on 7 and 8, and bristly ones on genitalia,

brown; sternite 2 with some separated whitish hairs; hairs on legs only a little

shorter than in g; tomentum on head and thorax as in g, but without any

tomentum on the smooth abdomen.

Head as in 3, with the interocular space on vertex subequal in width to

space below head; vertex not very deeply sunk in; postvertical spines more

distinct; antennae as in J, the joints similar, with the same proportions; pro-

boscis as long (about 2,8 mm) as in 4g, though slightly thicker and

more darkened above; palps also slender, but with more dark hairs.

Wings differ from those of ¢ in being faintly infuscated brownish, and with

indications of faint fuscous borders to basal cross vein and basal veins of first

posterior cell and vein between it and second submarginal cell; latter with a

longer appendix; first posterior cell however less widely opening on costal

margin; hind border of wings in apical half less narrower than posterior cells

than in 9; halteres similarly shaped.

Legs very similar, but hind femora less thickened, armed below with a

few more spines (7-8 in inner row and 7-9 in outer one); spicules on tibiae

longer than in 3; claws of tarsi shorter, more sickle-shaped, not so hook-shaped;

pulvilli smaller.

Genital segments with 7 narrow yellowish brown spines on each acantho-

phorite of tergite 9; anal lobes prominent, pincushion-like on each side, with

[22 ANNALS OF THE SOUTH AFRICAN MUSEUM

projecting bristly hairs.

From a ¢ holotype and 9 allotype in the South African Museum.

Length of body: about 16,5 mm

Length of wing: about 11-12 mm

Distribution

South West Africa: Kaokoveld: Orupembe (H. D. Brown, 1/5/70) (3

holotype); 21 km south of Orupembe (H. D. Brown, 1/5/70) (9 allotype).

This species is very near some dark forms of mauricer Beq. The 3 may at

once be distinguished by the much darker or black abdomen and venter,

the longer antennae, distinctly very much longer third antennal joint, on the

whole slightly longer proboscis, a first posterior cell in wings which opens more

broadly on costal margin, more inflated or convex tergite 1, more widely-

spaced setiferous puncturation (hairs) on abdomen, and distinctly much

longer and relatively narrower epimere.

The 2 may also be distinguished from that of maurice: by the antennal

characters, the longer and denser hairs on thorax, pleurae, metanotum, and

base of abdomen, the very broad yellow spot on apical half of tergite 7, the

slightly less infuscated wings, and the less hairy legs.

Afroleptomydas (Crossoprosopus) cognatus n.sp.

Another dark-bodied 3 from the same geographical region in South West

Africa is so closely related specifically to the 3 of nigrescens that it may almost

be considered as a subspecies of the latter. This 3 specimen however agrees

and differs from the 3 of nigrescens in the following respects:

Body longer, distinctly more robust, with more robust thorax and first

and second abdominal segments, with tergite 1 however also markedly inflated

and convex; abdomen distinctly stouter; body also mainly black; antennae

with joints 1, 2 and 3 entirely black (not with 3 brown as in nigrescens) ; clypeus

very dark blackish brown, not reddish brown; proboscis more darkened above;

thorax with the humeral tubercles and postalar calli similarly coloured; abdo-

men including venter also mainly black, with the hind margin of tergite 1 also

dark reddish brown, and ring-like hind margins of rest of tergites also broadly

ivory white; bullae also black, but much longer in vertical length and longer

than the interocular space; venter and hind margins of sternites similarly

coloured; legs with the front and middle femora darker posterolaterally and

below, the hind ones more extensively blackened on sides and apically above

than in nigrescens, and hind tibiae also more darkened along outer part towards

apex.

Integument also mainly shining, the mesonotum with coarse ‘nadelrissig’,

setiferous puncturation; abdomen with similar dull, discal, saddle-shaped

patches on tergites 2-6, but with distinctly coarser, more ‘nadelrissig’, com-

paratively less widely-spaced, setiferous puncturation on tergites; integument

of legs subshining, slightly more transversely striate.

NEW MYDAIDAE (DIPTERA) 123

Vestiture distinctly longer and denser, but also mainly snow white, dis-

tinctly longer and denser on pleurae, sides of tergites 1 and 2, with more long

hairs on sides basally of tergites 3 and 4; the hairs discally on mesonotum not

arranged in such well-defined streaks, more or less only as a broad central

streak; the shorter, more sericeous-gleaming, decumbent hairs on abdomen

above distinctly much longer than in nigrescens and also less widely spaced,

those discally on tergite 2 and basally on 3 distinctly very much denser; hairs

on venter also longer and denser; those on legs very similar, but hind femora

with longish white hairs on nearly basal half above; tomentum present on same

parts as in nigrescens.

Head with the vertex distinctly more markedly sunk in, with no distinct

postvertical spines detectable; antennae distinctly much stouter (the clubs

missing in specimen), with joint 1 slightly more thickened, but also about 24

times length of 2, with joint 3 more thickened, stouter, very much shorter than

in nigrescens; proboscis much stouter, slightly longer, about 3 mm.

Wings also greyish hyaline, comparatively broader, the hind border

scarcely or not markedly narrower than first and third posterior cells; second

submarginal cell with a long appendix; first posterior cell more narrowed

apically, more narrowly opening on costal margin; discoidal cell in this speci-

men not stalked apically; halteres gradually broadened to knob on inner side

as in nigrescens.

Legs comparatively longer than in nigrescens; front and middle femora

also unarmed below, the hind ones slightly more thickened, armed below with

the usual double row of spines, but with more numerous, shorter and stouter

ones (8 along inner row and g in outer one), beginning much nearer base;

basitarsus of hind legs longer than, not subequal to, claw-joint.

Hypopygium (cf. Fig. 2, bottom left) differs from that of nigrescens in having

the processes of sternite 9, in side view, less sinuous, slightly longer, appearing

more slender, in dorsal view less rapidly narrowed to apex, the latter more

broadly rounded and with 2 distinct punctures; aedeagal apparatus with the

epimere comparatively shorter and, in ventral view, more rapidly broadened to

apex.

From the single 3 holotype in the South African Museum.

Length of body: about 17,5 mm

Length of wing: about 13,5 mm

Distribution

South West Africa: Kaokoveld: 21 km south of Orupembe (H. D. Brown,

1/5/70).

From damarensis n.sp., described further on, which it also resembles, this

species differs in being much larger, more robustly built, with longer and

broader wings, longer first antennal joints, much shorter third antennal joints,

much longer proboscis, stouter, more bluntly rounded sternite 9 of which the

processes (cf. Fig. 2, bottom left and Fig. 1, bottom right), in side view, are

I24 ANNALS OF THE SOUTH AFRICAN MUSEUM

comparatively longer, more slender and, in dorsal view, do not diverge slightly

outwards.

Afroleptomydas (Crossoprosopus) aridicolus n.sp.

Still another almost entirely black-bodied J specimen from the Namib

Desert in the collections before me constitutes another new species which

closely resembles both nigrescens and cognatus, but more so the latter. Unfor-

tunately the antennae of this specimen are missing. It agrees with and differs

from the two species mentioned in the following respects:

Body tending to be more robustly built than in nigrescens, more like that of

cognatus, the abdomen as stout as in the latter; body also mainly very dark

blackish brown to black; clypeus very dark blackish brown as in cognatus;

proboscis very dark reddish brown as in the latter species; humeral tubercles

yellowish as in both nigrescens and cognatus, but the area immediately around

them black, not yellowish brown or reddish brown; postalar calli slightly

darker reddish brown; sclerites below wing-bases also dark reddish brown as in

cognatus; sides of tergite 1 very dark, not yellowish brownish or reddish brown

as in the other two species; abdomen with the hind margin of the distinctly

less inflated tergite 1 dark, the hind margins of tergites 2-8 also broadly ring-

like ivory yellowish, also becoming narrower posteriorly from tergite 5; bullae

blackish red, black in the other two species, their vertical length sub-equal to

interocular space on vertex, longer than in mgrescens, but shorter than in

cognatus, fairly widely separated; venter yellowish brown in at least basal half,

darker in the other species, the hind margins of sternites 2-6 broadly ivory

whitish, but 7 very narrowly so, as in cognatus; apical parts of tergite 9, anal

lobes, processes of sternite 9g and aedeagus yellowish brown; legs more yellowish

than in the other two species, the upper and outer surfaces of the front and

middle femora appearing darker, more brownish, the hind femora with more

than apical half dark brown and the basal part contrastingly yellowish, more

conspicuously delimited than in the other two species (in cognatus almost entire

outer face is dark brown); all the tibiae paler yellowish, the hind ones not

darkened or darkened towards apex; last tarsal joint also darkened.

Integument with the same parts shining as in the other two species; dorsum

of mesonotum however appearing duller; that of pleurae very similar; abdomen

with the same saddle-shaped, dull, discal, basal patches on tergites, but slightly

more extensive, leaving less of the sides shining, these patches also covered

with greyish to greyish yellow tomentum, visible in certain lights, the dorsum of

abdomen also with setiferous puncturation, distinctly less widely separated

than in nigrescens, more like that of cognatus, but slightly less coarse and with

shorter hairs; venter also shining, finely setiferously punctured, though less

dense than in cognatus and less coarsely transversely striated in apical half;

integument of legs as dull as in cognatus and also with indications of transverse

striation.

Vestiture very similar to that of the two other species, also mainly snow

NEW MYDAIDAE (DIPTERA) 125

white, though distinctly longer and denser than in nigrescens, but distinctly

shorter than in cognatus, more of the clypeus dorsally covered with hairs which

are slightly longer than in cognatus, the tufts on vertex tinted slightly more

yellowish in certain lights; hairs on mesonotum not so distinctly and demarcately

present as distinct streaks as in nigrescens, more like those of cognatus, the broad

medial streak however broader than in the latter, leaving narrower, more or

less hairless streaks on sides discally; these decumbent hairs also gleaming

sericeous yellowish; hairs on pleurae present on the same sites, but longer and

denser than in nigrescens, more like those of cognatus; long and dense white hairs

on sides of tergites 1, 2 and 3 and 4 as in cognatus, but slightly shorter ; decumbent

ones on tergites longer and less widely separated than in nigrescens, but dis-

tinctly shorter than in cognatus, also gleaming slightly sericeous yellowish;

hairs on venter distinctly shorter and slightly less dense than in cognatus; hairs

on coxae as in the latter, but comparatively shorter; hairs on legs with those on

front and middle femora also long and dense, but slightly shorter than in cogna-

tus, those on hind femora however distinctly less dense and shorter than in

either of the other two species.

Head with the vertex less deeply sunk in than in cognatus, but slightly

more so than in nigrescens; interocular space on vertex also subequal to space

on head below; postvertical spines, as in cognatus, not detectable; antennae

with joint 3 and club missing in specimen, but joint 1 slightly thickened, quite

21 times length of 2; proboscis about 2,7 mm long, distinctly shorter than that

of cognatus, longer than vertical length of eyes, its labella shorter and more

rounded apically than in the other two species; palps also slender.

Wings very faintly yellowish hyaline; veins yellowish, not brownish as in

cognatus; second submarginal cell with an appendix; first posterior cell narrowed

apically, even more so than in cognatus, narrowly opening on costal margin;

discoidal cell shortly stalked as in nigrescens; halteres pale yellowish white, the

knobs darkened above.

Legs slightly longer than in nigrescens, more like those of cognatus; front

femora armed below along inner aspect with a row of distinct, short, spine-

like bristly hairs, not detectable in the other two species; hind femora thickened,

subspindle-shaped, armed below with an inner row of 10-11 and an outer row

of about 9 whitish spines, beginning near base, thus with 2 or 3 more spines in

outer row than in cognatus; spicules on tibiae longer, as long as or longer than

width of tibiae, shorter than this width in the other two species; basitarsus of

hind legs longer than claw-joint.

Hypopygium (cf. Fig. 2, bottom middle) with the lateral lobes of tergite g

slightly angularly produced; sternite g conical, bluntly rounded apically,

obliquely grooved basally on each side and then continued longitudinally,

its dorsal processes projecting much beyond apex of sternite, in side view long

and slender, only slightly curving upwards near apex, in dorsal view grooved

above along inner aspect, slightly bent outwards near apex, the latter relatively

bluntly pointed; aedeagal apparatus with the shoe-horn-shaped epimere

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

relatively shorter than in the other two species, about 1,6 times length of

phallic tubes (in nigrescens and cognatus it is quite twice length of the tubes).

From the single ¢ holotype in the South African Museum.

Length of body: about 17,5 mm

Length of wing: about 12,5 mm

Distribution

South West Africa: Arechadamab, east of Swakopmund in the Namib

Desert (H. D. Brown, 11/5/1959).

Afroleptomydas (Crossoprosopus) damarensis n.sp.

Another black-bodied 3 specimen from South West Africa in the collec-

tions of the Stuttgart Museum and submitted by Professor Lindner is apparently

a new species resembling maurice: Beq. from the Kalahari in Botswana, and also

resembles nigrescens n.sp. It has two identification labels ‘Gen. nov. pr. ?

Mydaselpis Bezzi’ and ‘? rufithorax Wied. det. H. Oldroyd, 1957’ pinned under

the locality label. It is however not related to either the genus Mydaselpis or to the

Cape species Afroleptomydas rufithorax.

When compared with 39 of the variable species mauricei, especially the

darker forms, it agrees and differs from the latter in the following respects:

Body almost entirely black; proboscis entirely black, not infused with

reddish brown or reddish below; clypeus much darker; antennal joints 1 and 2

dark blackish brown, not so yellowish as in mauricez, the broad middle part of

clubs however also orange yellowish; humeral tubercles, which are compara-

tively smaller, also yellowish; rest of thorax, excepting obscure reddish brown

postalar calli and yellowish brown sclerites below wing-bases, entirely black;

abdomen mainly black, without any yellowish or brownish as in mauricei, the

extreme sides of tergite 1 also black, but hind margin of tergite 1 however also

reddish brown; bullae black anteriorly and reddish posteriorly (entirely black

in mauricet), smooth, without any punctures; ivory yellowish hind margins of

tergites 2-7 broader, more broadened on extreme sides, and with those of 6 and

7 broader than in mauricei; hind margins of sternites very broadly and con-

spicuously ivory yellowish, even those of 6 and 7 comparatively broad; lobes

of tergite 9, sternite 9, and processes of latter yellowish brown; legs mainly

very pale yellowish brown, the front and middle femora not darkened above

or below, the hind ones conspicuously black above in apical half, the black

patch-like and well marked off, not merely infused to a variable extent as in

maurice. and other species, with the hind tibiae darkened apically and the

claw-joints of all the tarsi darkened.

Integument very similar to that of maurice: and related species of the subgenus

Crossoprosopus.

Vestiture very similar to that of mauricec and equally long, also mainly

white, the longish hairs on head, mesonotum, pleurae, and abdomen basally

however appearing to be slightly more dirty whitish than snow whitish (the

NEW MYDAIDAE (DIPTERA) 127

tps

toon Sean

Ny Yr. aN

Biget

Left: Dorsal view of right antenna, proboscis, and below, left view of hypopygium of $ Afrolepto-

mydas lindneri n.sp.

Right: Proboscis, side view of left antenna, and below left view of hypopygium of 3 Afroleptomydas

(Crossoprosopus) damarensis n.sp.

eee —aedeasus. Al, — anal lobes; IX.S. = sternite 9; EX.F. — tergite 9; Pr. = left

process of sternite 9; VIII.S. = sternite 8; VIII.T. = tergite 8.)

specimen however appears to be slightly greasy) ; hairs on mesonotum similarly

disposed in streaks; those on sides of tergite 4 also long as on sides of preceding

tergites, not shorter than on sides of 3 as in mauricei; longish hairs on venter also

present on sternite 4; fine, decumbent hairs in fine setiferous punctures on rest

of abdomen above appearing slightly more sparsely disposed than in maurice;

greyish white tomentum on head, thorax above, metanotum, pleurae and on

abdomen above similar to that of mauricei and most other species of the sub-

genus; that on abdomen above in this somewhat greasy specimen however

appears dark in part; hairs on legs as in mauricei, long, dense and white on coxae,

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

slightly long and dense on upper and outer faces of front and middle femora,

and comparatively sparse and short on hind ones.

Head with the antennae (cf. Fig. 1, top right) comparatively longer than in

mauricet, distinctly much longer than mesonotum; joint I proportionally shorter

than in latter, less slender and only about twice length of 2; joint 3 propor-

tionally longer, with a slightly longer apical thickened part, the club also

elongate amphoriform, broadest beyond middle, its apical sensory area dis-

tinctly more conical all round, not sunk in on inner lower part, the apical

crater-like tubercle distinctly more rounded, not so transverse as in mauricei;

proboscis (cf. Fig. 1, top right) proportionally much shorter, only about 1,8 mm

long and subequal in length to third antennal joint, its labella proportionally

short, apically bluntly rounded (not sharply pointed), and with slightly fewer

striae.

Wings, in proportion to body-length, relatively shorter than in mauricet,

more glassy hyaline, without any distinct indication of a faint yellowish tint;

veins pale yellowish brown; first posterior cell slightly narrowed apically,

subtending angularly on costal margin; halteres like those of mauricev.

Legs proportionally distinctly shorter than in mauricet; hind femora simi-

larly thickened, but more subspindle-shaped, the thinner basal part being

proportionally shorter, armed below with the usual double row of pallid spines

on tubercles which are however closer together, due to the shorter femora, but

also with about g spines in both inner and outer rows; hind tibiae as granular

below; spicules on tibiae and tarsi as in maurice.

Hypopygium (cf. Fig. 1, bottom right) similar to that of mauricez, but sternite

g more conically produced, its apex not so bluntly rounded and itself apparently

longer, its processes, in side view, distinctly less sinuously S-curved, straighter,

its apex scarcely or not bent downwards, in ventral view, the apex is more

curved outwards; epimere of aedeagal complex comparatively and propor-

tionally shorter relative to phallic tubes.

From the single ¢ holotype in the Stuttgart Museum.

Length of body: about 15,5 mm

Length of wing: about 9,5 mm

Distribution

South West Africa: Khomashochland (17/5/1953). No collector’s name

appears on the locality label, but the handwriting is that of Herr F. Gaerdes

of Okahandja.

From the ¢ of nigrescens, which is also mainly dark-bodied and with dark

abdomen, it may be distinguished by the entirely dark and much shorter pro-

boscis, shorter and rounded labella, absence of yellowish brown on extreme

sides of tergite 1, half reddish bullae, comparatively shorter wings, hind femora

which have a contrasting, patch-like, black infusion above, denser hairs on

abdomen above, and relatively shorter epimere in the aedeagal apparatus.

NEW MYDAIDAE (DIPTERA) 129

Afroleptomydas (Crossoprosopus) aquilus n.sp.

A somewhat damaged, markedly bare 9 specimen from Damaraland in

the collections before me cannot be allocated to any of the known 33 from

South West Africa. Superficially it resembles some 2 forms of mauricei, but is

entirely distinct. It is characterized as follows:

Body mainly very dark blackish brown to black; clypeus very dark, dark

blackish brown; proboscis dark reddish brown, more black above and on

labellar part; antennal joints 1 and 2 dark reddish brown, joint 3, almost

basal half of clubs and apical tubercle part of latter very dark blackish brown

or almost black, the greater middle part of clubs however orange yellowish;

humeral tubercles yellowish; extreme sides of mesonotum very obscurely

reddish brownish in certain lights, the postalar calli more yellowish, and extreme

base of mesonotum reddish brown; scutellum black; metanotum black, its

extreme base behind scutellum and two submedial streaks obscurely yellowish

brown; pleurae very dark blackish brown to black; sclerites below wing-bases

and sutural parts of pteropleuron yellowish brown, and the sclerite below

halteres yellowish; extreme sides of tergite 1 also yellowish; abdomen with

transverse depressed hind margin of tergite 1 reddish brown, the broad hind

margins of tergites 2—7 yellowish, those of 6 and 7 broad discally, that of 7 even

extending basalwards to more than half of the tergite, and tergite 8 mainly

reddish brown discally, becoming paler apically; bullae comparatively small,

black, but orange yellowish across hind margin, not punctured, very widely

separated; venter dark or black, its base yellowish and hind margins of ster-

nites 2-7 broadly yellowish, the last one the narrowest; legs mainly pale

yellowish, the front and middle coxae, basal halves externally of hind ones

dark, very dark blackish brown, and an inner subapical infusion and also an

outer more obscure or fainter apical one on hind femora dark brownish.

Integument of head in front, greater part of sides of propleuron, the meso-

pleuron, sternopleuron, part of hypopleuron behind and below halteres, and

to a large extent abdomen above and below shining; rest of pleurae slightly

duller, covered with fine greyish bloom or tomentum; mesonotum duller,

more subshining, more leathery, covered with fine separated, setiferous punc-

tures, more or less in streaks; scutellum and metanotum dull, leathery; tergites

1 and 2 discally subshining, covered with fine, separated, setiferous punctures,

denser on sides; rest of tergites with comparatively sparse, fine, setiferous

punctures discally, their sides transversely striate, becoming coarser on sides

of 5-7; tergites 7 and 8, especially latter, more coarsely striate or grooved, even

discally ; integument of venter mainly smooth, shining, more transversely striate

on last three sternites; legs subshining, the femora transversely striate, the hind

tibiae more evidently so below.

Vestiture very poorly and sparsely developed, markedly short; that on

head in front relatively short and sparse, snow white, comparatively shorter

than in other mainly bare species; that on sides of clypeus short, not extending

across genal furrows; those on head below sparse; hairs on occiput also sparse

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

and white, with whitish postvertical spines on each side; sides of occiput with

comparatively sparse greyish white tomentum; hairs on thorax above very short,

decumbent, slightly sericeous yellowish in certain lights, more or less arranged

in streaks; those on sides of mesonotum also relatively short, not as long as in

some other species; metanotum with very short, sparse, yellowish hairs discally;

posterior margin of mesopleuron with scarcely detectable, short, yellowish

gleaming hairs; propleural knob however with sparse, longish, white hairs;

rest of pleural sclerites without any detectable, short hairs, only with fine,

rather sparse, greyish bloom or tomentum; discal part of tergite 1 with markedly

short, sparse, yellowish or golden gleaming hairs, even shorter, sparser, scarcely

detectable on sides of the tergite; tergite 2 with similar, very short, yellowish

or golden gleaming hairs, slightly denser and shorter on sides; tergite 3 with

even sparser, short, yellowish gleaming hairs; tergites 4 and 5 with very short,

very sparse, widely-separated, golden gleaming hairs mainly discally; tergite 6

with sparse, forwardly-directed, very short, brownish hairs; tergites 7 and 8

with longer, widely-spaced, forwardly-directed, brownish hairs, slightly longer

and denser on 8; venter mainly smooth and hairless, only last sternite with

some forwardly-directed, sparse hairs on sides; hairs on legs markedly short and

comparatively widely spaced, gleaming sericeous yellowish or golden, even

the denser ones on front and middle femora also very short.

Head with the vertex somewhat sunk in; the interocular space on vertex

slightly broader than on head below; antennae about subequal in length to

mesonotum, with joint 1 only slightly thickened, about 24 times length of the

rather short transverse joint 2, joint 3 slightly longer than club, a little more

than its apical 4 thickened, the clubs elongate pyriform, broadest beyond middle

across a little more than apical 4, their bases constricted neck-like, the apical

sensory area more rapidly narrowed below, the rim below of crater-like terminal

prominence rather prominently projecting; proboscis about 2,4 mm long,

longer than vertical length of eyes, its labella broad, obtusely pointed apically;

palps small, but distinct.

Wings (left wing of this specimen unfortunately missing) relatively long,

faintly tinted yellowish brown throughout, the veins yellowish brown; first

posterior cell broadly opening on costal margin, scarcely or not narrowed

apically; second submarginal cell with an appendix; discoidal cell shortly

stalked apically; halteres racket-shaped, their broadened apical part darkened

above.

Legs moderately long; hind femora slightly thickened, subspindle-shaped,

broadest slightly beyond middle, armed below with the usual double row of

relatively short, yellowish white spines on tubercles, about 7 in both rows;

front and middle femora unarmed below, front and middle tibiae slightly

curved, the spicules on former not so well developed as on middle ones; spicules

on tubercles in outer lower row in apical half of hind tibiae stout and spine-

like; basal joint of hind tarsi slightly longer than claw-joint; hind claws more

strongly developed, sickle-shaped, those of front tarsi least developed, more

NEW MYDAIDAE (DIPTERA) 131

hook-like; pulvilli well developed, but not reaching apices of claws.

Genital segments with tergite 8 slightly pointed apically; acanthophorites

each with 6 detectable brownish, dorsally-grooved and apically bluntly-pointed

spines, the last (or apical) one being spine-like; anal lobes with some stoutish

spine-like bristles basolaterally.

From the single ¢ holotype in the South African Museum.

Length of body: about 15,3 mm

Length of wing: about 12 mm

Distribution

South West Africa: Damaraland: Karibib (H. D. Brown, 20/5/1959).

The @ of this species may be easily recognized by its marked bareness

and feebly-developed vestiture on body. Superficially this 9 resembles the 29

of some similarly-coloured forms of maurice: Beq., but may at once be distin-

guished by the absence of longish hairs on ptero- and metapleurae, sides

basally of tergite 1, and basolaterally on hind coxae, distinctly longer antennae,

more broadly opening first posterior cell, and fewer spines on acanthophorites.

From the 9 of matetsiensis Beq., which is also very bare, it may be dis-

tinguished by the mainly black body, absence of even, sparse longish hairs on

ptero- and metapleurae, sides of metanotum and first tergite, slightly longer

antennae, broadly yellow discal hinder part of tergite 7, dark infusion laterally

on apical part of hind femora, etc.

The possibility that this 2 may be that of the 3 damarensis n.sp., which is

also very dark bodied, with mainly dark clypeus, antennae, and proboscis,

and also with very pale legs, is however ruled out by the distinctly very much

longer antennae, much shorter proboscis, apically much narrowed first posterior

cell, and more numerous spines on hind femora below of the 3 damarensis.

Afroleptomydas (Crossoprosopus) femoralis n.sp.

A pale-coloured 3 specimen from South West Africa, with comparatively

thickened hind femora and belonging to the pale-coloured matetsiensis-section,

is a new species which is characterized as follows:

Body rather stout and robust, with broad and robust thorax and base of

abdomen, with the abdomen thickened and cylindrical; the body and legs

mainly pale coloured, pale yellowish red to yellowish brown; frons and vertex

black; proboscis reddish, its labella mainly black; a middle streak on anterior

half of mesonotum and a broadish sublateral streak on sides of mesonotum

darkened or black, ending posteriorly on each side in a smooth dark spot;

mesopleural and sternopleural parts obscurely darkened, more brownish or

castaneous; metanotum also darkened, more brownish in hinder half; hind

margins of tergites, including tergite 8 broadly and conspicuously ring-like,

yellowish white, becoming narrower on posterior tergites; hind margins of

sternites 1—7 also broadly yellowish white; sides of transverse basal depression

of tergite 2 slightly darkened; tergites 2-5 basally obscurely darkened, and

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

extreme sides of 5-7 and more or less basal half of sternite 8 also darkened,

more blackish brown; anal lobes tending to be paler, more yellowish; bullae

transversely elongate, widely separated, their posterior halves orange yellowish

and anterior halves black and shining, the extreme hind margins also darkened,

their anterior margins with a row of granules; legs mainly pale yellowish brown,

the anterior and middle trochanters darkened behind, the outer faces of

femora, especially hind ones, showing through very slightly dark in certain

lights, with the spines on hind femora below pallid on reddish yellow tubercles,

the granules below hind tibiae shining dark reddish brown to almost black,

and apical halves of claws black.

Integument of central part of frons, clypeus and head below shining; that

of mesonotum dull, leathery, with setiferous puncturation lodging the short

decumbent hairs; scutellum smooth, shining; metanotum mainly dull; pleurae

mainly dull, but propleural and sutural parts, as well as sclerites below wing-

bases, more or less shining; abdomen mainly dull, but lateral basal parts

of tergites and ring-like hind margins subshining, the tergites mainly setiferously

punctured, lodging the hairs; sides of exposed parts of tergite 8 and processes

of sternite 9 shining; venter subshining, the pale hind margins of sternites more

shining; legs with the trochanters shining, the femora subshining, more shining

below, the femora and tibiae with fine setiferous puncturation, lodging the hairs.

Vestiture rather dense, dense, long and creamy whitish on head, sparser

and more white on head below; that on thorax above dense, creamy white,

longer on sides and posteriorly in front of scutellum; that on disc of mesonotum

shorter, directed backwards, gleaming sericeous yellowish, arranged in a

narrow central denser streak and two broader submedial denser streaks, and a

broadish sublateral, less dense, barer streak, ending posteriorly in an entirely

bare spot on each side; pleurae with faint tomentum on dull parts, the long

hairs across hind part of mesopleuron, on pteropleuron, and metapleural parts

also creamy white; longish dense hairs on metanotum, tergite 1 (and its sides),

tergite 2, on sides especially, and to a feebler extent basally on sides of 3 also

creamy white; short, backwardly-directed hairs in setiferous punctures on rest

of abdomen above not very dense, gleaming sericeous yellowish to pale golden,

slightly longer and sparser on sides of tergites; hairs on venter slightly longer

than on dorsum, sparser, also gleaming sericeous yellowish to pale golden;

hairs on hypopygium rather sparse, pale golden; those on legs gleaming pale

sericeous yellowish, slightly longer and denser on femora above, comparatively

dense on front and middle femora above; tomentum on head dense, greyish

white, present on sides of frons, vertex and behind eyes; greyish tomentum also

present on mesonotum along the streaks of hairs; metanotum also with faint

greyish tomentum.

Head broader than thorax, with the vertex sunk in, interocular space on

vertex subequal to length of central part of frons to antennae and also subequal

to narrowest part of width of interocular space on head below; some slender

whitish postvertical spines present; antennae with only joints 1 and 2 present

NEW MYDAIDAE (DIPTERA) 133

in specimen, the former quite 2} times length of the comparatively short

joint 2 and not much thickened; clypeus with an indication of a central groove;

proboscis about 1,96 mm long, much shorter than vertical length of eyes, its

labella a little less than twice length of rest of proboscis; palps small.

Wings glassy hyaline, with a scarcely perceptible milky whitish tint in

certain lights; costal vein, basal part of second vein and parts of veins at base

of wings yellowish, the rest darker, more brownish; second submarginal cell

with an appendix; first posterior cell slightly narrowed apically, opening (or

sessile) on costal margin; discoidal cell shortly stalked apically; halteres

whitish, their knobs slightly darkened medially above.

Legs stoutish; hind femora markedly thickened, more so than in most

other species, armed below with a double row of rather short, stout, whitish

spines on tubercles (11-12 in outer row and I1 in inner row, beginning near

base) and in right femur also with a small sublateral, subapical spine on each

side; front and middle femora unarmed; front and middle tibiae slightly

curved, the front ones with an outer lower row of large spicules, those on dorsum

small, the middle tibiae with more or less two rows of longer spicules below and

two irregular rows of shorter ones above; hind tibiae with irregular rows of

setiferous granules below, the outermost ones in apical part with longer and

stouter spicules; basal joint of hind tarsi dorsally subequal in length to claw-joint.

Hypopygium with the lateral lobes of tergite 9 rather sharply angular;

sternite 9 relatively rather small, conical, bluntly rounded apically and there

medially sulcate, its processes projecting much beyond apex of sternite, their

slender part, in side view, almost straight, more narrowed and rounded apically

in dorsal view; aedeagal apparatus with the phallic tubes much shorter than

the shoe-horn-shaped dorsal epimere.

From the single ¢ holotype in the South African Museum.

Length of body: about 19 mm

Length of wing: about 12 mm

Distribution

South West Africa: Kaokoveld: Otjivakandu (sometimes spelled Otjiva-

kondo) (H. D. Brown, 27/4/1970).

From the ¢ of matetstensis Beq., which it resembles superficially, it may

however be distinguished by the distinctly stouter body, shorter proboscis,

denser hairs on head in front, denser and longer hairs on the usual hairy parts

of body, broader ring-like hind margins of tergites, more orange-coloured

discal part of bullae, more thickened hind femora, more numerous granules on

hind tibiae below, and straighter, less sinuous processes of sternite 9.

Afroleptomydas (Crossoprosopus) browni n.sp.

Some ¢¢ and 99 from the Okovango River region in the collections

constitute a new species belonging to the mainly pale-coloured matetsiensis-

section. With great pleasure this new species is named after the collector and

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

donor, Dr H. D. Brown, of the Department of Agricultural Technical Services.

The 3¢ are characterized as follows:

Body with the head mainly black; clypeus, buccal cavity, and to a certain

extent lower parts of genae and head below, yellowish brown to dark reddish

brown; proboscis yellowish brown below, usually darkened above, the labella

mainly dark to black; antennal joints 1 and 2 yellowish to yellowish brown

and basal part or half of 3 also yellowish brown to reddish brown to a variable

extent, rest of 3 and basal part of clubs dark or black; greater part of clubs

conspicuously orange yellow, the sensory area sometimes also darkened;

thorax above dark or black, the humeral tubercles, area around them, noto-

pleural sides of mesonotum, postalar calli, narrow base of mesonotum, and

scutellum yellowish brown to reddish brown, the humeral tubercles themselves

more conspicuously pale yellowish, and sometimes with an indication of a faint

narrow, sublateral, reddish or yellowish streak discally on each side of mesono-

tum; connection between postalar calli and scutellum with a conspicuous black

spot, continuous with the black hollow on each side of scutellum; metanotum

reddish brown to dark reddish, usually darker centrally and to a variable

extent in apical half, sometimes paler on extreme sides; pleurae with the

anterior and lower parts darkened or black to a variable extent, the upper

anterior part of mesopleuron, area below wing-bases, pteropleuron (or apical

half of it), and metapleural parts around and below halteres, and hypopleural

part being yellowish brown, with a black spot or streak anteroventrally to

posterior spiracle, extending down, sometimes broadly, to metasternum, the

sternal parts also mainly dark or black to a variable extent; abdomen cylindri-

cal, mainly reddish brown, the discal basal half or more of tergite 1 and to a

variable extent the extreme lateral margins of the tergites black or darkened;

tergites 3-7 sometimes discally appearing slightly darkened in certain lights;

hind margins of tergites 1-8 broadly yellowish white, the bases of these rings

tinted orange to a variable extent; venter yellowish to yellowish brown, the

hind margins of sternites also broadly yellowish, more evident on sides, the

medial part shining through more brownish; bullae black, shining, not pitted;

legs mainly reddish brown, the tibiae and tarsi sometimes appearing more

yellowish, the base of hind tibiae sometimes more yellowish, the apices of claws

black.

Integument of central part of frons, clypeus, and head below as usual smooth

and shining; that of proboscis and labella, antennal joints 1 and 2, and basal

part of 3 also shining; mesonotum dull, leathery, with some setiferous punctures

lodging the vestiture where present; postalar calli and scutellum more shiny;

metanotum transversely slightly rugulose on sides, mainly dulled by tomentum;

pleurae mainly dulled by fine bloom or tomentum; abdomen dulled by a slight

greyish bloom which in these mounted specimens gives it a slight oily appear-

ance, the underlying integument very finely microareolate and with fine,

separated, setiferous puncturation lodging the hairs, the transverse basal

depression of tergite 2 and much narrower one across 3 shining, and the extreme

NEW MYDAIDAE (DIPTERA) 135

sides of tergites (not covered with bloom) also shining; ring-like hind margins

of the tergites more or less subshining; tergites 3—7, in certain lights, showing

shallow, transverse striae on sides; venter smooth, shining, with indications of

transverse striation on sides of sternites and on posterior sternites, the integu-

ment also with fine, separated, setiferous puncturation where hairs occur; legs

subshining, the femora more shining below, with indications of transverse

striation on both femora and tibiae, the surface also covered with fine setiferous

puncturation lodging hairs.

Vestiture moderately long, mainly white; hairs in tufts on head in front

very dense, snow white, leaving bases of antennae, space below antennae, and

dorsum of clypeus bare; rest of long white hairs on vertex, occiput, and head

below not so dense; whitish or pale yellowish white postvertical spines present,

shortish and bristle-like; sides of frons and face narrowly, sides of vertex, and

most of occiput to lower part of eyes densely covered with greyish white tomen-

tum; hairs on mesonotum more or less arranged in 5 streaks, the central one

narrowest, the broad one on each side composed of dense, decumbent, longish,

snow white hairs, the submedial streaks begin with a patch of dense, longish,

white hairs which become shorter discally and also less dense, sometimes with

a scarcely perceptible sericeous yellowish tint, the narrow central streak of

short, decumbent, whitish or pale sericeous yellowish hairs end in a patch of

denser, longer, snow white ones in front of scutellum, the broad lateral and

submedial streaks also with greyish tomentum; hairs on metanotum fairly

dense, long, snow white, directed forwards and absent from a broad central

streak, the surface also with greyish tomentum; pleurae mainly with fine,

slightly greyish yellow bloom or tomentum, the hind margin of mesopleuron

with or without a few longish white hairs, the pteropleuron and area in front

of halteres with sparse, longish, white hairs, but hypopleuron and area below

halteres conspicuously bare; prosternal and metasternal parts however with

longish white hairs like the dense white ones transversely across in front of

hind coxae; abdomen with fairly long, dense, snow white hairs basally on

tergites 1 and 2, especially on sides, with some long ones also on sides of tergite 3

basally and some shorter ones on sides of 4; rest of abdomen above with short,

decumbent, somewhat separated, slightly sericeous yellowish gleaming hairs

in fine, setiferous punctures, slightly denser discally on tergite 2 and slightly

longer on sides of tergites; decumbent hairs on venter very slightly longer than

above, also gleaming slightly sericeous yellowish, absent from extreme sides

of sternites; hairs on hypopygium as separated as on tergites, more or less

equally spaced, very slightly longer, but also gleaming pale sericeous yellowish;

hairs on legs comparatively short, long and snow white only on outer side of

hind coxae, gleaming slightly sericeous yellowish in certain lights on rest of

legs, those ventro-laterally on outside of femora slightly longer than dorsally.

Head broader than thorax; vertex sunk in, the interocular space on vertex

subequal in width to, or only very slightly broader than, space below head, the

margins of eyes converging above and below; antennae subequal in length to

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

Big, 2:

Side view of left lateral lobe of tergite 9 and side and dorsal views of left process of sternite 9

of §¢ (all to same scale) of:

Afroleptomydas (Crossoprosopus) angolensis n.sp. (top left).

Afroleptomydas (Crossoprosopus) ovamboensis n.sp. (top middle).

Afroleptomydas (Crossoprosopus) nigrescens n.sp. (top right).

Afroleptomydas (Crossoprosopus) cognatus n.sp. (bottom left).

Afroleptomydas (Crossoprosopus) aridicolus n.sp. (bottom middle).

Afroleptomydas (Crossoprosopus) browni n.sp. (bottom right).

mesonotum, with joint 1 only slightly thickened, about 2-24 times length of

2, joint 3 about, or usually a little more than, 3 times combined length of 1

and 2, usually a little shorter than club, the latter elongate pyriform, broadest

beyond middle nearer apex, sometimes slightly constricted at about the middle,

with a distinct, well-marked-off, basal, neck-like constriction, its apical sensory

area demarcated by a ridge below, rapidly conically narrowed below to the

crater-like tubercle; proboscis comparatively short and stout, slightly upcurved,

shorter than vertical length of eyes, about 1,8-2 mm, its labella about a third,

or a little more, of its length; palps small, but distinct, with some brownish hairs.

Wings clear hyaline, with a very faint yellowish tint in certain lights;

costal and subcostal veins yellowish, the rest of veins yellowish brown, those in

hinder part more brownish; base of second submarginal cell more often acute,

sometimes however truncated, with an appendix; first posterior cell narrowed

apically, more often narrowly opening on costal margin, occasionally just

subtending on it or even broadly opening on it; discoidal cell usually shortly

stalked apically; halteres racket-shaped, pallid, the broadened apical part

usually darkened above to a variable extent.

Legs moderately long, the front and hind femora subspindle-shaped, but

hind ones much thickened; front and middle ones unarmed below, but thickened

NEW MYDAIDAE (DIPTERA) EY y

hind ones armed below with the usual double row of whitish to yellowish white

spines on dark tubercles (about 9-11 in outer row, beginning near base and

7-11 in inner row); tibiae slightly curved, the spicules along inner and lower

aspect of front ones not developed and those along outer lower aspect less

developed and shorter than on middle tibiae as are also the apical spurs; spicules

on hind tibiae on small tubercles, those from middle towards apex in both the

inner and outer rows becoming longer; basal joint of hind tarsi subequal to,

or only very slightly longer than, claw-joint; claws well developed, the hind

ones longer, more sickle-shaped; pulvilli well developed.

Hypopygium (cf. Fig. 2, bottom right) with the lateral lobes of tergite g

slightly produced and angularly pointed; sternite 9g navicular, longitudinally

striated, apically with a vertical sulcus, in side view conical, bluntly or roundly

pointed apically, its dorsal process or prong on each side projecting much

beyond the sternite, in side view slender, slightly upcurved near apex, its sharp

apex slightly bent down, in dorsal view slightly hollowed, narrowed

apically and fairly sharply pointed, the apices slightly diverging; aedeagal

apparatus with the shoe-horn-shaped epimere slightly broadened apically,

projecting beyond and longer than the divergent phallic tubes.

The 2 specimens, caught at the same time as the ¢¢ and at the same

localities, which I take to be the 99 of this species, are characterized as follows:

Body mainly or almost entirely reddish brown to dark brown; frons black

and shining; clypeus and head below dark reddish as in dg, the former also

brilliantly shining; proboscis as in 3 yellowish brown below, the lower half and

apex, or entire labella, darkened; antennae similarly coloured, joint 3 tending

to become darkened, the greater part of clubs sometimes even more conspicu-

ously orange yellow; thorax above more dark reddish brown to blackish brown,

a middle streak and a broader sublateral one on each side tending to be slightly

darker than the rest; humeral tubercles yellowish as in ¢; scutellum, metano-

tum, and pleurae similarly coloured, but latter sometimes more uniformly

reddish brown; abdomen stouter, sometimes broad, dorsoventrally depressed,

mainly reddish brown to dark reddish brown, the anterior transverse depression

and anterior discal half of tergite 1 dark or blackish brown as in 3; hind margins

of tergites 1-5 discally very broadly yellowish, narrower on sides, that of 5—7

extending basalwards discally, and tergites 6 and 7 discally extensively infused

yellowish, with 7 almost entirely so discally, their extreme hind margins discally

not yellowish but narrowly darkened; basal part and sides of tergite 7 and to

a variable extent extreme sides of 2-6 sometimes much darkened; bullae black

and shining as in g; tergite 8 mainly dark reddish brown, more yellowish

apically; venter mainly yellowish brown to dark reddish brown, the hind

margins of sternites shining through brownish as in 3, only extreme sides and

sometimes extreme hind margins of 1-6 narrowly yellowish to a variable

extent; legs coloured as in ¢, though sometimes darker, the tibiae and tarsi

sometimes appearing more obviously yellowish, sometimes basal part of hind

tibiae more distinctly yellowish.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

Integument of head in front, antennal joints 1 and 2, proboscis, and abdomen

above and below mainly smooth and shining; thorax above, metanotum and

pleurae dull as in ¢, covered with faint bloom or tomentum, the metanotum

transversely grooved on sides; mesonotum dull, leathery, with streaks of fine

setiferous puncturation where fine hairs are present; tergites 7 and 8, especially

the latter, transversely striated or grooved, and sides of 3-6 sometimes also

showing transverse striation to a variable extent; legs subshining as in 4, the

fine transverse striation on femora more evident.

Vestiture, unlike that of ¢g, poorly developed; short hairs on vertex,

anteriorly across frons, across sides of clypeus, and on head below shorter

than in g and with a distinct sericeous yellowish tint, those on sides of clypeus

sometimes even more yellowish or golden; those on occiput also shorter than

in 4; postvertical spines more evident and sometimes more yellowish brown;

greyish yellowish tomentum narrowly present along eye-margins and dense,

ereyish white tomentum also present on occiput as in ¢; hairs on mesonotum

very fine, short, gleaming sericeous yellowish to slightly golden and arranged

in streaks as in 3, those on sides of mesonotum very fine, short, not dense and

whitish as in 3; hairs on pleurae almost absent and very short; those on meta-

notum very short, fine, scarcely detectable; tergites 1 and 2 discally with fine,

greyish bloom or tomentum; tergite 1 discally also with fine, short hairs; sides

of tergite 1 with fine, short, sericeous yellowish hairs; tergites 3-6 mainly

smooth; tergites 7 and 8, especially latter, with forwardly-directed, dark brown

or blackish brown hairs; hairs on genital segments also blackish brown; venter

smooth, bare and shining, with fine, short, reversed, widely-spaced, yellowish

brown hairs present only on last two sternites; hairs on legs as in g, but only

shorter, those on coxae also much shorter and sparser and more sericeous

yellowish.

Head similar to that of 3; proboscis similar, about 1,5—1,8 mm long; anten-

nae similar, with the same shape and the same relationship between the

joints.

Wings tinted slightly more yellowish brownish, appearing darker, the

veins however with distinct fuscous borders, especially in middle part of wings,

rendering the wings darker and making the costal cell, basal half of first sub-

marginal cell, and almost entire first basal cell dark, and the discoidal and

second basal cells with clear streaks in middle; other venational characters as

in g; halteres very similar.

Legs like those of 3; hind femora also thickened, but slightly less so, not

so distinctly subspindle-shaped, armed below with the usual double row of

similar spines (in this case 8-12 in outer row and 8-9 in inner one) ; front femora

however with some distinct, spicule-like or hair-like spines along inner side

below; spicules on tibiae longer and more developed than in 3, and those on

hind tibiae distinctly more so.

Genital segments with about 8 relatively slender, yellowish brown to dark

or blackish brown spines on each acanthophorite, the apical last one being

NEW MYDAIDAE (DIPTERA) 139

more slender or spine-like.

Described from 10 3, including the holotype, and 4 929, including the

allotype, in the South African Museum.

Length of body: about 14-18 mm (¢9) and 15-17 mm (99)

Length of wing: about 10,5-12,5 mm (9) and 11-12,5 mm (99)

Distribution

Northern and north-eastern South West Africa: Kungveld: 48 km north

of Rooidag Gate (H. D. Brown, 7/4/1970) (3 holotype, 9 allotype, 1 3 paratype

and 1 9 paratype) ; Kungveld: 96 km north-east of Rooidag Gate (H. D. Brown,

8/4/1970) (1 3g paratype); 6 km south of Mukambo, Okavango (Okovango)

River (H. D. Brown, 17/4/1970) (1¢ paratype and 1 9 paratype); 30 km west

of Kurenkuru (Kuringkuru), Okavango (Okovango) River (H. D. Brown,

16/4/1970) (6 3 paratypes); Ovamboland: Border Beacon 24 between South

West Africa and Angola (H. D. Brown, 19/4/1970) (1 2 paratype).

This species differs from matetsiensis Beq. by the slightly darker reddish

abdomen, shorter antennae, more slender third antennal joint, distinctly much

shorter proboscis, shorter vestiture, comparatively shorter legs, shorter claws,

apically less broadly open first posterior cell in wings, and more slender and

narrower processes of sternite 9 in <.

Genus NAMADYTES Hesse

Namadytes Hesse, 1969: 278.

Namamydas Hesse, 1969: 284 (n.syn.).

The discovery of two additional species of Namadytes from South West

Africa, described below, and of which one is represented by both sexes, proves

without doubt that the 3 sex of Namadytes (unknown at the time of description)

is identical generically with the ¢ described by me as Namamydas. ‘The latter

genus thus falls away as a synonym of Wamadytes.

The genus Namadytes, now represented by 5 known species (2 by 3 only,

2 by 99 only and 1 by both sexes), all from Great Namaqualand in the southern

semi-arid half of South West Africa, may be redefined as follows:

Body with the abdomen tending to be pointed apically, more so in 99,

also rather markedly broad in basal half in 99, its dorsum on certain tergites

infused with yellowish or yellowish ochreous to a variable extent, sometimes

mainly or almost entirely yellowish; mesonotum discally sometimes with pale or

yellowish streaks; pleurae sometimes also extensively yellowish; tergite and

sternite 8 tending to be shortened.

Integument mainly dull, dull and with leathery microsculpture on mesono-

tum and metanotum; abdomen dulled towards apex, with dense, transverse,

‘nadelrissig’ puncturation in $4; terminal abdominal segments in 9° rather

coarsely, transversely, rugulosely striate or grooved; middle part of frons,

postalar calli, scutellum to a variable extent, tergite 1, and legs to a variable

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

extent shining; femora distinctly transversely finely striate.

Vestiture in some 44 fairly long and dense on head, thorax, base of abdo-

men, and legs, rather sparse in others; in 29 more poorly developed, short and

sparse on thorax above, pleurae, and tergites 1 and 2; mesopleuron with some

hairs along posterior margin in both sexes, though very sparse in 99; metanotum

bare or with scarcely discernible, very short, and fine ones on sides.

Head relatively broad, markedly broad across eyes in $3, broader than

across broadest part of thorax; interocular space on vertex very broad, much

broader than interocular space on head below, being nearly, or only a little

less than, twice distance between antennae and vertex; vertex a little sunk in;

eyes comparatively large and convex; antennae (cf. Fig. 3, left and middle;

Hesse 1969: 280, 284, figs 15-16) shortish, joint 3 about as long as, or shorter

than, rarely longer than, club, the latter elongate pyriform, more dilated

beyond middle, its apical sensory area terminal, subtruncate, or truncate,

or obliquely crater-like, sometimes appearing sunk in; proboscis much reduced,

vestigial, though the structures are still discernible in miniature, the stem being

shorter than labellar part; palps minute, not longer than the vestigial proboscis;

buccal cavity reduced and shallow.

Wings rather short, not projecting much beyond apex of abdomen, hyaline

in §¢ and with a feeble milky white sheen, in 99 usually infuscated in middle

parts, especially along the veins; alula broad, lobe-like; second submarginal

cell usually with a basal appendix; first posterior cell either opening narrowly

on costal margin or much narrowed apically and angularly subtending on it,

or with a short apical stalk (thus apparently opening on apex of second vein) ;

apex of discoidal cell either meeting posterior veins at a point or with a short

stalk.

Abdomen cylindrical in $4, slightly flattened in 99, with only 7 segments

clearly visible in 3, segment 8 not, or scarcely, visible, and only so on sides;

a projecting appendage, process or lobe (cf. Fig. 3, right AP) on each side

between lobe of tergite 8 and sternite 8 (probably arising dorsally from base

of sternite 8) present in $4 of some species; bullae comparatively small,

smaller in 99, widely separated.

Legs comparatively short, the tarsi, especially front and middle ones, also

comparatively short; tibiae slightly curved; hind femora slightly clavately

thickened, armed below with a double row of slender spines in apical half

beyond middle which become progressively longer towards the apex; spicules

on tibiae slender, bristle-like, sometimes very short, the apical ones on hind

tibiae much longer than those of other tibiae; basal tarsal joint of hind legs

slightly thickened, longer than claw-joint; pulvilli in QQ distinctly reduced,

not, or scarcely, extending beyond middle of claws, the latter in 99 also

less rapidly curved down apically.

Hypopygium of 3.3 (cf. Fig. 3) with the combined anal lobes well developed;

lateral lobes of tergite g only a little produced, only slightly lobe-like or obtuse-

angular, not sharply or angularly produced as in some other genera and species,

NEW MYDAIDAE (DIPTERA) 141

the hind margins of the lobes with markedly long bristly hairs on sides; lateral

lobes of concealed tergite 6 sometimes discernible or projecting; sternite g

inflated, rounded apically, centrally slightly grooved, not keeled, apically;

processes on sides of the sternite slightly curving inwards apically, bluntly

rounded apically and directed slightly upwards, the apical half below usually

with dense tooth-brush-like hairs; aedeagal apparatus in form of an apically-

obliquely-truncated, columnar or triquetrous process in the truncated part of

which, resembling the apex of an elephant’s trunk, there are the gonoporal

structures of the two parallel, contiguous or subcontiguous, phallic tubes, and

dorsally (anteriorly) to these the central-rod-like, slightly more projecting

egmienc (ch rig, 3. centre and right; Elesse 1969: fig. 17).

Genital segments in 992 with the last vertical genital segment markedly

shortened, the vertical carina shorter than half the length of lower margin of

acanthophorite; the latter armed with a row of rather slender spines.

The type-species of this genus is the 2 of Namadytes vansonit Hesse (Hesse

1969: 280) and the typical characters of the 3 are embodied in the generically

synonymic 4 Namamydas maculiwentris Hesse (Hesse 1969: 284, 285).

This genus resembles the North African genus Syllegomydas Beck. and

the South West African representative (subgenus Wotobates Hesse) of it super-

ficially in the much reduced and vestigial proboscis, vestiture, and wing-vena-

tion, but in the ¢ sex it may at once be distinguished by the entirely different

aedeagal apparatus. The 99 differ by the distinguishing characters given by me

(Hesse 1969: 278, 279), but chiefly by the broader interocular space on vertex,

less reduced buccal cavity and proboscis, the less consistently stalked first

posterior cell, which, if stalked, is not stalked on second vein but on costal

margin, by the much shorter vertical carina on genital segment, and by the

more reduced pulvilli.

The five known species may be separated as follows:

1(a) Males a Ae aX cae aus 2s it ie a ay so PF

(b) Females ae a Ae oA i a it ie si ie sn

2(a) Mesonotum and pleurae mainly dark or black; abdomen with more extensive dark or

black markings discally and laterally; greater part of femora and at least front and middle

tibiae darkened; hind femora distinctly more clavately thickened; pulvilli shorter, falling

short of apices of claws; antennal clubs darker or with only apical part paler, yellowish;

base of second submarginal cell with a longer appendix; lateral lobes of tergite 9 more

obtusely rounded apically; processes of sternite g relatively shorter; no projecting lobe

on each side from base of sternite 8 discernible; smaller species, about 10-15 mm long,

with a wing-length about 7,7-11 mm __.. : 3

(6) Mesonotum with the sides and 2 submedial ee pale lees Peddisny and eee

part of pleurae mainly yellowish; abdomen mainly, or more extensively, yellowish, or

with less extensive dark markings, only discally on tergites 1-3; legs entirely or mainly

yellowish; hind femora more slender; pulvilli longer, broader, reaching apices of claws;

at least apical half of clubs orange yellowish; base of second submarginal cell with a very

short appendix; lateral lobes of tergite g more sharply angular; processes of sternite 9

longer; a projecting lobe on each side from base of sternite 8 present; larger species,

about 17—17,5 mm long, with a wing-length about 12-12,5 mm .. .. & pallidus n.sp.

3(a) Larger species, about 15 mm long, with a wing-length of about 9,5 mm; abdomen above

more extensively yellowish, the apical half discally mainly or extensively yellowish; entire

142

ANNALS OF THE SOUTH AFRICAN MUSEUM

hind tibiae yellowish; tarsi mainly yellowish; vestiture distinctly longer and denser, that

on head infront not tending to be arranged awning-like in tiers, that on abdomen, especially

base, and legs much longer and denser; first posterior cell in wings with a longer apical

stalk; lateral lobes or sides of tergite g more subangular; processes of sternite g longer;

aedeagal apparatus thicker, more triquetrous .. is .. 6 maculiventris (Hesse)

Smaller species, about 10-13 mm long, with a wing-length of about 7,7-11 mm; abdomen

above less extensively yellowish, more black being evident, the apical half with yellowish

to a variable extent only discally posteriorly and across hind margins of tergites; more than

apical half of hind tibiae blackish brown like the rest; middle joints of tarsi tending to be

darkened; vestiture distinctly shorter and sparser, long only on head in front where it

tends to be arranged awning-like in tiers, that on abdomen and legs distinctly very much

shorter; first posterior cell apically subtending angularly on costal margin, or with a very

short stalk; sides of tergite 9 more rounded; processes of sternite 9 relatively shorter;

aedeagal apparatus much smaller, more columnar... .. 6 cimbebasiensis n.sp.

Clypeus more yellowish; mesonotum yellowish, with 3 broad, nearly contiguous, black

streaks; abdomen more extensively and broadly ochreous yellowish above, including

greater parts of tergites 1 and 7, only 8 entirely dark; legs paler, yellowish brown, basal

parts of tibiae more extensively yellowish; vestiture sparser, shorter, the metanotum bare,

that on legs shorter and sparser; first posterior cell in wings distinctly shortly stalked

apically; acanthophorites of last genital segment each with about 6 spines

2 cimbebasiensis n.sp.

Clypeus brownish; mesonotum mainly dark or black, only sides obscurely paler, densely

covered with greyish tomentum; leaving only 3 widely-separated dark bare streaks;

abdomen above less broadly yellowish and, if extensively yellowish, at least apical half of

tergite 1 and tergites 6-8 dark; legs darker, more brownish, only knees yellowish; vestiture

slightly denser, longer, the metanotum with some short hairs on sides, hairs on legs dis-

tinctly longer, denser; first posterior cell either broadly sessile, subtending angularly,

or narrowly open, on costal margin; acanthophorites each with about 7-9 spines .. 5

Tergites 3-5 broadly ochreous yellowish discally, to a lesser extent also on 2 discally,

central discal part of 6 reddish; hind margins of tergites 1-3 and 4 and 5 yellowish white

discally; last tergite more coarsely grooved; antennal joint 3 much shorter, it plus 1 and 2

as long as club, the latter more rapidly thickened from base; proboscis evident only as a

small globular labella; infusion in wings less conspicuous along veins; apex of first posterior

cell narrowly open; acanthophorites each with about g spines; larger species, about

18 mm long ig ee ee oe oe ue + os 2 vansoni Hesse

Tergites 3-5 yellowish discally only on posterior two-thirds; hind margins of tergites 1-4

and discally on 5 and 6 more whitish; last tergite less coarsely grooved; antennal joint 3

much longer, only a little shorter than club, the latter more gradually thickened to beyond

middle; proboscis rudimentary, but with a distinct short stem and labella; infusion in

wings more evident in middle part, darker there, more blackish brown along veins; apex

of first posterior cell angularly subtending, or sessile, on costal margin; acanthophorites

each with about 7 spines; slightly smaller species, about 15 mm long .. Q prozeskyi Hesse

Namadytes maculiventris (Hesse) n.comb.

Namamydas maculiventris Hesse, 1969: 285, figs 16-17 (n.syn.).

This species, represented by a unique ¢ from Vioolsdrif in the Richters-

veld, was described by me as the type-species of my new genus Wamamydas

which at the time was however represented by the male sex only. The subse-

quent discovery of a new species of Namadytes, represented by both sexes and

described below, shows without doubt that this 3 Namamydas maculiventris

belongs to the other new genus Wamadytes described from 99 only a few pages

ahead.

NEW MYDAIDAE (DIPTERA) 143

Namadytes cimbebasiensts n.sp.

This comparatively smallish new species* from South West Africa,

represented by both sexes collected at the same locality and on the same date,

shows that there are distinct differences between the 3 and Q as far as the

colour of the body, the vestiture, the infusions along the wing-veins, and the

length of the pulvilli are concerned and proves without doubt the generic

identity of the 99 of Namadytes and the 33 of Namamydas.

The species is characterized as follows:

Body in 3 with the head, thorax above, metanotum, pleurae, and basal

half, or greater part, of tergite 1 mainly black; discal basal patches on tergites

2-4 (or sometimes 2-7), lateral patches on these segments, sometimes entire

segments 5—7 above and below, and medial infusions on ventral sternites 2-4

blackish brown; vestigial proboscis yellowish and lower part of rims of buccal

cavity yellowish white; the following parts yellowish: shoulders to a variable

extent, extreme sides of mesonotum, postalar calli, hind border of scutellum,

pleural part below wing-bases, pteropleuron to a variable extent, metapleural

part below halteres, halteres, sides of tergites 1-4 (or 5), and sternites 1-4

(or 5); hind margins of tergites and sternites comparatively broadly more

yellowish white; bullae relatively small, blackish, widely separated; legs

brownish, the apices of femora, basal parts of middle and hind femora, bases,

posterior and apical parts of hind coxae, bases and extreme apices of tibiae,

and bases and apices of tarsi yellowish; antennae brownish, articulations of

joints 1 and 2, bases and apices of joint 3 and, to a variable extent, apical half

or part of clubs yellowish; hypopygium yellowish to pale yellowish brown.

Body in 2 mainly ochreous yellowish; antennal joints below, part of face

below antennae, and the clypeus also yellowish; mesonotum with 3 broad,

almost contiguous, posteriorly-abbreviated, black streaks; pleurae mainly

yellowish, with slight brownish infusions on meso-, sterno-, and hypopleurae,

and sternal parts in front of coxae; metanotum blackish brown; base of tergite 1,

the transverse groove across base of 2, patches on sides of tergites 2—7, and entire

tergite 8 also blackish brown, the extreme lateral margins of these tergites

being darker, almost black; venter mainly yellowish except for some slight

brownish infusions along middle; hind margins of tergites broadly yellowish

white; bullae smaller than in 3, blackish; lobes below genital segment yellowish ;

legs almost entirely pale yellowish, only the coxae in front, subapical parts of

front and middle femora above, at least apical half of hind femora above,

apical half of hind tibiae, and middle parts of rest of tibiae with a slight touch

of brownish.

Integument with the middle part of frons, face and clypeus shining; mesono-

tum and metanotum dull, leathery; pteropleuron somewhat shining; abdomen

above with the basal parts and especially basal transverse groove across tergite

2 also shining, rest of abdomen in 3 with fine ‘nadelrissig’ puncturation, in

* As coming from ‘Cimbebasia’ an old name for South West Africa.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

® smoother and somewhat shining, with a lateral intramarginal streak of

fairly coarse rugose puncturation along sides of tergites 2-5, coarser in Q;

tergites 7 and 8 in Q transversely grooved, the latter more coarsely so; legs

subshining, the femora finely transversely striate.

Vestiture denser and longer in 4, in 9 very sparse, absent from greater part

of abdomen, that on head in front in 3 long and dense, sericeous white, arranged

awning-like across vertex, sides of frons, above antennae, and across clypeus,

in 2 arranged similarly, but very much sparser, the sides of frons and face with

more evident, denser, grey tomentum; hairs on mesonotum sericeous white,

arranged more or less in 4 streaks, longer, denser, more evident in front half

and medially posteriorly, much shorter and sparser in Q; hairs on pleurae

sparse in both sexes, more so in 9, whitish, present only on mesopleuron pos-

teriorly, on pteropleuron, part of metapleuron anterior to halteres, and on

sternum in front of posterior coxae, and very sparse in 9; hairs on abdomen of 3

fairly dense, but short, long and white on sides of tergite 1, fine and short,

decumbent, located in the fine puncturation, gleaming more sericeous yellowish,

slightly longer on lateral lobes of tergite 9 and on anal lobes, sparse and short

on venter, but as dense and as long in puncturation on sternites 5—7 as above;

hairs on abdomen in @ with a few sparse longish ones on sides of tergite I, very

short and sparse and almost absent from rest of tergites except for sparse,

Fh T Hy

(47 f

Pig.

Left: Inner view of right antenna of 3 Namadytes pallidus n.sp. Centre: Right antenna of g

Namadytes cimbebasiensis n.sp. (left); hypopygial structures (centre) of 3 of same species, with

dorsal view of anal lobes (top) and much enlarged aedeagal apparatus (below).

Right: Hypopygial structures, from right side, of ¢ Namadytes pallidus n.sp., with enlarged

posterior view of the aedeagal apparatus (top).

(Ae = aedeagal apparatus; AL = anal lobes; AP = appendage or lobe of ?sternite 8; IX.S. =

sternite 9; TXT. lateral lobe or tergite 9; Pr process of sternite 9; VII.S. = sternite 7;

VILE: tergite 7; VILL: sternite 8; VIII.T. = exposed lateral lobe of hidden tergite 8.

NEW MYDAIDAE (DIPTERA) 145

short, reversed, slightly golden yellow ones on tergites 6 and 7, and sparse,

but slightly longer, ones on 8; venter in 9 bare, but with longer yellowish hairs

on lobes below acanthophorites; hairs on legs comparatively short, gleaming

sericeous yellowish in certain lights, sparser in 9, with the spines and spicules

yellowish.

Head with the interocular space on vertex broad, about as wide as distance

from apex of ocellar tubercle to apex of clypeus, appearing wider in 92 due to

the smaller and less convex eyes; interocular space on head below much

narrower, the inner margins of eyes converging below; antennae (cf. Fig. 3,

second from left) in ¢ about as long as, or a little longer than, mesonotum,

distinctly much shorter than mesonotum in Q; joint 1 about, or a little more

than, twice length of 2, not much thickened; joint 3 shorter than club, the latter

elongate pyriform, with a distinct bottle-necked base, broadest beyond middle,

obliquely subtruncate apically, the sensory area being fairly large; clypeus

roundly convex, more so in 9; buccal cavity shallow, apparently slightly

broader in 9; proboscis vestigial, rudimentary, ending in two small, rounded

lobes and a median, dorsal, finger-like lobe; palps minute, scarcely evident.

Wings not projecting, or not projecting much, beyond apex of abdomen,

the membrane markedly wrinkled, hyaline in g, but with a slight milky

whitish tint in certain lights; veins yellowish brown, but costal and subcostal

veins and those at base of wings yellowish; veins in middle part of wings in 9

bordered with fuscous; second submarginal cell with an appendix basally;

apex of first posterior cell with a very short stalk or angularly subtending on

costal margin; discoidal cell rather acute apically, meeting first and third

posterior cells at a point (or third posterior cell very shortly stalked apically) ;

halteres yellowish white, sometimes slightly darkened above near inner apical

angle.

Legs relatively rather short, slightly longer in 9; hind femora clavately

thickened, with a double row of yellowish spines below in apical half, beginning

at about just before middle, 5-6 in outer row and 5-6 (more often 5) in inner

row, on slight tubercles, those subapically at thickest part of femora the longest;

tibiae in ¢ slightly curved, but hind ones in 9 substraight, the spicules on tibiae

short, except apical ones (spurs) and especially hind ones; tarsi relatively short,

the hind ones longer, the tarsi in 9 appearing much shorter owing to the slightly

longer tibiae; basal joint of hind tarsi longer than claw-joint, much longer in

©, the basal joints of rest of tarsi shorter than claw-joints; claws more rapidly

bent down apically in ¢ than in 9; pulvilli much reduced in Q, scarcely, or not,

reaching middle of claws.

Aypopygium of 3 like that of maculwentris (cf. Fig. 3, centre; Hesse 1969: 286,

fig. 17), with the sides of tergite g not projecting much, broadly rounded

apically; no projecting lobe on each side between lateral lobe of tergite 8 and

sternite 8 discernible; processes of sternite g bent slightly upwards near apex,

their apices curving slightly inwards; aedeagal apparatus more slender than

that of maculiventris, more columnar.

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genital segments of 2 with 6 golden yellow spines on each acanthophorite.

From 3 ¢6, including the ¢ holotype, and 1 Q allotype in the South

African Museum.

Length of body: about 10-13 mm

Length of wing: about 7,7-11 mm

Distribution

South West Africa: Excelsior, No 127, Maltahohe District (H. D. Brown,

7/8/1969).

The ¢ of this species is very near the ¢ of maculwwentris (Hesse), but is

smaller, with less extensive yellowish on the abdomen above, with distinctly

less dense and shorter vestiture on body and legs, with the lateral lobes of

tergite 9 more rounded and not so subangular, and with the aedeagal apparatus

more columnar and more slender.

The @ differs from both the 99 of vansont Hesse and prozesky: Hesse in its

more extensively and broadly ochreous yellowish abdomen above, its paler

legs, shorter and sparser vestiture, bare metanotum, apically shortly-stalked

first posterior cell, and the fewer spines, only 6, on each acanthophorite.

Namadytes pallidus n.sp.

This mainly yellowish species, represented by the ¢ sex only, is charac-

terized as follows:

Body mainly ochreous yellow; sides of face and head below dark reddish

brown; frons, vertex, occipital part, and eyes black; shoulders pale yellowish

white; the following parts dark or black: three broad, longitudinal streaks on

mesonotum, not reaching posterior margin and lateral ones not extending to

shoulders, a posterior, discal infusion on scutellum, an infusion below anterior

spiracle on pleurae, continuous towards sternum with a large infusion on lower

half of sternopleuron, a spot or infusion in middle of mesopleuron, an infusion

on lower half of hypopleural part, hinder part of metanotum, basal part of

tergite 1, basal transverse groove across tergite 2, a basal, discal, M-shaped

mark on tergite 2, the bullae (which are widely separated), and a small, basal,

discal spot on tergite 3; sides of abdomen, especially on tergites 4-7, intra-

marginally with a faint brownish streak in one specimen; hind margins of

tergites and to a certain extent those of sternites fairly broadly whitish; antennae

with joints 1 and 2 yellowish brown, joint 3 and basal part, or half, of club

brownish, rest of club orange yellowish, and the junctions between joints and

extreme base and apex of joint 3 yellowish; hypopygial parts yellowish like

rest of abdomen; legs almost entirely pale yellowish, the front and middle

femora infused with brownish to a variable extent, and apices of claws black.

Integument with the central and ocellar part of frons shining; that of meso-

notum and metanotum dull and leathery; scutellum more or less shining;

pleurae dull, with leathery microtexture; abdomen above subshining with

fine, setiferous, ‘nadelrissig’ puncturation; venter more smooth and shining;

— Fr

NEW MYDAIDAE (DIPTERA) 147

legs more or less shining, finely transversely striate, especially femora.

Vestiture fairly long and dense, about as long and dense on anterior part as

in maculwentris, entirely snow white; that on head in front and on clypeus very

dense, the sides of face also with greyish tomentum; that on mesonotum even

slightly denser than in maculwentris, leaving 3 broad, bare, longitudinal streaks,

the lateral ones not reaching shoulders; hairs on sides of mesonotum and

medially in front of scutellum longer and denser than discal ones; metanotum

entirely bare; hairs on pleurae, where present on propleuron, along upper and

hind margins of mesopleuron, on metapleural part in front of and below halteres,

and on hind coxae, long and dense; hairs on tergite 1, especially laterally, to a

slightly lesser extent those on tergite 2 (more so laterally), and to a much lesser

extent sides of 3 also long and dense; decumbent hairs in puncturations on rest

of abdomen above much shorter, but also dense; those on anal lobes and pro-

jecting sides of tergite g longer again; hairs on venter sparse; those on legs

fairly dense, sericeous white, those on dorsal surfaces of femora markedly long;

spines on femora and spicules on tibiae pale yellowish white.

Head with the interocular space on vertex about subequal in width to

distance between top of ocellar tubercle to anterior rim of clypeus and much

broader than space between eyes on head below which is however relatively

proportionally broader than in maculwentris; vertex slightly sunk in; antennae

(cf. Fig. 3, left) much shorter than mesonotum, joint 1 thickened, quite three

times as long as 2, joint 3 rather stoutish, rod-like, much shorter than club;

latter a little shorter than rest of joints combined, elongate pyriform, broadest

beyond middle, its base slightly constricted bottle-neck-like and articulating

part between it and apex of joint 3 rather long, the sensory area at apex of club

slightly obliquely truncate; clypeus prominently roundly convex; buccal

cavity shallow; proboscis much reduced, rudimentary, much shorter than

antennal joint 1, evident as a short base and a short, or much shorter, bilobate

labellar part; palps comparatively less reduced than in the other species and in

one specimen quite as long as the reduced proboscis.

Wings just about reaching apex of abdomen, hyaline, with a slight milky

white tint in certain lights, especially along costal half in basal half; membrane

distinctly wrinkled; veins very pale yellowish white, those between first sub-

marginal and first basal cells, posterior vein of first, third posterior and

anal cells appearing darker in certain lights; second submarginal cell with a

very short appendix basally; apex of first posterior cell with a very short stalk

to costal margin; acute apex of discoidal cell with a short stalk of variable

length; halteres whitish.

Legs relatively slender, more so than in the other known species; hind

femora only very slightly subclavately thickened, armed below with a double

row of short spines on slight tubercles (2 or 3 in outer row and 2~4 in inner

one) in more or less apical third, the inner row beginning before the outer one;

tibiae slightly curved, with bristle-like spicules, only the apical ones (spurs)

being stoutish; tarsi with the hind ones in 4, as in the 39 of the other species

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the genus, longer than the rest, its basal joint longer than the claw-joint;

claws rapidly bent downwards apically and the pulvilli well developed, broad,

extending to apices of claws.

Hypopygium (cf. Fig. 3, right) with the anal lobes well developed; projecting

sides of tergite 9 slightly obtusangularly prominent apically; a distinct pro-

jecting lobe-like appendage or process present on each side between lateral

lobe of concealed tergite 8 and base of sternite 8 (cf. Fig. 3, right AP); sternite 9

inflated and lobe-like, rounded apically, its processes, in side view, compara-

tively less slender apicalwards and more rapidly narrowed from base below

than in the case of maculiventris (cf. Hesse 1969: 286, fig. 17), also comparatively

shorter and, in dorsal view, distinctly broader; aedeagal apparatus very similar

to that of maculiventris, only stouter.

From 2 3 specimens (holotype and paratype) in the South African

Museum.

Length of body: about 17-17,5 mm

Length of wing: about 12—12,5 mm

Distribution

South West Africa: Great Namaqualand: 48 km south-east of Keetmans-

hoop (J. G. Rozen and E. Martinez, 30/10/1968).

This species is very near maculwventris (Hesse), but, apart from the differences

already mentioned in the text, may be easily distinguished by its larger size,

extensively yellowish body, pleurae, and legs, its much longer wings, more

slender and longer legs, and presence of a distinct projecting lobe-like process

posteriorly between lateral lobe of concealed tergite 8 and base of sternite 8.

Other species of NAMADYTES

The other two known species of Namadytes, described from South West

Africa, are Namadytes vansoni Hesse (Hesse 1969: 280, fig. 15) and Namadytes

prozeskyt Hesse (Hesse 1969: 282) and both known from the Q sex only.

Halterorchini n.tribe

The new species of Mydaidae acquired by the South African Museum and

those submitted to me for description since the publication of my revision of

this family as represented in Southern Africa in 1969, and more especially

representatives of the 99 of two of the species described previously or now as

new genera in this paper from the male sex only, necessitate a new approach to

the classification of the Mydaidae.

The genus Halterorchis Bezzi, based on 99 only, occupies an anomalous

position among the genera of the subfamily Syllegomydainae and in 1969

(Hesse: 10, 17, 287) it was provisionally referred to a new tribe Syllegomydaini

which at the time was erected to accommodate a long list of genera. At that

time the 9° of Halterorchis were the only ones with a reduced number of normal

NEW MYDAIDAE (DIPTERA) 149

abdominal segments, namely only 7, the eighth being modified to form a distinct

hood over the genital segment (modified segment 9, with its acanthophorites)

and sternite 8 is also modified, partially concealed, not scoop-like. Both the

2 of a new species of Nothomydas Hesse and that of the new genus Namibimydas,

described in this paper, agree with the 99 of Halterorchis in having the same

anomalous type of abdomen and modified tergite 8 and sternite 8. It is quite

evident that this modification of the tergite and sternite is a constant character

of group value and, as the ¢4, not only of Nothomydas but also of the other two

new genera Mimadelphus and Namibimydas, described below, also have only 7

normal tergites and sternites (or sternite 7 sometimes concealed), the eighth

being entirely concealed and not discernible or even half concealed, this

abdominal anomaly, together with certain other secondary characters which

all these genera share, should be considered as of distinct tribal value.

A new tribe Halterorchini is therefore proposed to accommodate the

genera Halterorchis Bezzi (1924), Nothomydas Hesse (1969), Mimadelphus n.gen.

and Namibimydas n.gen.

This new tribe of the Syllegomydainae is characterized as follows:

Abdomen of both g¢¢ and 992 with only 7 normal, unmodified tergites

and sternites, the eighth in ¢¢ being entirely concealed and not discernible, or

with only the lateral apical part exposed as a lobe-like process; a projecting pro-

cess or lobe connected with the concealed sternite 8 or tergite 8 may be present in

some 4 (cf. Fig. 6, AP.); abdomen in 99 distinctly modified when compared

with the usual 8 normal, not greatly-modified, segments in the 99 and the half

concealed and discernible eighth segment in the ¢¢3 of the other 3 tribes;

tergite 8 in known 99 more distinctly modified in the form of a dorsal hood

over the spine-bearing acanthophorites and sternite 8 half concealed under

sternite 7, its hind margin or hinder part (cf. Fig. 7, right) indented or emargi-

nate V-like, not entirely exposed and scoop-like as in 99 of the other

tribes; tergite 9 in ¢¢ either produced apically on each side into a curved,

flattened, dorsal process which most likely participates in copulation (cf. Figs 4,

6; Hesse 1969: fig. 18 B), or the tergite is divided into 2 lobes from the apex of

each of which there extends ventrally a flattened sheet which is fused or con-

tinuous with the dorsal process on each side of sternite 9 (cf. Fig. 6); tergite

g in 99, as in genera of the two tribes Syllegomydaini and Cephalocerini,

constitutes the genital segment or central carinate ridge and spine-bearing

acanthophorites; sternite 9 in g¢ (cf. Figs 4, 6) either navicular, hollowed

scoop-like, or very much enlarged and shell-shaped, indented or sulcate

apically, with the edges of the indentation on each side raised carinately and

externally to this the apical part is produced on each side into a flattened,

curved, bluntly-pointed process, prong or a curved spine (cf. Figs 4, 6; Hesse

1969: fig. 18 B); aedeagal apparatus in ¢J, as far as this is discernible, either

rod-like, in form of a forked rod, or as a forwardly-curved process.

Other supplementary characters shared by these genera are:

Head with relatively broad to markedly broad interocular space on vertex;

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

antennae (cf. Figs 4, 5) comparatively short and stoutish, with much inflated,

pyriform clubs which are longer than the relatively short third antennal joints;

proboscis vestigial or long and very slender.

Vestiture with the hairs markedly granulated; tomentum on head, thorax,

and abdomen, especially on latter, in known 92° extensively developed.

Wings comparatively narrow, but with an Afroleptomydas-type of venation.

Legs comparatively hairy, with the apical spicules or spurs of tibiae long

or markedly long.

The four known genera may be separated as follows:

1(a) Proboscis rudimentary or vestigial; processes of sternite g more slender, spine-like; lateral

processes of tergite 9 shorter, lobe-like; body above with dense, woolly or vellose vestiture

Mimadelphus n.gen.

(6) Proboscis weil developed, long to elongate, slender, with the labella broad or spear-

blade-shaped; processes of sternite g broader, more flattened, strap-like; lateral processes

of tergite 9 either longer, more straplike or in form of a ventral, downwardly-directed,

flattened sclerite fused to processes of sternite 9; body above only hairy or densely hairy,

not densely woolly or fur-like ae : Pe age

2(a) Sternite 9 of hypopygial part much eee epetige its processes feo ie curved,

flattened, spine-like; lateral processes of tergite 9 projecting, curved, strap-like; sternite 7

normally feosesui: concealed segment 8 without a visible projecting, lobe-like appendage

on each side; legs comparatively shorter, with the front and middle trochanters small,

normal; apical spicules of tibiae and especially those of tarsi, not markedly elongate;

smaller, less densely hairy forms .. ; oie Nothomydas Hesse

(6) Hypopygial part greatly developed; aoe 9 pepe developed, Argonauta-shell-

shaped, much laterally compressed, keeled below, its processes broad, curved inwards

and then slightly outwards; lateral processes of tergite g in form a broadened, downwardly-

extending sheet united with inner part of processes of sternite 9; sternite 7 almost con-

cealed, pushed far forwards under sternite 6 by the greatly developed sternite 9;

concealed segment 8 with a distinct projecting lobe-like appendage on each side; legs

comparatively much longer, the front and middle trochanters with a flattened lobe-like,

basal extension; apical spicules of tibiae and those of tarsi conspicuously and markedly

elongated; larger, very densely-haired Asilid-like forms 7. Namibimydas n.gen.

YrY

a a

1(a) Tergite 9 with the raised, central, carinate ridge very short, less elevated, the acantho-

phorites smaller, with fewer spines, only 5 or 6, on each side, the spines slender, rod-like;

trochanters of front and middle legs small, without a lobe-like extension basally; vestiture

on body and legs less developed, less woolly, the hairs on abdomen above shorter and

sparser, directed forwards on most of the segments, and the tomentum denser; wings

much narrower; smaller forms, less than20mmlong .. ie

(6) Tergite 9 with the raised central carinate ridge longer, more ae oe anche like, the

acanthophorites with more than 6 (about 8) spines on each side, the spines much broader,

shoe-horn-shaped and hollowed above; trochanters of front and middle legs larger, with a

distinct, flattened, lobe-like extension basally; vestiture on body and legs distinctly much

denser, more woolly, that on abdomen above longer and denser and only that on last two

segments directed forwards, and the tomentum relatively less dense; wings broader;

larger forms, about 21-24 mm long 2 ue Namibimydas n.gen.

2(a) Proboscis rudimentary, vestigial ; half poised sternite ° 8 appearing carinate or sulcate

centrally, its hind margin more shallowly emarginate or indented V-shaped; hairs on

head sparser, coarser, those on mesonotum leaving 3 broad, entirely bare, tomented

streaks, abdomen above with a Ses of crescent- or []-shaped markings; bullae small

or minute .. <¢ Halterorchis Bezzi

(6) Proboscis well dev eae eee! danpate spear- ae half concealed sternite 8 more

broadly and deeply indented or emarginate (V-shaped) posteriorly, not appearing cen-

trally carinate or longitudinally sulcate; hairs on head distinctly denser and finer, those

ee ee

NEW MYDAIDAE (DIPTERA I5!1

on mesonotum denser, finer, leaving narrower, not entirely bare, streaks: abdomen above

with fimer, shghily demser, short hairs, and only partly densely tomen t

peculiar pattern; bullae larger -. ae ef fe - .. Nothom

iS)

“ep

Genus NoTHOMYDAS Hesse

Nothompdas Hesse, 1969: 290

This genus, which is now placed in the new tbe Halterorchini and of

which the orginal description was based on a single J specimen from the

South West African side of the Orange River at Vioolsdrif, has since the

publication of the description, been augmented by representatives of a closely

related new species. The latter, composed of $j and a 2, was collected at

Port Nolloth in Namaqualand. To complete the description of the genus, the

diagnostic characiers of the © representative, as well as some supplementary

characters of the J3 of the new species, are as follows:

Head much broader than thorax; interocular space on vertex moderately

broad, broader than inierocular space on head below, in J quite 12-12 as

wide, in © the space below is relatively broader relative to the mterocular

space above; eyes markedly large m 3, markedly convexly globular, smaller

in 9; antennae as described and figured in 1969 (Hesse: 290, 294, fig. 18 B),

but clubs m known © more globular; ceniral ocellar ridge on frons prominent

and shining in both sexes; clypeus short, convex; buccal cavity relatively deep,

in both sexes; rest of cephalic structures as described in 1960.

Wings comparatively shori in both sexes, not reaching tp of abdomen,

comparatively narrow; first posierior cell ather narrowly opening on second

vem or more broadly so and thus appearing very shortly stalked or sessile on

costal margin; rest of wing characiers as described in 1969 for the type-species.

Abdomen of 3 3 as described for the type-species, but the modified lobe-like

extension on each side of the partly hidden tergite 9 either strap-like or narrower

and spme-like; abdomen m known © broader, more dorso-venitrally depressed,

also with 7 normal iergites and siernites, but partly hidden tergite 8 hoodlike

over genital segment (iergite g), and sternite 8, as in O° of the other Halter-

orchini, parily hidden, its hmder part indented V-shaped, the sides or limbs of

the V somewhat inflated.

Flypopygium of 33 as described and figured for the type-species in 1069

(Hesse: 293, fig. 18 B).

Gemial segment, or modified tergite 9, of ° as described for O° of Halter-

orchini above, with a short, raised, ceniral, carinate ndge and a spine-bearing

acanthophorite on each side, and medially below these well-developed anal

lobes.

Legs relatively shori and as described for the genus in 1960, but front and

middle femora with or without fine, brisile-like spmes along inner surfaces

below; front and middle tibiae with or without short, fine, bristle-like spicules,

more distinctly present in 2; hind tibiae with a row of spicules along outer part

below, these m © longer and more strongly developed; crown of spicules at

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

apices of tibiae rather markedly long, especially the hind ones; spicules on tarsi,

especially on hind ones, also rather long; claws as described for type-species,

but pulvilli sometimes narrower.

Integument of head with the central frontal ridge, head below and to a

certain extent clypeus shining; pleurae either entirely shining or in part dull;

integument of abdomen either mainly shining above or mainly dull in 34, dull

discally above in 9, in 39 finely setiferously punctured above; integument of

rest of anatomy as described in 1969.

Vestiture in $3 as described for 3 of type-species, but in known Q shorter

and sparser on front half of body, and fine hairs on abdomen above directed

apicalwards on tergites 2-7 and sternites 3~7, those on venter much shorter

and sparser than in 44; hairs on legs either short or long and dense, especially

on front and middle ones, slightly less so in known Q, otherwise as described for

the type-species in 1969.

From other Halterorchini this genus differs by the characters given in the

key to the known genera.

The type-species 1s Nothomydas gariepinus Hesse and the other species is

Nothomydas namaquensis n.sp. described below.

Nothomydas namaquensts n.sp.

This species, which was collected on the coast more to the south of the

Orange River in Namaqualand, is very near the type-species gariepinus from

which it differs in the following respects:

Body mainly black in 3, in ¢ garepinus the postalar calli, abdomen above,

and to a variable extent the pleurae are more obscurely reddish brown; 2

of new species with the sides of mesonotum, humeral region, 2 submedial, dis-

cal, posteriorly-abbreviated, narrow streaks, the postalar calli, and discal part

of scutellum yellowish brown, the pleural parts, metanotum, tergites 1 and 2,

discal parts of 3 and 4, and venter to a variable extent also yellowish brown to

reddish brown; narrow yellowish white hind margins of tergites and sternites

in g more sharply defined than in gariepinus; humeral tubercles in ¢ dark,

more yellowish in 9, markedly tomented greyish white in both sexes; antennae

in ¢ of type-species more yellowish brown, in 3 of new species darker brown to

blackish brown, paler yellowish brown in Q; legs in ¢ of type-species yellowish

brown, the front and middle femora appearing dark above, and tibiae and

tarsi more yellowish, in ¢ of new species the legs are darker brownish to blackish

brown, and slightly paler, more yellowish brown in Q.

Integument as described for the genus; that of abdomen in 3 more exten-

sively shining than in ¢ of type-species, the dull, to a variable extent greyish-

white-tomented parts less extensive, the hinder half of abdomen tending to be

more shining, the abdomen above, apart from setiferous puncturation, also

more finely transversely striated; in 9 greater discal parts of tergites to tergite

5 dull and greyish white tomented, the sides more coarsely transversely striated,

and intramarginal, longitudinal depression along tergites 2—5 rugulose; legs,

NEW MYDAIDAE (DIPTERA) 153

especially femora, distinctly finely transversely striated in both sexes, and hind

ones more coarsely so basally and apically and slightly more so than in type-

species.

Vestiture in ¢ distinctly longer and denser, but also mainly white on greater

part of head, body and legs; that on head distinctly much longer and denser on

vertex, behind antennae, on clypeus, and sides of buccal cavity, that on vertex

and frons with a slight yellowish tint in certain lights, that on occiput and head

below denser than in type-species; dense snow-white hairs along notopleural

part, posterior margin of mesopleuron, pteropleuron and metapleural part as

dense, but slightly longer; dense white ones on sides of tergite 1 and sides

basally of 2 also slightly longer, appearing slightly more creamy yellowish in

certain lights; fine, short, decumbent hairs in small punctures on abdomen

above distinctly darker, more blackish, not so gleaming sericeous yellowish, and

the longer ones along sides of abdomen above also darker, more brownish;

hairs on venter very slightly denser than in type-species, darker, gleaming more

brownish or blackish brown, and those on hypopygium as long and as dense,

but gleaming more brownish; coxae, as in the type-species, mainly smooth

below, but with the longish white hairs on sides of hind coxae denser; hairs on

legs, especially on femora above, and more especially front and middle ones,

conspicuously longer and denser, but also snow white, those on hind femora

above with more long ones than in type-species, composed of whitish and dark

ones, and those below also denser, sometimes gleaming slightly more yellowish

brown; hairs on tibiae, especially front and middle ones, also much longer and

denser, also snow white, those on hind tibiae with more numerous long ones

along inner aspect, and rest of short hairs on hind tibiae distinctly darker, more

brownish or blackish brown.

Vestiture in 9 also mainly snow white, slightly shorter than in J, but also

long and dense on head, humeral tubercles, notopleural part, hind margin of

mesopleuron, on pteropleuron, metapleural part, and sides of tergite 1; fine,

short, decumbent hairs in small punctures on abdomen above from tergite 2

apicalwards sparser than in 4, directed forwards and gleaming pale sericeous

yellowish or whitish; hairs on venter also sparser, gleaming more sericeous

whitish and directed forwards on sternites 3-7; hairs on legs distinctly less

dense, the long ones sparser, and all the hairs on femora and tibiae gleaming

sericeous whitish; tomentum in @ denser, greyish white, more extensively

present, especially on head, humeral tubercles, metanotum, ptero- and meta-

pleurae, hinder part of hypopleuron, and broadly and conspicuously on

abdomen discally above.

Head with the interocular space on vertex comparatively broader than in

type-species, as 7:6, and space on head below proportionally also broader, the

interocular space on head below in Q relatively broader than in gy as 5:4;

antennae proportionally shorter than in gariepinus, slightly shorter in Q than in

6, shorter than width of interocular space on vertex in both sexes, slightly

longer than this width in the type-species, with the joints proportionally also

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

shorter than in the type-species, the inflated club, apart from its apical process,

less truncated apically, the process itself more conically prominent; pro-

boscis, though slender and similarly shaped (cf. Hesse 1969: fig. 18 B, top)

distinctly much shorter, relatively shorter in 9 than in 3, only about 2-2,

4 mm long (as against 3 mm in type-species), with the labella slightly more

rounded apically in 3 than in Q; palps distinctly shorter and thicker than in

type-species.

Wings clear hyaline as in the type-species; veins yellowish brown to

brownish; second submarginal cell without any appendix; first posterior cell

broader apically than in type-species, not sharply narrowed, more angularly

subtending, or only shortly stalked, on costal margin, not so distinctly stalked

apically as in type-species; axillary lobe relatively narrower; knobs of halteres

slightly darker and darker in 3 than in 9.

Legs proportionally much shorter, the tibiae also curved, but basal part of

hind femora slightly more curved; front femora without any distinct spines

below (in 3 of type-species there are a few slender spines); hind femora in 3

with fewer and shorter spines below, only about 2 or 3 distinct ones in an apical

outer row and 2 in the apical inner row, but in 9 with 2 or 3 apically on inner

aspect and 5 or 6 along outer aspect from about middle; spicules on hind tibiae

in ¢ shorter than in ¢ of the type-species, in 2 longer and stouter than in J;

tarsi proportionally much shorter, the hind ones shorter, stouter and thicker

in ¢ than in Q, its basal joint proportionally longer; claws proportionally much

shorter than in type-species, the pulvilli narrower and apparently slightly

shorter, more so in 9.

Hypopygium of 3 similar to that of the ¢ of type-species (cf. Hesse 1969:

fig. 18 B), but differs in having the curved, modified lobes or extensions of

tergite g distinctly shorter, slightly narrower, less broadly strap-like, and with

the hind margin of the truncated apex even more incised or emarginate; curved

prongs or processes of sternite 9 comparatively shorter, more or less subequal

in length (from basal bend to apex in side view) to dorsal length of tergite 7

(in type-species it is distinctly longer than tergite 7).

Genital segments of 2 as described for the genus, with 5 detectable, slender,

reddish golden spines on each acanthophorite, the third from base appearing

longer, and the last two more slender; hind margin of sternite 8 indented

U-shaped; tergite 8 with separated, setiferous punctures, and exposed sides of

the sternite transversely substriate.

From 1 ¢ holotype, 4 4 paratypes, and 1 9 allotype in the South African

Museum.

Length of body: about 9-10 mm

Length of wing: about 5,5-7 mm

Distribution

Namaqualand: Port Nolloth (South African Museum Expedition, October

1967).

NEW MYDAIDAE (DIPTERA) 155

MIMADELPHUS n.gen.

A single ¢ specimen from South West Africa submitted to me is referable

to the new tribe Halterochini and is very near and in the same category as the

genus WNothomydas. As certain, often slight, morphological differences, such as

antennal characters, reduction of the proboscis, differences in venational

characters of the wings, the degree of the development of the vestiture, and

differences and anomalies in the structure of the genital structures of the 34,

have been found to differentiate genera in the Mydaidae, the occurrence of

such differences in the 3 specimen concerned points to a distinct and separate

generic status for it.

The ¢ specimen of this new genus differs from §¢ of Nothomydas s. str.

in having a much reduced and rudimentary proboscis, more deeply sunk in

vertex, very much denser, fur-like vestiture on body, sparser and shorter hairs

on each side just behind antennae, absence of hairs along hind border of

mesopleuron, hairs on mesonotum separated by 3 distinct, longitudinal, bare

streaks, very much shorter anal lobes, shorter, curved, lobe-like extensions of

tergite g (cf. Fig. 4), longer, navicular sternite 9 with its longer, more slender,

sharply-pointed, spine-like processes, and a very much smaller, shorter, aedea-

gal, rod-like structure which is not bifid or forked apically.

The probability that this § may represent the unknown ¢ of Halterorchis

Bezzi, which was described from the 2 only and which also has a vestigial, or

much reduced, proboscis and no hairs across hind border of the mesopleuron,

is not excluded. The fact that antennal joint 3 is much longer, the alula of wings

is much larger and more lobe-like, the third posterior cell is much more acute

apically and its posterior vein more in line with that of first posterior cell, and

the legs are proportionally shorter, excludes it from Halterorchis.

The type-species of this new genus is Mimadelphus vellosus n.sp.

Mimadelphus vellosus u.sp.

Body mainly black; antennae pale yellowish brown, the apex of joint 3

and apical part of clubs darker, more blackish brown; clypeus showing brownish

through the hairs; buccal cavity and vestigial proboscis pale yellowish brown;

following parts also yellowish brown: shoulders, postalar calli, hind border of

mesopleuron, area below wing-bases, including most of pteropleuron, sutural

parts between sterno- and hypopleurae, sides broadly of tergite 1, extreme sides

of tergite 2, to a more obscure extent extreme sides of rest of tergites, and genital

structures; sides of tergites 2—7 broadly and venter slightly more reddish brown;

hind margins of tergites on sides more yellowish; sutures between tergites 1 and

2 and 2 and 3 more reddish; bullae shining black; legs with the coxae dark

brownish, their apical parts more yellowish, the femora yellowish brown,

darker above, subapically, paler below and apically, the tibiae more yellowish,

hind ones darker apically, the tarsi brownish above, more yellowish below;

claws yellowish, black-tipped.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

AL eee MULTE

. ‘

IT ee Te

AN Speen S27 Oia

Py E So | 2

oo” scat 7 ~

Fig. 4.

Left: Right antenna of § Mimadelphus vellosus n.gen.et n.sp. Middle: Side view of apex of abdo-

men and hypopygium of same species.

Right: Ventro-posterior view of hypopygial structures. (Ae = aedeagus; AL = anal lobes;

IX.S. = sternite 9; 1X.T. = lateral lobe of tergite 9; Pr = process of sternite 9; VII.S. = ster-

nite 7; VII.T. = tergite 7; VIII.S. = part of hidden sternite 8.)

Integument of central frontal part, clypeus, greater part of pleurae, hinder

part of scutellum, sutural parts between tergites 1 and 2, sides of abdomen

broadly, venter, and legs shining; mesonotum duller, with fine, microscopical,

leathery texture; metanotum dull, and a broad, central, dorsal, partially-bare

and partly dark-haired, black streak on abdomen above also dull.

Vestiture on head and body above dense, fur-like, vellose, reminiscent of

that of the genus Agaperemius Hesse; that on sides of frons, clypeus and sides

of face very dense, long, directed downwards, snow white; sides of frons

anteriorly also with whitish tomentum; hairs on frons on sides medially just

behind antennae sparser than in Nothomydas, erect and white; those on vertex

anteriorly erect, white, and a dense cluster on each side of occipital part partly

on vertex dense, white, directed more inwards and backwards; hairs discally

on mesonotum not very dense, very slightly pale sericeous yellowish, arranged

in 2 rows in anterior half, separated by a central, bare streak and by a

sublateral, anteriorly-abbreviated, bare streak on each side, these 3 streaks

coalescing behind middle to form a broad, discal, bare area; hairs on

sides of mesonotum and on shoulders very dense, snow white, becoming

sparser above wing-bases and across base of mesonotum; scutellum with

2 backwardly-directed, snow white tufts; metanotum with greyish white

tomentum and no hairs; pleurae mainly bare, the hinder part of ptero-

pleuron and metapleural part with snow white hairs, and the propleuron

with some scattered white hairs; hairs on abdomen above granular, very dense,

vellose, fur-like, mainly decumbent, those on tergite 1 submedially and on sides

anteriorly more erect, snow white, as are also those on extreme sides of 2 and 3,

and to a much lesser extent on extreme sides of rest of tergites; those across

hind margin of tergite 1, on greater part of 2 discally, broadly on sides of 3-7

above more decumbent, directed backwards, dense, more creamy whitish,

leaving a broad, but posteriorly-narrowing, central, black or dark streak, mainly

covered with black, decumbent, granular, scale-like hairs, and with some

scattered, pale ones, especially across hind margins of tergites; sides anteriorly

NEW MYDAIDAE (DIPTERA) 157

of tergites 3-5 also appearing dark, but covered with dark, fulvous brownish

hairs; tergite 2 discally on each side also with a submedial, dark-haired patch;

hairs on venter slightly less dense, whitish on sides of sternites, especially 1-4, the

rest of the hairs appearing slightly fulvous in certain lights; hairs on legs mainly

pale, long, dense and white on femora above, especially so along inner upper

apical half of middle femora, and basally above on hind ones, and along hinder

parts of front and middle tibiae; rest of hairs on legs shorter, gleaming more

pale sericeous yellowish in certain lights; spines on hind femora below and

spicules on hind tibiae more pale yellowish to pale fulvous yellowish, the rest

of spicules on tibiae more whitish.

Head across eyes broader than thorax; eyes markedly large, convex;

interocular space on vertex broader than interocular space on head below, as

30:20; vertex sunk in; antennae (Fig. 4, left), very close together, much shorter

than thorax, joint 1 thickened, about 14-12 times length of 2, only a little

longer than broad, joint 3 comparatively short, only a little longer than 1 and

2 combined, slightly thickened apically; club dilated, pyriform, somewhat

laterally compressed, in side view broadest subapically much beyond middle,

a little longer than rest of the joints combined, its apical sensory area, around

the crater-like tubercle, sunk in, the tubercle itself rather long and prominent,

quite as long as antennal joint 2, the inner margin of sensory area and inner

apical part of club somewhat emarginate and depressed; buccal cavity small;

proboscis much reduced, only about 0,32 mm long; palps not detectable.

Wings rather narrow, the membrane wrinkled, entirely hyaline, but with

a scarcely perceptible milky whitish tint in certain lights; costal vein, subcostal

and anterior vein of anal cell yellowish, the rest of veins yellowish brown;

second submarginal cell with a very short appendix; first posterior cell slightly

narrowed apically, opening on second vein (or very shortly stalked on costal

margin); discoidal cell sharply acute apically and shortly stalked; alula fairly

well developed, almost quadrate; halteres yellowish.

Legs comparatively short and hairy, as described under vestiture; hind

femora slightly clavately thickened, armed below with a single row of 5 or 6

distinct spines from a little before middle to a little beyond middle, ceasing

before broadest part of femora, those towards the apex the longest; front and

middle femora subspindle-shaped, without any spines; front and middle tibiae

curved, the spicules on anterior aspect longer, more hair-like on middle ones

where the hairs on posterior part are also longer as they are on upper, posterior,

apical part of middle femora; hind tibiae only slightly curved in apical part,

its row of spicules (4) along lower outer aspect fairly long and stoutish, spine-

like, its crown of apical spicules also longer and stouter than those on other

tibiae; tarsi rather short, progressively longer from front tarsi to hind ones,

the spicules below hind ones also much longer and stouter, the hind basi-

tarsus a little longer than claw-joint; claws curved down apically; pulvilli

well developed.

Hypopygium (cf. Fig. 4, middle and right) as described for the genus;

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

navicular sternite g with some stoutish setae on slight tubercles ventrally and

anterolaterally; aedeagal apparatus comparatively short, rod-like, not forked

apically.

This unique 3 holotype is in the Stuttgart Museum.

Length of body: about 11 mm

Length of wing: about 7,5 mm

Distribution

South West Africa (H. Walter, 1/3/1953). (Without any reference to

geographical region, but judging from the distribution of species of the related

genus Nothomydas, probably also from some sandy desert part in Great

Namaqualand. )

NAMIBIMYDAS n.gen.

Two specimens, a g and a 9 and both somewhat damaged, from the Namib

Desert, submitted by Professor E. Lindner, represent a remarkable new genus

which differs from all known genera of Mydaidae, not only in the peculiar

genital structures of the ¢, but also in other morphological characters.

As the abdomen in both sexes has only 7 normal discernible tergites and

sternites and the usual eighth tergite and sternite of other genera concealed,

reduced or modified, this genus is also to be placed in the new tribe

Halterorchini.

It is characterized as follows:

Body moderately large, elongate, asilid-like; abdomen stoutish, elongate,

cylindrical in 3, slightly dorso-ventrally depressed up to penultimate segment

in 9.

Head (cf. Fig. 5) markedly broad, much broader than thorax, broader

than in most other genera; interocular space on vertex proportionally very

broad, displacing the eyes laterally to a very marked extent, proportionally

broader in 3 than in 9; interocular space on head below more or less equally

broad in both sexes, the inner margins of eyes in ¢ thus more convergent below;

eyes proportionally large for the broad head, larger in ¢ than in Q, the facets

along anterior inner aspect only slightly larger than the rest, slightly larger in

3; median ocellus present; antennae of 9 missing in the specimen, but in g

comparatively short and stout, joint 1 slightly thickened, joint 3 columnar,

broadened apically, the club elongate pyriform, longer than joint 3 as in the

other Halterorchini, its sensory area however more latero-apical; clypeus

prominent; medial lower part of buccal cavity rather prominently jutting out

lip-like; proboscis very characteristic, spear- or lance-shaped, very long and

slender, like that of species of Cephalocera, the labella of 3 narrower, lance-

blade-shaped and that of 2 more spear-blade-shaped (cf. Fig. 5, right and left) ;

palps distinct, thickened apically.

Thorax with the integument, including that of the pleurae, dull; pro-

pleural part rather inflated, prominent, the central prosternal sulcus rather

NEW MYDAIDAE (DIPTERA) 159

deep and distinct throughout; mesopleuron with hairs on upper posterior

part; metapleuron dull, with hairs.

Wings well developed, with strong veins; venation like that of most Sylle-

gomydaini and especially that of the genus Afroleptomydas; alula well developed,

lobe-like.

Abdomen with 7 normal, unmodified, unconcealed tergites and sternites

in 9 and also 7 tergites in g, but only 6 discernible sternites in 3, sternite 7

being concealed and displaced forwards by the greatly developed sternite g

(cf. Fig. 6); tergite 8 in § concealed, only partly exposed on sides as a lobe at

base of tereite 9 (cf. Fig. 6, left, VIII.T.), in 9 as in other genera of the new

tribe, modified to form the hood over the genital segments; sternite 8 in J

concealed or not developed and, in 9, partially or half concealed above hind

margin of sternite 7 and modified to form a ventral plate the hind margin, or

Dorsal view of head of g of Namibimydas gaerdesi n.gen. et n.sp.

Left: Proboscis of 2 of same species.

Top right: Apical part of proboscis of 3.

All drawn to the same scale.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

hinder part, of which is indented or emarginated V- or U-shaped (cf. Fig. 7,

right); tergite 2, as in other genera, broadly transversely depressed across base;

bullae on segment 2 minute in Q.

Legs (much damaged in the specimens, some of the legs and tarsi missing)

in both sexes however fairly long; trochanters (cf. Fig. 7, left) of front and middle

legs distinctly produced more lobe-like in outer basal part than in most genera,

and slightly smaller in 9 than in J; hind femora slightly thicker than the other

femora, armed with a double row of spines below; front and middle femora also

with some spines below; tibiae with long and conspicuous spicules, those in the

apical crown (spurs) markedly and conspicuously long as are also those on

tarsi (probably an adaptation to the sandy and dune-sand environment) ;

claws and pulvilli very strongly developed in 3, less so in 9, in which sex the

pulvilli are also relatively shorter.

Vestiture well developed, with markedly dense, long, bushy, and finely

granulate hairs on head (cf. Fig. 5), sides of mesonotum, metanotum, hind

margin of mesopleuron, on pteropleuron, metapleuron, base of abdomen, to a

lesser extent on rest of abdomen, hypopygium of J, on coxae, and femora;

hairs on disc of mesonotum slightly shorter; those on abdomen above in 92

shorter and less dense beyond tergite 2; tomentum on head and body well

developed in both sexes.

Hypopygium of 3 (cf. Fig. 6) remarkable, large; tergite 9 deeply indented

U-like, divided dorsally into a lateral part or lobe on each side, with the well-

developed anal lobes in between, and each lateral lobe depressed apically, the

depression extending ventrally as an infolded sheet which is connected with

the dorsal process or prong on each side of sternite 9; the latter enormously

developed, Argonauta-shell-shaped, laterally compressed, ventro-laterally

grooved on each side, keeled below, the entire structure displacing sternite 7

far forwards under sternite 6 so that only 6 abdominal sternites are discernible;

sternite g itself centrally sulcate in posterior (or apical) vertical part, and con-

nected on each side with a broad, flattened, dorsally slightly hollowed, apically

slightly outwardly- and downwardly-directed, bluntly rounded process or

prong which curves apicalwards from about middle of the sternite and each

of which, on dorsal apical part of the sternite, is connected to the latter by

more transparent chitin; tergite 8 almost entirely concealed, only an exposed

lobe-like lateral part discernible, a projecting appendage or lobe (cf. Fig. 6, AP.)

on each side of sternite 9 may represent an appendage or process on each side

of the concealed sternite 8 as in some species of Namadytes; aedeagal part

entirely hidden, discernible from a posterior view as a forwardly-bent hook,

from the base of which there extend apicalwards 2 filament- or strand-like

structures (cf. Fig. 6, right top); hairs on hypopygium dense and long on

extreme sides of tergite 9, below anal lobes, on processes of sternite 9, posteriorly

on latter, and on each side of concealed tergite 8 (projecting from under tergite

7), with the hairs posteriorly on hypopygium curved inwards and downwards

hiding posterior vertical part of sternite 9.

NEW MYDAIDAE (DIPTERA) 161

Genital segments of 2 (cf. Fig. 7, right) as in other genera of the Syllegomy-

dainae, but with tergite 8 in form of a hood over the terminal, central, carinate

ridge and acanthophorites; sternite 8 however half hidden under sternite 7,

its hind margin indented, V-shaped, and with the sides of the V swollen and

rugose; central carinate ridge of sternite 9 well developed, arched; acantho-

phorites each with comparatively short, broad, shoe-horn-shaped, blunt spines,

hollowed dorsally.

The type-species of this new genus is Namibimydas gaerdesi n.sp. which

I have great pleasure in naming for Herr F. Gaerdes of Okahandja who col-

lected it, who is well known as a keen and enthusiastic collector of insects in

South West Africa and who, from his extensive private collection, has contri-

buted much interesting insect material from that part of southern Africa to

museums in South Africa and in Germany.

Namibimydas gaerdesi n.sp.

This interesting species is characterized as follows:

Body-colour mainly black, with the lower apical half of antennal club

appearing obscurely reddish, the sensory area more yellowish brown; proboscis

black, its extreme base, lower part of buccal cavity, and the palps yellowish;

thorax with the anterior spiracular area behind humeral tubercles, pleural

part just below spiracle, the fossa between propleural and prosternal parts,

sutural part above between humeral tubercles and mesonotum, notopleural

part, outer part of postalar calli, middle part of hind margin of scutellum,

scleritic area around wing-bases (partly encroaching upon or embracing also

mesopleural, pteropleural, and metapleural parts), sutural parts on rest of

pleurae and, in 9, also 2 central streaks on metanotum reddish yellow to pale

yellowish brown; these coloured parts, in 9, tending to be more extensive,

affecting even greater part of pleurae which tends to be more reddish brown;

disc of mesonotum in 9 also tending to show an obscure central and an obscure

lateral reddish brown streak on each side; abdomen above mainly dark or

black, in 9 tending to be more dark brownish, becoming paler, more yellowish

brown, posteriorly, the sides in both sexes also more yellowish or pale brownish,

more so in 9; hind margins of tergites yellowish white on sides, more broadly

so across posterior lateral angles; bullae black in both sexes, small in 9, very

widely separated, surrounded by pale yellowish brown; venter yellowish

brown, the hind margins of sternites 1-4 in 9 and 1-6 in ¢ broadly yellowish

white; genital structures of 9 pale yellowish brown; extreme sides of tergite 9

and narrow hind margin of its lateral part in g, as well as greater part of

hypopygial structures also pale yellowish brown, excepting posterior, shining-

black, lateral and vertical parts of sternite 9; a small spot near posterior apical

angle of lateral part of tergite g in ¢ yellowish; legs mainly yellowish in both

sexes, the coxae in g, excepting the broadish pale yellowish brown hinder part,

dark or blackish, the front and middle femora on upper outer aspect to beyond

middle, and inner face of hind femora darkened, more brownish, especially in

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

g, and apical parts of tarsal joints also darkened, and apical parts of claws

black.

Integument of head and body mainly dull, covered with greyish white

tomentum; sides of tergites, especially in 9, more subshining, transversely

rugulose and punctured, tergites 6 and 7 more extensively transversely rugulose;

transverse basal depression across tergite 2 shining; hood-like tergite 8 in 9

more shining, more coarsely punctate-rugose, as are also the semi-inflated

hinder parts of sternite 8 in 9; modified tergite 9 in 9 coarsely punctured; venter

shining, transversely grooved or wrinkled laterally, more so in Q; sternite 9

and its dorsal processes in ¢ shining, its lower keel-like part, from just before

middle, longitudinally striate; legs shining, the femora finely transversely

striate.

Vestiture on head, body and legs mainly white, that on head long and

dense, that on frons in front, antennal joints 1 and 2, clypeus, and genae more

straw-coloured, more so in 9; postvertical spines distinct, slender, bristle-like,

yellowish; hairs on occipital part laterally behind eyes white, shorter than rest

of cephalic hairs; palps with a tuft of long white hairs; base of proboscis with a

pencil of long white hairs; hairs on mesonotum slightly less dense and shorter

than rest of hair on thorax, but denser and slightly longer in ¢ than in 9, witha

faint straw-coloured tint in certain lights, inclining to be arranged in 5 streaks,

corresponding with more or less 5 streaks of whitish tomentum, of which the

Fig. 6.

Side and posterior views of hypopygium of ¢ Namibimydas gaerdesi n.gen. et n.sp. The posterior

view is drawn with the vestiture only on one side.

(Ae = aedeagal structure as far as this can be made out in specimen, the top right is a side

view with all the parts not discernible; AL = anal lobes; AP = appendage of concealed segment

8; IX.S. = Argonauta-shell-shaped sternite 9; IX.T. = tergite 9; Pr = processes of sternite 9;

VIS. = stermte 6s VIET, = tergite 7 VILL. — tensite oe)

NEW MYDAIDAE (DIPTERA) 163

central one is narrow, the 3 middle hair-streaks, separated by 2 almost bare

streaks, more evident in g; tuft on humeral angles, hairs along notopleural

part, on postalar calli, and in front of base of scutellum dense, long, and snow

white; those on metanotum also long, snow white; pleurae mainly bare,

tomented, but hairs on upper hinder part of mesopleuron, on pteropleuron,

and metapleural part in dense snow white tufts; those on sternal parts also

long and snow white; hairs on abdomen long, snow white, long and dense

basally on sides of tergites 1 and 2 and, in J, fairly long and dense on rest of

tergites, below anal lobes, along upper lateral part of processes of sternite 9,

sides below processes, posterior part of sternite 9, and on sides of concealed

tergite 8; hairs on abdomen in @ shorter and sparser beyond tergite 2, slightly

denser and directed forwards on last 2 tergites; hairs on venter much longer

and slightly denser in 3, shorter and sparser in 9; hairs on legs also mainly

white, long and dense on femora above, much more so in 4, those in tibiae

shorter, but longer in ¢ than in 9.

Head (cf. Fig. 5) distinctly much broader than thorax, broader in 3 than

in 9, quite 5,75 mm in ¢g and 4,96 mm in Q; interocular space on vertex

markedly broad, proportionally broader in 3 (+3 mm) than in 9 (+ 2,2 mm);

vertex on the whole not much sunk in, but central ocellar ridge ends on vertex

in a fairly deep sulcus, the boss-like part on each side very prominent; antennae

(missing in the 9 specimen) nearly as long as width of interocular space on

vertex in 4, with joint 1 thickened, quite 14 times length of 2, joint 3 stout,

columnar, longer than combined length of 1 and 2, shorter than club, its

apical part broadened; club elongate pyriform as shown in Figure 5; clypeus

convex, somewhat shining, its apex slightly notched; buccal cavity fairly deep,

its lateral margins sharply carinate, the middle part on lower margin pro-

jecting prominently ledge-like, more broadly so in 4; proboscis slender,

markedly elongate, javelin-shaped in ¢ (Fig. 5, right), spear-shaped in 9

(Fig. 5, left), about 4,64-4,68 mm long, the labella narrow in ¢ and lance-

blade-shaped, slightly broader and spear-blade-shaped and more sharply

pointed in 9; palps in § more roundly inflated apically, in 9 more subspindle-

shaped.

Wings reaching tip of abdomen, clear, hyaline, with a faint milky whitish

tint in certain lights; membrane much wrinkled; venation similar to that of the

genus Afroleptomydas; veins rather stoutish, yellowish; first posterior cell slightly

narrowed apically, either opening on second vein in 9 (thus very shortly stalked

apically on costal margin) or partly opening on second vein and partly on

costal margin in ¢ (thus apically narrowly sessile on costal margin); second

submarginal cell with an appendix basally; discoidal cell acute apically,

meeting third posterior cell at a point; axillary lobe broad; alula well developed,

lobe-like; halteres whitish.

Legs (much damaged in @ specimen and some tarsi missing in 3 specimen)

moderately long, hairy, the hairs on front and middle femora above and,

especially, along inner upper aspect of hind femora markedly long and dense,

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

more so in 4, and the hairs on tibiae also longer in 3 than in 9; front femora

with 3 yellowish spines below in 3 and 2 pallid ones in 9; middle femora more

slender, unarmed below; hind femora slightly thickened, with a double row of

reddish yellow spines below (4—5 in inner row and 6—7 in outer one) on slight

tubercles, the outer row ending in 3 or 4 more closely-spaced and shorter ones

along outer margin of the apical femoro-tibial articulating cavity; tibiae with

more or less a double row of markedly long yellowish red to reddish spicules

along outer lower part, ending in a crown of conspicuously long apical ones;

tarsi also with markedly long reddish spicules on joints below, the basal joint

of hind tarsi longer than claw-joint; claws strongly developed, more so in g;

pulvilli in ¢ strongly developed, broad, about reaching curved apices of claws,

in @ distinctly less strongly developed, narrower, shorter than in 3, not reaching

apices of claws.

Hypopygium of 3 (cf. Fig. 6) as described for the genus.

Genital segments of Q (cf. Fig. 7, right) as described for the genus; acantho-

phorites each with about 8 spines of which the first basal spine is slender and

more rod-like and the other 7 (only 2 on each side in the specimen are still

present, the slightly hollow bases of attachment of the lost ones however indicate

the original total) are broad, shortish, bluntly rounded, cuneiform or shoe-

horn-shaped, and hollowed above; hinder V-shaped emarginated part of

semi-concealed sternite 8 thickened or inflated, rugulose, and sparsely punctured

ike Be

1 vA ee pene i

siete t7/s

Left: Ventral view of coxa, trochanter and part of femur of middle right leg of g Namibimpdas

gaerdesi n.gen. et n.sp.

Right: Posteroventral view of genital segments of 2 Namibimydas gaerdesi n.gen. et n.sp.

(Ac = acanthophorite; AL = anal lobes; Ca = central carina (ventral and side views) of last

Sp = spines of acanthophorite; ‘Tr trochanter; VII.S. Stennmute 7-3 V Lin. tergite 7;

VINES. = stermmite ss Vili — terote a.)

NEW MYDAIDAE (DIPTERA) 165

on each side, each puncture lodging a short hair.

From the 1 g holotype and 1 9 allotype in the Stuttgart Museum.

Length of body: about 21-24,5 mm

Length of wing: about 16-19 mm

Distribution

South West Africa: Namib Desert: Walvis Bay (F. Gaerdes, 24/1/1939

(Q) and 20/1/1940 (3)).

The ecology and biology of this interesting genus are unknown, but the

various morphological features of the type-species, such as its excessive hairi-

ness, the hairy legs with the markedly long tibial and tarsal spicules, and the

comparatively broad, shortish, shoe-horn-shaped spines on the acanthophorites

of the female, indicate adaptive responses to a hot and sandy environment.

Subfamily Mydainae

PARECTYPHUS n.gen.

One of the specimens of Mydaidae from South West Africa, kindly sub-

mitted to me by Professor E. Lindner of the Stuttgart Museum, belongs to a

new genus of Mydainae and very near to the genus Ectyphus Gerst. Up to now

the genus Ectyphus appears to be restricted to the southern and eastern Cape,

Natal, Transvaal, and possibly East Africa. It is an eastern genus of which the

species occur in semi-wooded and forested parts. This new genus may be

considered as representing Ectyphus in the Namib Desert of South West Africa.

Though not stated on the locality label this mydaid, like Ectyphus, probably

also inhabits a wooded environment, even in the treeless Namib, namely that

found along the banks of the Kuiseb River.

It differs from the genus Ectyphus in the following respects:

Body comparatively and markedly broader, the head and thorax propor-

tionally much broader, the abdomen in the male sex less cylindrical, more

flattened dorso-ventrally. Its general appearance suggests that of a cricket.

Integument very similar to that of Ectyphus, but the disc of the mesonotum

more finely leathery or rugulose, not so coarsely rugulose and also less dull;

that of abdomen above smoother, though also setiferously punctured and with

an intramarginal streak of coarse puncturation on sides of abdomen.

Vestiture with the hairs on head in front, thorax above, and on pleurae

in the § on the whole distinctly longer and denser than in even the most hairy

species of Ectyphus known.

Head markedly broad, only slightly broader than across broadest part

of the broad thorax, the latter proportionally less broad in Ectyphus making the

head appear much broader; interocular space on vertex proportionally much

broader, much broader than distance from base of ocellar ridge on vertex to

base of clypeus which distance, in Ectyphus, is equal, subequal to, or even a

little shorter than, interocular space; interocular space on head below pro-

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

portionally also broader; clypeus flat-nose-shaped as in Ectyphus; proboscis

also reduced and short, resembling that of Ectyphus; antennae (cf. Fig. 8, left)

proportionally shorter, much shorter than mesonotum, the clubs more pyri-

form, not so elongate-pyriform or spindle-shaped as in Ectyphus, the apical

sensory area and apical prominence less produced.

Wings with 3 submarginal cells, not 2 only as in Ectyphus, the normal second

has the basal appendix (which is present in all known species of Ectyphus) joined

to the posterior border vein of the marginal cell thus dividing the first sub-

marginal cell into 2 cells and thus forming the 3 submarginal cells, of which the

third is an enclosed cell; first posterior cell broadly opening on costal margin

apically.

Legs as in Ectyphus, but hind trochanters without any spines or processes;

hind femora also markedly thickened, but more clavately so, their apices more

narrowed, armed below in broadest subapical part and also sublaterally in

apical part with stoutish spines on tubercles; front and middle femora unarmed

below; front and middle tibiae armed with longer, stouter, spine-like spicules

than in Ectyphus, and apically with much longer apical spicules; hind tibiae

centrally carinate below as in Ectyphus, the carina also ending apically in a

blunt process, but the spicules on hind tibiae more developed and with 2 long

apical spicules on outer side of apical tibial process; tarsi with much stiffer

short hairs and longer spicules; claws more strongly developed and longer, and

the pulvilli well developed.

Abdomen markedly broad and flattened dorso-ventrally, even in 3, more

so than in some 36 of Ectyphus, otherwise as in latter genus; tergite 8 concealed,

but sternite 8 exposed as a scoop-like sclerite below and taking the place of

the normal sternite 9 of most Mydaidae; lateral lobes of tergite 9, unlike the

ES is

Fig. 8.

Right antenna of $ Parectyphus namibiensis n.gen. et n.sp.

Middle: Left view of posterior end of abdomen and hypopygium of same species.

Right top: Dorsal view of left lobe of tergite g and anal lobes of same species.

Right bottom: Posterior view of hypopygium.

(Ae = aedeagus; AL = anal lobes; IX.T. = lobes of tergite 9; PL = plate bearing process (Pr.)

of hidden sternite 9; VII.S. = sternite 7; VII.T. = tergite 7; VIII.S. = sternite 8.)

NEW MYDAIDAE (DIPTERA) 167

much rounded or subangular, half-concealed, lateral lobes of Ectyphus, well

developed, broad, prominent, and produced spinelike posteriorly on each side

(cf. Fig. 8); sternite 9 not discernible, probably modified and in part repre-

sented by the transverse, shield-like plate (cf. Fig. 8, Pl.).

Hypopygium (cf. Fig. 8, middle and right) of ¢ differs from that of 33 of

Ectyphus in having the process or prong on each side of the concealed sternite 9

not curved inwards or tongs-like, but produced posteriorly on each side as a

downwardly-bent hook, without any projecting appendage below, the base

inside of each hook, from a posterior view, arising from a transverse plate

(cf. Fig. 8, right below) which in part may represent sternite 9; aedeagal

apparatus columnar and, as far as can be seen in the unique ¢ specimen, in

form of a downwardly-directed, slightly laterally-compressed tube (cf. Fig. 8

Ae), without any cowl-like or inverted cup-like cap as in Ectyphus.

This new genus of South African Mydainae, with 3 submarginal cells

in the wings, alters my definition of the subfamily (Hesse 1969: 9, 19, 364, 365)

in which it was stated that the wings have only 2 submarginal cells and that

the second opens either on the costal margin or on the second vein, and that a

posterior cross vein is present or not.

In this new genus there are 3 submarginal cells, the second opens, as in

Ectyphus, on the second vein, and a posterior cross vein is also present.

To accommodate this new genus and to correct statements as to the

characters of the known African genera of the Mydainae the following revised

part of the key to the subfamilies and genera of African Mydaidae is proposed.

1(a) Wings with 2 or 3 submarginal cells, the third, if present, formed by bisection of the first

by the extension of the basal appendix of the second .. 2 (Subfamily Mydainae Beq.)

(6) Wings with 3 submarginal cells of which the second is in form of an enclosed and apically-

stalked cell between the first and third .. Subfamily Dzochlistinae Beq. (Non-African)

2(a) Second submarginal cell opening on second vein and with an appendix, or extended and

bisecting appendix (vein) at base; hind border of wings with a distinct cross vein; clypeus

broad, convex, flat-nose-shaped; antennal joint 3 much longer than 1 and 2 combined;

hind femora markedly thickened, incrassate, usually with more than 2, usually 4, rows

of spines below; hind tibiae compressed, sia carinate below, the carina ae in an

apical process or spine in gg iM 3

(b) Second submarginal cell opening iinet on “easel aenein. seeil 7 thew a paca

appendix; hind border of wings without a cross vein; clypeus not flat-nose-shaped; antennal

joint 3 short, or very short, shorter than 1 and 2 combined; hind femora usually less

markedly incrassate, with fewer, or only 2, rows of peed below; hind tibiae usually

without a carina below 4

3(a) Wings with only 2 submarginal elle the base of econ fii an 7m appeadlix: Read andl body

narrower, the abdomen in §¢ more cylindrical or subcylindrical; hind trochanters with

I or 2 spines below; spicules on tibiae shorter, less developed; lateral lobes of tergite 9

in gd less produced, more rounded, lobe-like, or only subangular; processes of sternite 9

in gg curved more inwards, blunter, with a distinct palp-like appendage below; aedeagal

apparatus stoutish, conical, ending apically in a cowl-like cap; terminal part of oviscape

in 9 usually withdrawn into segment 7 .. ae Ectyphus Gerst.

(b) Wings with 3 submarginal cells of which the third (enclosed) i is foamed by the bisection of

first by the basal appendix of the second joining anterior vein of normal first sub-marginal;

head and body markedly broad, the abdomen in g broader, more dorso-ventrally flattened ;

hind trochanters without spines; spicules of tibiae more developed, longer; lateral lobes

of tergite 9 in ¢ well developed, prominent, produced spine-like; processes of sternite 9

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

directed posteriorly, bent down hook-like apically, without any projecting appendage;

aedeagus columnar, slightly laterally compressed, without a cowl-like cap

3 Parectyphus n. gen.

4(a) Antennae shorter, the clubs normally dilated or inflated, not constricted at middle, not

appearing deformed Ae .. Rhopalia Macq. (from Egypt)

(b) Antennae longer, the clubs ssleneedl coms gied at middle, their basal part broadened

or extending lobe-like laterally, or with lobe-like extensions, appearing deformed or

malformed .. So: a. i Ae: a Pen eacean Gerst. (North African)

The type-species of the new genus Parectyphus is Parectyphus namibiensis n.sp.

Parectyphus namibiensts n.sp.

The 3 holotype of the type-species is characterized as follows:

Body mainly black; antennae blackish brown, the clubs more yellowish

brown, becoming paler in apical third, but apical tubercles blackish; buccal

cavity and proboscis yellowish brown; anterior thoracic or shoulder spiracles,

sides of mesonotum above wing-bases, and postalar calli more reddish brown

and so is the area below wing-bases; hind margins of sternites 2 and 3 slightly

yellowish brown; hypopygial parts yellowish brown; legs with the femora dark

reddish black, the knees yellowish brown, the front and middle tibiae and tarsi

also reddish black, the hind tibiae more dark reddish brown, their tarsi also

reddish black; claws yellowish brown, black-tipped.

Integument of central part of frons and clypeus smooth and shining; that

of mesonotum dull, finely rugulose; scutellum dull, finely rugulose, its hind

border, posterior to a slight transverse depression, finely longitudinally rugu-

lose; metanotum dull, finely rugulose, but with a few transverse rugae, especially

on sides; pleurae mainly shining, but posterior border of mesopleuron, parts

of sterno- and hypopleurae, opposite middle coxae (pore region), upper part

of pteropleuron, and to a certain extent metapleural part, wrinkled; inflated

lateral part of tergite 1 longitudinally striated; abdomen above transversely

depressed just behind base of tergite 2, rest mainly smooth, but not too densely

setiferously punctured, denser so posteriorly, but discal part of lateral lobes of

tergite g mainly smooth, though showing fine, shallow, transverse striation in

certain lights; venter smooth, shining, with sparse, setiferous puncturation in

basal half, becoming denser posteriorly, sternite 7 with some transverse rugae

posteriorly and 8 more coarsely transversely rugose and setiferously punctured;

legs mainly shining, the femora transversely striate, the hind ones more coarsely

so basally.

Vestiture with very dense and long, silvery-gleaming, snow white hairs on

head, those on sides of frons in front and on clypeus directed downwards

awning-like, also with whitish tomentum on sides of frons, face, and occiput;

hairs on vertex and less dense and shorter ones on head below also silvery-

gleaming snow white; vertex without any distinguishable, post-vertical spines

or setae; first antennal joint bare, the second with a crown of a few hairs;

hairs on mesonotum arranged in punctures also entirely silvery-gleaming snow

white, but much shorter than on head, denser and longer only along noto-

NEW MYDAIDAE (DIPTERA) 169

pleural part and in front of scutellum, discally absent from more or less 5

longitudinal streaks, the middle 3 more evident, hairs also absent from postalar

callar region and along middle of scutellum; metanotum bare; hairs across

hind margin of mesopleuron, on pteropleuron, sides of metasternum, outer

sides of middle and hind coxae, tergite 1, especially sides, and on anal lobes,

long and silvery-gleaming snow white; those on rest of abdomen above not very

dense and in setiferous punctures, those on lateral lobes of tergite 9, sparse ones

on venter, dense, stouter and longer ones on sternite 8, those on prosternal

part, front coxae, on discal parts of middle and hind coxae, and on legs, inclu-

ding the spines and spicules, dark or black.

Head with joint 1 of antennae (cf. Fig. 8, left) only slightly thickened, only

about 1,4 times length of 2, joint 3 only a little more than twice length of 1

and 2 combined, slightly thickened in the joint-like apical part; club slightly

longer than joint 3 (as 6:5), dilated pyriform, its base slightly constricted neck-

like, broadest at about apical third, with short, scattered, black hairs, arranged

more or less transversely and more on outer side from about basal third to

apical third, the apical part of club subtruncate on inner aspect and with the

tubercular prominence on outer apical aspect rounded, crater-like and with a

short central style; clypeus narrowly separated from inner margins of eyes;

lower middle part of buccal cavity rather prominent, lip-like; proboscis shorter

than antennal joint 3, its labellar part subequal in length to rest or basal stem,

covered with fine, short spinules; palps minute.

Wings projecting a little beyond apex of abdomen, hyaline, with a very

faint milky white tint, the membrane much wrinkled; veins yellowish brown,

more yellowish in basal part, the costal vein appearing dark due to the short

black setae along its course to about level of first posterior cell; venation as

described for the genus; alula broad, lobe-like; first posterior cell broadly

opening on costal margin; halteres dark, the knobs black.

Legs with the hind femora clavately thickened, slightly curved basally,

broadest part at about apical third and, apart from the rather stiff setae-like

and short hairs, are armed below with 2 rows of stoutish spines on slight tuber-

cles, mainly concentrated in apical broadest part, the outer row beginning

as isolated spines from near base, the inner row from about or beyond middle,

in addition there are an outer lateral row of 3 or 4 spines in apical part, a few

irregularly-situated ones on inner lateral aspect, and a few dorsal ones in apical

part or half; front and middle femora covered above with rather long stiff

hairs and longer, setae-like hairs, the middle femora more curved than front

ones; front and middle tibiae with fairly dense, stiff, black hairs and long,

spine-like spicules, the apical ones below markedly long; hind tibiae with much

finer, shorter, and sparser hairs, denser towards apex, and with fewer, shorter,

and finer spicules, the apical ones on outer side of apical tibial process longest;

apical process on hind tibiae below only moderately long and stout; tarsi

with dense, stiff, black hairs and long spicules, the front and middle tarsi

shorter than their tibiae and also shorter than hind tarsi, the latter also shorter

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

than their tibiae and with the basal joint longer than the claw-joint (minus

claws); hind claws longer than the others, and all claws sharply bent down

apically.

Hypopygium (cf. Fig. 8, right) as figured and described for the genus.

The unique ¢ holotype is in the Stuttgart Museum.

Length of body: about 15 mm

Length of wing: about 12,5 mm (the tips of both wings broken off

in the specimen).

Distribution

South West Africa: Gobabeb (23.3 S. 15. O E. 408 m.). This locality

name is not shown on maps, but is about 112 km south-east of Walvis Bay on

the Kuiseb River in the Namib Desert and, at present, is the site of the Namib

Desert Research Station.

SUMMARY

In this paper, an addendum to my ‘Revision of the Mydaidae (Diptera)

of southern Africa’, three new genera and sixteen new species from the Namib

Desert and other parts of South West Africa are described. The genus Wamamp-

das Hesse, described in 1969, is sunk as a synonym of Namadytes Hesse and a

key is given to separate the known species of the latter. A new tribe Halter-

orchini is erected to accommodate the anomalous genera Halterorchis Bezzi,

Nothomydas Hesse, and the two new genera Mimadelphus and Namibimpdas.

A short descriptive key for the separation of these genera is appended. A revised

key to the subfamily Mydainae, to include the new genus Parectyphus, is also

given. In 8 text-figures certain genital structures which help to distinguish

some of the new species and the more important structures of the new genera

are illustrated. A short list of the more important bibliographical references is

added.

ACKNOWLEDGEMENTS

In the preparation of this paper my thanks are due to the following persons

for kindly submitting valuable specimens for identification and description,

or for collecting Mydaidae in South West Africa and generously donating the

material to the South African Museum: Professor E. Lindner of the Staatliches

Museum fiir Naturkunde in Stuttgart who very kindly submitted the repre-

sentatives of the new genera and species housed in the said museum; Herr

F. Gaerdes of Okahandja in South West Africa who collected some of these

specimens; Dr H. D. Brown of the Department of Agricultural Technical

Services in Pretoria who, in his pursuit of Orthoptera in South West Africa,

found some time to collect most of the new species of Afroleptomydas and to

present them to the South African Museum; Dr J. G. Rozen and Mr E.

Martinez of the American Museum of Natural History who collected one of the

NEW MYDAIDAE (DIPTERA) 171

new species in the southern part of South West Africa and kindly presented it to

the South African Museum; and to Mr F. Gess of the Albany Museum who,

while still on the staff of the South African Museum, collected two new species

at Port Nolloth and on the South West African bank of the Orange River

respectively.

REFERENCES

BEQuAERT, M. 1959. Diptera (Brachycera). Jn HANSTROM, B., BRINCK, P. & RUDEBECK, G.,

eds. South African animal life. 6: 356-372. Stockholm: Almqvist & Wiksell.

BEQUAERT, M. 1961. Contribution a la connaissance morphologique et la classification des

Mydaidae (Diptera). Bull. Inst. r. Sci. nat. Belg. 37 (19): 1-18.

BreQuarErT, M. 1963. Contribution nouvelle a connaissance des Mydaidae de la région éthio-

pienne. Mém. Inst. r. Sci. nat. Belg. (2) 71: 1-68.

Bezz1, M. 1924. The South African Mydaidae (Diptera) as represented in the South African

Museum. Ann. S. Afr. Mus. 19: 191-232.

Hesse, A. J. 1969. The Mydaidae (Diptera) of southern Africa. Ann. S. Afr. Mus. 54: 1-388.

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for diological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins, arranged in the following order:

(1) Heading, consisting of informative but brief title, name(s) of author(s), address(es) of

author(s), number of illustrations (figures, enumerated maps and tables) in the article.

(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-

headings to be used sparingly and enumeration of headings to be avoided. (4) Summary

(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

ILLUSTRATIONS

To be reducible to 12 cm x 18 cm (19 cm including caption). A metric scale to appear

with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their

proper place in the text), with arabic numbering; items of composite figures to be designated

by capital letters (A, B, C etc.).

REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

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

Fiscuer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

FiscHer, P.-H., Duvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.

Archs Zool. exp. gén. 74: 627-634.

Konun, A. J. 19604. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region

of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Koun, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

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

Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature issued

by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).

The Harvard system of reference to be used in the synonymy lists, with the full references

incorporated in the list at the end of the article, and not given in contracted form in the synonymy

list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, 6; Liste: 11. Turton, 1932: 80.

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NEW MYDAIDAE (DIPTERA)

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60 Band

November 1972 November

Part 2) Deel

SEIrOCENE MARINE INVERTEBRATES FROM

LANGEBAANWEG, CAPE PROVINCE

BRIAN KENSLEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

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Wi T=, 5» 7s b=\D!0))- 24(2), 27; Bile), BS)

Price of this part/Prys van hierdie deel

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Trustees of the South African Museum © _ Trustees van die Suid-Afrikaanse Museum

1972

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In Suid-Afrika gedruk deur

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Courtweg, Wynberg, Kaap

PLIOCENE MARINE INVERTEBRATES FROM LANGEBAANWEG,

CAPE PROVINCE

BRIAN KENSLEY

South African Museum, Cape Town

(With g figures)

[MS. accepted 12 Fune 1972]

CONTENTS

PAGE

Introduction ‘ : ; : : = 78

Systematics : ‘ P : : A

Discussion ; ; ; : : ea iol7/

Brief ecological comments . : : 3) aLGS

Summary : : : : : Sites)

Acknowledgements. ‘ : ; LOO

References : : : : : =) 190

INTRODUCTION

The deposits which are being commercially exploited for their phosphate

content at Langebaanweg, 105 kilometres north-north-west of Cape Town,

are of palaeontological significance mainly because of the unique assemblage

of Pliocene mammals which they have yielded (Hendey 1970a). Non-mamma-

lian vertebrate remains are not uncommon (e.g. Simpson 1971), but until

recently invertebrate fossils were rare, although the occurrence of marine

molluscs in the deposits had been noted (Hendey 1970a). During 1971 two

assemblages of marine invertebrates were recovered from ‘E’ quarry, and it is

this material which is here described.

The present paper is in the nature of an inconclusive report, mainly on

the molluscs. In some cases specific identification is given with almost no

uncertainty. In several cases, however, the generic and even the familial

position of the specimens is in doubt. Where these have been speculated upon,

it is quite possible that with further and more complete material becoming

available, the taxonomic position of the specimens will be altered. In some

cases, certainty may never be reached, as structures such as radulae and oper-

culae, which are essential for an accurate identification, are not preserved.

Whether further material will in fact become available is doubtful, considering

the difficulties involved in obtaining fossiliferous phosphatic matrix, and in

removing the fossils from this extremely resistant matrix.

The first series of specimens (South African Museum L14187) were

recovered from an exposure near the south-western corner of ‘E’ quarry. These

Uae

Ann. S. Afr. Mus. 60(4), 1972: 173-190, 9 figs.

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

fossils, which are very well preserved and show almost no sign of being beach-

worn, were embedded in a cemented, fine-grained brown phosphate rock that

was adhering to an older, water-worn and heavily indurated phosphate rock.

A few isolated shark’s teeth (Odontaspis sp.) occurred in association with the

invertebrates.

The second assemblage (S.A.M. L14259) was recovered from a trench in

the eastern part of the quarry (BDT 2/1971), and these specimens, of which

only internal casts and external impressions were preserved, came from a con-

solidated non-phosphatic light-coloured silt that was also adhering to boulders

of heavily indurated phosphate rock.

At both localities the indurated phosphate rock occurs as rounded water-

worn boulders, cobbles, and pebbles, and these together with the deposit con-

taining the invertebrate fossils, occur in a loose and non-phosphatic sand and

silt. The latter deposit also contains the remains of marine vertebrates

such as sharks, teleosts, and whales, and grades upwards into a non-marine

deposit from which most of the ‘E’ quarry terrestrial fossils are recovered.

SYSTEMATICS

CEMENTED PHOSPHATIC MATRIX (L14187)

Phylum MOLLUSCA

Class GASTROPODA

Order ASPIDOBRANCHIA

Family Patellidae

Cellana capensis (Gmelin)

Cellana capensis: Barnard, 19636: 315.

Description

Shell somewhat domed, with apex anterior to midpoint, circumference

narrower at anterior end. Apex eroded in all specimens. Sculpturing consisting

of very fine radiating ridges, sometimes slightly granular, often reaching almost

to apex.

Material

Numerous examples of external and internal casts. Several incomplete

specimens up to 29 mm in diameter. S.A.M. L14187/A.

Remarks

Both sculpturing and general proportions, which have been assessed in

some cases by means of silicone casts taken from external casts of shells, agree

well with living material. The characteristic colour pattern of the species is

naturally entirely lacking. The species occurs alive on the east coast of southern

PLIOCENE MARINE INVERTEBRATES FROM LANGEBAANWEG 175

Africa from East London to Mocambique, Malagasy, and has also been recorded

from Pakistan.

Patella granularis Linnaeus

Patella granularis: Koch, 1949: 503. Barnard, 1962: 161.

Material

Several internal casts and several incomplete specimens up to 48 mm in

diameter. S.A.M. L14187/B.

Remarks

The domed shape of the larger specimens and the fine radiating ribs

bearing low blunt spines characterize this species, which has been recorded

from the Pleistocene of the Namaqualand coast and the Saldanha Bay area.

The species occurs alive from southern Angola to Natal.

Family Fissurellidae

Diodora parviforata (Sowerby)

Risa &

Diodora parviforata: Barnard, 1962: 191; 19630: 294.

Description

Shell oval in circumference, narrower at posterior end. Foramen small,

circular, situated posterior to and slightly below apex. Sculpture consisting of

numerous radiating lirae reaching almost from apex to circumference; in

upper region growth lines form cancellate pattern with lirae.

Material

One complete specimen, longer diameter 22,8 mm, shorter diameter

15,6 mm, altitude 11,0 mm. §.A.M. L14187/C.

Remarks

No differences can be detected between the present specimen and living

material taken from the west coast. The species has been recorded from the

Pleistocene of the Alexander Bay/Port Nolloth area, and living from the west

coast of the Cape Peninsula to Port Alfred.

Fic 1. Diodora parviforata.

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Haliotidae

Haliotis saldanhae n.sp.

Pigs 92

Description

Outer lip of shell margin slightly convex. Four or five foramina present on

distinct ridge between upper surface of body whorl and region between foramina

and columella ridge. Distinct groove between columella and rest of shell. Body

whorl with about 15 smoothly-rounded spiral lirae, some stronger than others.

One or two lirae present between larger foramina, nine or ten lirae between

foramina and columella, two stronger than rest. Faint growth-lines visible.

Several strong smoothly-rounded undulations at angle of about 40° to growth-

lines.

Material

HOLOTYPE: an almost complete specimen, 87 x 63 mm, S.A.M. L14187/D,

2 incomplete and eroded specimens, one with entire body whorl, 117 x 88 mm.

From the second assemblage (see p. 186) a complete internal cast, agreeing

well with present species 74 X 53 mm, S.A.M. L14259/D.

Remarks

Halwotis saldanhae bears some resemblance to both of the present-day species

of Haliotis which occur around the Cape. In general shape, H. saldanhae more

closely resembles H. midae which is a broader shell than H. sanguineum. The

outer lip margin is slightly convex as in H. midae, rather than slightly concave

as in H. sanguineum. The foramina of the present species would seem to be fewer

in number (4 or 5) and larger than in H. midae or H. sanguineum of comparable

size (8 or 9). The most obvious difference between H. saldanhae and the two

present-day species lies in the nature of the sculpture; H. sanguineum possesses

spiral lirae both above and below the foramina (but these are often evanescent

in large specimens). This spiral sculpture is rather similar to the present

material where it is somewhat stronger. H. sanguineum lacks any sculpture other

than growth-lines and spiral lirae. H. midae characteristically lacks spiral

sculpture, but possesses strong oblique undulating raised ridges, whereas the

i, ae Bs

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KE . ae say wks v N\

; NN

Fic. 2. Haliotis saldanhae n.sp.

PLIOCENE MARINE INVERTEBRATES FROM LANGEBAANWEG Ae

present species possesses more widely-spaced oblique undulations, which never

reach the raised condition of H. midae. A groove is present between the columella

and the body whorl in H. saldanhae, as in H. midae. This is never found in H.

sanguineum, where the columella merges smoothly with the rest of the shell.

H. tuberculata of the Mediterranean and West Africa is a more elongate

species, possesses numerous spiral striae, and lacks any oblique sculpture other

than strong growth-lines.

As the present species cannot satisfactorily be reconciled with any fossil or

living species of Haliotis, it is afforded separate specific status. H. saldanhae could

perhaps be regarded as ancestral to H. midae, given a loss of spiral sculpture, or

its obliteration by the oblique undulations, which would become stronger and

more closely spaced. The specific name ‘saldanhae’ is derived from Saldanha

Bay, the Pliocene shore having been an extension of this bay.

Family Trochidae

Oxystele tigrina (Chemnitz)

Oxystele tigrina: Barnard, 1962: 191; 1963): 267.

Description

Shell of four whorls, convex in profile, apex damaged or eroded in all

specimens. Umbilicus closed. Outer shell layer almost smooth, but broken

away in several cases, exposing 10-12 spiral lirae. Latter more distinct in

upper whorls, second whorl with seven to nine, third with fewer.

Material

Numerous examples ranging in diameter from 10,0 mm to 31,5 mm

(altitude difficult to measure due to damage). S.A.M. L14187/E.

Remarks

Because of the strength of and similarity in the number of the spiral lirae,

the present material is thought to be closest to O. tigrina. The material also

agrees in general proportions, i.e. altitude to diameter ratio, with this species.

The possibility exists that the smaller specimens may belong to a different

species, but these too agree well with living material of similar size of O. tigrina.

The latter has been recorded from the Pleistocene of the Algoa Bay area, and

occurs alive from Saldanha Bay to Natal.

Family Phasianellidae

Tricolia neritina (Dunker)

Tricolia neritina: Barnard, 1963): 210.

Material

Numerous examples, largest specimen with diameter of 4,5 mm. S.A.M.

L14187/F.

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

No differences can be detected between the present material and living

specimens, which occur from Liideritzbucht to Algoa Bay.

? Family Littorinidae

? Genus LITTORINA

Fisz 2

Description

Shell squat, profile evenly convex, with simple protoconch of one whorl,

plus 34—4 postnatal whorls. Aperture broadly oval. Sculpturing either entirely

absent, or with very faint spiral lirae visible on body whorl.

Remarks

The present species, with its lack of characters, could be placed in several

families with equal lack of certainty, but would seem closest to Littorina

africana in overall proportions, in the lack of a strong ridge on the body whorl,

and in the faint spiral lirae. There is some superficial resemblance to Phasianella

capensis, but the greater height of the spire in larger specimens of the latter

species seems to discount this identification.

Material

Numerous examples ranging in altitude from 2,5 mm to 7,0 mm. S.A.M.

L14187/G.

Fic. 3. ? Littorina sp.

Family Muricidae

Ocenebra scrobiculata (Dunker)

Tritonalia scrobiculata: Barnard, 1959: 212.

Material

One specimen, 6,3 mm altitude. S.A.M. L14187/H.

Remarks

The characteristic cancellate sculpture formed by strong spiral lirae,

with nodules formed where these are crossed by axial ridges, agrees perfectly

PLIOCENE MARINE INVERTEBRATES FROM LANGEBAANWEG 179

with the shells of living examples of the species. The nodules do not show the

squamae found in the ‘crawfordi’ form of the species. 7. scrobiculata has been

recorded alive from Saldanha Bay to Natal.

Family Thaididae

Thais dubia Krauss

Thais dubia: Barnard, 1959: 221; 1962: 190.

Material

One specimen, apex damaged, altitude 17 mm, diameter 11,0 mm.

S.A.M. L14187/I.

Remarks

The general form and proportions, the extent of the callus, shape of the

columella and the aperture, along with the faint spiral sculpture and obscured

axial and growth lines agree well with many specimens of living material of

this species. 7. dubia has been recorded from the Pleistocene deposits of the

Namaqualand coast, Knysna, and the Algoa Bay area.

Family Nassariidae

Bullia sp.

Material

Several damaged specimens and casts. Largest example altitude 26,8 mm.

S.A.M. L14187/J.

Remarks

The present species belongs to the group of slender species in the genus,

and is very close to B. digitalis. Unfortunately, as a callus is not visible on any

of the specimens, further identification is not possible.

Family Turridae

‘Crassispira’ sp.

Fig. 4.

Description

Protoconch 14 whorls, five postnatal whorls. Basal whorl bearing 15-16

slightly oblique smooth axial ridges, those on outer lip reflecting slight sinuosity

of outer lip near suture. Fourth whorl with 12-13 axial ribs, reaching from

suture to suture, third whorl with 9-11 axial ribs. Latter difficult to distinguish

on earlier whorls due to matrix. Two or three very faint spiral lirae visible on

base below axial ribs.

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 4. ‘Crassispira’ sp.

Material

Two complete specimens, altitude 8,9 mm and 7,6 mm, diameter 3,7 mm

and 3,5 mm respectively, one with damaged apex. S.A.M. L14187/K and A.

Remarks

In general proportions, number of whorls, and in the number of axial

ribs per whorl, the present material agrees well with Crassispira hottentota (see

Barnard 1958: 120, and Kilburn 1970: 40). Shells of C. Aottentota have been

collected from False Bay to East London.

‘Clavatula’ sp.

Fig. 5

Description

Shell of six whorls plus protoconch. Outer lip with broad, moderately

deep sinus. Sculpturing consisting of oblique, smooth, axial ribs, 12 on basal

whorl, 12-13 on fifth whorl, thereafter obscure. Spiral lirae faintly visible on

base.

Fic. 5. ‘Clavatula’ sp.

PLIOCENE MARINE INVERTEBRATES FROM LANGEBAANWEG 181

Material

Two complete specimens, altitude 9,4 mm and 9,1 mm, diameter 4,9 mm

and 4,5 mm respectively. $S.A.M. L14187/L.

Remarks

The generic position of the present material, as with the previous species,

is speculative, only the shell being available. Nevertheless, the character of the

outer lip sinus suggests a species of Clavatula close to C. sinuata and its variety

sigillata (see Barnard 1958: 99).

Naas sp.

Description

Shell evenly conical, aperture less than half total length. Five whorls

plus protoconch. No axial sculpture, basal whorl with 8-10 strong spiral lirae,

following whorls each with seven spiral lirae.

Material

One specimen, base damaged, altitude 10,0 mm, diameter 4,4 mm.

8.A.M. L14187/M.

Remarks

With only the shell available, the generic position of the present specimen

cannot be verified. The specimen is, however, reminiscent of several species of

the genus Turris, especially T. cingulifera, which it resembles in the spiral

sculpture and the aperture shape.

Order OPISTHOBRANCHIATA

Family Pyramidellidae

Turbonilla krausst Clessin

Turbonilla kraussi: Barnard, 1962: 190; 1963a: 84.

Description

Shell slender, tapering, of seven whorls, apical whorl absent. Each whorl

bearing numerous smooth axial ridges, about 20 on base, 18-20 on previous

whorl.

Material

Two specimens, altitude 5,0 mm and 4,8 mm, diameter 1,9 mm and 1,7 mm

respectively, S.A.M. L14187/N.

Remarks

Because of the small size of the specimens and the hardness of the encrusting

matrix, an accurate count of the axial ribs is difficult. Nevertheless, the speci-

mens agree well with living material of similar size. The species has

been recorded from the Pleistocene of Sedgefield and the Algoa Bay area, and

alive from Saldanha Bay to Natal.

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

? Family Pyramidellidae

? Genus PYRAMIDELLA

Fig. 6

Description

Shell elongate, smoothly conical, protoconch of one whorl, six postnatal

whorls, profile of latter very slightly convex, sutures well marked. Aperture

wide, columella simple, bearing well-defined callus. Two or three spiral lirae on

base, rest of shell lacking sculpture.

Material

Numerous complete specimens up to altitude 9,0 mm, diameter 4,9 mm.

§.A.M. L14187/O.

Remarks

In the simple columella and the general proportions the present material

most closely approaches the genus Turbonilla, although the number of postnatal

whorls (6) is low for the genus. The lack of sculpturing is also not characteristic

of the genus, while several species of Pyramidella such as P. ima Bartsch, and

P. hera Bartsch, do lack both axial and spiral ornamentation. None of the species,

however, has spiral lirae on the base, as occurs in the present material.

Fic. 6. ? Pyramidella sp.

Order PULMONATA

Family Siphonariidae

Stphonaria sp.

Bis 7

Description

Circumference of shell oval, outer surface bearing eight or nine strong

radial ribs, with weaker ribs between them. Ribs extend beyond circumference

of shell. Growth lines faintly visible. Ribs faintly reflected in internal casts,

interior with central portion dome-like, with slight groove on one side.

PLIOCENE MARINE INVERTEBRATES FROM LANGEBAANWEG 183

Material

Five specimens, of similar size, largest with longest diameter 13,2 mm,

altitude 5,0 mm. S.A.M. L14187/P.

Remarks

The possibility exists that the present material belongs to the Patellidae,

but several factors give an impression of Szphonaria rather than Patella. These

factors include the small size, the raised dome-like central portion with its

faint groove possibly indicating the position of the siphon. Of the Patellidae

the present material resembles juveniles of P. longicosta (and less obviously

P. oculus) particularly in the external ribbing. The internal doming, however, is

unlike these species which are very flat when juveniles. It is also unusual, if

these are Patellas, that only juveniles of a particular growth size should occur,

with no sign of adult specimens, although adults of another species of Patella

do occur. Of the known species of Szphonaria, the present material most closely

resembles S. deflexa and particularly the variety cyanomaculata. This latter species

also has strong radial ribs, but usually these are more numerous (15-20 as

opposed to 9-12). S. deflexa has a present-day distribution from Natal to the

Cape Peninsula and has been recorded from the Pleistocene of the Algoa Bay

area.

Fic. 7. Siphonaria sp.

Class PELECYPODA

Family Donacidae

Donax cf. serra (Chemnitz)

Fig. 8

Description

Shell typically donaciform; posterior face set at angle to rest of shell, but

not separated by strong ridge, with numerous fine concentric serrulations. Few

very faint radial grooves just anterior to posterior face, further radial lines

visible below surface layer, margin slightly undulate. Apex smoothly rounded.

Maximim apex-to-posterior margin distance only slightly shorter than distance

of apex-to-anterior margin.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

One complete right valve 27,5 mm x 16,9 mm. S.A.M. L14187/Q.

Remarks

In shells of Donax serra of similar total length to the present specimen, the

apex-to-posterior distance is obviously shorter than the apex-to-anterior dis-

tance. The present specimen is close to being equilateral. The serrulations of

the posterior face in similar sized specimens of D. serra are stronger and fewer

in number than in the present specimen. In spite of these differences, D. serra

is the species most closely approached. D. rogerst (Haughton 1931: 36) from

the Pleistocene of the west coast, although almost equilateral, has the posterior

area demarked from the rest of the shell by a strong angular ridge. D. haughtonz,

also from the Pleistocene of the west coast (Carrington & Kensley 1969: 213)

is strongly inequilateral.

The distribution of living D. serra is from Walvis Bay to Durban (Barnard

1964) and the species has been recorded from the Pleistocene deposits of

Liideritzbucht, Orange River, Velddrift, and Sedgefield.

Fic. 8. Donax cf. serra

Donax sp.

Fig. 9

Description

Posterior area demarked from rest of shell by rounded ridge, strong radial

grooves and ridges in this region. Numerous faint growth lines visible. Margin

slightly serrulate.

Material

One incomplete left valve (posterior portion only). S.A.M. L14187/R.

Remarks

In the strength of the ridges in the posterior region, the angle of the latter

to the rest of the shell, and the pattern made by the radial lines and growth

lines on the posterior face, the present specimen most closely resembles Donax

aemulus, a semitropical/tropical species, occurring from Natal to Mogambique

PLIOCENE MARINE INVERTEBRATES FROM LANGEBAANWEG 185

Fic. 9. Donax sp. Right valve, external view; right valve, hinge.

and Malagasy. This species has also been recorded from the Pleistocene of

Durban.

UNCONSOLIDATED SEDIMENTS (L14259)

Phylum BRACHIOPODA

Kraussina rubra (Pallas)

Kraussina rubra (Pallas), Jackson, 1952: 22.

Material

Numerous external and internal casts, particularly of dorsal valve. $.A.M.

L142590/A.

Remarks

Few casts of the external surface of the shell are present, and these are

incomplete. The number of radial striae agrees well with this species. The

internal casts of the dorsal valve are abundant and quite characteristic, with

the branchidium situated between two large eye-like depressions of the umbonal

callosity. A. rubra occurs alive from Lambert’s Bay to the Transkei.

Phylum MOLLUSCA

Class GASTROPODA

Family Patellidae

? Cellana sp.

Material

Several complete internal casts and fragments of external casts. S.A.M.

L14259/B.

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

The external casts show very fine striations as in Cellana capensis, but the

material is too incomplete for further comment.

Family Haliotidae

Haliotis sp.

Material

Fragments of shell casts, including apex of one shell. S.A.M. L14259/C.

Entire internal cast $.A.M. L14259/D.

Remarks

See ipsa 70:

Family Turbinidae

Turbo sarmaticus Linnaeus

Turbo sarmaticus: Barnard, 1963: 214.

Material

Numerous examples of casts of the operculae. Diameter of operculae up

to 24 mm, indicating a shell with a diameter of about 65 mm. Also numerous

fragments of internal casts of shells with diameter of individual whorls of up to

28 mm, indicating a shell of about 70 mm diameter. S.A.M. L14259/D.

Remarks

The external, 1.e. papillose, surface of the operculum of this species can be

deduced as the sandy matrix has filled all the spaces between the papillae. The

species has been recorded from the Pleistocene of the Saldanha Bay area, and

occurs living from False Bav to the Transkei, and very occasionally from Table

Bay and the west coast of the Cape Peninsula.

Class PELECYPODA

Family Donacidae

Donax sp.

Material

Two internal casts of left valves, 34 mm and 22 mm in length. S.A.M.

L14259/E.

Remarks

The general shape, structure of the hinge (determined from silicone casts)

and position of the adductor muscle scars, all agree well with living material

of this genus. Without external casts, the specific position is difficult to

determine.

PLIOCENE MARINE INVERTEBRATES FROM LANGEBAANWEG 187

Phylum ECHINODERMATA

Class EGHINOIDEA

Parechinus angulosus (Leske)

Material

One almost complete internal cast, 23 mm in diameter, $.A.M. L14259/F.

Remarks

No differences can be detected between the characteristic internal

sculpturing of the ambulacral plates and pores of the present material and that

of living material. P. angulosus is the most common echinoid living around the

Cape, and is usually found on rocky shores.

DIscussION

The age of the Langebaanweg marine invertebrate fossils

The relationships of the deposits containing the marine fossils to the rest

of the ‘E’ quarry succession has been one of the more problematical aspects of

the local stratigraphy, and it is not intended to enter into a discussion of this

in the present paper.

The marine deposit is overlain by an horizon of phosphatic rock, and a

layer of non-phosphatic sand and silt, bearing mammalian fossils. The mamma-

lian fauna of this deposit dates from the late Pliocene, and has an inferred age

of about 4 million years (Hendey 19705). Should further work necessitate a

change in the age of this mammalian fauna, the inferred age of the marine

deposits will also need to be altered.

The loose non-phosphatic sand and silt in which the marine vertebrates

occur cannot readily be distinguished from the overlying deposits. These

two sets of deposits are regarded as possibly having been accumulated during

a single geological episode, with their respective faunas indicating a change in

the environment from a marine shoreline to an estuary. The most commonly

occurring terrestrial vertebrate (a tortoise) in the non-phosphatic sand and

silt is also found in association with the marine vertebrates of the marine

deposits, while shark’s teeth similar to those which occur in such abundance

in the marine deposits are also found. There is no reason to believe that the

marine invertebrate assemblage from the site BDT 2/1971 is not contempora-

neous with the associated marine vertebrates at the same level. This assemblage

is thus regarded as broadly contemporaneous with the mammalian fauna from

the overlying deposits.

It is less certain that the marine invertebrates from the cemented phos-

phatic matrix (L14187) are contemporaneous with the marine vertebrates

which occur in the unconsolidated sediments at the same level. The shark’s

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

teeth recovered from the cemented matrix, however, are similar in type and

preservation to those of the unconsolidated sediments. (In how far these teeth

may be used to indicate contemporaneity is uncertain, as these forms cover a

time span from the Cretaceous to recent times.) The two invertebrate assem-

blages do have certain genera and possibly also species in common. The

indications are that these two assemblages are similar in age, and that they

both date from the Upper Pliocene. The differences in preservation and in the

nature of the matrices may simply reflect deposition under differing conditions

rather than geologically significant temporal separation. The two assemblages

have nevertheless been treated separately as their contemporaneity has not

been proved.

The marine deposits of ‘E’ quarry are underlain by a clay of undertermined

thickness, the age and origin of which is unknown, and thus sheds no additional

light on the age of the marine fauna.

Reports on the Tertiary of southern Africa are few in number and of a des-

criptive nature. Haughton (1926, 1932), in discussing the fossil mollusc faunas

of the west coast, came to the conclusion that these were of late Tertiary age

and correlated them with the Alexandria Beds of Redhouse, in the Zwartkops

area of the east coast. These latter beds were described by Newton (1913) as

being of Mio-Pliocene age. It is now considered that the west coast deposits

discussed by Haughton are of Pleistocene age (Carrington & Kensley 1969:

190-191). None of the forms mentioned by Newton have been found in the

present material.* Cox (1939) reported on the Tertiary and Post-Pliocene

deposits of Inhambane, Mocambique. ‘The material mentioned is very typically

of Indian Ocean affinity, and has nothing in common with the present material.

BRIEF ECOLOGICAL COMMENTS

From the species list (Table 1) it may be seen that both assemblages

contain rock- as well as sand-dwelling forms. Amongst the phosphatic matrix

assemblage, only the two species of Donax and the single species of Bulla are

sand-dwellers. All the other species are either permanently attached to a rocky

substrate (the acorn barnacles, which are too fragmentary for identification),

or grazers found on rocky shores or on algae attached to rocks (the patellids

Oxystele, Haliotis, Littorina, Tricolia, Siphonaria, Diodora), or are carnivores

found in rock pools or under stones (Tritonalia, ‘Clavatula’, ‘ Turris’, “Crassispira’,

Thais). Of these, the species tentatively placed in Littorina is the most abundant

(although little quantative importance can be attached to a single ‘cluster’ of

fossil shells). It seems quite likely that the Pliocene shore in the area under

discussion had both rocky and sandy components. The second most abundant

species is that tentatively placed in the Pyramidellidae. Comparatively little

is known of the ecology of these forms, many of which are probably commensals

or parasites of other invertebrates.

* Engelbrecht e¢ al. (1962) record Donax serra from the Tertiary Alexandria Beds.

PLIOCENE MARINE INVERTEBRATES FROM LANGEBAANWEG 189

TABLE I

List of species from both assemblages

CEMENTED PHOSPHATIC UNCONSOLIDATED SEDIMENTS

MATRIX (L14187) (£14259)

MOLLUSCA: GASTROPODA BRACHIOPODA

Bullia sp. S Kraussina rubra R

Cellana capensis R MOLLUSCA: GASTROPODA

‘Clavatula’ sp. R ? Cellana sp. R

‘Crassispira’ sp. R Haliotis sp. R

Diodora parviforata R Turbo sarmaticus R

Haliotis saldanhae R MOLLUSCA: PELECYPODA

? Littorina sp. R Donax sp.

Ocenebra scrobiculata R ECHINODERMATA: ECHINOIDEA

Oxystele tigrina R Parechinus angulosus R

Patella granularis R

? Pyramidella sp. R

Stphonaria sp. R

Thais dubia R

Tricolia neritina R

Turbonilla kraussi R

*Turris’ sp. R

MOLLUSCA: PELECYPODA

Donax cf serra Ss

Donax sp. S)

ARTHROPODA:

CRUSTACEA: cirRIPEDIA

Acorn barnacle fragments R

R = rock-dwelling S = sand-dwelling

Amongst the unconsolidated sediment assemblage, only the Donax is a

sand-dweller. The other species are sessile on rocks (Avaussina) or are herbivo-

rous forms found on rocks (Haluotis, Cellana, Turbo). Judging from the number of

casts of Turbo operculae, this species must have been fairly abundant.

Three genera, viz. Haliotis, Cellana and Donax, appear in both assemblages.

Whether these included the same species is difficult to decide, due to the poor

state of the sandy matrix material.

It is interesting to note that Cellana capensis at present only occurs northwards

from East London, and that Turbo sarmaticus (with the rare exceptions mentioned

above) occurs eastwards from False Bay. This may indicate a somewhat higher

sea temperature on the west coast during the Pliocene than obtains at present.

Although several of the fossil forms have been identified with extant

species, it is possible that some of the specimens are representatives of extinct

species. Further material and more detailed work will clarify the situation.

SUMMARY

22 fossil mollusc forms including a new species of Halzotis, one species of

brachiopod and one species of echinoderm are recorded from two assemblages

from Langebaanweg, Cape Province. The age of the fossils is deduced by

correlation of the overlying mammalian fauna with that of East Africa, as

dating from the Upper Pliocene.

I90 ANNALS OF THE SOUTH AFRICAN MUSEUM

ACKNOWLEDGEMENTS

My grateful thanks are due to Mr Q. B. Hendey, of the South African

Museum, for providing the material for the present paper, and also for supply-

ing the information concerning the geological position and age of the fossils.

I am indebted to Mr G. Branch and Professor A. C. Brown, both of the

Department of Zoology, University of Gape Town, for comments on the patel-

lids and Bullia respectively, to Dr P. A. Hulley of the South African Museum

for information concerning the fossil shark teeth, and to Mr A. Tankard for

comments on the manuscript. Finally, my sincere thanks are due to Mrs lone

Rudner, without whose careful and very thorough preparation of the fossils

this work would not have been possible.

REFERENCES

BARNARD, K. H. 1958. Contributions to the knowledge of South African marine Mollusca. Part I.

Gastropoda: Prosobranchiata: Toxoglossa. Ann. S. Afr. Mus. 44: 73-163.

BARNARD, K. H. 1959. Contributions to the knowledge of South African marine Mollusca. Part

II. Gastropoda: Prosobranchiata: Rhachiglossa. Ann. S. Afr. Mus. 45: 1-237.

BARNARD K. H. 1962. Revised list of South African late Tertiary and Pleistocene marine

Mollusca. Trans. R. Soc. S. Afr. 36: 179-196.

BARNARD K. H. 1963a. Contributions to the knowledge of South African marine Mollusca.

Part III. Gastropoda: Prosobranchiata: Taenioglossa. Ann. S$. Afr. Mus. 47: 1-199.

BARNARD, K. H. 1963. Contributions to the knowledge of South African marine Mollusca.

Part IV. Gastropoda: Prosobranchiata: Rhipidoglossa, Docoglossa. Tectibranchiata.

Polyplacophora. Solenogastres. Scaphopoda. Ann. S. Afr. Mus. 47: 201-360.

BARNARD, K. H. 1964. Contributions to the knowledge of South African Marine Mollusca.

Part V. Lamellibranchiata. Ann. S. Afr. Mus. 47: 361-593.

BuLLEN-NEwToNn, R. 1913. On some Kainozoic shells from South Africa. Rec. Albany Mus. 2:

315-352.

CarrINcTON, A. J. & KENsLEy, B. F. 1969. Pleistocene molluscs from the Namaqualand coast.

Ann. S. Afr. Mus. 52: 189-223.

Cox, L. R. 1939. Estudo paleontolégico das respectivos faunas de moluscos. Bolm Servs Ind.

Minas Geol., Lourenco Marg. 3: 65-105.

ENGELBRECHT, L. N. J., CoERTzE, F. J.. & Snyman, A. A. 1962. Die geologie van die gebied tussen

Port Elizabeth en Alexandria, Kaapprovinsie. (The geology of the area between Port Elizabeth

and Alexandria, Cape Province.) Pretoria: Geological Survey.

Haucuton, S. H. 1926. On some new Mollusca from Tertiary beds in the west of the Cape

Province. Trans. R. Soc. S. Afr. 13: 159-162.

Haucuton, S. H. 1931. The late Tertiary and recent deposits of the west coast of South Africa.

Trans. geol. Soc. S. Afr. 34: 19-57.

HeEnpey, Q.B. 1970a. A review of the geology and palaeontology of the Plio/Pleistocene deposits

at Langebaanweg, Cape Province. Ann. S. Afr. Mus. 56: 75-117.

HEnpDEyY, Q. B. 1970). The age of the fossiliferous deposits at Langebaanweg, Cape Province.

Ann. S. Afr. Mus. 56: 119-131.

Jackson, J. W. 1952. A revision of some South African Brachiopoda, with descriptions of new

species. Ann. S. Afr. Mus. 41: 1-40.

Kitpurn, R. N. 1970. Taxonomic notes on South African marine Mollusca. I. Ann. Cape Prov.

Mus. (Nat. Hist.) 8: 39-48.

Kocu, H. J. 1949. A review of the South African representatives of the genus Patella Linnaeus.

Ann. Natal Mus. 11: 487-517.

Smpson, G. G. 1971. Fossil penguin from the late Cenozoic of South Africa. Science

171: 1144-1145.

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins, arranged in the following order:

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(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-

headings to be used sparingly and enumeration of headings to be avoided. (4) Summary

(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

ILLUSTRATIONS

To be reducible to 12 cm x 18 cm (19 cm including caption). A metric scale to appear

with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their

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REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, 5, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

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

Fiscuer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

Fiscuer, P.-H., Duvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.

Archs Zool. exp. gén. 74: 627-634.

Konan, A. J. 19602. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region

of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konan, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

Ture, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn scHULTZE, L.

Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 260-270.

ZOOLOGICAL NOMENCLATURE

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by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).

The Harvard system of reference to be used in the synonymy lists, with the full references

incorporated in the list at the end of the article, and not given in contracted form in the synonymy

list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, 5; Liste: 11. Turton, 1932: 80.

Brian Kensley

PLIOCENE MARINE INVERTEBRATES

FROM LANGEBAANWEG,

CAPE PROVINCE

VOLUME 60 PART 5 —- NOVEMBER 1972

, mo 7.65

OF THE SOUTH AFRICAN

MUSEUM

CAPE ‘TOWN

eNNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60 Band

November 1972 November

Part 5 Deel

ie, KARE PLEGITOGNATE Fish,

MACRORHAMPHOSODES URADOI (KAMOHARA)

(TRIACANTHODIDAE) IN SOUTH AFRICAN WATERS ~

PaA) HULEEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE RARE PLECTOGNATH FISH, MACRORHAMPHOSODES URADOI

(KAMOHARA) (TRIACANTHODIDAE) IN SOUTH AFRICAN

WATERS

1 ek, lolloieieiang

(South African Museum, Cape Town)

(With 2 figures)

[Ms. accepted 20 Fune 1972]

CONTENTS

PAGE

Introduction : : a LO

Description of material . ; 5 LOR

Summary . : : ‘ 2 Oe:

Acknowledgements : : 5 Oy

References . : : : 5. YL

INTRODUCTION

Among a large quantity of material donated to the South African Museum

in 1971 by the skipper of the Cape Town-based trawler Lobelia, was a single

example of a long-snouted triacanthodid fish. Unfortunately, during transport

at sea, the tip of the snout of the specimen was damaged, being compressed

between the lid and side walls of the canister, so that characters associated

with this region (mouth width, degree of twisting of the mouth) are difficult

to interpret. However, the specimen has been identified as Macrorhamphosodes

uradot (Kamohara, 1933), and as such represents not only the first record of

the species in the southern African region, but also the first record outside the

waters of Japan. It therefore sheds new light on the factors affecting the geo-

graphical isolation of the species as suggested by Tyler (1968).

Two genera of long-snouted triacanthodid fishes, Macrorhamphosodes and

Halimochirurgus, are recognized at present (Tyler 1968), and are easily dis-

tinguished from each other by snout width, twisting of the mouth, teeth shape,

and the length of the third dorsal spine. The specimen falls within the genus

Macrorhamphosodes by virtue of its flattened spatulate teeth, especially in the

lower jaw, and because the third dorsal spine is well developed, its length being

0,76 the length of the second dorsal spine (Figs 1, 2). |

The specimen has been identified as M. uradoi, and may be distinguished

from the closely related M. platycheilus Fowler, 1934 by its extremely short gill

opening, which extends ventrally to a level of about one-third down the

pectoral base (Fig. 1), dental formula and fin count. Furthermore, the pre-

maxillary pedicels are elongated into thin filaments (Fig. 2) and the maxillaries,

IQI

Ann. S. Afr. Mus. 60 (5), 1972: 191-195, 2 figs

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

which are difficult to distinguish in the crushed snout, are comparatively

narrow (Fig. 2).

In view of this record, it would appear that the distribution of M. uradoi

is much wider than previously thought, and that like its congener M. platycheilus,

it bridges the Indonesian region to achieve an Indo-West Pacific distribution

pattern. M. platycheilus has recently been reported from East African waters

(Tyler 1970).

bem RF :

bea eee

‘: 4 2 3g 4 & é ~ a ; nee “4 <: tz A eK

He ay 3

3 ,

Fic. 1. Macrorhamphosodes uradoi (SAM 26308). A. Lateral view; B. Ventral view.

THE RARE PLECTOGNATH FISH 193

Fic. 2. Macrorhamphosodes uradoi (SAM 26308). A. Premaxillaries;

B. Left maxillary; C. Lower teeth (dorsal view); D. Lower tooth

(vertical cross-section).

DESCRIPTION OF MATERIAL

Macrorhamphosodes uradoi (Kamohara, 1933)

Halimochirus uradoi Kamohara, 1933: 392, figs 1-3.

Macrorhamphosodes uradoi: Kamohara, 1937: 7, pl. 1, fig. 2; 1938: 43; 1952: 61; 1958: 51;

19614: 5; 19615: 309, pl. 39, fig. 5; 1964: 67. Fraser-Brunner, 1950: 5. Tomiyama & Abe,

1958: 29, fig. Tyler, 1968: 194, figs 73-79, 204.

Material

One specimen, SAM 26308 (S.L. c. 165 mm), trawled off Bird Island,

Port Elizabeth (33°58’S; 25°40’E), in 220-240 fms (400-450 metres).

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Proportional dimensions are not given, as the snout of the specimen is

badly damaged.

Dl, 14 Aree re Te iC 12, Vertebral) coum oa

Body elongate and laterally compressed; caudal peduncle long, its width

1,7 in peduncle length. Eye large, interorbit 1,7 in largest diameter of eye.

Snout elongate, forming tube (anterior region of snout damaged); premaxil-

laries with long, filamentous pedicels, maxillaries comparatively narrow. 2

teeth in upper jaw (1 broken), flattened but tapering to blunt point; 8 spatu-

late teeth in lower jaw (+ 3 missing), compressed and slightly curved distally.

Gill slit small, extending ventrally to about one-third level down pectoral base,

its length 4,6 in largest diameter of eye. Origin of dorsal above pectoral base;

Ist spine with asperities to about one-half its length, extending a little beyond

tip of 2nd spine, but falling short of origin of soft dorsal; 3rd dorsal spine well

developed, its length 76,7°% the length of 2nd spine. Pectoral situated below

midline of body, its rays extending posteriorly beyond origin of 3rd dorsal

spine. Ventral origin below posterior base of pectoral; pelvic equal in length

to 1st dorsal spine, with minute filament in axil; pelvis much wider anteriorly

and tapering to a point posteriorly, its ventral surface broad and flattened.

Anal origin below 5th soft dorsal ray; caudal rounded. Scales with asperities.

Colour: (in alcohol) yellowish-tan, somewhat darker above than below;

edges of fins tinged pink.

Distribution

Southern Japan to South Africa in 100-240 fms (183-450 m).

SUMMARY

A specimen of the rare triacanthodid fish, Macrorhamphosodes uradot (Kamo-

hara, 1933) is described from the southern African region. This represents not

only the first record in this region, but also the first record outside Japanese

waters. Its depth distribution is increased to between 400 metres and 450 metres.

ACKNOWLEDGEMENTS

I should like to express my thanks to Capt. Olivari and crew of the M/T

Lobelia for the many interesting specimens they have donated to the South

African Museum, and to Mr S. X. Kannemeyer, of this Department, for his

assistance.

REFERENCES

FRASER-BRUNNER, A. 1950. Studies in plectognath fishes from the ‘Dana’-Expeditions. 1. An

interesting new genus of triacanthodid fishes from the Celebes Sea. Dana Rep. 35: 1-8.

Kamonara, TT’. 1933. On a new fish from Japan. Zool. Mag., Tokyo 45: 389-393.

Kamonara, T. 1937. A review of the triacanthodid fishes found in the waters of Japan. Annotnes

Zool. jap. 16: 5-8.

*KAMOHARA, T. 1938. On the offshore bottom-fishes of Prov. Tosa, Shikoku, Japan. Tokyo: Mazuren.

Kamonara, T. 1952. Revised descriptions of the offshore bottom-fishes of Prov. Tosa, Shikoku,

Japan. Rep. Kochi Univ. nat. Sci. 3: 1-122.

THE RARE PLECTOGNATH FISH 195

Kamonara, T. 1958. A catalogue of fishes of Kochi Prefecture (Province Tosa), Japan. Rep

Usa mar. biol. Stn Kochi Univ. 5 (1): 1-76.

Kamouara, T. 1961a. Notes on the type specimens of fishes in my laboratory. Rep. Usa mar.

biol. Stn Kochi Univ. 8 (2): 1-9.

Kamonara, T. 19610. Coloured illustrations of the fishes of Japan. 2. Osaka: Hoikusha.

Kamouara, T. 1964. Revised catalogue of fishes of Kochi Prefecture, Japan. Rep. Usa mar. biol.

Sin Kochi Univ. 11 (1): 1-99.

*TomiyaMa, I. & Ase, T. 1958. Encyclopaedia zoologica illustrated in colours. Pisces and Cyclosto-

mata. Tokyo.

Tyer, J. C. 1968. A monograph on plectognath fishes of the superfamily Triacanthoidea.

Monogr. Acad. nat. Sci. Philad. 16: 1-364.

TYLER, J. C. 1970. New records of triacanthoid plectognath fishes. Notul. Nat. 435: 1-7.

* References not seen by the author.

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins, arranged in the following order:

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headings to be used sparingly and enumeration of headings to be avoided. (4) Summary.

(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

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To be reducible to 12 cm X 18 cm (19 cm including caption). A metric scale to appear

with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their

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REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

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World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

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

FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

FiscHer, P.-H., Duvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.

Archs Zool. exp. gén. 74: 627-634.

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

of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn scHULTZE, L.

Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature issued

by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).

The Harvard system of reference to be used in the synonymy lists, with the full references

incorporated in the list at the end of the article, and not given in contracted form in the synonymy

list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, 6; Liste: 11. Turton, 1932: 80.

P. A. Hulley

THE RARE PLECTOGNATH FISH,

MACRORHAMAMPHOSODES URADOI

(KAMOHARA) (TRIACANTHODIDAE)

IN SOUTH AFRICAN WATERS

ANNALS

‘OF THE SOUTH AFRICAN

MUSEUM

CAPE ‘TOWN

aNNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60 Band

November 1972 November

Part, 9 6 Deel

Peek OoRT ON THE MESOPELAGIC FISHES

COLLECTED DURING THE DEEP-SEA CRUISES

OF R.S. ‘AFRICANA IT’, 1961-1966

P. A. HULLEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

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Price of this part/Prys van hierdie deel

R3,50

Trustees of the South African Museum © _ Trustees van die Suid-Afrikaanse Museum

1972

ISBN 0 949940 20 8

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica Press, Pty., Ltd, Die Rustica-pers, Edms., Bpk.

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A REPORT ON THE MESOPELAGIC FISHES COLLECTED DURING

THE DEEP-SEA CRUISES OF R.S. ‘AFRICANA II’, 1961-1966

Raa EoriEy

South African Museum, Cape Town

(With 3 figures and 1 table)

[Ms. accepted 20 Fune 1972]

CONTENTS

PAGE

Introduction : : , a Oy

Station list . : 5 : = §100

Species list . ’ ; : 200

Systematic discussion . : 2038

Summary . : ‘ : 5 BRP

Acknowledgements ‘ : 5) BED

References . : ; ; 5 AG

INTRODUCTION

In 1960, an investigation on the composition of the mesopelagic fauna of

the southern African region was instigated by Dr F. H. Talbot, formerly of the

South African Museum, as part of the survey of the biology of tuna. The original

intention was to build up a reference collection of possible forage organisms of

tunas, but the survey was expanded by the Division of Sea Fisheries, as part

of their biological programme, to incorporate an intensive survey of the meso-

pelagic fauna of the region.

As far as the ichthyofauna is concerned, it appears that the southern

African region has been fairly well sampled during cruises of the S.S. Preter

Faure, S.S. Pickle and R.V. Discovery, while stations in the area were occupied

by the Valdivia, Dana and Deutsche Siidpolar Expedition. Recently, an investi-

gation of the entire area was carried out by the South African Museum (Grindley

& Penrith 1965), while the western South Indian Ocean was covered during

Cruises 3 and 6 of R.V. Anton Bruun, and the eastern South Atlantic Ocean by

the R.V. Walther Herwig and the SRTM Vyandra.

This paper is intended only as a general supplement to the above work,

so that detailed taxonomic descriptions and synonymies are not given. The

hydrography of the southern African region is comparatively well known and

has been adequately summarized by Grindley & Penrith (1965).

The midwater collections, on which this paper is based, were obtained

197

Ann. S. Afr. Mus. 60 (6), 1972: 197-236, 3 figs, 1 table.

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

during six cruises of R. 8. Africana I, from 1961 to 1966 (Fig. 1). A total of

146 stations were occupied during the cruises and fishing was undertaken at 60

of these. However, material from only 48 stations was sent to the South African

Museum for identification. Details of the position, estimated fishing depth,

fishing time, time of day and gear for the relevant stations are given below in

the Station List. The majority of hauls were made during the day, either with

an N2ooB net (1961-3) or Isaacs-Kidd Midwater Trawl (1964-6), but at

several stations during the 1964 Cruise, an Isaacs-Kidd Midwater Trawl

adapted with an N200 ring at the codend was employed.

SOUTHERN

AFRICA

Fic. 1. Stations occupied during the Deep-sea Cruises of R.S. Africana IT from 1961 to 1969

(®@ fishing stations).

The material, which included mostly juveniles to adults and only a few

larval fishes, was given to the South African Museum for identification, and

has now been incorporated into the collection of this institution.

Fishes of the family Paralepidae have not been included in this paper,

while specific identifications of the Melamphaeidae, Scopelosauridae and

Cyclothone have not been undertaken.

The species list, based on these collections, has been divided into three

regions for convenience:

(1) South East Atlantic—stations west of 19°E (St A 2945-53; 2958-63;

2967-8; 3616-43; 4218-34).

(11) South of Agulhas—stations between 19°E and 26°E (St A 2386-94; 2957;

2965-6).

(111) South West Indian—stations east of 26°E (St A 1225-54; 1877-96).

Year

1961

1962

1963

1964

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES

Date

14-VI

15-VI

16-VI

17-VI

19-VI

20-VI

21-VI

Pip |

23-VI

24-VI

25-VI

26-VI

6-VII

7-VII

8-VII

10-VII

11-VII

12-VII

13-VII

14-VII

23-VI

24-VI

25-VI

26-VI

27-VI

3-VII

7-VII

9-VII

11-VII

3-IV

Station

A 1225*

A 1226*

eT22 7

A 1228*

A 1229

A 1230

A 1232

A 1233

A 1234

Fei 2or,*

A 1236

PEST

A 1238

A 1239

A 1245*

A 1246

A 1247

A 1250

A 1251

A 1252

A 1253

A 1254

EO 77

A 1879

A 1880*

A 1881*

A 1882*

A 1888

A 1892

A 1894

A 1896

A 2386

A 2387

STATION LisT

Position

25°55'S, 33°33'E

25°50’S, 34°06’E

26°00’S, 34°30’E

26°08’S, 35°06’E

26°12'S, 36°12’E

26°42’S, 37°41'E

27°10'S, 41°45 E

27°22’S, 43°35 E

27°A7'S, 45°41'E

27° 48'S, 47°19'E

28°11’S, 49°20’E

29°20'S, 50°30’E

30°50’S, 51°36’E

32°07'S, 52°49'E

36°16’S, 56°45'E

37°43, 95 57 27 ©

37°17'S, 54°36'E

35 48'S, 47°37'E

34°57'S, 44°00’E

34°14'S, 40°46’E

33°35'S, 37°51'E

3325'S, 37°07'E

3301'S, 34°49’E

33 01'S, 34°49’E

40°44'S, 33°36'E

43°39'S, 35°14'E

4525'S, 36°32’E

46°40’S, 38°00’E

46°58’S, 37°56’E

43°17'S, 48°55'E

36°28/S, 41°22’E

36°02’S, 35°38’E

B4°12'S, 26°24'E

36°33’S, 20°01’E

37°36'S, 19°34’E

A 2389 ca3Q°o1’S, 20°04’E

A 2390

A 2301

A 2393

A 2394

A 2945

A 2948

A 2950*

A 2951

A 2953

A 2957

A 2958

A 2061

A 2962

A 2963

A 2965

A 2966

A 2967

40°18’S, 20°26’E

41°59'S, 21°59'E

43°50’S, 25°00’E

44°40’S, 26°05’E

S6711'S, 14,14 5

39°00’S, 10°12’E

41°05'S, 11°41’E

42°28'S, 14°25'E

44°24’8, 20°16’E

A2°T 1S, 19 26 EF

41°40’S, 17°17’E

40°12’S, 14°41’E

39° 06'S, 13°34’E

38°00’S, 15°00’E

40°17'S, 15°54’E

38°09’S, 19°50’E

37°45'9, 18°00’E

Gear

N200B

IKMT+

IKMT

IKMT+

IKMT

IKMT+

Fishing

time

(mins)

24,

Time of

day

1250-1314

1616-1640

0958-1021

1430-1453

1725-1752

1835-1850

1908-1927

1630-1653

1720-1745

1750-1808

1635-1755

1625-1830

1620-1825

1600-1730

1404-1459

1715-1815

1621-1748

1637-1745

1635-1740

1500-1555

III5—-1235

1640-1745

II 10-1250

1630-1730

1330-1500

1635-1845

13550 O35

1000-1045

1630-1715

1615-1805

1348-1545

1611-1706

1520-1640

1357-1453

1625-1740

1917-1947

1345-1415

1330-1435

1832-1945

1315-1345

2000-2130

1952-2145

1415-1700

1340-1600

I 100-1400

1930-2130

1425-1715

1840-2025

1300—

1500-1715

1615-1830

1255-1726

1400-1600

199

Depth

(metres)

300-0

I 000—500-—0

500-0

I 500-0

500-0

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fishing

Year Date Station Position Gear time Time of Depth

No (mins) day (metres )

29-III A20968 36°38’S, 16°28’E 35 — — a

1965 7-IV A 3616 §«931°19'S, 10°08 EE =IKMT 225 1525-1910 I 000-0

13-IV <A 3630 8 32°31’S, 01°55/E % 167 1313-1600 a

1I5-IV A 3632 8 33°41’S, 05°54’E . 135 1700-0915 1 000—500—0

16-IV A 3634 ~33°23’S, 09°31'E me 110 1610-1800 =

23-1V A3643)" Y36°51°S, 12°43 5 45 120 0930-1130 ‘I 000-0

1966 8-VII A 4218 29°30’S, 10°03’E op 60 1325-1425 1 400-0

I6-VII -A 4229 §©629°25’S, 11°30 E i 60 OQ10O—I010 600-0

22-VII <A 4234* 26°06'S, 13°02’E 3) 60 1050-1150 375-0

IKMT+t = IKMT adapted with N2o0 ring

* — material not received from these Stations

SPECIES LIsT

South East South of | South West

Atlantic Agulhas Indian

Searsiidae

Persparsia kopua

-{.

Gonostomatidae

Valenciennellus tripunctulatus .

Vincinguerria nimbaria .

Vincinguerria attenuata .

Photichthys argenteus

Gonostoma atlanticum

-—

—

Gonostoma elongatum

Gonostoma bathyphilum .

Cyclothone spp.

Sternoptychidae

Arg yropelecus gigas

Arg yropelecus aculeatus .

Arg yropelecus hemigymnus

Sternoptyx diaphana

Stomiatidae

Stomias boa boa

Chauliodontidae

Chauliodus sloani

Astronesthidae

Astronesthes indicus

Neonesthes capensis , -+-

+ + FHt+H+ 44+ 44+ 4+

+++ +4

4 +

Melanostomiatidae

Echiostoma barbatum

Leptostomias gladiator

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 201

South East South of South West

Atlantic Agulhas Indian

Eustomias sp. 4"

Opostomias gibsonpacer . an

Flagellostomias bouree .

Bathophilus ater

Pachystomias microdon .

Malacosteidae

Malacosteus niger .

Idiacanthidae

Idiacanthus fasciola

Idiacanthus atlanticus :

Idiacanthus sp... +

Bathylagidae

Bathylagus antarcticus

Bathylagus bericotdes

++ 4+ +44

Scopelarchidae

Neoscopelarchoides elongatus .

Scopelosauridae

Scopelosaurus sp.

Myctophidae

Protomyctophum (Protomyctophum)

andersson.. A =e

Protomyctophum (Protomyctophum)

normant 5 i : +

Protomyctophum (Protomyctophum)

bolin . : i : 2 oh a =F

Protomyctophum (Protomyctophum)

andriashevi.

Protomyctophum (HMierops) paral-

lelum ‘ :

Protomyctophum (Hierops) sub-

parallelum . ! +

Electrona antarctica

+ + +4

Electrona rissot h

Benthosema suborbitale .

Diogenichthys atlanticus .

Aygophum proximum

Aygophum hygom

Hygophum hansen :

Symbolophorus boops . : +

Notolychnus valdiviae

Lampadena notialts

++++ 4

-{-

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

Lampanyctus alatus

Lampanyctus australis

Lampanyctus pusillus

Lampanyctus ater .

Lampanyctus cf. achirus

Lepidophanes indicus

Ceratoscopelus warmingi

Lobianchia dofleini.

Diaphus cf. theta .

Diaphus ostenfeldi .

Scopelopsis multipunctatus

Bregmacerotidae

Bregmaceros mcclellandt.

Melanonidae

Melanonus gracilis

Trachipteridae

Trachipterus sp.

Melamphaeidae (sensu latu)

Diretmidae

Diretmus argenteus .

Scombropidae

Howella brodiet

Chiasmodontidae

Chiasmodon niger .

Bramidae

Pterycombus cf. petersi .

Brotulidae

Neobythites macrops

Gempylidae

Epinnula orientalis .

Ceratiidae

Cryptopsaras couesi

Scorpaenidae

Helicolenus dactylopterus dactylop-

terus : . : :

Nemichthyidae

Borodinula infans .

Nemichthys scolopaceus .

Serrivomeridae

Serrivomer beanit

South East

Atlantic

+ + + FH + fF H4+t44+44+4+4+4+4

South of

Agulhas

South West

Indian

—:

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 203

SYSTEMATIC DiscUssION

Family Searsiidae

Persparsia kopua (Phillips, 1942)

Fic. 2. Persparsia kopua (Phillips, 1942).

Bathytroctes kopua Phillips, 1942: 49, pl. 16, fig. 1.

Persparsia kopua Parr, 1960: 48, fig. 33. Matsui & Rosenblatt, 1971: 447.

Bathytroctes rostratus Norman, 1930: 268, fig. 1, pl. 2, fig. 3.

Persparsia taningi Parr, 1951: 18; 1960: 50, figs 35, 36. Tucker, 1954: 208 (partim).

Material

SAM 26143, 1 (18,5), St A 1247, I 500-0 m.

SAM 26144, 1 (30), St A 2968, 600-0 m.

SAM 26145, 1 (64), St A 4218, I 400-0 m.

Description

D 21; A14; P 20; V 8 (9); gill-rakers 34-35. Photophore distribution

given in Table r.

Remarks

Although the specimens fit the description of P. taningi, it is now held that

this species is synonymous with P. kopua, and that differences in photophore

distribution and the calcified support to the shoulder organ represent onto-

genetic changes (Matsui & Rosenblatt 1971). These specimens possess laterally

directed teeth on the premaxilla, so that it is considered that this is a character

which is lost with growth.

204. ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE I

Persparsia kopua. Photophore distribution (terminology according to Parr 1960).

SAM SAM SAM

26143 26144 26145

(SL 18,5mm) (SL 30 mm) (SL 64 mm)

b++-4

Jot] +++t4+] | 4+4+4+414)441

Sl etl ttl] +i | | 4444 |] el tor

| ++++4+4+4+4+4+4+4+4+4+44

——

—

SAM 26143 represents the first record of the species in the south-western

Indian Ocean.

Family Gonostomatidae

Valenciennellus tripunctulatus (Esmark, 1871)

Maurolicus tripunctulatus Esmark, 1871: 489.

Valenciennellus tripunctulatus : Goode & Bean, 1895: 513. Jordan & Evermann, 1896: 578. Murray

& Hjort, 1912: 612, fig. 478. Pappenheim, 1914: 182. Norman, 1930: 300. Bruun, 1931: 290.

Koefoed, 1958: 4. Grey, 1964: 219, fig. 59. Grindley & Penrith, 1965: 282.

(For full synonymy see Grey 1964: 224.)

Material

SAM 26209, 1 (21), St A 1894, 500-0 m.

SAM 26210, 3 (24-26), St A 3630, 1 000-0 m.

SAM 26211, 1 (24), St A 3634, 1 000—500—-0 m.

Description

D7; A24; P17; V 7; gill-rakers 3 + 12 (2 in angle). Branchiostegal

rays 9-10. Eyes tubular, directed upward. Interorbital less than snout length.

Photophores: ORB 1, in front of eye; OP 3-4, upper smaller, lower two level

with end of maxillary; BR (6); IV (3) + (4) + (17) = 24; VAV 5, the first

elevated and double; OA (2) +3 =5; AC (3) + (3) + (3) + (2) + (4) = 15.

SAM 26211: AG (3) + (3) + 3) Gi 4) =

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 205

Vineinguerria nimbaria (Jordan & Williams, 1895)

Kalarges nimbarius Jordan & Williams, 1895: 793, pl. 76.

Vineinguerria nimbaria: Norman, 1930: 292. Marshall, 1954: 342. Grey, 1955: 273; 1964: 130,

figs 29-32.

Vincinguerria sanzoi Jespersen & Taning, 1919: 218, pl. 17, figs 2, 5; 1926: 22, figs 13-17. Norman,

1930: 292. Fraser—Brunner, 1931: 218. Fowler, 1936: 366, fig. 8, 1206. Smith, 1958: 131;

1961: 106, fig. 156. Grindley & Penrith, 1965: 282 (partim).

(For full synonymy see Grey 1964: 137.)

Material

SAM 26212, 1 (19,5), St A 2386, 500-0 m.

Remarks

Distinguished from the related species, V. attenuata, by the presence of

SO photophores and 18 gill-rakers on the first arch, of which 13 are on the lower

limb. Grindley & Penrith (1965) referred all their Vincinguerria specimens to

V. sanzoi, but one of these (SAM 23663) is identified as V. nimbaria. The others

should be referred to V. attenuata.

Vincinguerria attenuata (Cocco, 1838)

Maurolicus attenuatus Cocco, 1838: 193, pl. 8, fig. 13.

Vincinguerria attenuata Jordon & Evermann, 1896: 577. Goode & Bean, 1895: 513. Norman,

1930: 291. Fowler, 1936: 1205. Marshall, 1951: 3; 1954: 109, figs 5, 14, xi, 8; 1960: 7,

pl. 2, figs 1, 2. Grey, 1955: 2733 1964: 143, figs 29-31, 34.

Vineinguerria sanzoi: Grindley & Penrith, 1965: 282 (partim).

(For full synonymy see Grey 1964: 148.)

Material

SAM 26213, 1 (31), St A 1246, 500-0 m.

SAM 26214, 1 (24), St A 1877, 850-0 m.

SAM 26215, 1 (22), St A 2389, 820-0 m.

SAM 26216, 2 (13-15), St A 3616, 1 000-0 m.

SAM 26217, 2 (21-23), St A 3630, 1 000-0 m.

SAM 26218, 1 (24), St A 3632, 1 000—500-0 m.

SAM 26219, 1 (24), St A 3634, 1 000—-500-0 m.

Remarks

Distinguished from V. nimbaria by the absence of SO photophores and

from V. poweriae by a higher gill-raker count. In the above specimens, the gill-

raker count of 19-22 is somewhat higher than the 18-19 range given by Grey

(1964). Six specimens (SAM 23338, 23359, 23653) identified as V. sanzor by

Grindley & Penrith (1965) should be referred to V. attenuata.

Photichthys argenteus Hutton, 1873

Phosichthys argenteus Hutton, 1873: 269 (lapsus calami).

Photichthys argenteus Hutton, 1873: 269, pl. 15, fig. go. Giinther, 1887: 178, pl. 45, fig. A. Goode

& Bean, 1895: 104, pl. 32, fig. 122. Brauer, 1906: 92, fig. 37. Gilchrist, 1922: 55. Barnard,

1925: 150. Norman, 1930: 292, pl. 2, figs 1, 2. Smith, 1961: 104, fig. 153. Grey, 1960:

100; 1964: 84. Grindley & Penrith, 1965: 282.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM 26220, 2 (19-21), St A 1894, 500-0 m.

SAM 26221, 2 (59-64), St A 2968, 600-0 m.

SAM 26222, 1 (235), St A 3616, 1 000-0 m.

SAM 26223, 3 (98-241), St A 3630, 1 000-0 m.

SAM 26224, 2 (15,5-16), St A 3632, 1 000—-500-0 m.

SAM 26225, 11 (108-160), St A 4229, 600-0 m.

Gonostoma atlanticum Norman, 1930

Gonostoma denudatum atlanticum Norman, 1930: 282. Marshall, 1960: 100, fig. 42A.

Gonostoma atlanticum: Grey, 1960: 105; 1961: 462; 1964: 166, fig. 38.

(For full synonymy see Grey 1964: 171.)

Material

SAM 26206, 2 (20-40), St A 2389, 820-0 m.

Remarks

Distinguished from the related G. denudatum by the lack of an adipose dorsal

fin, presence of vomerine teeth, and a gill-raker count of 11 + 6 = 17.

These two specimens constitute the most southerly record of the species in

the Atlantic.

Gonostoma elongatum Ginther, 1878

Gonostoma elongatum Ginther, 1878: 187; 1887: 173, pl. 45, fig. B. Brauer, 1906: 75, pl. 4, fig. 4,

fig. 27. Murray & Hort, 1912: 604, pl. 2, text fig. 490. Norman, 1930: 283. Fowler, 1936:

230, 1204, fig. 105. Marshall, 1951: 3; 1954: 209, pl. 4, figs vii, 12, ix, 4. Koefoed, 1958: 12.

Grey, 1964: 171, figs 39, 40. Grindley & Penrith, 1965: 282.

(For full synonymy see Grey 1964: 178.)

Material

SAM 26297, 1 (157), St A 2961, 700-0 m.

SAM 26208, 1 (142), St A 2968, 600-0 m.

SAM 26299, 1 (190), St A 3616, 1 000-0 m.

SAM 26300, 1 (169), St A 3643, 1 000-0 m.

Gonostoma bathyphilum (Vaillant, 1888)

Neostoma bathyphilum Vaillant, 1888: pl. 8, fig. 1.

Cyclothone bathyphilum: Gilchrist, 1913: 66. Fowler, 1936: 227, fig. 103.

Gonostoma grandis Barnard, 1925: 143.

Gonostoma bathyphilum: Norman, 1930: 285. Smith, 1961: 104, fig. 151. Grey, 1964: 180, figs 41, 42.

(For full synonymy see Grey 1964: 183.)

Material

SAM 26301, 2 (35-47), St A 3632, 1 000—500—-0 m.

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES

Cyclothone spp.

Material

SAM 26252, St A 1230, 250-0 m.

SAM 26253, St A 1234, 250-0 m.

SAM 26254, St A 1236, 250-0 m.

SAM 26255, St A 1237, I 000—500—0 m.

SAM 26256, St A 1238, 1 000—500-0 m.

SAM 26257, St A 1239, I 000—500-0 m.

SAM 26258, St A 1247, I 500-0 m.

SAM 26259, St A 1250, 500-0 m.

SAM 26260, St A 1251, 500-0 m.

SAM 26261, St A 1252, 500-0 m.

SAM 26262, St A 1253, 500-0 m.

SAM 26263, St A 1254, 500-0 m.

SAM 26264, St A 1877, 820-0 m.

SAM 26265, St A 1879, 1 000-0 m.

SAM 26266, St A 1888, 1 000-0 m.

SAM 26267, St A 1892, 1 000-0 m.

SAM 26268, St A 1896, 1 000-0 m.

SAM 26269, St A 2386, 500-0 m.

SAM 26270, St A 2387, 823-0 m.

SAM 26271, St A 2389, 820-0 m.

SAM 26272, St A 2390, 823-0 m.

SAM 26273, St A 2391, 820-0 m.

SAM 26274, St A 2393, 820-0 m.

SAM 26275, St A 2394, 500-0 m.

SAM 26276, St A 2945, 700-0 m.

SAM 26277, St A 2951, 700-0 m.

SAM 26278, St A 2953, 600-0 m.

SAM 262709, St A 2958, 600-0 m.

SAM 26280, St A 2961, 700-0 m.

SAM 26281, St A 29063, 600-0 m.

SAM 26282, St A 2965, 600-0 m.

SAM 26283, St A 2966, 600-0 m.

SAM 26284, St A 2967, 600-0 m.

SAM 26285, St A 2968, 600-0 m.

SAM 26286, St A 3616, 1 000-0 m.

SAM 26287, St A 3630, 1 000-0 m.

SAM 26288, St A 3632, 1 000—500-0 m.

SAM 26289, St A 3634, 1 000—500-0 m.

SAM 26290, St A 3643, 1 000-0 m.

SAM 26201, St A 4218, 1 400-0 m.

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

‘This genus was represented at almost every station (40 out of 48) and forms

the bulk of the material collected. While Grindley & Penrith (1965) reported

on the complete absence of Cyclothone specimens in their samples, several

specimens have been found amongst their myctophid material (Wisner, personal

communication).

Family Sternoptychidae

Argvropelecus gigas Norman, 1930

Arg yropelecus gigas Norman, 1930: 302, fig. 10. Fowler, 1936: 1208. Parr, 1937: 39. Schultz,

1938: 147; 1961: 600, fig. 5; 1964: 250, fig. 64. Blache, 1964a: 74, fig. Baird, 1971: 38,

fig. 24.

Arg yropelecus affinis: Jespersen, 1915: fig.

Material

SAM 26156, 1 (77), St A 2968, 600-0 m.

SAM 26157, 1 (38), St A 3634, 1 000—500—0 m.

SAM 26158, 4 (41-71), St A 4229, 600-0 m.

Arg yropelecus aculeatus Cuvier & Valenciennes, 1849

Arg yropelecus aculeatus Cuvier & Valenciennes, 1849: 406. Sauvage, 1891: 483, pl. 48, fig. 5.

Goode & Bean, 1895: 127. Brauer, 1906: 110, fig. 47. Murray & Hjort, 1912: 642. Norman,

1930: 303, fig. 11. Schultz, 1961: 607, fig. 9; 1964: 256, fig. 62. Grindley & Penrith, 1965:

282. Baird, 1971: 48, fig. 35.

Arg yropelecus acanthurus Fowler, 1936: 246, 1207.

(For full synonymy see Schultz 1961: 607.)

Material

SAM 26159, 1 (27), St A 1230, 250-0 m.

SAM 26160, 1 (10), St A 2387, 823-0 m.

SAM 26161, 1 (12), St A 2966, 600-0 m.

SAM 26162, 1 (23), St A 3630, 1 000-0 m.

Argyropelecus hemigymnus Cocco, 1829

Arg yropelecus hemigymnus Cocco, 1829: 146. Brauer, 1906: 106, fig. 45. Zugmayer, 19110: 51.

Murray & Hjort, 1912: 612. Gilchrist, 1913: 66. Pappenheim, 1914: 182. Barnard, 1925:

153. Norman, 1930: 303, pl. 2, fig. 4. Schultz, 1937: 4; 1961: 601, fig. 6; 1964: 251,

fig. 65. Smith, 1961: 107, fig. 160. Blache, 1964a: 76. Grindley & Penrith, 1965: 282.

Baird, 1971: 42, fig. 28.

(For full synonymy see Schultz 1961: 601.)

Material

SAM 26175, 1 (12), St A 1239, I O0OO—500—0 m.

SAM 26176, 10 (9-15), St A 1246, 500-0 m.

SAM 26177, 1 (21), St A 1250, 500-0 m.

SAM 26178, 2 (damaged), St A 1253, 500-0 m.

SAM 26179, 2 (9-42), St A 1254, 500-0 m.

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 209

SAM 26180, 3 (11-19), St A 1892, 1 000-0 m.

SAM 26181, 8 (8-33), St A 1894, 500-0 m.

SAM 26182, 3 (10-11), St A 1896, 1 000-0 m.

(

SAM 26183, 1 (22), St A 2389, 820-0 m.

SAM 26184, 1 (damaged), St A 2390, 823-0 m.

SAM 26185, 3 (13-25), St A 2945, 700-0 m.

SAM 26186, 1 (31), St A 2958, 600-0 m.

SAM 26187, 1 (14), St A 2965, 600-0 m.

SAM 26188, 5 (10-25), St A 2966, 600-0 m.

SAM 26189, 3 (30-33), St A 2968, 600-0 m.

SAM 261990, 1 (25), St A 3616, 1 000-0 m.

SAM 261091, 10 (16-27), St A 3630, 1 000-0 m.

SAM 26102, 1 (17), St A 3632, 1 000—500-0 m.

SAM 26193, 7 (17-27), St A 3634, 1 000—500—-0 m.

SAM 26194, 1 (21), St A 4229, 600-0 m.

Remarks

Schultz (1961) distinguished two species in the hemigymnus-complex:

A. hemigymnus, which he confined to the Mediterranean and Atlantic, and is

replaced by A. intermedius in the Indo-Pacific region. Although he did not

examine specimens from the Cape of Good Hope, Schultz (1961) placed

A. hemigymnus recorded off South Africa by Barnard (1925) and Smith (1961)

in the synonymy of A. intermedius, referred Norman’s (1930) specimens from

similar localities to A. hemigymnus, and placed the single specimen recorded by

Gilchrist (1913) under both synonymies. Blache (1964a) suggests that A.

hemgymnus and A. intermedius may only be geographical subspecies, and he

restricts A. hemigymnus to localities north of the equator and A. intermedius to

localities south of about 11°S. Grindley & Penrith (1965) refer all their specimens

to A. hemigymnus.

Examination of some 60 specimens from both the eastern South Atlantic

and western South Indian Ocean has revealed that the diagnostic characters

for separating the two species used by Schultz (1961) are extremely variable:

specimens were found to have a barbed dorsal blade and smooth pectoral shield

(i.e. A. hemigymnus); smooth dorsal blade and scalloped pectoral shield (i.e.

A. intermedius); and specimens with combinations of the two characters. It is

therefore probable that there is a single species, and all specimens have con-

sequently been referred to A. hemig ymnus. Baird (1971) has included A. zntermedius

as a junior synonym of A. hemigymnus.

Sternoptyx diaphana Hermann, 1781

Sternoptyx diaphana Hermann, 1781: 33. Cuvier & Valenciennes, 1849: 415. Giinther, 1887: 169,

pl. 45, figs D, D!. Goode & Bean, 1895 :124, pl. 39, fig. 146. Brauer, 1906: 115, figs 56-63.

Pappenheim, 1914: 183. Barnard, 1925: 154. Norman, 1930: 305. Fowler, 1936: 241,

1207, fig. 113. Smith, 1961: 106. Schultz, 1961: 617, fig. 2; 1964: 262, fig. 69. Blache,

1964a: 83, fig. Baird, 1971: 75, fig. 58.

(For full synonymy see Schultz 1961: 617.)

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM 26163, 1 (damaged), St A 1254, 500-0 m.

SAM 26164, 2 (29-34), St A 2945, 700-0 m.

SAM 26165, 2 (22-25), St A 2948, 700-0 m.

SAM 26166, 1 (40), St A 2958, 600-0 m.

SAM 26167, 1 (15), St A 2965, 600-0 m.

SAM 26168, 1 (18), St A 2966, 600-0 m.

SAM 26169, 2 (14-16), St A 2967, 600-0 m.

SAM 26170, 1 (17), St A 2968, 600-0 m.

SAM 26171, 6 (13-16), St A 3616, 1 000-0 m.

SAM 26172, 1 (13), St A 3630, 1 000-0 m.

SAM 26173, 2 (11-15), St A 3632, 1 000—500-0 m.

SAM 26174, 1 (32), St A 3643, 1 000-0 m.

Family Stomiatidae

Stomias boa boa (Risso, 1810)

Esox boa Risso, 1810: 330, pl. 10, fig. 34.

Stomias boa: Cuvier, 1817: 184. Risso, 1826: 440, pl. 14, fig. 40. Goode & Bean, 1895: 108,

pl. 35, fig. 128. Brauer, 1906: 49. Zugmayer, 19115: 71, pl. 4, fig. 1. Barnard, 1925: 137.

Smith, 1961: 98, fig. 131. Gibbs, 1969: 4, fig. 5.

Stomias boa boa Morrow, 1964a: 293.

Material

SAM 24640, 1 (102), St A 3630, 1 000-0 m.

SAM 24641, 1 (210), St A 1888, 1 000-0 m.

SAM 24642, 1 (252), St A 1888, 1 000-0 m.

SAM 24657, 1 (162), St A 2963, 600-0 m.

SAM 262092, 1 (75), St A 3634, 1 000—500—-0 m.

Family Chauliodontidae

Chauliodus sloani Bloch & Schneider, 1801

Chauliodus sloani Bloch & Schneider, 1801: 430. Cuvier & Valenciennes, 1849: 382. Goode &

Bean, 1895: 96, fig. 115. Gilchrist, 1913: 66; 1922: 42. Fowler, 1936: 219. Smith, 1961: 102,

fig. 145. Morrow, 1964c: 283, fig. 74. Grindley & Penrith, 1965: 282. Gibbs & Hurwitz,

1967: 798, figs 1-3.

Chauliodus sloaniti Giinther, 1887: 179.

Chauliodus sloanei: Brauer, 1906: 40, figs 7-9. Pappenheim, 1914: 167. Barnard, 1925: 141.

Regan & Trewavas, 1929: 32, fig. 24. Norman, 1930: 308; 1939: 21.

(For full synonymy see Morrow 1964c: 287.)

Material

SAM 24408, 1 (damaged), St A 1239, I 000—500—0 m.

SAM 26195, 3 (55-58), St A 2945, 700-0 m.

SAM 26196, 1 (229), St A 2948, 700-0 m.

SAM 26197, 1 (245), St A 2961, 700-0 m.

SAM 26108, 3 (41-204), St A 2966, 600-0 m.

SAM 26199, 1 (30), St A 2967, 600-0 m.

SAM 26200, 6 (125-246), St A 2968, 600-0 m.

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 2Aan

SAM 26201, 1 (194), St A 3616, 1 000-0 m.

SAM 26202, 4 (35-124), St A 3630, I 000-0 m.

SAM 26203, 2 (113-119), St A 3634, 1 000—500-0 m.

SAM 26204, 1 (195), St A 3643, 1 000-0 m.

SAM 26205, 2 (109-150), St A 4229, 600-0 m.

SAM 26206, 1 (59), St A 3632, 1 000—500-0 m.

Family Astronesthidae

Astronesthes indicus Brauer, 1902

Astronesthes indicus Brauer, 1902: 287; 1906: 33, fig. 5, pl. 2, fig. 3. Regan & Trewavas, 1929: 23.

Norman, 1930: 306. Gibbs, 1964: 322, fig. 82.

(For full synonymy see Gibbs 1964: 323.)

Material

SAM 24654, 1 (31,0), St A 2966, 600-0 m.

Description

DRO the 6;,.V 6: OV 5; VAL 8; IP 5; PV 6; VAV 8; AC &,

Remarks

This represents the first record of the species in the eastern South Atlantic.

Neonesthes capensis (Gilchrist & Von Bonde, 1924)

Astronesthes capensis Gilchrist & Von Bonde, 1924: 5. Barnard, 1925: 134. Smith, 1961: 101.

Neonesthes capensis: Gibbs, 1964: 346, fig. 91.

Neonesthes macrolychnus Regan & Trewavas, 1929: 30. Norman, 1930: 308.

(For full synonymy see Gibbs 1964: 348.)

Material

SAM 24636, 1 (152,4), St A 3616, 1 000-0 m.

SAM 24646, 1 (140,0), St A 3632, 1 000—500-0 m.

SAM 24650, I (151,5), St A 2957, 700-0 m.

Description

D 10(9); A 25 (24); P8 (7); V7.

Family Melanostomiatidae

Echwostoma barbatum Lowe, 1843

Echiostoma barbatum Lowe, 1843: 88. Smith, 1961: 99, fig. 135. Morrow & Gibbs, 1964: 482,

fig. 135.

Echiostoma tanneri: Norman, 1930: 314. Barnard, 1937: 48.

(For full synonymy see Morrow & Gibbs 1964: 486).

Material

SAM 23203, 1 (28,0), St A 1896, 1 000-0 m.

Remarks

This specimen represents the first record of the species in the Indian Ocean.

Leptostomias gladiator (Zugmayer, 1911)

Nematostomias gladiator Zugmayer, 19114: 5; 19115: 76, pl. 3, fig. 5.

Leptostomias gladiator Regan & Trewavas, 1930: 61, fig. 41B. Morrow & Gibbs, 1964: 441, fig. 131

A, G-K.

212 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 3 Leptostomias gladiator (Zugmayer, 1911); head and barbel. Fixed teeth black,

depressible teeth unshaded.

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 213

Material

SAM 26294, 1 (95), St A 1246, 500-0 m.

Description

DP - 9; V 7. IP 11; PV 42; VAV 17 + 4; OV 40; VAL 17;

AC 17; suborbital 1; postorbital absent.

Remarks

The specimen falls closest to L. gladiator, in having a filament at the base

of the stem of the barbel, one pair of strongly developed filaments at the base

of the bulb, and two tubercules at the distal end of the bulb. The form of the

bulb (Fig. 3) differs from those figured by Morrow & Gibbs (1964: fig. 131,

G-—K), but this feature appears to be highly variable in different specimens.

This specimen represents the first record of the species in the Indian Ocean.

Eustomias;sp.

Material

SAM 26295, 1 (+ 73), St A 1254, 500-0 m.

Remarks

The specimen is damaged and the barbel is missing. One species, Eustomias

filferum, is at present known from southern African waters (Smith 1961).

Opostomias gibsonpace: Barnard, 1948

Opostomias gibsonpacei Barnard, 1948: 344, figs 1, 2. Smith, 1961: 100.

Material

SAM 24649, 1 (264), St A 2957, 700-0 m.

Description

D 23; A 25; P1+5; V 9. First mandibular tooth piercing premaxillary.

Suborbital 1; postorbital 1; IP 10; PV 28; VAV 24; OV 26; VAL 21; AC 18.

Remarks

Gibbs (personal communication) considers that O. gibsonpacei may be a

junior synonym of O. micripnus.

Flagellostomias boureet (Zugmayer, 1913)

Eustomias boureet Zugmayer, 1913: 3.

Flagellostomias boureei Regan & Trewavas, 1930: 57, figs 8 D, 9 B, 11 A, 12 A, 33, 34, pl. 2, fig. 2.

Norman, 1930: 310. Morrow & Gibbs, 1964: 430, fig. 130.

(For full synonymy see Morrow & Gibbs 1964: 433.)

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM 24639, 1 (223), St A 3616, 1 000-0 m.

Description

D5 VA. Ogee ol 8Ose Nae

Bathophilus ater (Brauer, 1902)

Dactylostomias ater Brauer, 1902: 286; 1906: 57, pl. 3, fig. 3.

Bathophilus ater Regan & Trewavas, 1930: 68, fig. 46. Morrow & Gibbs, 1964: 461. Barnett &

Gibbs, 1968: 831.

Material

SAM 24656, 1 (90,0), St A 2962, 600-0 m.

Description

D 16; A 15; P 6 (the first two rays longer and a little separated); V to.

Postorbital photophore with luminous patches in front and below.

Remarks

While dorsal and anal counts are somewhat higher, and pectoral and pelvic

counts somewhat lower, than the values given by Regan & Trewavas (1930),

the specimen falls closest to B. ater in arrangement of the pectoral fin rays, the

presence of two luminous patches associated with the postorbital photophore,

and an JA count of 38.

Pachystomias microdon (Giinther, 1878)

Eustomias microdon Giinther, 1878: 180.

Pachystomias microdon Giinther, 1887: 210, pl. 53, fig. C. Regan & Trewavas, 1930: 70, figs 14 B,

15. Morrow & Gibbs, 1964: 375, fig. 97.

Pachystomias atlanticus Regan & Trewavas, 1930: 70, pl. 6, fig. 1.

(For full synonymy see Morrow & Gibbs 1964: 377.)

Material

SAM 246, 35, 1 (183,9), St A 3616, 1 000-0 m.

Remarks

The specimen has been identified by Mr R. H. Goodyear.

Family Malacosteidae

Malacosteus nger Ayres, 1848

Malacosteus niger Ayres, 1848: 69; 1849: 53, pl. 5. Giinther, 1887: 214, pl. 54, fig. C. Murray &

Hjort, 1912: 612. Regan & Trewavas, 1930: 142, figs 25, 138. Norman, 1930: 317. Koefoed,

1956: 16. Morrow, 19646: 545, fig. 144.

Malacosteus indicus Giinther, 1878: 181; 1887: 214, pl. 54, fig. B. Brauer, 1906: 65, figs 23-25,

pl. 4, fig. 1. Gilchrist, 1922: 54. Barnard, 1925: 139. Regan & Trewavas, 1930: 143. Smith,

1961: 100, fig. 137.

(For full synonymy see Morrow 19646: 547-)

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES mes

SAM 24660, 1 (159,0), St A 2967, 600-0 m.

SAM 26226, 1 (152,0), St A 2393, 820-0 m.

SAM 26227, 1 (24,0), St A 3630, 1 000-0 m.

Material

SAM 24644, 1 (110), St A 3634, 1 000—500-0 m.

SAM 24645, 1 (112,5), St A 3634, I 000—500-0 m.

SAM 24651, 1 (72,4), St A 2966, 600-0 m.

SAM 24652, 1 (84,4), St A 2966, 600-0 m.

SAM 24653, 1 (75,7), St A 2966, 600-0 m.

SAM 24655, 1 (94,5), St A 2962, 600-0 m.

SAM 24659, 1 (149,0), St A 2948, 700-0 m.

(

Remarks

Two species have been distinguished by Morrow (19646) on the basis of

the size of the postorbital photophore. However, in the above specimens, the

size of this photophore appears to vary with age and ranges from 24,0%

(SAM 24655) to 31,89% (SAM 24660) of the eye diameter. It is probable

therefore that M. danae is a junior synonym of M. niger.

Family Idiacanthidae

Idiacanthus fasciola Peters, 1877

Idiacanthus fasciola Peters, 1877: 847. Giinther, 1887: 215. Brauer, 1906: 60, figs 17—20, 22, pl. 4,

figs 2-3. Pappenheim, 1914: 168. Regan & Trewavas, 1930: 129. Fowler, 1936: 1199.

Koefoed, 1956: 8. Smith, 1961: 102. Gibbs, 1964: 514, figs 137-140. Grindley & Penrith,

1965: 282.

Idiacanthus ferox Giinther, 1887: 216, pl. 52, fig. D. Brauer, 1906: 59. Murray & Hort, 1912: 86.

Barnard, 1925: 136.

(For full synonymy see Gibbs 1964: 520.)

Material

SAM 24658, 1 (85), St A 2953, 600-0 m.

SAM 26239, 1 (61), St A 2968, 600-0 m.

Idiacanthus atlanticus Brauer, 1906

Idiacanthus atlanticus Brauer, 1906: 62, fig 21, Smith, 1961: 102. Krueger, 1967. Novikova 1967,

Idiacanthus niger Regan, 1914: 14; 1916a: 378. Regan & Trewavas, 1930: 128, fig 124. Smith,

1961: 102 fig. 142. Grindley & Penrith, 1965: 282.

Material

SAM 24637, 1

SAM 24638, 1

SAM 24643, 1

SAM 24644, 1

SAM 24661, 1

SAM 26240, |

SAM 26241, 1

SAM 26243, 2

345), St A 3616, 1 000-0 m.

375), st A 3616, I 000-0 m.

314), St A 3634, I 000-0 m.

43), St A 3630, I 000-0 m.

423), St A 2967, 600-0 m.

415), St A 2391, 820-0 m.

423), St A 4218, I 400-0 m.

42-44) St A 3613, I 000-0 m.

LDN DN LDN NN NNN

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Distinguished from the related species J. fasciola by the anterior position

of the pelvic fins and by 16 photophores situated between the pelvic and anal

origins. SAM 24661 with pelvic origin just in front of dorsal origin. Juvenile

males with long intromittent organ incorporating anterior anal rays.

Idiacanthus sp.

Stylophthalmus paradoxus Brauer, 1906: 67, pl. 5, figs 1-7. Regan, 19164: 136. Barnard, 1925: 140.

Smith, 1961: 100, fig. 138.

Material

SAM 26242, 3 (no SL), St A 1234, 250-0 m.

Remarks

Although described as a separate species by Brauer (1906), specimens of

this type are now known to be larval stages of Idiacanthus (Beebe 1934; Gibbs

1964).

Family Bathylagidae

Bathylagus antarcticus Giinther, 1878

Bathylagus antarcticus Giinther, 1878: 248; 1887: 220. Goode & Bean, 1895: 55. Brauer, 1906: 12,

fig. 2. Barnard, 1925: 129. Norman, 1930: 276; 1937: 81. Smith, 1961: 97, fig. 130.

Bathylagus glacialis Lonnberg, 1905): 68. Norman, 1930: 276. Blache, 19640: 11, fig.

Material

SAM 26245, 2 (48-60), St A 1888, 1 000-0 m.

SAM 26246, 1 (37), St A 2953, 600-0 m.

SAM 26248, 1 (27), St A 4218, 1 400-0 m.

Remarks

Norman (1930) distinguished between B. antarcticus and B. glacialis mainly

on the basis of anal fin ray count: 18-21 in B. glacialis; (21) 22-25 in B. antarcticus.

This has been followed by Blache (19645). However, Barnard (1925) and Smith

(1961) hold that the two species are synonymous. There is a definite overlap

of anal ray counts in the above specimens and therefore all material has been

referred to B. antarcticus. SAM 26245 represents the first record of the species in

the Indian Ocean.

Bathylagus bericoides (Borodin, 1929)

Scopelus bericoides Borodin, 1929: 110.

Bathylagus microcephalus Norman, 1930: 275, fig. 5. Grindley & Penrith, 1965: 282.

Bathylagus bericoides Parr, 1937: 39, fig. 16. Cohen, 1964: 46, fig. 15.

Material

SAM 26247, 4 (51-61), St A 3616, 1 000-0 m.

SAM 26249, 1 (212), St A 2961, 700-0 m.

SAM 26250, 1 (189), St A 2963, 600-0 m.

SAM 26251, 1 (114), St A 3616, 1 000-0 m.

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 2r7

Family Scopelarchidae

Neoscopelarchoides elongatus (Norman, 1937)

Scopelarchus elongatus Norman, 1937: 86.

Neoscopelarchoides elongatus: Marshall, 1955: 310, fig. 3b, b’, pl. 19, figs 1, 2.

Material

SAM 26228, 1 (106), St A 2953, 600-0 m.

Remarks

This specimen represents the first record of the species in South African

waters, and completes the circumpolar distribution pattern of the species given

by Marshall (1955: fig. 4).

Family Scopelosauridae

Scopelosaurus sp.

Material

SAM 23605, 1 (damaged), St A 2966, 600-0 m.

SAM 23606, 1 (37,5), St A 2945, 700-0 m.

SAM 23607, 1 (31,5), St A 2963, 600-0 m.

Remarks

The specimens have been sent to Bertelsen, Krefft and Marshall, and are

to be described in a revision of the family by these authors.

Family Myctophidae

Protomyctophum (Protomyctophum) andersson (Lonnberg, 1905)

Mpyctophum anderssoni Lonnberg, 1905a: 763; 19055: 61. Norman, 1930: 320.

Myctophum (Myctophum) anderssoni Brauer, 1906: 172, fig. 84.

Electrona (Protomyctophum) anderssoni Fraser—Brunner, 1949: 1046, fig.

Protomyctophum (Protomyctophum) anderssoni Andriashev, 1963: 226, figs 6, 7. Becker, 1963: 2

(transl.) ; 1967: 89.

(For full synonymy see Andriashev 1962: 226.)

Material

SAM 26073, 2 (26,9-27,0), St A 1888, 1 000-0 m.

Description

P 14; V 8; gill-rakers 7 + 1 + 19 (20); SAO 2.

Remarks

Both specimens are damaged, so that dorsal, anal and AO counts are not

possible.

Protomyctophum (Protomyctophum) norman (Taning, 1932)

Myctophum normani Taning, 1932: 127, fig. 2.

Electrona (Protomyctophum) normani Fraser—Brunner, 1949: 1046, fig.

Protomyctophum (Protomyctophum) normani Andriashey, 1962: 231, fig. 10. Becker, 1963: 4 (transl.)

fie b> 1967: 89:

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM 26074, 3 (14,8-18,0), St A 2393, 820-0 m.

Description

D 13; A 23 (22); P14; V 8; gill-rakers 6 + 1-4 19; AG apie

Remarks

Although resembling P. tenisoni in that the Prc, and Prc, are almost touch-

ing, the specimens have been identified as P. normani because of the number of

AO photophores and the position of SAO, above VO,. It appears that

Andriashev’s (1962) key character regarding the origin of the anal fin in rela-

tion to the dorsal may be open to question. In these specimens, the origin of

the anal fin is below the middle of the dorsal fin. ‘The pectoral count, further-

more, appears to be somewhat higher than that of Andriashev’s two specimens.

Protomyctophum (Protomyctophum) bolin. (Fraser—Brunner, 1949)

Electrona (Protomyctophum) bolint Fraser—Brunner, 1949: 1045, fig.

Protomyctophum (Protomyctophum) bolint Andriashev, 1962: 232, figs 11, 12. Becker, 1963: 3

(transl.) ; 1967: go.

Material

SAM 26075, 3 (15,9-16,2), St A 1879, 1 000-0 m.

SAM 26076, 11 (15,5-18,0), St A 2393, 820-0 m.

SAM 26077, 2 (14,0-17,2), St A 2394, 820-0 m.

SAM 26078, 1 (15,0), St A 2958, 600-0 m.

Description

D 12 (13); A 24 (23); P15; V 8; gill-rakers 5 + 1 + 15 (16); AO 17 (78).

Protomyctophum (Protomyctophum) ? andriashevi Becker, 1963

Protomyctophum (Protomyctophum) andriashevi Becker, 1963: 6 (transl.), fig. 2; 1967: 89.

Material

SAM 26079, 1 (47,3), St A 4218, 1 400-0 m.

Description

D 13; A 24; P 16; V 8; gill-rakers 4 + 1 + 14; AO 16.

Remarks

Four species of primitive Myctophinae are known in which the distance

between Prc, and Prec, is less than an organ diameter, namely P. andriashevt,

P. normani, P. anderssoni and P. tenisoni. This specimen has 3 SAO photophores

and can be distinguished from P. anderssoni, and has tentatively been identified

as P. andriashevi according to Becker’s (1963) table for the separation of the

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 219

other named species. In this specimen, the pectoral count, AO count, gill-

raker count and the position of SAO, correspond to that of P. andriashevi rather

than P. norman or P. tenisoni, but the line SAO,/SAO, passes through VO,

rather than VO).

Protomyctophum (Hierops) parallelum (Lonnberg, 1905)

Myctophum parallelum Lonnberg, 19054: 764.

Myctophum (Myctophum) parallelum Brauer, 1906: 174, fig. 86.

Electrona (fierops) parallela Fraser—Brunner, 1949: 1047, fig.

Protomyctophum (Hterops) parallelum Andriashev, 1962: 237, fig. 16. Becker, 1963: 9 (transl.);

1967: 90. Nafpaktitis & Nafpaktitis, 1969: 8, figs 3, 4.

Material

SAM 26080, 1 (27,7), St A 2961, 700-0 m.

SAM 26081, 1 (41,0), St A 2963, 600-0 m.

Description

D 12; A 23-24; P 14; V 8; gill-rakers 3 + 1 + 13; AO 17-109.

Remarks

Both specimens have AO counts varying on the two sides: SAM 26080,

AO 17-18; SAM 26081, AO 18-109.

Protomyctophum (Hterops) subparallelum (Taning, 1932)

Myctophum arcticum subparallelum Taning, 1932: 128.

Electrona (Hterops) arctica subparallela Fraser—Brunner, 1949: 1047.

FMierops subparallela Bolin, 1959: 5-

Protomyctophum (Hierops) subparallelum Andriashev, 1962: 243, figs 14, 15. Nafpaktitis & Nafpaktitis,

1969: 8, figs 4-6.

(For full synonymy see Andriashey 1962: 243.)

Material

SAM 26082, 1 (24,2), St A 2394, 820-0 m.

Description

Dye 22: P15; V 8; gill-rakers 4 + 1 + 135 AO 15.

Electrona antarctica (Ginther, 1878)

Scopelus antarcticus Ginther, 1878: 184; 1887: 196, pl. 51, fig. D.

Myctophum antarcticum Norman, 1930: 322, fig. 28; 1937: 85. Smith, 1961: 120.

Myctophum (Myctophum) antarcticum Brauer, 1906: 168. fig. 82 a—c.

Electrona (Electrona) antarctica Fraser—Brunner, 1949: 1048, fig.

Electrona antarctica: Andriashev, 1962: 240, figs 18, 19.

(For full synonymy see Andriashev 1962: 240.)

Material

SAM 26083, 2 (22,2-23,3), St A 1888, 1 000-0 m.

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

D 13; A 21; P-; V 8; gill-rakers 4 + 1 + 13; AO 18.

Remarks

The low gill-raker count, position and size of the PLO and the number

of AO photophores, which occur in an even line, separate this species from the

related E. carlsbergi, E.. subaspera, E. paucirastra and E. ventralis.

SAM 26083 represents the most northern record of the species in the

Indian Ocean (43°17'S, 48°55’E).

Electrona rissot (Cocco, 1829)

Scopelus rissoi Cocco, 1829: 144; 1838: 15, pl. 2, fig. 5.

Myctophum rissot: Brauer, 1906: 170, fig. 83. Fowler, 1936: 380, fig. 184. Norman, 1930: 320.

Electrona (Electrona) rissoi Fraser—Brunner, 1949: 1048, fig.

Electrona rissoi Bolin, 1959: 5. Becker, 1967: 92. Trunov, 1968: 745, fig. 1. Nafpaktitis & Nafpak-

titis, 1969: 10, figs 4, 6, 8.

(For full synonymy see Bolin 1959: 5.)

Material

SAM 26084, 2 (52,5-53,0), St A 4229, 600-0 m.

Description

D 13; Aig; P15; V 8; gill-rakers 8 + 1 + 19; AO 11.

Benthosema suborbitale (Gilbert, 1913)

Material

Myctophum suborbitale Gilbert, 1913: 82.

Myctophum (Benthosema) suborbitale Bolin, 1939: fig. 3.

Benthosema suborbitale Fraser-Brunner, 1949: 1051, fig. Bolin, 1959: 10. Becker, 1967: 95.

Nafpaktitis & Nafpaktitis, 1969: 11, figs 9, 10.

SAM 26085, 2 (21,0-27,0), St A 1253, 500-0 m.

SAM 26086, 1 (24,5), St A 2966, 600-0 m.

Description

D 13 (12); A 17; P 14; V 8; gill-rakers 3 + 1 + 9 (10); AO 5 +6.

Remarks

Distinguished from B. fibulatum, also known from the Indian Ocean

(4°N-18°N), by gill-raker count and by the presence of a luminous organ on

the ventral border of the orbit. Grindley & Penrith (1965) report the occurrence

of B. fibulatum off the Natal coast, but on the basis of distribution, it seems likely

that these specimens should be referred to B. suborbitale.

Diogenichthys altanticus (Taning, 1928)

Myctophum laternatum atlanticum 'Taning, 1928: 56.

Myctophum laternatum Norman, 1930: 324.

Diogenichthys scofieldi Bolin, 1939: 22 figs 3, 16.

Diogenichthys atlanticus Fraser-Brunner, 1949: 1054, fig. Bolin, 1959: 11. Becker, 1967: 95.

Nafpaktitis & Nafpaktitis, 1969: 13, figs 12, 13.

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES Pa

Material

SAM 26087, 1 (21,0), St A 1239, 1 000—500-0 m.

SAM 26088, 3 (11,5-17,5), St A 1254, 500-0 m.

SAM 26089, 1 (13,5), St A 1894, 500-0 m.

SAM 26099, 1 (20,0), St A 2966, 600-0 m.

SAM 26001, 1 (17,0), St A 3630, 1 000-0 m.

SAM 26002, 1 (16,0), St A 4229, 600-0 m.

Description

Diaeewnaro (177); P19 (12); V8; gill-rakers 2 (9) + 1-412 (11)

AO 7 +3 (4).

Aygophum proximum Becker, 1965

Hygophum proximum Becker, 1965: 81, fig. 7 a. Nafpaktitis & Nafpaktitis, 1969: 17, figs 16, 17.

(For full synonymy see Becker 1965: 81.)

Material

SAM 26003, 1 (17,3), St A 1230, 250-0 m.

Description

D 13; A 18; P 13; V 8; gill-rakers 4 + 1 + 13; AO5 +6.

Remarks

Both Becker (1965) and Nafpaktitis & Nafpaktitis (1969) suggest that the

distinctive characters of H. proximum may only be of subspecific importance

and that the species probably consists of a series of geographically varying

populations of H. microchir. SAM 26093 has tentatively been identified as

H. proximum because of the position of Pol, and because the line connecting

Pol, with Pol, passes through the anterior margin of the last AO,. The number

of AO photophores (5 + 6) is somewhat lower than the values (7 + 7; 6 + 7)

found in southern specimens (Nafpaktitis & Nafpaktitis 1969).

Aygophum hygom (Litken, 1892)

Scopelus hygomi Liitken, 1892: 256 (partim), fig. 15.

Hygophum hygomi Fraser—Brunner, 1949: 1050, fig. Bolin, 1959: 6. Grindley & Penrith, 1965: 292.

Becker, 1965: 68, fig. 3; 1967: 92. Nafpaktitis & Nafpaktitis, 1969: 19, figs 17, 18.

(For full synonymy see Becker 1965: 68.)

Material

SAM 26094, 1 (16,7), St A 1251, 500-0 m.

SAM 26095, 1 (18,5), St A 1252, 500-0 m.

SAM 26006, 4 (13,0-14,5), St A 1254, 500-0 m.

SAM 26097, 6 (13,4-21,0), St A 1894, 500-0 m.

SAM 26008, 4 (14,0-15,0), St A 1896, 1 000-0 m.

PLP ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

D 14; A 21; P15; V 8; gill-rakers 5 + 1 + 15 (14); AO 7 + 6 (5).

Hygophum hanseni (Taning, 1932)

Scopelus hygomi Liitken, 1892: 256 (partim).

Myctophum (Myctophum) benoiti var. reinhardti Barnard, 1925: 242.

Myctophum hanseni Taning, 1932: 132, fig. 4.

? Myctophum reinhardti: Smith, 1961: 121.

Hygophum hanseni Fraser—-Brunner, 1949: 1050, fig. Bolin, 1959: 7. Becker, 1965: 95, fig. 11 a;

1967: 94. Nafpaktitis & Nafpaktitis, 1969: 19, figs 17, 18.

Material

SAM 26099, 1 (28,2), St A 2391, 820-0 m.

Description

D 13; A 21; P15; V 8; gill-rakers 4 + 1 + 12; AO5 +8.

Symbolophorus boops (Richardson, 1844)

Myctophum boops Richardson, 1844: 39, pl. 27, figs 6-12.

Myctophum humboldti (non Risso) Norman, 1930: 325. Smith, 1961: 120, fig. 195.

Myctophum humboldti barnardi 'Taning, 1932: 128.

Symbolophorus boops: Andriashev, 1962: 252, figs 24, 25. Becker, 1967: 96. Nafpaktitis & Nafpak-

titis, 1969: 27, figs 29, 30.

(For full synonymy see Andriashev 1962: 252.)

Material

SAM 26100, 2 (19,2—22,8), St A 4229, 600-0 m.

Description

D 14; A 20-22; P 12; V 8; gill-rakers 5 + 1 + 12-13; AO 7 + 7-8.

Remarks

Confusion appears to exist in the identification of S. boops and S. veranyi

from southern African waters. Bolin (1959) has referred specimens from the

Cape of Good Hope to S. veranyi, while Andriashev (1962) refers specimens

from this region to S. boops. Becker (1965) records both species from the South

Atlantic. Until the situation becomes clearer, SAM 26100 is referred to S. boops.

Notolychnus valdwiae (Brauer, 1904)

Myctophum valdiviae Brauer, 1904: 398, fig. 6. Fowler, 1936: 375, fig. 182.

Myctophum (Myctophum) valdiviae Brauer, 1906: 206, fig. 127.

Vestula valdiviae Bolin, 1946: 146, fig. 4.

Notolychnus valdiviae Fraser-Brunner, 1949: 1077, fig. Bolin, 1959: 23. Becker, 1967: 108.

Nafpaktitis & Nafpaktitis, 1969: 33, figs 37, 38.

Material

SAM 26101, 2 (13,0-17,0), St A 1234, 250-0 m.

SAM 26102, 1 (16,0), St A 1236, 250-0 m.

SAM 26103, 1 (20,1), St A 2966, 600-0 m.

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 22

Description

D 11; A 13 (12); P 11-12; V 6; gill-rakers 2 + 1 + 7 (8).

Remarks

AO counts were not possible as the specimens had been badly rubbed.

The absence of records from the eastern South Atlantic would confirm the

distribution pattern of the species. SAM 26103 represents the southernmost

record for the species in the Indian Ocean.

Lampadena notialis Nafpaktitis & Paxton, 1968

Lampadena notialis Nafpaktitis & Paxton, 1968: 13, fig. 5. Nafpaktitis & Nafpaktitis, 1969: 34,

figs 40, 43.

Material

SAM 26104, 2 (20,2-28,0) St A 2391, 820-0 m.

Description

D 14; A 14; P 14; V 8; gill-rakers 7 4+ 1 + 16-17; AO 6 + 3; Prc2 + 1.

Remarks

Although closely resembling L. speculigera, with which it overlaps in the

region 40°—-45°S in the southern Indian Ocean, these specimens have a gill-

raker count 24-25, which is typical of L. notzalis. This record therefore represents

the first in the southern African region. One specimen with AO,5 (all level)

on the one side.

Lampanyctus alatus Goode & Bean, 1895

Lampanyctus alatus Goode & Bean, 1895: 79 (partim). Norman, 1930: 330. Grindley & Penrith,

1965: 283. Becker, 1967: 109. Nafpaktitis & Nafpaktitis, 1969: 53, figs 67, 68.

Lampanyctus (Lampanyctus) alatus Fraser—Brunner, 1949: 1090, fig.

non Myctophum (Lampanyctus) alatus: Barnard, 1925: 240 (= L. australis).

Lampanyctus pusillus: Smith, 1961: 123 (partim).

Macrostoma alatum Fowler, 1936: 1232.

Macrostoma pusillum: Fowler, 1936: 1231 (partim).

Material

SAM 26105, 1 (50,0), St A 1236, 250-0 m.

SAM 26106, 1 (damaged), St A 1251, 500-0 m.

SAM 26107, 4 (16,1-25,0), St A 1896, 1 000-0 m.

SAM 26108, 4 (26,5-50,5), St A 2391, 820-0 m.

SAM 26109, 1 (21,0), St A 2965, 600-0 m.

(

SAM 26111, 2 (damaged), St A 2968, 600-0 m.

SAM 26110, 6 (19,0-24,0), St A 2966, 600-0 m.

2 (45,1I-51,0), St A 4218, I 400-0 m.

SAM 26112,

Description

Bea) Act6 (17)5 P12 (73); V 8; gill-rakers 3 (4) + 1 +9 (8);

AO,6 (5); AO,6 (7); total AO 12 (11-14).

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

Lampanyctus australis ‘Taning, 1932

Lampanyctus alatus australis 'Taning, 1932: 145.

Myctophum (Lampanyctus) alatus: Barnard, 1925: 240.

Lampanyctus australis: Becker, 1967: 109. Nafpaktitis & Nafpaktitis, 1969: 54, figs 68, 69.

Material

SAM 26113, 2 (49,0—-103,5), St A 2394, 820-0 m.

SAM 26114, 1 (25,0), St A 2958, 600-0 m.

SAM 26115, I (31,5), St A 2967, 600-0 m.

SAM 26116, 2 (50,0—-92,0), St A 4218, 1 400-0 m.

SAM 26117, 1 (71,0), St A 4229, 600-0 m.

Description

D 13; A118 (17-19); P14; V 8; gill-rakers 6 (7) 1 - > 19)(e =e

Remarks

Although ZL. australis resembles L. alatus, it can easily be distinguished by

its higher AO and gill-raker counts and by the higher number of pectoral rays.

While these counts for the above specimens fall within the range given by

Nafpaktitis & Nafpaktitis (1969), SAM 26117 had a higher gill-raker count.

As in the Indian Ocean, the distribution of L. australis and L. alatus overlaps

between 30°S and 40°S in the eastern South Atlantic. ZL. australis extends further

north to about 23°S on the west coast than on the east coast of South Africa.

Lampanyctus pusillus (Johnson, 1890)

Scopelus pusillus Johnson, 1890: 457.

Lampanyctus pusillus Taning, 1928: 66. Smith, 1961: 123 (partim), fig. 204. Bolin, 1959: 28.

Becker, 1967: 109. Nafpaktitis & Nafpaktitis, 1969: 52, figs 51, 66.

Macrostoma pusillum Fowler, 1936: 1231 (partim).

Lampanyctus (Lampanyctus) pusillus Fraser—Brunner, 1949: 1099, fig.

(For full synonymy see Bolin 1959: 28.)

Material

SAM 26118, 4 (18,0-30,5), St A 1239, 1 000—500—0 m.

SAM 261109, 1 (21,0), St A 1253, 500-0 m.

SAM 26120, 3 (30,0-31,0), St A 1254, 500-0 m.

SAM 20121502755), ot 30280) 1 O00—Ommn:

NAM 2619978165731 9), ot Aneto, 1 400-Oim,

Description

D 13; A 15; P 14; V 8; gill-rakers 3 + 1 + 8; AO 4-5 + 6.

Lampanyctus ater-complex

As has been pointed out (Andriashev 1962; Nafpaktitis & Nafpaktitis

1969), the taxonomy of these species is confused and requires further investiga-

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 225

tion, especially with regard to the validity of L. niger and to clinal variations

within species. The following identifications are therefore only tentatively

advanced.

Lampanyctus ater ‘Taning, 1928

Lampanyctus ater Taning, 1928: 68. Parr, 1928: 104, fig. 17. Bolin, 1959: 33. Becker, 1967: 116.

Nafpaktitis & Nafpaktitis, 1969: 44, figs 53, 54.

Lampanyctus (Lampanyctus) ater Fraser—Brunner, 1949: 1086, fig.

(For full synonymy see Bolin 1959: 33.)

Material

SAM 26123, 1 (110,5), St A 4218, I 400-0 m.

Description

D 16; A 19; V 8; gill-rakers 5 + 1 + 11; AO 6-7 + 13-14.

Remarks

The specimen falls closest to L. ater in dorsal, anal and gill-raker counts.

However, the upper Pol is anterior to the vertical from the last anal ray and

SAO, is above the third anal ray. There are 5 + 1 infracaudal scales, the

infracaudal gland extending about half-way along the peduncle.

Lampanyctus cf. achirus Andriashev, 1962

Lampanyctus achirus Andriashev, 1962: 256, fig. 27. Becker, 1967: 116. Nafpaktitis & Nafpaktitis,

1969: 45, figs 54, 55-

Material

SAM 26124, 4 (59,9-67,8), St A 4218, I 400-0 m.

Description

D 15; A 18; V 8; gill-rakers 4 + 1 + 11 (10); AO 13-15.

Remarks

Gill-raker count, AO count and dorsal and anal counts separate these

specimens from L. lineatus and L. cuprarius, and they differ from L. ater in the

Prc/AO, interspace, the position of AO,1 and the length of the infracaudal

gland. They most closely resemble L. achirus: pectoral fins are absent; the infra-

caudal gland is more than 90% of the length of the caudal peduncle; AO,r is

above the last anal ray; and SAO, is above the fifth anal ray, behind the vertical

from AQO,I.

Lepidophanes indicus Nafpaktitis & Nafpaktitis, 1969

Lepidophanes indicus Nafpaktitis & Nafpaktitis, 1969: 61, figs 72, 74, 75.

? Lampanyctus pyrosobolus: Grindley & Penrith, 1965: 283.

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM 26125, 1 (20,6), St A 1236, 250-0 m.

SAM 26126, 12 (12,5-22,2), St A 1239, 1 000—-500-0 m.

SAM 26127, 2 (21,5-24,2), St A 1254, 500-0 m.

SAM 26128, 1 (26,5), St A 4229, 600-0 m.

Description

D 13; A13 (14); P13; V 8; gill-rakers 4 (3) -- 1 4+ a @e)}eeee

(5-7) + 4 (3-5).

Remarks

All specimens have been referred to L. indicus because of the presence of

a luminous patch above the pectoral fins, absence of a patch ventral to PVO,,

absence of luminous scales at the dorsal, AO count and gill-raker count.

SAM 26128 from the Atlantic falls closer to L. indicus on the basis of the above,

but possesses 4 luminous scales at the base of the anal fin and AO 7 + 3.

Ceratoscopelus warming: (Litken, 1892)

Scopelus (Nyctophus) warmingu Liitken, 1892: 259, fig. 19.

Myctophum (Lampanyctus) warmingi: Barnard, 1925: 237.

Lampanyctus warmingi: Fowler, 1936: 385, fig. 185.

Ceratoscopelus warmingi: Nafpaktitis & Nafpaktitis, 1969: 63, figs 76, 77.

Ceratoscopelus townsendi Fraser—Brunner. 1949: 1093 (partim), fig. Grindley & Penrith, 1965: 283.

Lampanyctus townsendi: Smith, 1961: 123.

Material

SAM 26129, 3 (17,0-20,5), St A 1239, I 000—500—-0 m.

SAM 26130, 2 (19,2-24,0), St A 4218, 1. 400-0 m.

Description

D 14 (13); A 14; P 14; V 8; gill-rakers 4 + 1 + 10 (11); AO 7 (6) + 5.

Lobianchia doflen (Zugmayer, 1911)

Myctophum (Lampanyctus) dofleint Zugmayer, 1911a: 3; 1911b: 35.

Myctophum dofleini Zugmayer, 1911b: 149, pl. 1, fig. 9.

Diaphus dofleini: Fowler, 1936: 402, fig. 200. Grindley & Penrith, 1965: 283.

Lobianchia dofleini: Bolin, 1959: 19. Becker, 1967: 98.

(For full synonymy see Bolin 1959: 19.)

Material

SAM 26131, 1 (35,2), St A 4218, I 400-0 m.

SAM 26132, 2 (29,8-30,0), St A 4229, 600-0 m.

Description

D 16; A 14; P 12; V 8; gill-rakers 5 + 1 + 13; AO 5 + 5.

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 227

Remarks

The Pre photophores are equally spaced and Prc, is located on the extreme

base of the caudal rays.

Diaphus cf. theta Eigemann & Eigemann, 1890

Diaphus theta Eigemann & Eigemann, 1890: 4. Fraser—Brunner, 1949: 1074, fig. Fowler, 1936:

407. Grindley & Penrith, 1965: 283. Becker, 1967: 102, fig. 6.

Material

SAM 26133, 1 (51,5), St A 2391, 820-0 m.

SAM 26134, 1 (damaged), St A 2394, 820-0 m.

SAM 26135, 19 (27,0—-71,0), St A 4229, 600-0 m.

Description

D 14 (13-15); A 13 (12-14); P11 (10-13); V 8; gill-rakers 8 + 1 + 16 (17);

AO 4 (5) + 5 (6).

Remarks

The specimens fall closer to D. theta than to other species in this complex

because of the gill-raker count, a separate AO,/Prc series, and because AO,!I

is not elevated. Wisner (personal communication) is of the opinion that speci-

mens referred to D. theta from the South Atlantic may prove to be specifically

distinct from those off southern California, the type locality.

Diaphus ostenfeldi Taning, 1932

Diaphus ostenfeldi Taning, 1932: 142 fig. 15. Becker 1967: 107. Trunov, 1968: 747, fig. 2.

Diaphus (Lamprossa) ostenfeldi Fraser-Brunner, 1949: 1072, fig.

Aethopora ostenfeldi Bolin, 1959: 22.

Material

SAM 26136, 7 (61,4-73,0), St A 4229, 600-0 m.

Description

D 16 (15-17); A 15 (16); P11; V 8 (7); gill-rakers 8 (g-10) + 1 + 15 (16),

total 24-26; Pol 3; AO 5 + 5 (6).

Remarks

The 3 Pol organs are not included in the AO, count. One specimen with

AO,6 on the one side, the sixth AO, reduced in size.

Scopelopsis multipunctatus Brauer, 1906

Scopelopsis multipunctatus Brauer, 1906: 146, fig. 71. Barnard, 1925: 246. Norman, 1930: 318.

Fraser—Brunner, 1949: 1099, fig. Smith, 1961: 118, fig. 191. Grindley & Penrith, 1965: 283.

Nafpaktitis & Nafpaktitis, 1969: 67, figs 47, 82.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM 26137, 9 (16,9—-19,3), St A 1254, 500-0 m.

SAM 26138, 1 (62,0), St A 4218, 1 400-0 m.

SAM 26139, 8 (63-67), St A 4229, 600-0 m.

Family Bregmacerotidae

Bregmaceros mcclellandu ‘Thompson, 1840

Bregmaceros mcclellandit Thompson, 1840: 185, figs. Fowler, 1936: 1254.

Bregmaceros maclellandi: Gilchrist & Thompson, 1914: 87; 1916: 319. Norman, 1930: 339.

Smith, 1961: 137, fig. 251. Grindley & Penrith, 1965: 283.

Bregmaceros macclellandi: Barnard, 1925: 325.

Bregmaceros macclellandii : D’Acona & Cavinato, 1965: 66, figs 1, 54-58.

Material

SAM 26147, 1 (32), St A 2961, 700-0 m.

SAM 26148, 2 (26-30), St A 3643, 1 000-0 m.

Family Melanonidae

Melanonus gracilis Giinther, 1878

Melanonus gracilis Ginther, 1878: 19; 1887: 84, pl. 14, fig. B. Brauer, 1906: 277, pl. 12, fig. 5.

Barnard, 1925: 332. Norman, 1930: 340. Fowler, 1936: 1252, fig. 538. Smith, 1961: 140,

fig. 261. Grindley & Penrith, 1965: 283.

Material

SAM 26149, 1 (132), St A 1888, 1 000-0 m.

SAM 26150, 1 (68), St A 2957, 700-0 m.

SAM 26151, 1 (95), St A 2958, 600-0 m.

SAM 26152, 1 (48), St A 2958, 600-0 m.

SAM 26153, 1 (104), St A 2961, 700-0 m.

SAM 26154, 1 (41), St A 2963, 600-0 m.

SAM 26155, 1 (62), St A 3643, 1 000-0 m.

Remarks

Although a second species, MM. zugmayert Norman, has been described, all

specimens have been referred to M. gracilis because of the narrower body depth

and the comparatively feeble dentition.

Family Trachipteridae

Trachipterus sp.

Material

SAM 26302, 1 (4,0), St A 2963, 600-0 m.

Remarks

This specimen is a larval form. At present three species, 7. iris, 7. cristatus

and 7. arcticus, are known from the southern African region (Smith 1961).

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES

ie)

\<e)

Family Melamphaeidae

Material

SAM 26580, 1 (17,0), St A 1239, I 00O—500-0 m.

SAM 26581, 2 (14,0-15,2), St A 1247, I 500-0 m.

SAM 26582, 1 (13,4), St A 1247, I 500-0 m.

SAM 26583, 6 (13,0-22,8), St A 1254, 500-0 m.

SAM 26584, 1 (damaged), St A 1254, 500-0 m.

SAM 26585, 1 (41,5), St A 1879, I 000-0 m.

SAM 26586, 6 (12,0-22,2), St A 1892, I 000-0 m.

SAM 26587, 2 (12,2-12,8), St A 1894, 500-0 m.

SAM 26588, 7 (12,5-17,0), St A 1894, 500-0 m.

SAM 26589, 8 (15,8—20,2), St A 1896, 1 000-0 m.

(

SAM 26590, 1 (16,9), St A 2945, 700-0 m.

SAM 26591, 1 (126,0), St A 2957, 700-0 m.

SAM 26502, 1 (28,8), St A 2958, 600-0 m.

SAM 26593, 1 (32,0), St A 2958, 600-0 m.

SAM 26594, 4 (10,0-18,0), St A 2966, 600-0 m.

SAM 26595, I (17,0), St A 2967, 600-0 m.

SAM 26596, 1 (103,2), St A 3616, 1 000-0 m.

SAM 26597, 2 (41,5-87,0), St A 3616, 1 000-0 m.

SAM 26508, 1 (32,2), St A 3632, 1 000—-500-0 m.

SAM 26599, 2 (22,0-24,1), St A 4218, I 400-0 m.

SAM 26600, 1 (24,6), St A 4218, 1 400-0 m.

Remarks

The following species are probably represented in the collection: Scopelobe-

ryx robustus, S. microlepis, Melamphaes microps, M. suborbitalis, Sio nordenskjoldiz,

Scopelogadus bean, Poromitra macrophthalma and P. atlantica.

Family Diretmidae

Diretmus argenteus Johnson, 1863

Diretmus argenteus Johnson, 1863: 403, pl. 36, fig. 2. Giinther, 1887: 45. Zugmayer, 19116: 107,

pl. 5, fig. 7. Barnard & Von Bonde, 1944: 237, fig. Smith, 1961: 150, fig. 285.

Diretmus aureus: Giinther, 1887: 45.

Material

SAM 26303, 1 (7), St A 1894, 500-0 m.

SAM 26229, 1 (51), St A 3634, I 000—500-0 m.

Family Scombropidae

Howella brodiei Ogilby, 1899

Howella brodiei Ogilby, 1899: 735.

Rhectogramma sherborni Norman, 1930: 348, fig. 39. Fowler, 1936: 1287, fig. 547.

Howella sherborni: Smith, 1961: 212, fig. 498. Grindley & Penrith, 1965: 284.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM 26140, 1 (31,7), St A 2394, 820-0 m.

SAM 26141, 3 (39-82), St A 3643, 1 000-0 m.

SAM 26142, 2 (55-74), St A 4218, I 400-0 m.

Description

D VIII + I, 9; AIT, 7; P 14-15; V I, 5; gill-rakers 7 + 1 +4 a1.

Family Chiasmodontidae

Chiasmodon niger Johnson, 1863

Chiasmodon niger Johnson, 1863: 408; 1864: 76. Jordan & Evermann, 1896: 2291. Norman,

1930: 349. Fowler, 1936: 1031, fig. 424. Smith, 1961: 176, fig. 377. Grindley & Penrith,

1965: 284.

Material

SAM 26146, 1 (52), St A 2958, 600-0 m.

Description

DX hes Ae rare:

Family Bramidae

Pterycombus cf. petersi

Material

SAM 26236, 1 (15,7), St A 1229, 300-0 m.

SAM 26237, 1 (19,5), St A 1232, 250-0 m.

SAM 26238, 1 (14,0), St A 1234, 250-0 m.

Remarks

Two species, P. falcatus and P. petersii, are at present known from the

southern African region. These specimens appear to fall closest to P. petersii.

Family Brotulidae

Neobythites macrops Ginther, 1887

Neobythites macrops Giinther, 1887: 102, pl. 20, fig. A. Gilchrist & Thompson, 1914: 89; 1916: 416.

Gilchrist & Von Bonde, 1924: 19. Norman, 1939: 84. Smith, 1961: 363, fig. 1019.

Neobythites analis Barnard, 1925: 879.

Material

SAM 26579, 1 (54,0), St A 2958, 600-0 m.

Family Gempylidae

Epinnula orientalis Gilchrist & Von Bonde, 1924

Epinnula orientalis Gilchrist & Von Bonde, 1924: 15, pl. 4, fig. 1. Barnard, 1925: 790. Smith,

1961: 311, fig. 865.

MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 231

Material

SAM 26304, 1 (damaged), St A 1229, 300-0 m.

Family Ceratiidae

Cryptopsaras couesi Gill, 1883

Cryptopsaras couesti Gill, 1883: 284. Giinther, 1887: 55. Smith, 1961: 428, fig. 1229.

Ceratias couesti Brauer, 1906: 317, pl. 15, fig. 7.

Cryptopsaras couest: Bertelsen, 1951: 139, figs 93-97.

(For full synonymy see Bertelsen 1951: 139.)

Material

SAM 26230, 1 (37,2), St A 3632, 1 000—500-0 m.

Family Scorpaenidae

Helicolenus dactylopterus dactylopterus (Delaroche, 1809)

Scorpaena dactyloptera Delaroche, 1809: 337, pl. 22, fig. 9.

Helicolenus maculatus: Gilchrist, 1922: 75. Barnard, 1925: 907. Davies, 1949: 26. Smith, 1961:

369, pl. 83, fig. 1034. Grindley & Penrith, 1965: 284.

Sebastes maculatus Cuvier & Valenciennes, 1829: 343.

Helicolenus dactylopterus dactylopterus Eschmeyer, 1969: 93.

(For full synonymy see Eschmeyer 1969: 93.)

Material

SAM 26578, 1 (16,0), St A 3634, 1 000—500—-0 m.

Family Nemichthyidae

Borodinula infans (Gunther, 1878)

Nemichthys infans Giinther, 1878: 251; 1887: 264, pl. 63, fig. B.

Avocettina infans: Barnard, 1925: 199. Fowler, 1936: 283, fig. 133. Smith, 1961: 391. Grindley

& Penrith, 1965: 283. Blache ef al., 1970: 223, fig. 607 a, b.

Material

SAM 26231, 1 (492), St A 2393, 820-0 m.

SAM 26232, 1 (385), St A 2958, 600-0 m.

SAM 26234, 1 (damaged), St A 2961, 700-0 m.

Nemichthys scolopaceus Richardson, 1848

Nemichthys scolopaseus Richardson, 1848: 25, pl. 10, figs 1-3. Brauer, 1906: 126, pl. 9, fig. I.

Roule, 1910: 1, pl. 1, figs 1-7. Barnard, 1925: 198. Blache et al., 1970: 224, fig. 608.

Nemichthys scolopacea Giinther, 1887: 263. Fowler, 1936: 286, fig. 134. Smith, 1961: 391, fig. 1107.

Grindley & Penrith, 1965: 283.

Material

SAM 26234, 1 (735), St A 2394, 820-0 m.

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Serrivomeridae

Serrivomer bean Gill & Ryder, 1883

Serrivomer beanti Gill & Ryder, 1883: 260. Fowler, 1936: 283, fig. 132. Smith, 1961: 392, fig. 1109.

Serrivomer bean: Barnard, 1925: 200. Grindley & Penrith, 1965: 283. Blache et al., 1970: 223,

fig. 606.

Material

SAM 26235, 1 (402), St A 3632, 1 000—500-0 m.

SUMMARY

A collection of mesopelagic fishes from the Deep-sea Stations of Africana IT

is described. The material is from the eastern South Atlantic and western South

Indian Oceans, and includes approximately 75 species, of which 10 are new

records.

ACKNOWLEDGEMENTS

I am grateful to the Director, Division of Sea Fisheries, Cape Town, for

the donation of the mesopelagic fish collection and to Dr A. de Dekker and

Mr A. Robertson, of the Division, for their help. My thanks are also due to

Dr G. Krefft, Institut fiir Seefischerei, Hamburg, and Mr R. Wisner, Scripps

Institution of Oceanography, La Jolla, for their helpful comments. I should

also like to thank Mr S. X. Kannemeyer, Department of Marine Biology,

South African Museum, for his assistance during all phases of this work.

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MESOPELAGIC FISHES COLLECTED DURING DEEP-SEA CRUISES 235

LONNBERG, E. 1905a. Pelagische von der schwedischen Siidpolar-Expedition 1901-1903 erbeutete

Fische. Zool. Anz. 28: 762—766.

LONNBERG, E. 1905). The fishes of the Swedish South Polar Expedition. Wiss. Ergebn. schwed.

Stidpolar-Exped. 5 (6): 1-69. ;

Lowe, R. T. 1843. Notices of fishes newly observed or discovered in Madeira during 1840-42

(Seriola gracilis, Brama longipinnis, Taractes asper, Pteraclis papilio, Luvarus imperialis, etc.).

Proc. zool. Soc. Lond. 1843: 81-95.

*LUTKEN, C. F. 1892. Spolia Atlantica. Bidrag til kundskab om det aabne havs laxesild eller

Scopeliner. Danske Vidensk. Selsk. Skrifter (6) 7: 221-297.

MarsuHa.t, N. B. 1954. Aspects of deep sea biology. London: Hutchinson.

MarsuHatt, N. B. 1955. Alepisauroid fishes. ‘Discovery’ Rep. 27: 303-336.

MarsuHa tt, N. B. 1960. Swimbladder structure of deep-sea fishes in relation to their systematics

and biology. ‘Discovery’ Rep. 31: 1-121.

Matsur, T. & RosENBLATT, R. H. 1971. Ontogenetic changes in patterns of light organs in

searsids and the taxonomy of Sagamichthys and Persparsia. Copeia 1971: 440-448.

Morrow, J. E. 1964a. Family Stomiatidae. In Fishes of the western North Atlantic. 4. Mem. Sears

Fdn mar. Res. 1 (4): 290-310.

Morrow, J. E. 19646. Family Malacosteidae. In Fishes of the western North Atlantic. 4. Mem.

Sears Fdn mar. Res. (4): 523-549.

Morrow, J. E. 1964c. Family Chauliodontidae. In Fishes of the western North Atlantic. 4. Mem.

Sears Fdn mar. Res. 1 (4): 274-289.

Morrow, J. E. & Gress, R. H. 1964. Family Melanostomiatidae. In Fishes of the western North

Atlantic. 4. Mem. Sears Fdn mar. Res. t (4): 351-511.

Morray, J. & Hyort, J. 1912. The depths of the ocean. London: Macmillan.

Narpaxtitis, B. G. & Narpaxtitis, M. 1969. Lanternfishes (family Myctophidae) collected

during Cruises 3 and 6 of the R/V Anton Bruun in the Indian Ocean. Bull. Los Ang. Cty

faus. nat. Hist. 5: 1-79.

Narpaxtitis, B. G. & Paxton, J. R. 1968. Review of the lanternfish genus Lampadena, with a

description of a new species. Contr. Sci. Los Ang. Cty Mus. 138: 1-29.

Norman, J. R. 1930. Oceanic fishes and flatfishes collected in 1925-27. ‘Discovery’ Rep. 2: 261-370.

Norman, J. R. 1937. Fishes. Rep. B.A.N.<. Antarct. Exped. (B) 1 (2): 50-88.

Norman, J. R. 1939. Fishes. Scient. Rep. John Murray Exped. 7: 1-116.

OcitBy, J. D. 1899. Additions to the fauna of Lord Howe Island. Proc. Linn. Soc. N.S.W. 23:

739°-745-

PAPPENHEIM, P. 1914. Die Fische der Deutschen Siidpolar-Expedition 1901-1903. II. Die

Tiefseefische. Deutsch. Stidpol.-Exped. 15 (2): 160-200.

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Bermuda Islands. Bull. Bingham oceanogr. Coll. 3 (3): 1-193.

Parr, A. E. 1951. Preliminary revision of the Alepocephalidae, with the introduction of a new

family Searsiidae. Am. Mus. Novit. 1531: 1-21.

Parr, A. E. 1960. The fishes of the family Searsiidae. Dana Rep. 51: 1-108.

Peters, W. C. H. 1877. Uebersicht der wahrend der von 1874 bis 1876 unter dem Commando

des Hrn. Capitan z.S. Freiherrn von Schleinitz ausgefiihrten Reise S.M.S. Gazelle gesam-

melten und von der Kaiserlichen Admiralitat der Koniglichen Akademie der Wissen-

schaften iibersandten Fische. Monatsber. Akad. Wiss. Berlin 1876: 831-854.

Puiturs, W. J. 1942. New records of bathypelagic fishes from Cook Strait. Rec. Dom. Mus.,

Wellington 1: 49-54.

Reean, C. T. 1914. Fishes. Nat. Hist. Rep. Br. Antarct. Terra Nova Exped. (Zool.) 1: 1-54.

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Recan, C. T. 19165. Larval and postlarval fishes. Nat. Hist. Rep. Br. Antarct. Terra Nova Exped.

(Zool.) 4: 125-155.

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dae. Oceanogrl Rep. ‘Dana’ Exped. 5: 1-39.

Recan, C. T. & Trewavas, E. 1930. The fishes of the families Stomiatidae and Malacosteidae.

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the years 1843-46, London: Reeve & Benham.

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

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de celles des environs de Nice et des Alpes maritimes. 3. Paris: Levrault.

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Sternoptychidae. Smithson. Misc. Collns 91: 1-5.

ScuuLtz, L. P. 1938. Review of the fishes of the genera Polyipnus and Argyropelecus (family

Sternoptychidae), with descriptions of three new species. Proc. U.S. natn. Mus. 86: 135-155.

ScHuLTz, L. P. 1961. Revision of the marine silver hatchet fishes (family Sternoptychidae).

Proc. U.S. natn. Mus. 112: 587-649.

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Taning (Myctophidae). Vop. Ikhtiol. 4: 745-748. (In Russian.)

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Atlantic, 1937-38. Part I. Rosaura Expedition Report, No. 4, Pt 1. Bull. Br. Mus. nat. Hist.

(Zool.) 6: 163-214.

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du ‘Talisman’ pendant les années 1880-83. 1: 1-406. Paris: Masson.

ZUGMAYER, E. 1g11a. Diagnoses des poissons nouveaux provenant des campagnes du yacht

‘Princesse-Alice’ (1901 a 1910). Bull. ,Inst. océanogr. Monaco 193: 1-14.

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Camp. scient. Prince Albert I 35: 1-159.

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‘Hirondelle II’ (1911 a 1912). Bull. Inst. océanogr. Monaco 253: 1-7.

* References not seen by the author.

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Examples (note capitalization and punctuation)

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

FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

FiscHER, P.-H., Duvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.

Archs Kool. exp. gén. 74: 627-634.

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

of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konun, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn scHULTZE, L.

Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

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Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

P. A. Hulley

A REPORT ON THE MESOPELAGIC FISHES

COLLECTED DURING THE DEEP-SEA CRUISES

OF R.S. ‘AFRICANA II’, 1961-1966

OF THE SOUTH AFRICAN

MUSEUM

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60 Band

November 1972 November

Part 7 Deel

MESOPELAGIC FISHES

FROM VEMA SEAMOUNT

(IK STATION 52)

PA, HULERY

Cape Town Kaapstad

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MESOPELAGIC FISHES FROM VEMA SEAMOUNT

(IK Station 52)

jee J\a Ilieibi eiay%s

South African Museum, Cape Town

(With 2 figures)

[Ms. accepted 20 June 1972]

CONTENTS

PAGE

Introduction : ‘ : : : : 5 287

Systematic discussion , : ; : : i POT

Summary : : ; : ; : : 3 242

Acknowledgements . : ‘ ; : NE 242

References 5 ; : ; , : 5 5 ey

INTRODUCTION

In 1960, a survey of the mesopelagic fauna of the seas around South Africa

was undertaken by the South African Museum (Grindley & Penrith 1965), as

part of an investigation of the forage organisms of tuna. A 1o ft Isaacs-Kidd

midwater-trawl was employed and a total of 45 stations were occupied between

October 1960 and April 1963 (IK 4-7, 10-48, 50-51).

In November 1966, during a survey of the Vema Seamount by the R.S.

Thomas B. Davey, a further station (IK 52: 31°38’S, 8°21.5’E, 5-XII-66/

6-XII-66, 2130-0530 (480 mins), 500-0 m) was occupied using the same

IKMT and starting from the south of the Seamount and working around the

west to the north side.

This paper represents the ichthyological results of that station.

SYSTEMATIC DiscussION

Family Searsiidae

Persparsia kopua (Phillips, 1942)

Bathytroctes kopua Phillips, 1942: 49, pl. 16, fig. 1.

Persparsia kopua Parr, 1960: 48, fig. 33. Matsui & Rosenblatt, 1971: 477. Hulley, 1972: 203, fig. 2;

Bathytroctes rostratus Norman, 1930: 268, fig. 1, pl. 2, fig. 3.

Persparsia taningi Parr, 1951: 18; 1960: 50, figs 35, 36. Tucker, 1954: 208 (partim).

Material

SAM 26601, 1 (28,0 mm).

Description

D 21; A 14; P 20; V g; gill-rakers 9 +1 +24. Photospores: 00 2; 100; POO

Pci Re) 7: JO; IPO; SPO; THO-s: SVO; IVO; AO; SAO; PAO;

CoO; PO; GO,,.

237

Ann. S. Afr. Mus. 60 (7), 1972: 237-244, 2 figs.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Gonostomatidae

Photichthys argenteus Hutton, 1873

Phosichthys argenteus Hutton, 1873: 269 (lapsus calami).

Photichthys argenteus Hutton, 1873: 269, pl. 15, fig. go. Giinther, 1887: 178, pl. 45, fig. A. Goode

& Bean, 1895: 104 pl. 32, fig. 122. Brauer, 1906: 92, fig. 37. Gilchrist, 1922: 55. Barnard,

1925: 150. Norman, 1930: 202, pl. 2, figs 1, 2. Smith, 1961: 104, fig. 153. Grey, 1960: 100;

1964: 84. Grindley & Penrith, 1965: 282. Hulley, 1972: 205.

Material

SAM 26603, 13 (28,2-46,0 mm).

Ichthyococcus sp.

Material

SAM 26602, 1 (64,3 mm).

Remarks

The specimen has been sent to Dr G. Krefft, and is to be described in a

revision of the genus by this author.

Cyclothone spp.

Material

SAM 26604, 48 (23,5-29,4 mm).

Family Sternoptychidae

Arg yropelecus gigas Norman, 1930

Arg yropelecus gigas Norman, 1930: 302, fig. 10. Fowler, 1936: 1208. Schultz, 1938: 147; 1961: 600,

fig. 5; 1964: 250, fig. 64. Blache,1964: 74, fig. Baird, 1971: 38, fig. 24. Hulley, 1972: 208.

Arg yropelecus affinis: Jespersen 1915: fig.

Material

SAM 26605, 2 (22,5-66,9 mm).

Arg yropelecus hemigymnus Cocco, 1829

Argyropelecus hemigymnus Cocco, 1829: 146. Brauer, 1906: 106, fig. 45. Zugmayer, I9II: 51.

Murray & Hort, 1912: 612. Gilchrist, 1913: 66. Pappenheim, 1914: 182. Barnard, 1925:

153. Norman, 1930: 303, pl. 2, fig. 4. Schultz, 1937: 4; 1961: 601, fig. 6. Smith, 1961:

107, fig. 160. Blache, 1964: 76. Grindley & Penrith, 1965: 282. Baird, 1971: 42, fig. 28.

Hulley, 1972: 208.

(For full synonymy see Schultz 1961: 601).

Material

SAM 26606, 5 (20,8-31,0 mm).

Family Chauliodontidae

Chauliodus sloani Bloch & Schneider, 1801

Chauliodus sloani Bloch & Schneider, 1801: 430. Cuvier & Valenciennes, 1849: 382. Goode &

Bean, 1895: 96, fig. 115. Gilchrist, 1913: 66; 1922: 42. Fowler, 1936: 219. Smith, 1961:

102, fig. 145. Morrow, 1964: 283, fig. 74. Grindley & Penrith, 1965: 282. Gibbs & Hurwitz,

1967: 798, figs 1-3. Hulley, 1972: 210.

Chaulidus sloanii: Giinther, 1887: 179.

Chauliodus sloanei: Brauer, 1906: 40, figs 7-9. Pappenheim, 1914: 167. Barnard, 1925: 141.

Regan & Trewavas, 1929: 32, fig. 24. Norman, 1930: 308; 1939: 21.

(For full synonymy see Morrow 1964: 287.)

Material

SAM 26607, 4 (144-192 mm).

MESOPELAGIC FISHES FROM VEMA SEAMOUNT 239

Family Scopelarchidae

Neoscopelarchoides sp.

Pig. 1

Neoscopelarchoides sp. Marshall, 1955: 311, fig. 3.

Material

SAM 26608 1 (49,9 mm).

Remarks

This specimen is a late larval stage. The lingual teeth resemble those

figured by Marshall (1955: fig. 3 a*).

Fic. 1. Neoscopelarchoides sp. (SAM 26608). A. Lateral view. B. Lingual teeth. Scale 10 mm.

Family Scopelosauridae

Luciosudis norman Fraser-Brunner, 1931

Luciosudis normani Fraser-Brunner, 1931: 220, fig. 2. Krefft, 1968: 95, figs 1-4.

Scopelosaurus normani Marshall, 1966: 196.

Material

SAM 26610, 1 (47,5 mm).

Description

D 11; A 18; gill-rakers 1 +1-+28; Vert 58.

Remarks

The specimen has been identified by Dr G. Krefft, Hamburg.

Family Myctophidae

Protomyctophum (Hierops) subparallelum (Taning, 1932)

Myctophim arcticum subparallelum Taning, 1932: 128.

Electrona (Hierops) arctica subparallela Fraser-Brunner, 1949: 1047.

Hierops subparallela Bolin, 1959: 5.

Protomyctophum (Hierops) subparallelum Andriashev, 1962: 243, figs 14, 15. Nafpaktitis & Nafpak-

titis, 1969: 8, figs 4-6. Hulley, 1972: 219.

(For full synonymy see Andriashev 1962: 243.)

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM 26609, 9 (damaged).

Description (composite)

D 11-12; A 21; P15; V 8; gill-rakers 4+1-+14 (13); AO 15.

Remarks

The heads of all the specimens are missing and the sides of the bodies are

badly scraped. ‘They have been identified as this species because:

(1) PLO not reduced in size and situated in front of PVQO,;

(2) anal origin below middle of dorsal;

(3) AO 15, in unbroken series, with 3 photophores behind anal base;

(4) Last AO separated from Prc, by distance greater than Prc,—Prcg.

Electrona rissot (Cocco, 1829)

Scopelus rissot Cocco, 1829: 15, pl. 2, fig. 5.

Myctophum rissoi: Brauer, 1906: 170, fig. 83. Norman, 1930: 320. Fowler, 1936: 380, fig. 184.

Electrona (Electrona) rissot Fraser-Brunner, 1949: 1048, fig.

Electrona rissoi Bolin. 1959: 5. Becker, 1967: 92. Trunov, 1968: 745, fig. 1. Nafpaktitis & Nafpak-

titis, 1969: 10, figs 4, 6, 8. Hulley, 1972: 220.

(For full synonymy see Bolin 1959: 5.)

Material

SAM 26611, 4 (47,1-64,6 mm).

Description

D13; A 18-20; P 15 (16); V 8; gill-rakers 8+1+418 (19); AO 11; Pre 2.

Diaphus effulgens (Goode & Bean, 1896)

Aethopora effulgens Goode & Bean, 1896: 87, pl. 27, fig. 103. Jordan & Evermann 1896: 566.

Myctophum (Nyctophum) effulgens Brauer, 1904: 393.

Diaphus effulgens Taning, 1928: 62. Nafpaktitis, 1968: 70, figs 44-46.

Diaphus (Lamprossa) effulgens Fraser-Brunner, 1949: 1071, fig.

Diaphus elucens: Smith, 1961: 122, fig. 201 (partim).

(For full synonymy see Nafpaktitis 1968: 70.)

Material

SAM 26612, 1 (99,8 mm).

Description

D 16; A 15; P 12; V 8; gill-rakers 6+1+13 = 20. Lateral line scales 36.

AO 6+5; Pre 4; PLO nearer pectoral base than lateral line; Dn extending

higher than level of dorsal margin of eye.

Lampanyctus australis Taning, 1932

Lampanyctus alatus australis Taning, 1932: 145.

Myctophum (Lampanyctus) alatus: Barnard, 1925: 240.

Lampanyetus australis : Becker, 1967: 109. Nafpaktitis & Nafpaktitis, 1969: 54, figs 68. 69. Hulley,

1972: 224.

Material

SAM 26613, 5 (89,1-103,4 mm).

Description

D 13 (14); A 18 (19); P 14 (15); V 8; gill-rakers 6 (5)+-1--13)0tae

AOa‘s (7); AOp 7 (a)> total AO ee.

MESOPELAGIC FISHES FROM VEMA SEAMOUNT 241

Lepidophanes supralateralis (Parr, 1928)

Big. 2

Lampanyctus supralateralis Parr, 1928: 94, fig. 12.

Lampanyctus (Lepidophanes) supralateralis Fraser-Brunner, 1949: 1091, fig.

Lepidophanes supralateralis Bolin, 1959: 34.

? Lampanyctus superlateratus: Grindley & Penrith, 1965: 283 (lapsus calami).

Material

SAM 26614, 1 (45,3 mm).

Description

D 13; A 14; P 14; V 8; gill-rakers 6-+1-+12 = 1g. Crescent of luminous

tissue on posterior rim of iris; PLO, SAO,, Pol and Prc, just above lateral line;

VLO well below lateral line; PO 5, the fourth elevated; VO 5, the posterior

four forming a straight descending series from elevated second; SAO 3, weakly

angulate; AO 5+4; Pol 2; Prec 3, well separated from AOP; 2 supracaudal and

3 infracaudal plates; 4 luminous scales on base of anal.

Remarks

This specimen represents the first record of the species in the eastern

South Atlantic. Grindley & Penrith (1965) have recorded specimens as L. super-

lateratus Parr from the south-western Indian Ocean.

Fic. 2. Lepidophanes supralateralis (Parr). Scale 10 mm.

Family Melamphaeidae

Melamphaes suborbitalis (Gill, 1883)

Plectromus suborbitalis Gill, 1883: 258. Norman, 1929: 157.

Melamphaes suborbitalis Giinther, 1887: 30. Fowler, 1936: 1265 (partim). Ebeling, 1962: 56, fig. 23.

(For full synonymy see Ebeling 1962: 56.)

Material

SAM 26615, 1 (76,0 mm).

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

D Ill, 16; A I, 8; P 15; V I, 7, inserted behind origin of pectoral; gill-

rakers (1st arch) 6+15; gill-rakers (4th arch) 11 on lower limb. 3 pores on

cheek inside angle. Sharp, antrose, posttemporal spine present.

Remarks

Although the possible existence of this species in the South Atlantic has

already been reported (Hulley 1972), this specimen represents the first record

of the species in the region.

Family Scombropidae

Howella brodiet Ogilby, 1899

Howela brodiet Ogilby, 1899: 735. Hulley, 1972: 229.

Rectogramma sherborni Norman, 1930: 348, fig. 39. Fowler, 1936: 1287, fig. 547.

Howella sherborni: Smith, 1961: 212, fig. 498. Grindley & Penrith, 1965: 284.

Material

SAM 26616, 1 (77,8 mm).

Description

D VIII-+I, 9; AIII, 7; P15; VI, 5; gill-rakers 7-+-1-+21.

Family ? Percichthyidae

Material

SAM 26617; 1 (31,1 mm), -

Remarks

The specimen has been badly damaged by the net, so that fin counts are

difficult to assess.

SUMMARY

A collection of mesopelagic fishes is described. ‘The material was taken at

31°38'S, 8°21.5’E (IK 52) during the survey cruise of R.S. Thomas B. Davey

to Vema Seamount, and includes 16 species, of which 2 are new records.

ACKNOWLEDGEMENTS

I am grateful to the Department of Oceanography, University of Cape

Town, and Prof. J. R. Grindley, formerly of the South African Museum, for

arranging the Vema cruise. I also wish to thank Mr M. J. Penrith for his help

on board, and Mr S. X. Kannemeyer for his assistance with sorting the material.

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MESOPELAGIC FISHES FROM VEMA SEAMOUNT 243

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Fow.er, H. W. 1936. Marine fishes of West Africa based on the collection of the American

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FRASER-BRUNNER, A. 1931. Some interesting West African fishes, with descriptions of a new

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FRASER-BRUNNER, A. 1949. A classification of the fishes of the family Myctophidae. Proc. zool.

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Gipss, R. H. & Hurwitz, B. A. 1967. Systematics and zoogeography of the stomiatoid fishes,

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Grey, M. 1960. A preliminary review of the family Gonostomatidae, with a key to the genera

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R.S.‘ Africana II’, 1961-1966. Ann. S. Afr. Mus. 60 (6): 197-236.

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catalogue of the species of fish-like vertebrates found in the waters of North America, north

of the Isthmus of Panama. Bull. U.S. natn. Mus. 47: 1-ccxiv, 1-3313.

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IV. Luciosudis Fraser-Brunner, 1931, ein valides Genus der Familie Scopelosauridae

(Osteichthyes, Alepisauroidei). Arch. Fisch Wiss. 19: 95-102.

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* Not seen by the author.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

MarsHaLL, N. B. 1966. Family Scopelosauridae. In Fishes of the western North Atlantic. 5 Mem.

Sears Fdn mar. Res. 1 (5): 194-204.

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searsids and the taxonomy of Sagamichthys and Persparsia. Copeia 1971: 440-448.

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NaFPakxtTITIs, B. G. 1968. Taxonomy and distribution of the lanternfishes, genera Lobianchia

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NarFpaktitis, B. G. & Narpaxtiris, M. 1969. Lanternfishes (Family Myctophidae) collected

during Cruises 3 and 6 of R/V Anton Bruun in the Indian Ocean. Bull. Los Ang. Cty Mus. nat.

Hist. 5: 1-79.

Norman, J. R. 1929. A preliminary revision of the berycoid fishes of the genus Melamphaés.

Ann. Mag. nat. Hist. (10) 4: 153-168.

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Norman, J. R. 1939. Fishes. Scient. Rep. John Murray Exped. 7: 1-116.

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Scuuttz, L. P. 1938. Review of the fishes of the genera Polyipnus and Arg yropelecus (family

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INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins, arranged in the following order:

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(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-

headings to be used sparingly and enumeration of headings to be avoided. (4) Summary

(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

ILLUSTRATIONS

To be reducible to 12 cm x 18 cm (19 cm including caption). A metric scale to appear

with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their

proper place in the text), with arabic numbering; items of composite figures to be designated

by capital letters (A, B, C etc.).

REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

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

FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

FiscHEr, P.-H., Duva, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.

Archs Kool. 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. 1960). Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn scHULTZE, L.

Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature issued

by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).

The Harvard system of reference to be used in the synonymy lists; with the full references

incorporated in the list at the end of the article, and not given in contracted form in the synonymy

list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

P. A. Hhulley

MESOPELAGIC FISHES

FROM VEMA SEAMOUNT

(IK STATION 52)

| VOLUME 60 PART 8 =— NOVEMBER 1972

meets

ANNALS

OF THE SOUTH AFRICAN

MUSEUM

CAPE ‘TOWN

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60 Band

November 1972 November

Part 8 Deel

mee CRETACEOUS STRATIGRAPHY OF

SAN NICOLAU AND SALINAS, ANGOLA

MICHAEL R. COOPER

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE CRETACEOUS STRATIGRAPHY OF SAN NICOLAU AND

SALINAS, ANGOLA

MicHAEL R. CoorerR

South African Museum, Cape Town

(With 2 figures)

[MS. accepted 29 Fune 1972]

CONTENTS

PAGE

Introduction . 5 - 245

Present investigation : 2 2A

Summary. ‘ ; s ) -250

Acknowledgements. : = QE

References. ; : Pe eae

INTRODUCTION

The paucity of ammonites within much of the Cretaceous of Angola has

led to the widespread use of other taxa, such as bivalves, echinoids and shark’s

teeth as stratigraphic indicators, often with somewhat spurious results.

Mouta & Borges (1926), in studying the stratigraphy of the Benguela and

Mocamedes basins, recognized the following stratigraphical succession:

Formations with Roundaireia forbesiana Stoliczka and R. drui Munier-Chalmas— Senonian

Beds of sandy and oolitic limestones with Acteonela anchietai Choffat and Nerineia capelloi

Choffat—Turonian

Strata with Nezthea tricostata Coquand—Cenomanian

Limestones and shales with ammonites (Dowvilleiceras mammillatum (Schlotheim) and

Stoliczkaia dispar (d’Orbigny)) — Albian

Strata with Pholadomya pleuromyaeiformis Choffat— Albian

Lower formations with gypsum— Aptian

These authors noted that at San Nicolau the Senonian overlies red arenites,

and they mentioned a basaltic layer within the former. ‘The Senonian beds

underlying the basalts were considered Coniacian on the basis of the similarity

of the fauna to that from the Coniacian of Tunis, including Pecten virgatus Nils,

Trigonarca cf. trichonopolytensis Forbes, Trigonia scabra Lamarck, Cyprina (Venili-

cardia) cf. barroist Coquand, Roundaireia forbesiana Stoliczka, Venus plana J.

Sowerby and Corbula elegans J. de C. Sowerby. They noted that the beds over-

lying the basalts contained Crassatella numidica Munier-Chalmas and Cardita

beaumonti d’Archiac, species characteristic of the Maastrichtian of ‘Tunis, and

Roundaireia drut, a species common in the Campanian and Maastrichtian.

Douvillé (1931) described eight species of ammonites collected at Salinas

(see Fig. 1), to which he assigned a Barremian to Turonian age. Spath (1931,

1932, 1951), in reviews of Douvillé’s paper, showed the fauna to be of Ceno-

manian age, an age supported by the fact that the entire fauna ‘. . . had all

been collected in a bed of only 4 m in thickness’ (Spath 1951: 129).

245

Ann. S. Afr. Mus. 60 (8), 1972: 245-251, 2 figs.

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fauna Studied by

Douville & Spath

Jombovacai

Fauna studied by

Howarth, 1968

Turon— L.Coniac.

fauna

=| Limestones & silts

ites etc

Limestones & silts fauna

“= Red arenites

Fic. 1. Locality map (after Carvalho 1961).

Mouta & O’Donnell (1933) and Mouta (1937) studied the Cretaceous

of the Mocamedes and Benguela basins and considered the Senonian to be

represented by the strata with Roundaireia forbesiana and R. drut at San Nicolau,

and by the ammonites of Salinas. At San Nicolau, Mouta & O’Donnell (1933)

considered the Senonian to be represented by two horizons, the Coniacian and

Maastrichtian, separated by a basaltic layer and characterized by R. forbesiana

and &. drut respectively.

Rennie (1929, 1945) described collections of bivalves and gastropods from

San Nicolau and Salinas. The presence of Trigonoarca angolensis Rennie and

CRETACEOUS STRATIGRAPHY OF SAN NICOLAU AND SALINAS, ANGOLA 247

Trgonocallista umzambiensis Woods at Salinas led Rennie to believe the beds

to be of Campanian-Maastrichtian age. The San Nicolau fauna, with Turritella

(Karta) boner Baily, Trigonoarca cf. trichonopolytensis (Forbes), Trigonia (Scabro-

trigonia) shepstoner. Griesbach, and Lima (Mantellum) sp., was correlated with the

Baba fauna and also included within the Campanian-Maastrichtian.

Spath (1951) described Eutrephoceras indicum (Spengler) and Baculites aff.

asper (Morton) from the beds overlying the basalt, placing them ‘. . . probably

also very high in the Senonian’.

In 1958 Carvalho reviewed the problems associated with the Cretaceous

formations of the Mocamedes basin and envisaged the following depositional

history:

During the Aptian—Albian a lagoonal basin was dominant over the region, slowly filling

with sediments, only to be overlain by thick torrential deposits marking an epoch of

regression and the proximity of high relief

Subsequently the Albian sea covered the north of the basin

During the Cenomanian and Turonian, the southern portion of the basin emerged, whilst

marine deposits were accumulating in the northern portion

The Coniacian and Santonian was an epoch of emergence, during which the region fell

under the influence of volcanic activity, with the eruption of the basalts

During the Campanian, there was a widespread transgression with the laying down of

sediments on a peneplained surface of Basement, early Cretaceous sediments and basalt

Within the southern region the sea regressed, whilst deposition continued in the north

The Maastrichtian saw a new phase of marine transgression, with the accumulation of

deposits throughout the region

In 1961 Carvalho mapped the marine beds below the basalts at San

Nicolau as Cenomanian-Turonian, and the overlying beds as Campanian,

considering the basaltic rocks, comprising analcime tephrites and basanites,

basalts and andesites (Andrade 1957: 742), as having occurred ‘. . . at the end

of the Turonian or during the Coniacian-Santonian’.

Howarth (1968) described an ammonite fauna from Ponta Grossa, about

3 km to the south of Salinas, in beds underlying the volcanics, to which he

assigned a Middle Turonian age.

Kennedy (1971), in referring to the faunas described by Douvillé and

Spath, considered ‘. . . all but the ‘“‘Stoliczkaia’” and “‘Pulchellia’’ indicate a high

Cenomanian or basal Turonian age. The Stoliczkaia is taken by Spath (1931)

and Howarth (1965) to be of Cenomanian age, but must surely be Lower

Cenomanian, whilst the ‘‘Pulchellia’? seems a misidentified acanthoceratid

(Spath 1931)’.

PRESENT INVESTIGATION

A recent examination of Douvillé’s type locality at Salinas, together with

exposures to the east of San Nicolau, has led to stratigraphical conclusions

somewhat different from those proposed by previous authors.

At Salinas, and along the beach to the south of Ponta Grossa, at least

35 m of unfossiliferous red sands and silts are exposed in the sea-cliffs and are

of pre-uppermost Cenomanian age. The first fossiliferous horizon is charac-

terized by the gastropod Pseudomelania salenasensis Rennie, together with

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

abundant Exogyra cf. columba (Lamarck), Protocardia hillana (J. Sowerby),

Exogyra olisipinensis Sharpe and Veniella forbesiana (Stoliczka). It was from these

beds that the collection described by Rennie (1945) as of Campanian-Maastrich-

tian age came. Unfortunately, ammonites are lacking. This horizon is succeeded,

a few metres higher up, by about 6 m of silts with abundant nodular limestone

horizons, the latter representing the source of Douvillé’s fauna. These beds are

characterized by the abundance of Exogyra cf. columba, together with the

ammonites Calycoceras naviculare (Mantell), Aanabiceras septemseriatum (Cragin),

Austiniceras dibleyi Spath and Pseudocalycoceras angolaense (Spath). Other ammo-

nite genera include Gaudryceras, Eucalycoceras, Protacanthoceras, Sciponoceras,

Tetragonites, Metoicoceras and ‘Stoliczkaia’. The proximity of the Turonian

boundary, suspected by Kennedy (1971), is confirmed by a single, fragmentary

specimen of Gombeoceras, found in surface scree.

The Salinas exposure, as exemplified by the faunas of Douvillé and Spath,

is therefore of uppermost Cenomanian age, and is conformably succeeded by

Turonian strata.

Exposures between those at Salinas and the coastal outcrop of the vol-

canics, 7 km to the south, are confined almost entirely to the Ponta Grossa

locality of Howarth (1968) which, at the time of my visit, was unfortunately

inaccessible. From this locality Howarth described a ‘mid-Turonian’ fauna

comprising eight ammonites belonging to the following six species: Anagaudry-

ceras involvulum (Stoliczka), Gaudryceras varagurense Kossmat, Mesopuzosia yubarense

(Jimbo), Damesites ainuanus Matsumoto, Mammites mocamedensis Howarth and

Prionocyclus carvalhoi Howarth. The fact that Mammites is predominantly a

Lower Turonian genus, while Damesites ainuanus is recorded from the Upper

Turonian of Japan and Prionocyclus carvalhoi is closest to the Middle Turonian

P. hyatti (Stanton), led Howarth to date the fauna ‘. . . with some certainty to

be very close to the mid point of the Turonian’,

In the sea-cliffs 6 km due west of San Nicolau (14°15’S, 12°23’E), and

about 5 m below the volcanics, fossiliferous strata are well-exposed in a deep

gully. This fauna is characterized by the abundance of Veniella forbesiana

(Stoliczka), together with Prionocyclus carvalhoi, Mammites mocamedensis and

Damesites ainuanus and the ammonite genera Proplacenticeras, Mesopuzosia,

Subprionocyclus, ?Subtissotia, Baculites, Gaudryceras, Scalarites, hossmaticeras,

Hauericeras and Hypophylloceras. Of these additional specimens the placenticera-

tids and the Mesopuzosia sp. closely resemble forms common in the Lower

Coniacian of Zululand, Subprionocyclus is characteristic of Upper Turonian

strata, Scalarites is known from the Turonian and Coniacian of Morocco

(Collignon 1966), the Hawericeras sp. is very close to H. antiquum Collignon

(Collignon 1961: 76) from the Lower Coniacian of Madagascar, while Sub-

lissotia is known from the Coniacian of Tunisia. The baculitids are, according

to Dr W. A. Cobban (United States Geological Survey, Denver), who kindly

examined them for me, closest to undescribed Upper Turonian forms from New

Mexico.

CRETACEOUS STRATIGRAPHY OF SAN NICOLAU AND SALINAS, ANGOLA 249

Immediately below this fossiliferous horizon is a bed rich in stromatolites.

This deposit has, therefore, all the features of a classical condensed sequence,

as recorded by Rod (1946), Jenkyns (1971) and others, viz. faunal enrichment

and faunal mixing, negligible thickness and widespread distribution, and

associated stromatolitic algae. Sedimentological evidence supports this, since

in the coastal cliffs west of San Nicolau about 25 m of limestones and silts are

seen to separate the volcanics from the unfossiliferous red sands, whereas the

measured section in the gully totalled 14 m, while the Upper Cenomanian at

Salinas was at least another 8 m. Howarth’s fauna is, without question, from

the same horizon which must be considered a condensed sequence spanning

Turonian to lowest Coniacian times.

Red sandstones

VVVV

= Basaltic rocks

2 Gata

VVvV Limestones

WIG? =P Ee

Fic. 2. Geological section through the west side

of the Jombovacai valley, about 2 km north of

the Posto de San Nicolau (for explanation see

text).

Deposition of this stratum was followed by a period of emergence and the

eruption of the alkali-basalts. The presence of a disconformity between these

beds and the basalts is evident in the section exposed in the western side of the

Jombovacai valley (see Fig. 2) about 2 km north of the Posto de San Nicolau.

Here, 3 m of highly fossiliferous white limestone, unfortunately lacking in

ammonites, are crammed with bivalves and gastropods with their recrystallized

shells preserved. These are overlain by 0,28 m of unfossiliferous maroon silts

succeeded by 1,05 m of hard red limestone, unlike any exposed in the coastal

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

section, in which the fossils are preserved only as internal moulds. At the base

of this horizon is a thin, small-pebble conglomerate. This red limestone horizon

is separated from the basanites by 1,54 m of unfossiliferous red silts.

The completely different mode of preservation of the fauna within this

latter limestone, together with its red colour, suggests the disconformity to lie

immediately above this horizon. The absence of pillow structures within the

lavas also testifies to their subaerial extrusion. In the Jombovacai valley at

least two distinct phases of lava extrusion are indicated by the occurrence of

1,33 m of interbedded red silts and sandstones.

During early Santonian times a widespread transgression resulted in the

deposition of marine beds on the newly laid down volcanics. That these beds

are, at their base, Lower Santonian is indicated by the occurrence of the

ammonite genera Texanites, Protexanites, Hauericeras and Damesites. This fauna

occurs in orange limestones characterized by the abundance of Cardium

(Trachycardium) reynoldsi Rennie.

The volcanics are, therefore, restricted to the Coniacian, probably only

the Upper Coniacian, and are consequently of much more limited duration

than previously thought.

With the exception of pyritized (now limonitic) worm burrows, fossils

are absent from the succeeding green silts until the occurrence of abundant

vertebrate remains and shark’s teeth about 20 m higher up. The most abundant

fossils are Anacorax pristodontus (Agassiz), together with Lamna_ biauriculata

Wanner, Rhombodus binkhorsti Dames, Enchodus sp. and Mosasaurus beaugei

Arambourg. On purely sedimentological evidence it seems unlikely that these

beds are indeed Maastrichtian, as suggested by Dartevelle (1942) from a study

of the shark’s teeth, since most of the Santonian and the entire Campanian

would have to be accommodated within 20 m of sediment. Furthermore, there

is no sign of the rich uppermost Campanian fauna of Egito, nor the Maas-

trichtian faunas of the Cuanza basin.

SUMMARY

At Salinas red arenites are overlain by limestones containing Calycoceras

naviculare, Kanabiceras septemseriatum and Austiniceras dibleyi, species characteristic

of the Sczponoceras gracile—Metorcoceras whitei zone of North America. The

absence of Mammites, present in higher beds to the south, suggests these beds

to be of uppermost Cenomanian rather than basal ‘Turonian age, although a

single specimen of Gombeoceras collected in surface scree indicates the proximity

of the latter stage. There is no evidence for Upper Albian or Lower Cenomanian

strata at Salinas. ‘These beds are conformably overlain by the ‘Turonian, repre-

sented by a condensed sequence as evidenced by the enriched and mixed faunal

assemblage, and the diminutive stratal thickness, with an admixture of basal

Coniacian forms. Deposition of these strata was followed by a period of

emergence and the eruption of the alkali-basalts, probably during Upper

Coniacian times. Following this period of volcanic extrusion, a marine trans-

CRETACEOUS STRATIGRAPHY OF SAN NICOLAU AND SALINAS, ANGOLA 251

gression deposited limestones with Protexanites and abundant Texanites, and

thus of Lower Santonian age. About 30 m above this horizon are beds with

abundant vertebrate remains and shark’s teeth. On purely sedimentological

evidence their Maastrichtian assignment is considered dubious.

ACKNOWLEDGEMENTS

I should like to express my thanks to Professor Virgilio Cannas Martins,

director of the Instituto de Investigagao Cientifica de Angola, for the assistance

given me by his institute, without which this study would have been impossible.

I should also like to express my thanks to Mr Joaquim Torquato of the same

institute for his help and assistance. To Dr Anténio Graca da Cruz, Director

of the Servicos de Geologia e Minas, I am indebted for his co-operation and

kindness.

I am especially grateful to Dr W. J. Kennedy of Oxford University for his

kind help and valuable suggestions.

REFERENCES

ANDRADE, M. M. de. 1957. Rochas vulcanicas da Orla Meso-Cenozéica entre Benguela e

Mocamedes. Garcia de Orta 5: 739-766.

CarvALHo, G. S. de. 1958. As formac6es cretacicas da bacia de Mocgamedes (Angola) e alguns

dos seus problemas. Publcées Mus. Lab. miner. geol. Univ. Porto (3) 75: 1-32.

CarvaAtuHo, G. S. de. 1961. Geologia do deserto de Mocamedes (Angola). Mems Fta Invest.

Ultramar (2), 26: 1-227.

Cotiicnon, M. 1961. Ammonites néocrétacées du Menabe (Madagascar). VII. Les Desmocerati-

dae. Annls géol. Madagascar 31: 1-115.

Coiiicnon, M. 1966. Les céphalopodes crétacés du bassin cotier de Tarfaya. Notes Mém. Serv.

Mines Carte géol. Maroc 175: 1-148.

DovuvitLe£, H. 1931. Contribution a la géologie de l’Angola. Bolm Mus. Lab. miner. géol. Univ.

Lisboa 1: 17-46.

HowartTH, M. K. 1965. Cretaceous ammonites and nautiloids from Angola. Bull. Br. Mus.

nat. Hist. (Geol.) 10: 335-412.

Howarth, M. K. 1968. A mid-Turonian ammonite fauna from the Mogamedes desert, Angola.

Garcia de Orta 14: 217-228.

Jenxyns, H. C. 1971. The genesis of condensed sequences in the Tethyan Jurassic. Lethaia 4:

327-352.

KENNEDY, W. J. 1971. Cenomanian ammonites from southern England. Spec. Pap. Palaeont.

8: 1-133.

Moura, F. 1937. Notice géologique sur |’Angola (Afrique Occidentale Portugaise). Comungdes

Servs geol. Port. 20: 19-37.

Mouta, F. & Borces, A. 1926. Sur le crétacé du littoral de Angola (Districts de Benguela

et Mossamedes). Bolm Ag. ger. Colén. Ultramar 14: 30-55.

Moura, F. & O’DonneELL, H. 1933. Notice explicative. Carte geologique de l’ Angola. 1 : 2 000 000.

Lisboa: Ministério das Coldénias.

Renniz, J. V. L. 1929. Cretaceous fossils from Angola (Lamellibranchia and Gastropoda).

Ann. S. Afr. Mus. 28: 1-54.

Rennig, J. V. L. 1945. Lamelibranquios e gastrépodos do Cretacico Superior de Angola. Mems

Jia Miss. geogr. Invest. Colon. (Geol.) 1: 1-141.

Rop, E. 1946. Uber ein fossillager im oberen Malm der Melchtaleralpen. Eclog. geol. Helv.

39: 177-198.

SpatH, L. F. 1931. A monograph of the Ammonoidea of the Gault. Palaeontogr. Soc. [Monogr.]

2 (3): 316.

SpaTH, L. F. 1932. Review of ‘H. Douvillé, 1931, Les ammonites de Salinas’. Geol. ZentBl. 1: 124.

Spatu, L. F. 1951. Preliminary notice on some Upper Cretaceous ammonite faunas from

Angola. Comungées Servs geol. Port. 32: 123-130.

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REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

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Examples (note capitalization and punctuation)

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

FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

FiscHER, P.-H., Duvat, M. & Rarry, A. 1933. Etudes surles échanges respiratoires des littorines.

Archs Zool. exp. gén. 74: 627-634.

Konun, 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. 1960. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): I-51.

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

Koologische und anthropologische Ergebnisse einer Forschungsreise 1m westlichen und zentralen Stid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

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’

‘ 7

of ; net

: ‘ ? }

? } ' ‘

a eo

Michael R. Cooper

THE CRETACEOUS STRATIGRAPHY OF

SAN NICOLAU AND SALINAS, ANGOLA

ANNALS

OF THE SOUTH AFRICAN

MUSEUM

CAPE ‘TOWN

aN AS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60 Band

November 1972 November

Part 9 Deel

mOINE VW SPECIES OF SOUTHERN AFRICAN

BREVIRAJID SKATE

HON DRICHTHYES, BATOIDEI RAJIDAE)

Pes AOL

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A NEW SPECIES OF SOUTHERN AFRICAN BREVIRAJID SKATE

(CHONDRICHTHYES, BATOIDEI, RAJIDAE)

LeeeNey eo hOie i Mo

South African Museum, Cape Town

(With 5 figures and 1 table)

[MS'. accepted 3 July 1972]

CONTENTS

PAGE

Introduction ; : ; 1 2 5e

Description of material . : . : Le AL

Discussion : : : : 4 : ; « » 262

Summary : : : : : : : / . 963

Acknowledgements , : : : . 263

References : : : ; ; ; j . 263

INTRODUCTION

In his revision of the southern African Rajidae, Norman (1935) held that

the two spinulose juveniles, reported as Raa plutonia Garman by Barnard (1925),

were specimens of Raja spinacidermis, since their dorsal surfaces were entirely

covered with small asperities and the enlarged thorns along the midline of the

back and tail showed signs of disappearance. Although these specimens were

not available to him, Hulley (1970), on the basis of two further specimens

(SAM 22911, 24450), tentatively accepted Norman’s synonymy. However, he

pointed out that, not only are there differences between these specimens and

Raja spinacidermis in the spination of the orbital region, the shape of the disc

and the tail length, but also there were marked differences in the tooth count

(36-38 rows in the upper jaw, compared with 54-60 rows in Raya spinacidermis).

The Division of Sea Fisheries, Cape Town, has recently collected 15 speci-

mens of these spinulose rajids, during the 1972 Hake Survey, and have donated

this material to the South African Museum. Examination of the rostral cartilage

and appendices and of the anatomy of the clasper of these specimens reveals

that they should be described as a new species, Breviraja stehmanni, and as such

represent the first record of the genus outside the waters of the western central

Atlantic. SAM 24450 can be identified as this species.

The genus Breviraja Bigelow & Schroeder, 1948, was constructed to

incorporate those rajids having ‘the tip of the rostral cartilage falling short of

the extremities of the pectoral rays and of tip of snout’, diagnostic characters

which were based on X-ray photographs. However, Ishiyama & Hubbs (1968)

point out that the interpretation of these photographs is erroneous, and that the

rostral cartilage is not foreshortened and extends virtually to the tip of the

snout, beyond the anterior tips of the pectoral rays. On the basis of the rostral

structure and the anatomy of the clasper, Ishiyama & Hubbs (1968) substan-

253

Ann. S. Afr. Mus. 60 (9) 1972: 253-263, 5 figs, 1 table.

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

tiate the validity of Brevraja and defined a further genus, Bathyraja, confining

it to the Pacific. Stehmann (1970) and Hulley (1970, 1972) agree with these

findings, but point out that the genus Bathyraja also exhibits a discontinuous,

antitropical distribution pattern in the Atlantic, while the genus Breviraja

appears to be confined to slope regions of the western central Atlantic. Further-

more, both Stehmann (1970) and Hulley (1972) have commented on the

phylogenetic position. of Breviraja.

Breviraja stehmanni is separated from all other southern African rajids by the

peculiar form of the rostral cartilage and rostral appendices, and is at present

the only species of this genus recorded from the region. It may easily be dis-

tinguished from all other species of the genus, except B. plutonia (Garman) and

B. cubensis Bigelow & Schroeder, by its long, cross-barred tail and dark dorsal

fins. It most closely resembles B. plutonia in that the dorsal fins are continuous,

but differs markedly from this species in tooth count, length of the anterior

lobe of the pelvic fin and number of scapular thorns. In these characters it

approximates B. cubensis, but may be distinguished from this species by its

continuous dorsal fins. Furthermore, B. stehmanni differs from both these species

in its comparatively larger disc, shorter tail, smaller eye and shorter anterior

pelvic lobe, and especially in the presence of a single, median row of larger

thorns on the back and tail, which extends to about one-half to two-thirds the

length of the tail.

The species is named in honour of Dr M. Stehmann, Institut fiir Seefis-

cherei, Hamburg.

DESCRIPTION OF MATERIAL

Breviraja stehmannzt n. sp.

Raia plutonia: Barnard, 1925: 68.

Raja spinacidermis: Norman, 1935: 46 (partim).

?Raja spinacidermis: Hulley, 1970: 173, pl. 4, fig. A.

Types

The type, an adult male (354,3 mm total length) and paratype, a female

(299,5 mm total length), trawled between 33°53,7'S, 17°23,9'E and 33°57,3'9,

17°22,2’E in 640 m (Division of Sea Fisheries, Station No. A 5854), in the

collection of the South African Museum (SAM 26636, 26637).

Material

The type and paratype, and 13 specimens of both sexes (132,5-340,0 mm

total length) trawled from two stations, A 5854 (10 April 1972: 33°53,7'S,

17°23,9'E-33°57,3'S, 17°22,2'E; 640 m; bottom temp. 5,55°C) and A 5871

(33°55,6'S, 17°25,0'E-33°56,1'S, 17°26,8’E; 600 m; bottom temp. 5,70 @);

and 1 specimen, a female (305 mm total length), trawled west of Cape Town in

160 fms (292 m). All specimens in the collection of South African Museum

(SAM 24450, 26638, 26639).

255

A NEW SPECIES OF SOUTHERN AFRICAN BREVIRAJID SKATE

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ANNALS OF THE SOUTH AFRICAN MUSEUM

256

‘UI PUL WD UI 9[VIG “MOTIA [RIJUIA “G ‘MOIA [eSIOG "VY ‘AdALVUVd ‘ds ‘u wmuvuyas vlog

i 8) |

A NEW SPECIES OF SOUTHERN AFRICAN BREVIRAJID SKATE 257

1,0cm

/———___}

Fic. 3:

Breviraja stehmanni n. sp. Neurocranium, rostral bar and appendices, and anterior extremities of

pectoral rays.

Description

Measurements for the type and paratype, respectively, are given, while

the figures in parentheses refer to the range of variation for 14 specimens.

Disc, 1,3; 1,2 (1,2-1,4) times as broad as long, its width 1,8; 1,9 (1,7—1,9)

in total length; obtuse in front, tip of snout marked by a low projection, with

maximum angle in front of spiracles 123°; 129° (115°—130°); anterior margins

slightly concave just behind tip of snout and again at level of spiracles, more

strongly so in adult males than in females and juveniles; outer angles broadly

rounded, posterior and inner margins strongly convex. Axis of greatest breadth

1,4; 1,5 (1,3-1,7) times as far from tip of snout as from posterior edge of disc.

Tail with narrow lateral folds along posterior third; its length from middle of

vent to origin of first dorsal fin 1,2; 1,3 (1,1—-1,3) times as long as distance from

middle of vent to tip of snout.

Entire upper surface of disc and tail with small asperities, except tip of

snout, narrow anterior margin of pectoral and anterior lobe of pelvic, and

mature males with naked area at pectoral base. Orbit with 4—5 (1-6) thorns at

anterior margin, well separated from 3 (1-4) thorns on posterior margin and

above spiracle; 1 very small interspiracular thorn on each side in some juveniles ;

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

Measurements expressed as permillage of the total length.

Type Paratype Range Mean

Total length .. ye oe: Ai pis a: 1000 1000

Disc width si ee a - = a 569 537 540-584 563

Disc length My, be ee te oe 444 437 423-471 456

Snout to greatest disc width .. is a si 261 261 265-300 279

Snout to middle of vent ie 3k 53 bye 403 340 393-414 402

Snout to axils of pelvics hae Be a Pe 352 410 338-365 352

Middle of vent to 1st dorsal origin .. of Se 488 456 470-498 486

Snout length .. We te of a - 93 94 89-115 104

Preoral length se ap _ ae ae 93 99 89-120 108

Prenasal length i es oe sth ae, 68 71 66-88 79

Eye-horizontal diameter ae ti 5 4 32 30 28-36 30

Eye + spiracle .. + ae 5% a : 45 47 45-56 48

Spiracle a He Bis a, o 29 27 21-36 28

Interorbital distance .. Ze os ais Mad 31 30 30-37 33

Interspiracular distance as Ba oe Me 67 72 67-75 71

Internasal distance a a - ne vy. 62 60 59-66 63

Mouth width .. a sf oe at <r 79 73 69-90 76

Gill slit lengths: Ist .. he ee is he II 16 II-14 13

BIEL | Ge oe te: a oh 12 16 11-16 14

5th Me i sé = II 10 9-13 II

Distance between inner ends of gill slits: 1st cB 129 131 I2I-I4I 13I

5th -: 69 79 67-90 |

ist dorsal fin: height .. ae ae 2M a Si 28 16-31 26

base length .. an ae he 54 56 44-53 49

and dorsal fin: height .. ee ee as Pra QI 33 16-34 28

base length .. ste aM 4 55 56 43-56 50

Interdorsal space i Be aE a ie fe) oO )

3-4 (1-4) median nuchal thorns; 1 scapular thorn (sometimes lost, but scar

present) on each side; a series of 26-39 (11-38) thorns along midline of back

and tail, usually interrupted between pectoral and pelvic girdles, extending

posteriorly to between one-half and two-thirds the length of the tail, showing

alternate development with increasing age; tail posterior to axis of pelvics

without additional rows of thorns or large prickles; anterior parts of dorsal fins

with small asperities. Sexually mature males with well-developed malar and

alar spines. Ventral surface smooth.

Snout rounded and barely produced, except somewhat so in adult males;

its length in front of orbits 3,0; 3,1 (2,9-3,8) times as long as distance between

orbits; its length in front of mouth 1,5; 1,6 (1,5-1,9) times as great as distance

between nostrils. Orbit 1,0; 1,0 (0,8-1,0) as long as distance between orbits

and 1,1; 1,1 (0,8-1,4) times as long as spiracle. Spiracle extending to below

middle of eye.

Neurocranium typically guitar-shaped and markedly constricted across

orbital region, with well-developed postorbital processes and poorly developed

jugal arches; nasal capsules directed obliquely forward to about 40° to median

axis, without ethmoidal nerve foramen; rostral cartilage projecting from

cranium to tip of snout as a thin, delicate rod, without a segment; rostral

appendices narrowly attached to extremity of rostral bar and extending pos-

A NEW SPECIES OF SOUTHERN AFRICAN BREVIRAJID SKATE 259

2,0cm

1,0 cm

Fic. 4.

Breviraja stehmanni n. sp. A. External view of right clasper from the dorsal side. B. Lateral view

of right clasper, opened to show structural features of the glans. i

ap—apopyle; cf—cleft; dd—dermal denticles; hp—hypopyle; rh—rhipidion; sh—shield;

sp—spike; st—sentinel.

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

1,0 cm

Fic. 5.

Breviraja stehmanni n. sp. Clasper cartilages. A. Dorsal terminal 1 (dorsal view). B. Axial,

dorsal marginal, dorsal terminal 2 and 3, and terminal bridge (dorsal view). C. Ventral ter-

minal (dorsal view). D. D1. Accessory terminal 1 (dorsal and lateral views). E. E*. Accessory

terminal 2 (dorsal and lateral views).

Ax—axial; dM—dorsal marginal; dT,, dT,—dorsal terminals 2 and 3; tb—terminal bridge.

A NEW SPECIES OF SOUTHERN AFRICAN BREVIRAJID SKATE 261

teriorly to level of anterior fontanelle, well separated from rostral bar; anterior

notch bridged with a thin cartilaginous connection; small additional appendix

on each side, with a foramen; radial cartilages of pectorals extending anteriorly

almost to appendices; anterior fontanelle well developed, with anteriorly

arched epiphysial bridge; 2 (sometimes 1) posterior fontanelles; orbito-nasal

canal foramen comparatively small, anterior cerebral vein foramen above

level of optic foramen.

Mouth weakly arched medially; nasal curtain fringed. Teeth arranged in

Al, 39 (38-44) rows in upper jaw, blunt and flat and in quincunx in females

and juveniles, but adult males with laterally directed, long, conical points in

centre of jaw. |

Anterior lobes of pelvics fin-like and continuously connected with posterior

lobes along outer margin of fin; anterior margin of pelvic about 70%, 78%

(68-84%) as long as distance from its own origin to rear tip of pelvic.

Dorsal fins similar in shape and about similar in size, confluent at base;

caudal membrane posterior to second dorsal about half as long as base of second

dorsal.

Vertebral count: Vtr 26; 26 (24-26); Vprd 69; 71 (65-74); V2 95; 97

(91-98). |

Claspers pointed; pseudosiphon absent; dermal denticles present on dorsal

border and ventral surface; inner dorsal lobe with two clefts separated by

terminal bridge; rhipidion fan-shaped and situated at hypopyle; shield well-

developed; laterally projecting sentinel and recurved spike situated medially.

Axial cartilage pointed terminally; dorsal marginal with short distal extension,

ventral marginal distally arched; dorsal terminal 1 cartilage with proximal

extension; dorsal terminal 2 and 3 cartilages simple and forming dorsal lobe

framework; ventral terminal cartilage with dorsally convex, outer, lateral

margin and with anterior notch at about one-third the length of the cartilage

from the proximal end; accessory terminal 1 cartilage U-shaped proximally

and with well developed Z-shaped lateral projection; accessory terminal 2

simple, with hooked, spatula-like distal extremity, closely attached to axial

cartilage along its inner lateral margin.

Colour

Upper surface of disc greyish, skin at bases of prickles and at rear sides

of larger thorns not especially pigmented; disc with irregular darker blotches

and scattered paler spots; tail with 6—7 irregular, dark crossbars, either con-

tinuous across dorsal surface or interrupted, the two most posterior crossbars

nearly black and passing through first and second dorsal fins. Lower surface

of disc pale but tip of snout with black spot; thin darker areas along anterior

margin of pectoral, wider along posterior margins and margins of pelvics;

dusky, irregular areas sometimes between nostrils, around mouth, between gills

and on belly; tail with dark crossbars encroaching from sides of tail, but mature

specimens with ventral surface of tail somewhat darker mottled in a few cases.

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

S7ze

Males mature at about a length of 340 mm, as ascertained from the degree

of calcification of the clasper cartilages and the presence of well developed

malar and alar spines. Females probably reach a slightly greater size at sexual

maturity.

Distribution

South of Agulhas Bank to west of Cape Town in 292~1 025 metres.

DISCUSSION

As has been pointed out, the genus Breviraja was previously thought to be

confined to the edges of the continental shelf and upper regions of the slope in

the western central Atlantic (Bigelow & Schroeder 1953; Stehmann 1970;

Hulley 1972) in depths of 200-727 fms (366-1 329 m). Breviraja stehmanni,

which is so far known only from the eastern South Atlantic, west of Cape Town,

appears to have a similar depth distribution, 160-560 fms (292-1 025 m).

Bottom temperatures at this depth and in this region may vary between 3°C

and 8°C (Hulley 1972). The species may be more widely distributed in the

southern African region and may not be confined only to the Atlantic, since

bottom temperatures are more or less uniform at depths greater than 100 m,

both east and west of Cape Point. Hulley (1972) has pointed out that the

majority of species are widely distributed throughout the entire southern

African region.

An association of Breviraja with the Dipturus-line of evolution (Hulley 1972)

appears to be substantiated by the anatomy of the claspers and neurocranium

of Breviraja stehmanni. The dorsal position of the foramen of the anterior cerebral

vein and the general arrangement of the clasper, especially the well developed

rhipidion, lack of a proximal shelf on the dT, and well developed vT, suggest

an association with the Dipturus/Amblyraja/Leucoraja/Raella-line rather than with

the Bathyraja/Raja/Rostroraja-line (Hulley 1972: fig. 56).

Breviraja might be considered to be ancestral to both the Dzpturus- and

the Amblyraja/Leucoraja/Raella-lines of evolution, since the predorsal caudal

vertebral count is high and the clasper exhibits primitive characteristics in the

retention of dermal denticles, a pointed distal tip to the Ax cartilage and the

well developed Z-shaped lateral projection of aT, (Fig. 5; Ishiyama & Hubbs

1968: fig. 2). However, the lack of an external pseudosiphon, arrangement of the

dT, and dT, cartilages and the more proximal position of the anterior notch on

the vT, point to a closer association with the Dipturus-line. The development of

a thin rostral bar, without anterior grooving, and of the characteristic rostral

appendages, indicate a specialized condition for the genus. It may therefore be

concluded that the genus Breviraja represents a very early split from the Dipturus-

line, which penetrated somewhat deeper regions and retained the neotonous

condition of the snout as an increased advantage in grubbing.

A NEW SPECIES OF SOUTHERN AFRICAN BREVIRAJID SKATE 263

SUMMARY

A new species of southern African skate, Breviraja stehmanni, is described

and represents the first record of the genus outside the waters of the western

central Atlantic. The phylogenetic position of Breviraja is briefly discussed.

ACKNOWLEDGEMENTS

I am deeply indebted to the Director and Staff of the Division of Sea

Fisheries, Cape ‘Town, for the collection and donation of the rajid material,

and especially to Mr L. Botha and Mr D. Chalmers of that Institute. I should

also like to thank Mr S. X. Kannemeyer, of the South African Museum, for his

assistance with the X-ray photography.

REFERENCES

BARNARD, K. H. 1925. A monograph of the marine fishes of South Africa. Ann. S. Afr. Mus.

21: 1-418.

BicELow, H. B. & ScHROEDER, W. C. 1948. New genera and species of batoid fishes. 7. mar. Res.

7: 543-566.

BicELow, H. B. & ScHROEDER, W. C. 1953. Fishes of the western North Atlantic. Part 2. Saw-

fishes, guitarfishes, skates and rays. Chimaeroids. Mem. Sears Fdn mar. Res. (2): i-xv, 1-588.

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

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

Huttey, P. A. 1972. The origin, interrelationship and distribution of southern African Rajidae

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

IsHtyAMA, R. & Huss, C. L. 1968. Bathyraja, a new genus of Pacific skates (Rajidae) regarded

as phyletically distinct from the Atlantic genus Breviraja. Copeia 1968: 407-410.

Norman, J. R. 1935. Coast fishes. Part I. The South Atlantic. ‘Discovery’ Rep. 12: 1-58.

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

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

Arch. Fisch Wiss. 21: 73-164.

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Style manual for biological journals. Washington: American Institute of Biological Sciences.

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REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

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

FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

FiscHER, P.-H., Duvat, M. & Rarry, A. 1933. Etudes surles échanges respiratoires des littorines.

Archs Kool. exp. gén. 74: 627-634.

Konn, A. J. 19604. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region

of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn scHULTZE, L.

Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature issued

by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).

The Harvard system of reference to be used in the synonymy lists, with the full references

incorporated in the list at the end of the article, and not given in contracted form in the synonymy

list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, 6; Liste: 11. Turton, 1932: 80.

P. A. Hulley

A NEW SPECIES OF SOUTHERN AFRICAN

BREVIRAJID SKATE

(CHONDRICHTHYES, BATOIDEI, RAJIDAE)

OF THE SOUTH AFRICAN

MUSEUM

| CAPE TOWN

aNINALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 60. Band

April 1973 April

Part 10 ©6Deel

Gee AMPHIPODA OF SOUTHERN, AFRIGA

yeas

THE GAMMARIDEA AND CAPRELLIDEA OF

SOUTHERN MOCGAMBIQUE

C. L. GRIFFITHS

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

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ISBN 0 949940 26 7

Printed in South Africa by In Suid-Afrika gedruk deur

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE AMPHIPODA OF SOUTHERN AFRICA

Pak

THE GAMMARIDEA AND CAPRELLIDEA OF SOUTHERN

MOCAMBIQUE

C. L. GriFFITHs

C.S.LLR. Oceanographic Research Umit, Zoology Department, University of Cape Town

(With 11 figures)

[MS. accepted 15 August 1972]

CONTENTS

PAGE

Introduction. : : : . 265

The collecting stations : : : «9 (266

Systematics ; : : ‘ : 58 7B

Gammaridea . : ; : : Seip!

Caprellidea : : 4 : : ae Zoo

Summary. : : : i “304

Acknowledgements. : ; ‘ 5 GO:

References ; : 3 i : a 2Or

INTRODUCTION

The first comprehensive works on the Crustacea of southern Africa were

those of T. R. R. Stebbing, culminating in his ‘General catalogue of South

African Crustacea’ in 1910, which was largely based on the extensive collections

of the S.S. Pieter Faure. K. H. Barnard continued to analyse the Pieter Faure

material and in 1916 published descriptions of many new species and records,

to be augmented in 1925 by a report on the last portion of the collection,

bringing the total number of recorded species to 207. Barnard went on to

publish more descriptions and a key to known species in 1940, and further

additions to the fauna were forthcoming in 1951, 1955 and 1957.

Despite this excellent background, South African Amphipoda have subse-

quently been almost completely neglected, although collecting has continued

unabated and many new records and undescribed species have undoubtedly

accumulated, especially in the extensive collection of the University of Cape

Town Ecological Survey.

It is in the light of these new data and of recent extensive taxonomic

revisions that the decision has been made to synthesize existing knowledge of

the South African amphipod fauna.

Mr Roy Dick has recently (1970) produced a key to, and distribution list

of the Hyperiidea of southern Africa and it is the aim of the present work, and

subsequent ones of a series, to cover the Gammaridea and Caprellidea of the

same area (Africa south of 20°S) on a regional basis.

265

Ann. S. Afr. Mus. 60 (10), 1973: 265-306, 11 figs.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

Mocambique has been selected as the subject of the first paper, since no

previous literature specifically on the amphipods of this area appears to exist,

although a checklist and key to the species of Inhaca Island (Macnae & Kalk

1958), descriptions of new species from Morrumbene estuary (Barnard 1916,

1955) and a key to common species of southern Africa (Day 1969) have

appeared. Moreover, the recent International Indian Ocean Expedition,

resulting in Ledoyer’s work (19674, 6) on the Amphipoda of Malagasy has

prompted the author’s interest in the fauna of the mainland of Mocambique.

Data for this paper were provided by the following:

(a) An extensive survey of Morrumbene estuary by the Zoology Depart-

ment of the University of Cape Town, involving visits in 1953, 1954, 1955 and

1968. During these expeditions to Mocgambique, exploratory collections were

also taken at Lagoa Poelela, Maxixe and Jangamo reef and the data from these

collections are also included.

(b) Collections from Inhaca Island and Ponta Zavora made by a team of

biologists from the South African Museum, Cape Town, in June 1971, and

kindly loaned by them to the author.

International Indian Ocean Expedition cruise in 1964.

(d) Information has also been drawn from collections taken at Inhaca

Island by. the University of the Witwatersrand (Macnae & Kalk 1958, 19624, 6).

THE COLLECTING STATIONS

Biorat

Morrumbene estuary

Pig. 2

Description

This rich estuary, fed by the Morrumbene, Inhanombe and other minor

rivers, opens into the northern end of Inhambane Bay (at 23°40'S/35°20’E).

The Inhanombe is the largest of the rivers, flowing northwards through man-

grove swamps before reaching a broad mixing basin between the San José

Mongué mission station and Morrumbene village. The shallow mixing basin

is about 2~3 kilometres wide at flood tide, its banks are mainly muddy in the

upper reaches, except for a few sandy areas around Tinga-Tinga; but from the

mission station to the mouth the banks are sand and the bottom sandy mud.

The tortuous Inhanombe channel is about two metres deep opposite the mission

station and reaches a maximum depth of 20 metres at Linga-Linga.

Tidal range at springs is about three metres at Mongué decreasing to two

metres at Tinga-Tinga. Salinity (at high and low tides respectively) varied

between 35%, and 31,2%, at Linga-Linga, 34,4%, and 22,9%, at Mongué, and

between 33,8%, and 11,8%, at Tinga-Tinga.

The banks of the estuary are lined with at least three genera of mangrove

THE AMPHIPODA OF SOUTHERN AFRICA 267

Mocambique

MORRUMBENE

MAXIXE

PONTA ZAVORA

ro) 3-70 o1-2

9-12 08

013-17

LOURENCO 520-21

MARQUES 022-23

Qinuaca ISLAND

Fig. 1. Collecting stations in southern Mocambique (20° to 27° S) with PED stations numbered.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

Morrumbene

Mongue «

a \:

Modified from chart 493

BARRAe PORTO de INHAMBANE

} =

MORRUMBENE ESTUARY

Fig. 2. Morrumbene estuary (after Day, unpublished).

including Avicennia, Rhizophora and Ceriops, these give way to Phragmites,

Barringtonia and Acrostichum where salinity falls below 10%,. In the mixing basin

sea grasses such as Halodule and Cymodocea are common near low water springs.

The survey of Morrumbene formed part of the University of Cape Town’s

estuarine survey series and a paper on the ecology of the estuary is in prepara-

tion by J. H. Day. Twenty-two species of amphipod were recovered during the

survey. Both the absolute number of individuals and the number of species

decreased in areas of low salinity.

Eighteen species were present in the mixing basin, of these only six were at

all common (Cheiriphotis megacheles, Grandidierella bonniert, Hyale inyacka, Laetmato-

philus purus, Lembos teleporus and Talorchestia australis). Above 'Tinga-Tinga seven

species were recorded, none of them common.

One particularly euryhaline species worthy of mention is Grandidierella

bonniert which was found from Linga-Linga to Tinga-Tinga and also recovered

from a fresh-water lake adjoining Lagoa Poelela.

THE AMPHIPODA OF SOUTHERN AFRICA

Station list

269

Catalogue number Date Position Substrate

MOR 30 19/1/54 Midchannel, Tinga-Tinga LWS Sandy mud

MOR 31 19/1/54 In dense mangroves, Tinga-Tinga Mud

MOR 37 20/1/54 Mid-tidal level, Linga-Linga Halodule marsh

MOR 40 20/1/54 Midtide rocks, Linga-Linga Rock

MOR 41 20/1/54 LWS, Linga-Linga Halodule marsh

MOR 45 20/1/54 Dredge 6-9 m, Linga-Linga ~

MOR 52 21/1/54 Netting in mouth of Rio Coche —

MOR 74 24/1/54 Netting at Mongué ferry Halodule beds

MOR 75 18/1/54 Midtide at Mongué ferry Sand

MOR 77 23/7/53 Diving 2-3 m, Linga-Linga Sand and weed

MOR 85 11/7/54 Above HWS at fern bank Mangrove roots

MOR 95 3/7/54 Handnetting (location ?) Aquatic vegetation

MOR 102 14/7/54 Netting at Linga-Linga Halodule and Cymodocea

MOR 108 14/7/54 Dredge 1,5-4 m, Linga-Linga Sand with Cymodocea

MOR 122 15/7/54 Dredge off Linga-Linga —

MOR 124 15/7/54 Plankton netting, Linga-Linga —

MOR 147 16/7/54 Netting, mouth of Rio Coche —

MOR 179 18/7/54 Mangroves at Mongué ferry —

MOR 180 18/7/54 Mongué ferry, HWS Sand

MOR 193 19/7/54 Linga-Linga Sand

MOR 212 15/7/54 LWS, Mongué ferry Mangroves?

MOR 218 13/7/54 From wreck, Linga-Linga Ship’s hull

MOR 232 12/7/54 Rio Coche —

MOR 238 12/7/54 Opposite Rio Coche Sand

MOR 240 12/7/54 Opposite Mongué ferry Mud

MOR 243 12/7/54 Mongué ferry =

MOR 244 12/7/54 Mongué ferry =

MOR 250 12/7/54 Tinga-Tinga Mangroves

MOR 253 12/7/54 Fern bank Acrostichum

MOR 255 12/7/54 Head of estuary of Inhanombe River a

Subsidiary collections made during the expeditions to Morrumbene

(a) Jangamo Reef lies 96 kilometres south of Inhambane Bay at

24°06'S/35°30’E. A party of biologists from the University of Cape Town, led

by Professor J. H. Day, visited the area from 7 to 10 July 1968. The reef was

found to be of flat sandstone transected by gulleys and covered by shallow pools,

the rock extending from high water neaps to low water springs. Ulva was domi-

nant at higher levels, below which there was an area of red algal turf running to

low water springs. At low tide Idanthyrsus formed large encrustations and

zoanthids, algae, sponges, ascidians and hydrozoans fringed the gulleys. Corals

were present but not abundant.

The dominant intertidal amphipods were Elasmopus affinis and Elasmopus

japonicus which were present in approximately equal numbers and together

accounted for over 75% of the collection. Of the 14 other species recorded,

Caprella equilibra and Cheiriphotis megacheles were the most common.

Station list

Position

Date

8/7/68

Catalogue number

JAN 12, JAN 14

General collection of amphipods

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

(b) Lagoa Poelela (24°33'S/35°05’E), one of a series of brack-water lagoons

forming a chain along the coast, was briefly visited by a team of University of

Cape Town zoologists in January 1954. The lagoon was 30 kilometres long and

5 kilometres wide, the shore clean sand and the water crystal clear without any

signs of aquatic vegetation and little sign of aquatic life. At the time of sampling

salinity near the shore was 6,5%, and the temperature at the surface was 19,5°C.

Only three amphipods were found in the lagoon, Orchestia ancheidos was

common in weed along the drift-line, Grandidierella bonniert in sand along and

above the watermark and Melita zeylanica in the weedy shallows. With the

exception of O. ancheidos, the same species were recovered from an adjoining

freshwater lake.

Station list

Catalogue number Date Position

BORa 26/1/54 Netting along the shore

ROE? 26/1/54 Digging between Juncus plants 30 cm above water

: level

POE 6 26/1/54 Digging in waterlogged sand on shore

POE 8 26/1/54 In weed along driftline

POE 10 26/1/54 Handnetting in weed on the margin of a shallow

freshwater lake south-west of the lagoon

POE 12 26/1/54 Collecting along margin of freshwater lake

Morrumbene estuary on the western shore of Inhambane Bay, was visited

briefly in July 1953.

A rapid examination of the shore revealed only two species of amphipod,

Lembos podoceroides and Chevalia aviculae, both at low tidal levels on the sand banks

and piles of the wharf.

Station list

Catalogue number Date Position

PEA 2 24/7/53 General collection from low tide sand banks and

piles, Maxixe

PEA 4 24/7/53 General collection from low tide sand banks and

piles, Maxixe

Inhaca Island

Bigs

Detailed ecological descriptions of Inhaca Island may be found in Kalk

(1958), Macnae & Kalk (1958) and Kalk & Macnae (1962a, 6). The island lies

at the tip of the Inhaca peninsula, forming an eastern boundary to Delagoa Bay.

It is centred at 26°o01'S/32°56’E with a maximum length of 11 kilometres and a

width of 6 kilometres. The mangrove fringed northern bay and the Saco da

Inhaca in the south contain extensive mud flats dominated by Halodule and

Cymodocea through which narrow channels run. The Saco is more rocky than

the northern bay and there is a young coral reef near Ponto Torres.

THE AMPHIPODA OF SOUTHERN AFRICA 271

| PORTINHO 25

aa INHACA 7

: aN |

\ /

BIOLOGICAL : HE eN :

STATION |g ae \ N

/ \

/ le /

\ hE aS ey, @ Na \

Noe i SNES \

AEN tA ve \ EA

: : yh ~ \\# gh a

AEN Vy rare *: TE

NN ey << “

a Ta rio \ !

~PONTA yo tee e \

PUNDUINI VA

i ae pe NX / a | Eee

y : i SANTA MARIA

Fig. 3. Inhaca Island, showing features referred to in the text (after Macnae & Kalk, 1958).

272 ANNALS OF THE SOUTH AFRICAN MUSEUM

The west coast, which borders shallow Delagoa Bay, exhibits intertidal

sand flats up to 1 kilometre wide. ‘These are often covered by coral debris or

Cymodocea and opposite Barreira Vermelha there is a coral reef.

The eastern shore is exposed to the surf and is rocky along its whole length,

though the rocks are usually only at intertidal levels. At Cabo da Inhaca there

are three rock terraces, the highest not now immersed at any state of tide.

The island provides an extremely wide range of habitats, rock and sand

being present in both exposed and sheltered areas at all tidal levels. Moreover,

the landward side of the island, bathed by the warm waters of shallow Delagoa

Bay, apparently represents an isolated area of true tropical fauna, while the

open coast is considered subtropical, with coral growing on the rocks, rather

than forming true reefs.

A collection of amphipods was made on the island by a team from the

South African Museum in June 1971 and these, together with the species

recorded by Macnae & Kalk, reveal a total of 22 species. Of the 16 species

recovered during the South African Museum’s brief collecting visit, 11 were

new to Inhaca, an indication of the number of species still to be discovered.

The fauna is best grouped according to area of occurrence. In the northern

bay, only one species is found, Orchestia anomala, which is common along the

drift line. The eastern seaboard is also sparsely populated, only Hyale grandi-

cornis, Stenothoe valida and a species of Podocerus having been found.

The southern bay is the richest area with 16 recorded species. Important

among these are Maera hamigera and Maera inaequipes, which are common in

coral. Cymadusa filosa, Ampithoe ramondi, Elasmopus affinis and Talorchestia australis

are common intertidally.

On the more muddy, sheltered west coast, 13 species have been found.

Cymodocea is a habitat favoured by Ericthonius brasiliensis, Cymodusa filosa, Caprella

scaura and Aora typica, while Orthoprotella mayeri is common on the hydroid

Lytocarpus. Maera hamigera and Maera inaequipes once again occur in coral. At

high intertidal levels Orchestia anomala occurs among Bostrychia, Hyale inyacka

under rocks on sand and Talorchestia australis under weed on the drift line.

Station list

Catalogue number Date Position

IN 158 June 1971 In a sponge, Saco da Inhaca

IN 159 June 1971 General collection, Santa Maria

IN 160 June 1971 General collection on mud, west coast

IN 161 June 1971 On Cymodocea, west coast

During their visit to Inhaca, biologists from the South African Museum

also briefly visited Ponta Zavora (30°02'S/35°10'E) where they found five

species of amphipod in the intertidal zone. Elasmopus japonicus and Cymodusa

THE AMPHIPODA OF SOUTHERN AFRICA 273

filosa were fairly common, while single specimens of Gammaropsis semichelatus,

Amaryllis macrophthalma and Maera sp. (female) were collected.

Dredge stations

Samples from 23 dredges forming part of a series taken by the S.S. Anton

Bruun in August 1964 were donated to the University of Cape Town and are

incorporated in the survey.

Station list

Cat. no. Date Position Depth Substrate Gear

(m)

PED 1-2 18/8/64 24°46'S/35°20’E 132 Rock and shelly sand Rock dredge

PED 3-7 18/8/64 24°46'S/35°18’E 110. Coarse sand and rock Rock dredge

PED 8 18/8/64 24°49/S/35°13’/E 73 Shelly sand and rock Rock dredge

PED 9-12 19/8/64 24°46'S/34°50’E 22 Shelly sand and rock Rock dredge

PED 13-17 19/8/64 24°53’S/34°56’E 55 Fine grey sand and rock Rock dredge

PED 18-19 _ 19/8/64 25°07’S/34°34’E 112 Dark sandy mud Agassiz dredge

PED 20-21 22/8/64 = 25°5,7’S/33°02’E 42 Shell and rock Rock dredge

PED 22-23 22/8/64 26°00'S/33°05’/E 135 = Shell and rock Rock dredge

These samples contained 27 species of amphipod, apparently separable

into two groups according to substrate preferences.

Areas with fine substrates (PED 13-18) revealed 16 species while the coarser

bottomed areas revealed 11 species, only five being common to both substrate

types.

On shell, sand and rock, the most important species were Gammaropsis

atlantica, Maera inaequipes, Maera serrata and Orthoprotella mayert.

In the fine sand and mud areas no one species was recovered in even

moderate numbers, indicating a sparse, mixed population of mud-loving

burrowing types (such as Ampelisca, Podocerus, Metaphoxus and Urothoe). The few

caprellids found here would have been living on the occasional rocks present.

SYSTEMATICS

The form of presentation used here follows that of J. L. Barnard and

J. C. McCain, in that families, and genera within families, are presented in

alphabetical order.

It will be noted that many major taxonomic changes have taken place in

the years since K. H. Barnard’s (1940) key to South African Amphipoda. The

reader is referred to J. L. Barnard (19690, 1970a) for diagnosis of gammaridian

taxa and to J. L. Barnard (1958) for a world species list. Information on the

revised taxonomy of Caprellidea may be found in McCain (1970) and McCain

& Steinberg (1970), the latter work also including bibliographies and syno-

nymies for all known species.

For the convenience of readers unfamiliar with recent developments,

relevant changes at family level since 1940 are listed below:

O74: ANNALS OF THE SOUTH AFRICAN MUSEUM

Old taxa New taxa

Atylidae Incorporated into Dexaminidae

Lepechinellidae

Hyalellidae Incorporated into Superfamily

Hyalidae Talitroidea

Talidae

Photidae Incorporated into Isaeidae

Metopidae Incorporated into Stenothoidae

Pontigeneiidae Incorporated into Eusiridae

Jassidae Becomes Ischyroceridae

Caprellidae Split to form

Phtisicidae

Caprellidae (revised)

Aeginellidae

Caprogammaridae

In the following account, no attempt has been made to provide a full list

of references or synonyms, but the reader is referred to works which will provide

good descriptions and synonymy lists, and to those which refer specifically to

the southern African region.

The holotypes of all new species have been placed in the South African

Museum, Cape ‘Town, the 8.A.M. Catalogue numbers given are museum

numbers, while other codes (MOR, PED, etc) refer to University of Cape

Town collections, the number in brackets after the code refers to the number of

individuals found.

Suborder GAMMARIDEA

Family Ampeliscidae

Ampelisca diadema (Costa, 1853)

Ampelisca assimilis: Sars, 1895: 168, pl. 58.

Ampelisca diadema: K. H. Barnard, 1916: 133.

Records: PED 15J (2).

Diagnosis: Article 3 of pereiopod 5 longer than article 4; pereon segments 5-7

with ventral hooks; third pleonal epimeron rounded; pleon segment 4 with a

high, evenly-rounded dorsal carina; antenna 1 slightly exceeding peduncle of

antenna 2.

Distribution: Cosmopolitan.

THE AMPHIPODA OF SOUTHERN AFRICA 2

~!I

Ampelisca fusca Stebbing, 1888

Ampelisca fusca Stebbing, 1888: 1052, pl. 105.

Records: PED 6K (1); PED 23Q (6).

Diagnosis: Article 4 of pereiopod 5 twice the length of article 3, produced

posteriorly to cover half article 5, produced portion setose, article 5 slightly

produced anteriorly ; four eyes, with corneal lenses, lower pair directed forward;

pleon segment 4 saddle-shaped dorsally; antennae 1 and 2 equal and as long as

body.

Distribution: Endemic, Cape Agulhas to Mocgambique.

Ampelisca palmata Barnard, 1916

Ampelisca palmata K. H. Barnard, 1916: 136, pl. 28, figs 30-31.

Records: PED 15H (2).

Diagnosis: Article 3 of pereiopod 5 longer than article 4, 4 with front apex

narrowly produced along half the length of 5; third pleonal epimeron quadrate,

scarcely produced; pleon segment 3 with a low subacute dorsal keel; antenna 1

® much longer than peduncle of antenna 2.

Distribution: Southern and west Africa.

Family Amphilochidae

Gitanopsis mariae n. sp.

Fig. 4

Diagnosis of male: Head with extremely large probosciform rostrum, lateral

cephalic lobe quadrate, eye of moderate size, black. Antenna I not setose, as

long as head, flagellum seven-articulate, accessory flagellum absent. Antenna 2

slightly longer than 1, flagellum seven-articulate. Mandible with large tritura-

tive molar, palp article 1 distally expanded, articles 2 and 3 subequal, article 3

distally curved. Maxilla 1, palp biarticulate. Outer plate of maxilla 2 narrower

than inner. Outer lobe of maxilliped slightly excavate, palp four-articulate,

articles 1 and 2 subequal and article 2 expanded distally.

Gnathopods 1 and 2 very weak and slender with gnathopod 2 slightly the

larger. Coxa 1 rounded and very small, partially concealed by coxa 2. Coxa 3

antero-distally produced, coxa 4 deeply excavate posteriorly.

Pereiopods 3-5, article 2 almost as wide as long. Third pleonal epimeron

rounded and somewhat produced postero-inferiorly.

Uropods 1-3, outer rami marginally shorter than inner. Telson dorsally

excavate, 70% length of peduncle of uropod 3, tapering distally, apically

tridentate.

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Gitanopsis mariae n. sp., holotype, male, 7 mm.

A. Lateral view. B. Mandible. C. Maxilla 1. D. Maxilla 2. E. Maxilliped. F. Gnathopod 1.

G. Gnathopod 2. H. Telson.

THE AMPHIPODA OF SOUTHERN AFRICA

ie)

“I

Holotype: Male, 7 mm, unique. $.A.M. A 13066.

Type-locality: Collected at Santa Maria, Inhaca Island, by biologists of the

South African Museum, June 1971.

Records: IN 159 J.

Remarks: ‘This species is easily recognized by its distinctive rostrum, which is

much larger than is usual in this genus, and by the extremely weak gnathopods,

which are unlike those of any other representatives of the genus.

Gitanopsis pusilla Barnard, 1916

Gitanopsis pusilla K. H. Barnard, 1916: 144. Stephensen, 1949: 8, fig. 1.

Records: PED 21E (1); IN 158C (1).

Diagnosis: Distinguished from other members of the family by the very short

pyriform telson which is less than half the length of the peduncle of uropod 3;

body smooth; process of article 5 of gnathopod 1 extending half the length of

article 6, palm transverse, convex, defined by two spines.

Distribution: ‘Tristan da Cunha, Kerguelen, South Georgia, Lambert’s Bay to

Mocambique.

Family Ampithoidae

Ampithoe ramond: (Audouin, 1826)

Ampithoe vaillanti: K. H. Barnard, 1916: 253.

Ampithoe ramond:: J. L. Barnard, 1970): 50, figs 18-19.

Records: IN 159C (8); JAN 12J (1); MOR 74J (1); MOR 212C (2).

Diagnosis: Accessory flagellum absent; gnathopods 1 and 2 with article 2

lobed; palm of gnothopod 2 defined by a conspicuous lobelike tooth, dactyl

serrate; outer ramus of uropod 3 terminating in two strongly recurved spines.

Distribution: Cosmopolitan in warm and temperate seas.

Cymadusa filosa Savigny, 1818

New synonymy:

Grubia australis K. H. Barnard, 1916: 258.

Grubia filosa: Shoemaker, 1935: 245, figs 4, 5-

Cymadusa australis: K. H. Barnard, 1940: 480.

Cymadusa filosa: J. L. Barnard, 1955: 29, fig. 15.

Records: MOR 218D (1); IN 160B (3); JAN 12C (2); PEA 24C (3).

Remarks: C. australis has previously been separated from C. filosa by the lack of

plumose setae on antenna 2 and coxae 1~4 in the adult male, and by the lack of

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

distal lobes on article 2 of gnathopods 1 and 2. Barnard’s types of C. australis,

however, included only one adult male, and subsequent samples (including

some from the type locality) identified by Barnard as C. australis did have

plumose setae. Re-examination of the type specimens has shown that small

pellucid lobes on the gnathopods are present, and that the females do not differ

in any respect from females of C. filosa.

Since no more material corresponding to the male type of C. australis has

been found, it seems more than likely that the one male type is aberrant, having

failed to develop the plumose setae of an adult male. (It is well known that

plumose setae develop with maturity and are fully developed only in aged

males.)

The largest specimen so far recovered, a male of over 30 mm, showed

plumose setae on the telson, fringing the peduncles of the uropods, on the

anterior edge of article 2 of the pereiopods and the gnathopods, on the peduncle

of antenna 2 and on the ventral margin of the head.

Distribution : Circumtropical.

Paragrubia vorax Chevreux, 1901

Fig. 5

Paragrubia vorax: J. L. Barnard, 1965: 541, fig. 35. Ledoyer ,1967: 135, fig. 23.

Records: JAN 12B (3) (the first record of this species from the southern African

mainland).

Diagnosis: Gnathopod 1 considerably larger than 2, article 6 broadly expanded

in adult males, palm slightly oblique and strongly concave. Young males show

a strong palmar spine but this is lost with age. Accessory flagellum present.

Distribution: ‘Tropical Indo-Pacific.

Family Aoridae

Aora typica Kroyer, 1845

Aora typica: Ledoyer, 1967: 131, fig. 15.

Records IN| 160 Ay):

Diagnosis: Easily recognized by gnathopod 1 which has a proximal tooth on the

anterior margin of article 2 and a long distal projection on article 4.

Distribution: Gosmopolitan.

Lembos podoceroides Walker, 1904

Lembos podoceroides Walker, 1904: 279, pl. 6.

Records: PED 6 QO (2); PEA 2 L (6).

THE AMPHIPODA OF SOUTHERN AFRICA

oe |

“ 7 \ }

ty gues

etes22 DUS Saas

Fig. 5. Paragrubia vorax Chevreux, male, 10 mm.

A. Antenna 1. B. Antenna 2. C. Upper lip. D. Mandible. E. Lower lip. F. Maxilla 1. G. Maxilla 2.

H. Maxilliped. I. Gnathopod 1. J. Gnathopod 2. K. Uropod 3.

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis: Gnathopod 2 of male with the base of the hand produced backwards

in a long pointed spur and the palm with a small semicircular sinus near the

finger hinge; the posterior and lower margins of the third pleonal epimeron are

convex and the postero-inferior corner is slightly produced with a diagonal

ridge running across the epimeron to it.

Distribution: Indian Ocean, Red Sea.

Lembos teleporus Barnard, 1955

Lembos teleporus K. H. Barnard, 1955: 94, fig. 47. Ledoyer, 1967: 135, figs 16-17.

Records: IN 160 C (2); MOR 77 W (7); MOR 102 P (2); MORSI7 ia

MOR 212 H (21).

Diagnosis: Pereon segment 3 with a strong ventral spine in adult males; article 6

of gnathopod 1 male three times the length of article 5, widening distally, palm

with a blunt tooth near finger hinge, palmar angle quadrate, palm and dactyl

crenulate.

Distribution: Malagasy, Mogambique, South West Africa.

JANICE, n. gen.

Generic diagnosis (male): Article 3 of antenna 1 shorter than article 1, accessory

flagellum absent; gnathopod 1 subchelate, articles 4—6 lacking teeth, article 6

slightly shorter than, and of subequal width to, article 5; gnathopod 2 heavily

setose, article 4 projecting to protect article 5 posteriorly, article 5 wider than

6 but subequal in length; uropod 3 uniramous, ramus equal to peduncle.

Remarks: ‘The lack of teeth on gnathopod 1 and the complete lack of accessory

flagellum are alone sufficient to demand the erection of a new genus. In addi-

tion, the structure of gnathopod 2 is unique among Aoridae.

Type-species: Janice spinidactyla n. sp.

Janice spinidactyla n. sp.

Fig. 6

Diagnosis of male: Head as long as two pereon segments; eyes small, round, dark.

Antenna 1 extending to end of pereon segment 5, ratio of peduncular articles

2:3: 1, flagellum subequal to peduncle and composed of 18 articles, acessory

flagellum absent. (Antenna 2 missing.) Mandible with large triturative molar

and three-articulate palp, lacinia mobilis 2-toothed with seven spines proxi-

mally.

Gnathopod 1 larger than 2, subchelate, articles 4-6 lacking teeth, setose

posteriorly, article 6 as wide as, but slightly shorter than, article 5, palm trans-

verse, convex, defined by two large spines and with a few small spines along its

edge, dactylus slightly longer than palm and cut into four teeth.

THE AMPHIPODA OF SOUTHERN AFRICA

281

male 6 mm.

n. sp., holotype,

Fig. 6. Zanice spindiactyla n. gen.,

Tip of article 6 and article 7 of

D. Gnathopod 2 without its setae.

Maxilliped.

B. Mandible.

A. Lateral view.

‘gnathopod 2.

Telson.

H. Uropod 3. I.

G. Uropod 2.

Uropod 1,

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

Article 4 of gnathopod 2 with a spoon-shaped projection extending nearly

half the length of article 5, with long, plumose setae along its margins; article 5

as long as, and wider than, article 6 and extremely setose posteriorly, article 6

narrow, palm transverse; four large and many small spines along its length;

dactylus matching palm with ten spines which become longer distally.

Pereiopods 1 and 2 glandular; pereiopods 3-5 with article 2 slightly

expanded, that of pereiopod 5 extremely setose posteriorly.

Uropod 1 projecting to the end of uropod 3, rami subequal and # length of

peduncle, each bearing three spines dorsally and five or six terminally. Uropod 2

half the length of 1, rami subequal and slightly shorter than peduncle, each with

one dorsal and five terminal spines. Uropod 3 half the length of 2, uniramous,

ramus slightly longer than peduncle, cylindrical, terminating in about 12 long

spines.

Telson not extending to end of peduncle of uropod 3, emarginate, fleshy,

two lateral and three terminal setae on each side.

Holotype: Male, 6 mm. $.A.M. A 13067.

Type-locality: Found amongst Acrostichum, Morrumbene estuary above Tinga-

Tinga, 12 July 1968. Salinity at capture site 10%,.

Records: MOR 253L.

Paratype: One male, MOR 253L.

Family Colomastigidae

Colomastix pusilla Grube, 1864

Colomastix pusilla: H. K. Barnard, 1925: 346. J. L. Barnard, 1955: 39-42, fig. 20.

Records: MOR 283 L (1).

Diagnosis: Distinguished from other members of the family by the smoothly

rounded telson; uropod 3 rami equal; eyes in live specimen with red lenses

outlined in white; flagellum of antenna 1 two-articulate.

Remarks: Probably considerably more common than the literature suggests

since, with its small size and slender body, it is likely to pass through the sorting

screens used by many workers.

Distribution: Cosmopolitan in tropical and temperate seas.

Family Corophiidae

Cerapus tubularis Say, 1818

Cerapus abditus: Stebbing, 1910: 616, pl. 55A.

Cerapus tubularis: J. L. Barnard, 1962: 61, figs 27-28. Ledoyer, 1967: 137, fig. 27.

Records: MOR 108 F (1); PED 7 Q (2); PED 11 X (4); PED 12 W (4).

THE AMPHIPODA OF SOUTHERN AFRICA 283

Diagnosis: Rostrum prominent; article 3 of antenna 1 as long as article 1,

article 1 with a sharp ventral projection; gnathopod 2 male extremely powerful,

article 5 with a large postero-distal triangular process and a smaller process near

the articulation with article 6; uropod 2 uniramous.

Remarks : C. abditus has been found to be the adult of C. tubularis, the main source

of misidentification having been Stebbing (1906) who restricted C. tubularis to

forms with only three flagellar articles on antenna 1, whereas the number is, in

fact, variable between two and five. Possibly development in colder regions is

retarded, making the terminal ‘abditus’ type rarer.

Distribution: Cosmopolitan in warm and temperate seas.

Corophium triaenonyx Stebbing, 1904

Corophium triaenonyx Stebbing, 1904: 25, pl. 6A. K. H. Barnard, 1940, 482.

Records: MOR 238 K (1); MOR 240 G (1); MOR 243 X (1); MOR 253 D (1);

IN 158 A (2).

Diagnosis: Antenna 2 male with article 4 distally produced into a large curved

- tooth, a smaller tooth on its inner side; article 7 of gnathopod 2 markedly

tri-dentate, the third and largest tooth forming the unguis, pleon segments 4—6

distinct.

Distribution: Tropical Atlantic, Indian Ocean, Mediterranean, Chilka Lake.

Ericthonius brasiliensis (Dana, 1853)

Ericthonius brasiliensis: Stebbing, 1910a: 463. J. L. Barnard, 1955: 37. Ledoyer, 1967: 137, fig. 30.

Records: MOR 218 E (1); JAN 12 H (8); PED to T (1); PED 15 M (1);

amongst Cymodocea at Inhaca (Macnae & Kalk, 1958).

Diagnosis: Head not rostrate; gnathopod 2 male very powerful, article 5

postero-distally with a large bidentate process, dactyl very strong, falciform;

uropods 1 and 2 sublamellar, minutely pectinate, uropod 3 uniramous, ramus

half the length of peduncle and bidentate at tip.

Distribution: Cosmopolitan in tropical and temperate seas.

Grandidierella bonniert Stebbing, 1908

Grandidierella bonnieri: Ledoyer, 1967: 137, fig. 28a.

Records: MOR 30 T (3); MOR 30 U (3); MOR 41 N (1); MOR 52 N (2);

MOR 74 H (7); MOR 77 Z (3); MOR 102 N (1); MOR 240 F (7); MOR 243

Y (1); POE 1 A (1); POE 3 D (common); POE 6 A (common) ; POE 10 F (1);

POE 12 G (5).

Diagnosis: Pereon segment 1 in male with a strong medio-ventral spiniform

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

process; gnathopod 1 male with article 5 not narrowing distally, a strong

spinose projection on inner apical corner, a smaller one on the distal margin,

and a spine on the lower margin.

Distribution: Atlantic and Indian Oceans, Caribbean.

Stphonoecetes ortentalis Walker, 1904

Siphonoecetes orientalis Walker, 1904: 294, pl. 7, fig. 49. K. H. Barnard, 1916: 270.

Records: MOR 124 F (2).

Diagnosis: Eyes poorly developed; flagellum of antenna 2 of one long and two

short densely setose articles with characteristic marginal and apical unguiform

spines; flagellum of antenna 1 subequal to peduncle and consisting of 10-14

articles; rostrum acute and deflexed.

Distribution: ‘Tropical Indo-Pacific.

Family Dexaminidae

Polycheria atolli Walker, 1905

Polycheria antarctica: K. H. Barnard, 1916: 211.

Polycheria atolli: Ledoyer, 1967: 131, fig. 13a.

Records: PED to P (4); PED 15 Q (1).

Diagnosis: Urosome segments with dorsal carinae, segments 2 and 3 fused;

pereiopods chelate; uropods 1 and 3 subequal, 2 much shorter with outer ramus

half the inner; telson cleft to base.

Distribution: Antarctic and southern Oceans, tropical Indian Ocean.

Family Eusiridae

Eusiroides monoculoides (Haswell, 1880)

Eusiroides monoculoides: K. H. Barnard, 1916: 174. J. L. Barnard, 1964: 221, fig. 1.

Records: PED 15 K (1).

Diagnosis: Gnathopods subchelate, article 5 lobate, shorter than 6; third

pleonal epimeron posteriorly convex with 10-12 upturned teeth; telson cleft

half its length, apices bidentate.

Distribution: Circumtropical.

Family Gammaridae

Elasmopus affinis Della Valle, 1893

Elasmopus a ae Sars, 1895: 521, pl. 183.

Records: JAN 12 K (1); JAN 12 M (105); IN 159 G (f.c.); PED 10 R (4);

PED 15 N (2).

THE AMPHIPODA OF SOUTHERN AFRICA 285

Diagnosis: Pereiopods robust, article 2 of pereiopod 5 two-thirds as wide as long;

outer ramus of uropod 3 larger than inner, two fascicles of spines on outer edge,

tips of both rami obliquely truncate and densely spinose; telson narrowly cleft,

tip of each lobe obliquely truncate with three or four apical spines. Gnatho-

pod 2 g with 3-spined tubercle at base of dactylus.

Distribution: Mediterranean, Atlantic, southern Indian Ocean.

Elasmopus japonicus Stephensen, 1932

Elasmopus spinimanus (non Walker, 1905): K. H. Barnard, 1925: 358.

Elasmopus japonicus: Sivaprakisam, 1968: 278, figs 3-5.

Records: JAN 12 A (98); PEA 24 E (8).

Diagnosis: Article 6 of gnathopod 2 large, a rounded process at the base of the

dactylus bears eight strong spines on its margin and three more at its base,

finger less than half the length of article 6.

Remarks : ‘The specimens identified by K. H. Barnard (1916) have a dorsal keel

on pleon segment 4 which excludes them from E. spinimanus. They agree closely

with Stephensen’s figures of E. japonicus.

Distribution: Indo-Pacific, extending to South West Africa.

Maera hamigera (Haswell, 1880)

Maera hamigera: K. H. Barnard, 1916: 196, pl. 27, figs 11-12. J. L. Barnard, 1965: 507, fig. 16.

Records: IN 158 B (4); IN 159 A (8); IN 160 E (1).

Diagnosis: Body not dorsally dentate, posterior edge of third pleonal epimeron

serrate; uropod 3 extending much beyond 1 and 2, rami equal; right gnatho-

pod 2 male larger than the left, palm defined by a strong tooth followed by a

marked concavity and a number of further teeth (six in these specimens but

variable).

Distribution: Indo-Pacific.

Maera inaequipes (Costa, 1851)

Maera inaequipes: K. H. Barnard, 1916: 193. J. L. Barnard, 1959: 25, pl. 5.

meecoras; JAN 12 D (1); IN 160 A (1); PED 23 S (1).

Diagnosis: Gnathopod 2 with a sinus in centre of transverse palm, the stout

dactylus having a rounded tooth which fits the depression; article 2 of pereio-

pod 5 with six or seven serrations; uropod 3, rami unequal, truncated; telson

cleft to base, each lobe bidentate, with four or five terminal spines.

Distribution: Cosmopolitan in tropical and temperate seas.

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

Maera serrata Schellenberg, 1938

Maera inaequipes serrata: Ledoyer, 1967: 127, fig. 9.

Maera serrata: J. L. Barnard, 19706: 155, figs 96-97.

Records: PED 6S (1); PED 10 N (19).

Diagnosis: Differs from M. inaequipes only by the serrated posterior border of the

third pleonal epimeron; the number of serrations is variable, the present

specimens having from three to seven teeth.

Distribution: Indo-Pacific (this is the first record from the southern African

mainland).

Mallacoota subcarinata (Haswell, 1880)

Elasmopus subcarinatus: Stebbing, 1910: 458.

Maera subcarinata: K. H. Barnard, 1940: 460, fig. 26.

Records: PED 21 D (1).

Diagnosis: Pleon segment 3 bicarinate, the two teeth apically inclined towards

one another; gnathopod 2 palm spinose with a central cavity containing a

strong tooth; telson with widely divergent lobes, their tips bidentate.

Distribution: Indian Ocean, Pacific, Mediterranean.

Melita appendiculata (Say, 1818)

Melita fresneli: Stebbing, 19106: 596. K. H. Barnard, 1916: 189, pl. 28, fig. 32.

Melita appendiculata: J. L. Barnard, 19706: 161, figs 103, 104.

Records: MOR 45 E (2); MOR 45 G (2); MOR 77 V (5); MOR 124 D (1);

PED 15 (6); PED 16) Us a) TIN nso) Di4):

Diagnosis: Pleonal tooth formula 7 : 7 : 7 : 5 : 2; pleonal epimeron 3 produced

into a long tooth; gnathopod 2 male unequal, either the larger, article 6 with a

characteristic spoon-shaped palm, three teeth near finger hinge, dactyl power-

ful, hind margin of article 6 longer than front margin; article 2 of pereiopods 4

and 5 narrowing distally, not produced postero-inferiorly.

Distribution: Gosmopolitan.

Melita zeylanica Stebbing, 1904

Melita inaequistylis: (part) K. H. Barnard, 1916: 191.

Melita zeylanica: Sivaprakasam, 1966: 112, fig. 12a-j.

Records: MOR 95 D (3); MOR 253 C (F.C.); POE 10 G (3); POE 12 F (F.C.).

Diagnosis: Pleon without dorsal teeth, segment 2 with two or three submedian

fascicles of spines on each side; gnathopod 1 with short palm distally produced

into a setose lobe into which the dactylus fits.

THE AMPHIPODA OF SOUTHERN AFRICA 287

Remarks: K. H. Barnard (1916) doubtfully synonymized Walker’s (1904)

M. ienuicornis but this has not been accepted since Walker’s specimens had

pleonal teeth while Barnard’s did not.

Distribution: A brackwater species found in India, Ceylon and southern Africa.

Family Haustoriidae

Urothoe elegans Bate, 1857

Urothoe elegans: Ledoyer, 1968: 23, pl. 5.

Records: PED 18 X (1).

Diagnosis: Distinguished from other southern African species by articles 4 and 5

of pereiopod 3, which are longer than broad; dactyl of pereiopod 3 minutely

denticulate.

Distribution: Atlantic and Indian Oceans.

Family Isaeidae

Chetriphotis megacheles (Giles, 1885)

Cheiriphotis durbanensis K. H. Barnard, 1916: 247.

Cheiriphotis megacheles: J. L. Barnard, 1962: 17, fig. 4.

Records: MOR 37 W (4); MOR 41 M (2); MOR 45 F (4); MOR 77 X (4);

MOR 77 Y (7); MOR 238 E (C.); JAN 12 N (29).

Diagnosis: A polymorphic species, young specimens having a moderately well-

developed inner ramus to uropod 3 which disappears in fully developed adults;

gnathopod 2 male changes from an oblique palmed form bearing three large

teeth to a transverse one bearing four or five small irregular teeth; article 3 of

antenna 1 shorter than 1 or 2, accessory flagellum tri-articulate.

Distribution: Indo-Pacific.

Chevalia aviculae Walker, 1904.

Chevalia aviculae: K. H. Barnard, 1916: 252. J. L. Barnard, 1970): 166, fig. 107.

Records: PEA 4 L (5).

Diagnosis: Accessory flagellum of antenna 1 uniarticulate; gnathopod 2 stout,

article 6 subquadrate, palm transverse, convex, defined by a strong tooth;

pereiopods 3 and 5, dactyl bifurcate; urosome segments 1 and 2 coalesced;

uropod 3 obliquely truncate and setose, inner ramus longer than outer.

Distribution: Indo-Pacific, west coast of South Africa.

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gammaropsis afra (Stebbing, 1888)

Eurystheus afer: K. H. Barnard, 1916: 249, pl. 28, fig. 11.

Gammaropsis afra: J. L. Barnard, 19706: 170, fig. 108.

Records: IN 159 H (2).

Diagnosis: Eyes not lageniform; gnathopod 2 palm of two irregular humps,

finely crenulate; otherwise resembling G. atlantica, of which it may eventually

prove to be a variety.

Distribution: Indo-Pacific and eastern Atlantic Oceans.

Gammaropsis atlantica (Stebbing, 1888)

Eurystheus atlanticus: Stebbing, 1908: 86.

Gammaropsts atlantica: J. L. Barnard, 19706: 174, figs 111-113.

PED 15 E (present).

Diagnosis: Cephalic lobes moderately projecting, eyes lageniform; accessory

flagellum thin, six-articulate in adults; gnathopod 2 article 6 14 times as long as

broad, palm oblique with a large cavity near the defining angle and a bulge

near the hinge.

Remarks: A highly variable species, especially as regards the eyes, which range

from lageniform (in South African specimens) to oval. J. L. Barnard (19700)

discusses two phenotypes found in Hawaii and suggests that G. atlantica and

G. afra (Stebbing) may form a single species complex.

Distribution: Eastern Atlantic and Indo-Pacific.

Gammaropstis inhaca n. sp.

Fig. 7

Diagnosis of male: Head equal to two pereon segments, ocular lobes extending

half the length of article 1 of antenna 1; eyes oblique, oval, black.

Articles 1 and 3 of antenna 1 equal, shorter than article 2, flagellum

fourteen-articulate and 70% the length of peduncle; antenna 2 slightly longer

than antenna 1, article 2 produced distally, article 3 curved, flagellum thirteen-

articulate.

Mandibular molar large, palp triarticulate, articles 2 and 3 equal, longer

than article 1.

Coxa 1 acutely produced forwards to base of antenna 2, distal portion

flared outwards; coxae 2—7 small, scarcely touching. Gnathopod 1, article 2

constricted near its origin, article 6 slightly larger than 5, setose on medial

surface, palm oblique, a marked concavity near the hinge followed by a large

THE AMPHIPODA OF SOUTHERN AFRICA 289

Fig. 7. Gammaropsis inhaca n. sp., holotype, male 7 mm.

A. Head. B. Antenna 1. C. Antenna 2. D. Mandible. E. Maxilla 1. F. Maxilla 2. G. Maxilliped.

H. Gnathopod 1. I. Gnathopod 2. J. Pleonal epimeron 3. K. Uropod 3. L. Telson.

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

tooth; remainder of palm concave, a small tooth at defining angle; dactyl cut

into ten teeth, closing onto inner surface of article 6. Gnathopod 2 slightly

larger than 1, inner surface of article 6 extremely setose, palm similar to that of

gnathopod 1 but more exaggerated, dactyl with four teeth, closing onto inner

surface of article 6.

Pereiopods 1 and 2 glandular; article 2 of pereiopods 3-5 not expanded.

Peduncle of uropod 1 with a large distal spine. Uropod 2 not extending

beyond uropod 1, peduncle lacking spine. Rami of uropod 3 slightly longer than

peduncle, outer ramus slightly exceeding inner, and having a small second

article.

Telson quadrate, two subterminal setae on lateral lobes.

Posterior edge of third pleonal epimeron concave proximally, convex

distally, postero-inferior corner faintly notched, a diagonal ridge running to the

notch.

Colour (as preserved): White, a black area distally on article 2 of gnathopod 1

and of pereiopods 4 and 5.

Holotype: Male, 7 mm, unique. S.A.M. A 13068.

T ype-locality: Recovered on intertidal mudflats off the west coast of Inhaca

Island by biologists of the South African Museum, June 1971. |

Records: IN 160 D.

Remarks: This species is easily distinguished from most others by the large teeth

on the palms of gnathopod 1 and 2. Species with similar gnathopods are

G. setiferous, which has a uniarticulate outer ramus to uropod 3, and G. kergueleni,

which is probably the closest relative, but which differs in the structure of coxa I

and by the lack of teeth on the dactyl of gnathopod 2.

Gammaropsis semichelatus (Barnard, 1957)

Eurystheus semichelatus K. H. Barnard, 1957: 8, Fig. 5.

Records: JAN 12 E (en AN 1247 As (6) PR ACoA i (a):

Diagnosis: Article 2 of gnathopod 2 distally lobed, article 3 strongly lobed,

article 6 elongate oblong, distally projecting forwards to form a chela with the

short stout dactyl; third pleonal epimeron postero-inferiorly quadrate, minutely

notched and with an oblique ridge running diagonally to the notch.

Distribution: Endemic to Natal and Mocambique.

Photis kapapa Barnard, 1970

Fig. 8

Photis kapapa J. L. Barnard, 1970b: 192, figs 124, 125.

Records: MOR 244 A (15); PED 15 P (8).

Diagnosis: Article 2 of gnathopod 2 male distally produced into a large sacklike

THE AMPHIPODA OF SOUTHERN AFRICA 2g!

Fig. 8. Photis kapapa Barnard

Male, 3 mm: A. Antenna 1. B. Gnathopod 1. C. Gnathopod 2. F. Uropod 3 and telson. Male,

5 mm: D. Gnathopod 2 (inner aspect). Female, 4 mm: E. Gnathopod 2.

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

lobe bearing stridulation ridges, article 5 cupshaped, with a tumid posterior

lobe, article 6 variable, palm slightly to extremely oblique, defined by a large

blunt tooth in young male which becomes an elongate curved process in termi-

nal form, bisinuate between defining tooth and finger hinge. Coxae 2 to 4 and

sometimes I bearing stridulation ridges.

Distribution: Hawai, Mogambique. This is the first record of the species outside

Hawaiian waters.

Remarks: Some of the males in the present collection are considerably larger

(5 mm as compared with 2,5 mm) and have more highly developed gnathopods

than those figured by Barnard (1970). In its terminal form (Fig. 8d) the palm of

gnathopod 2 male is very oblique and the defining tooth has become enlarged

into a curved lobe arising from the inner margin of the palm and curving

distally and outwards. Small males however show gnathopods (Fig. 8c) very

like those figured by Barnard and I have no hesitation in equating the present

material with that from Hawaii, the only consistent differences being the length

of the spines of uropod 3 and the number of segments of the flagellum of

antenna I (6 as against 5 in Hawaiian specimens).

Family Leucothoidae

Leucothoe spinicarpa (Abildgaard, 1789)

Leucothoe spinicarpa: K. H. Barnard, 1916: 148. Sivaprakasam, 1967: 384, fig. 1.

Records: JAN 14 B (2); ‘Portuguese East Africa’ (K. H. Barnard 1955).

Diagnosis: Article 6 of gnathopod 1 finely crenulate and spinose, article 7 about

half the length of 6; gnathopod 2, process of article 5 densely setose, article 6

massive, palm convex, minutely serrulate throughout; pleon segment 3 postero-

inferiorly quadrate.

Remarks: A rather variable species. J. L. Barnard (1962) points out that in

immature specimens article 3 of antenna 1 is longer in relation to articles 1 and 2

than in adults. In the past, this relationship has been used as an important

taxonomic characteristic and the observation of differential growth may

necessitate a taxonomic revision of the genus.

Distribution: Cosmopolitan.

Family Lysianassidae

Amaryllis macrophthalma Haswell, 1880

Amaryllis macrophthalma: K. H. Barnard, 1916: 114.

Records: PED 23 R (1); JAN 12 F (6); JAN 12 R (1); PEA 24 A (1).

Diagnosis: Eyes vertically elongate, subcrescentic; pleon segment 3 postero-

inferiorly squarely upturned with a little pocket above the point; uropod 2 rami

THE AMPHIPODA OF SOUTHERN AFRICA 293

subequal, the inner markedly constricted a third before the tip; telson extending

beyond the peducle of uropod 3, more than 50% cleft, apices not divergent.

Distribution: Indo-Pacific, extending around the South African coast to South

West Africa.

Aristias symbiotica Barnard, 1916

Aristias symbiotica K. H. Barnard, 1916: 122.

Records: PED 6 N (1).

Diagnosis: Eyes fairly large, oval to circular; third pleonal epimeron postero-

inferiorly quadrate, hind margin finely serrulate; telson as broad as long,

two-thirds cleft, each apex with a stout spine set in a notch; hind margins of

article 2 of pereiopods 3 and 4 with three to four serrations, pereiopod 5 with

six serrations; uropods with short spines at tips of their peduncles, rami minutely

spinulose; uropod 3 rami lanceolate, inner longer than article 1 of outer.

Remarks: All the species of this genus lead a semi-parasitic existence in the

branchial cavities of ascidians or sponges.

Distribution: Endemic, South West Africa to Mocambique.

Lysianassa cinghalensis (Stebbing, 1897)

Lystanassa cinghalensis: Ledoyer, 1968: 19, fig. 1.

iecrds JAN 12 G (2); IN 159 E (2).

Diagnosis: Eyes large, dark, reniform; article i of antenna 1 twice as long and

14 times as wide as articles 2 plus 3, accessory flagellum tri-articulate; gnatho-

pod 1 simple; coxa 1 with a small setiferous notch on the lower margin;

gnathopod 2 very long, article 2 as long as 4 to 6 combined; uropod 2 inner

ramus moderately constricted; uropod 3 peduncle slightly keeled; telson entire

and oval.

Distribution: Tropical Indian Ocean. This is the first record of this species from

the southern African mainland.

Microlysias indica Barnard, 1937

Fig. 9

Microlysias indica K. H. Barnard, 1937: 144.

Records: MOR 102 M (1); MOR 122 T (2); MOR 138 G (1).

Diagnosis: Antenna 2 male half as long as body; article 4 very tumid; gnatho-

pod 1 subchelate, article 6 longer than 5, 1,5 times as long as broad; gnatho-

pod 2, article 5 widening distally, article 2 of pereiopods 3-5 feebly serrate.

Remarks: Barnard’s original specimens were described as having an indistinct

fourth article to the maxillipedal palp but the present specimens have a distinct,

though small, fourth article.

Distribution: South Arabian coast, southern Africa.

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Microlysias indica Barnard, male, 6 mm.

A. Antenna 1. B. Antenna 2. C. Mandible. D. Maxilliped. E. Gnathopod 1. F. Gnathopod 2.

G. Uropod 3. H. Telson.

THE AMPHIPODA OF SOUTHERN AFRICA 295

Trischizostoma circulare Barnard, 1961

Trischizostoma circulare J. L. Barnard, 1961: 51, fig. 20.

Records: 25°36'S/35°21'E, 730 m.

Diagnosis: Article 6 of gnathopod 1 nearly circular, dactyl smooth; rostrum

elongate; telson entire; article 6 of gnathopod 2 asymmetrical, distal end

produced.

Distribution: The above record is the only one to date.

Trischizostoma sp.

icoords= PED) 22 T (1 juvenile).

Remarks: ‘The characteristic powerful gnathopods of this individual identify it

as a member of the genus Trischizostoma. The gnathopods were not, however,

sufficiently developed to allow a specific identification. (T. remipes Stebbing,

1908 and T. circulare are the only species previously recorded from the east

coast of Africa.)

Family Oedicerotidae

Pertoculodes longimanus (Bate & Westwood, 1868)

Perioculodes longimanus: Chevreux & Fage, 1925: 162, figs 163, 164. Ledoyer, 1967: 127, fig. 7.

Records: PED 18 W (1).

Diagnosis: Rostral projection short, lateral corners rounded, eyes broadest

dorsally, with about twelve lenses; gnathopods 1 and 2, process of article 5 to

tip of article 6, article 6 three times as long as broad; rami of uropod 3 very

narrow, unarmed; telson evenly rounded, twice as long as broad.

Distribution: Mediterranean, Atlantic, Indian Ocean:

Synchelidium haplocheles (Grube, 1864)

Synchelidium haplocheles: Sars, 1895: 318, pl. 112, fig. 1.

Records: MOR 212 K (1).

Diagnosis: Rostrum short, evenly curved; eyes large, round, bright red; process

of article 5 of gnathopod 1 produced well beyond hind margin of article 6;

article 6 medially widened, palm longer than hind margin and having six large,

blunt denticles; gnathopod 2 slender, chela one-fifth the length of article 6.

Distribution: North Atlantic, Mediterranean, Ceylon, southern Africa.

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Phoxocephalidae

Metaphoxus sp.

Fig. 10

Diagnosis: Rostrum longer than peduncle of antenna 1, evenly tapering, tip

rounded. Eyes consisting of about 12 ocelli. Antenna 1 as long as head, flagellum

ten-articulate, accessory flagellum eight-articulate. Antenna 2 equal to

antenna I.

Mandibular molar reduced to nine spines; mandibular palp with article 3

distally expanded. Maxilla 1 palp uniarticulate, inner lobe bearing three spines.

Maxillipedal palp article 4 elongated, bearing a strong spine at its tip. Gnatho-

pod 1 about half the size of gnathopod 2, palms of both gnathopods slightly

oblique, convex, defined by a rounded convexity.

Pereiopod 1, article 2 with four long setae posteriorly, article 4 14 times

longer than broad, anteriorly slightly produced to overlap article 5; article 5

almost as broad as long; article 6 slender with six heavy spines along posterior

margin, the last extending to the tip of the dactyl and comparable in width to

it. (Pereiopods 2 and 3 are missing.)

Pereiopod 5, article 2 serrate posteriorly and anteriorly, article 4 with two

fascicles of spines posteriorly. Second pleonal epimeron bearing a group of five

plumose setae on its outer surface. Third pleonal epimeron quadrate.

Uropod 1, rami subequal. Uropod 2, peduncle with two dorsal and one

terminal spine, inner ramus nearly twice as long as outer. Uropod 3 rami

foliaceous, fringed with long plumose setae.

Material: A single damaged male, 10 mm, recovered in two portions such that

pereiopods 3 and 4 were missing on both sides. Unique.

Records: PED 18 V (1).

Remarks: Distinguished from other members of the genus by the very long

rostrum, the setae on the second pleonal epimeron, and the third uropods. The

gnathopods resemble those of M. simillimus and M. pectinatus but the defining

lobes are more rounded and not spinose.

Family Podoceridae

Laetmatophilus purus Stebbing, 1888

Laetmatophilus purus Stebbing, 1888: 1198, pl. 132. K. H. Barnard, 1916: 274.

Records: MOR 243 Z (80).

Diagnosis: Article 2 of gnathopod 2 broad, channeled anteriorly, article 6 with

palm long, undefined, a broad lobe near the hinge followed by a narrow blunt

tooth; pereon transversely corrugated; articles 5 and 6 of gnathopod 1 subequal

and setose, 6 abruptly widening at its base, dactyl toothed.

Distribution: Endemic, South West Africa to Morrumbene.

THE AMPHIPODA OF SOUTHERN AFRICA 297

EEE

Fig. 10. Metaphoxus sp., male, 10 mm.

: A. Head. B. Antenna 1. C. Antenna 2. D. Mandible. E. Maxilla 1. F. Maxilliped. G. Gnatho-

pod 1. H. Gnathopod 2. I. Pereiopod 5. J. Pleonal epimeron 2. K. Uropod 3. L. Telson.

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

Laetmatophilus tridens Barnard, 1916

Laetmatophilus tridens K. H. Barnard, 1916: 275, pl. 28, fig. 22.

Records: PED 10 Q (1).

Diagnosis: Pereon segments 2—4 with small medio-dorsal tubercles; dactyl of

gnathopod 1 very thick and very convex, distally cut into ten closely-set

spine-teeth; article 2 of gnathopod 2 with two anterior keels, one ending

acutely, the other subacutely, palm setose, distally bearing three teeth, that

closest to the hinge triangular, the second cylindrical and the third broad and

denticulate.

Distribution: Endemic, Saldanha Bay to Mocambique.

Podocerus cristatus (Thompson, 1879)

Podocerus cristatus: K. H. Barnard, 1916: 276. J. L. Barnard, 1962: 67, fig. 31.

Records: PED 15 G (4).

Diagnosis: Pereon segments 6 and 7 and pleon segments 1 and 2 medio-dorsally

carinate (on large specimens small carinae may also appear on segments 5 or

even 4 and 3); gnathopod 2 palm bearing a denticulate lobe near the hinge

followed by a conical tooth.

Distribution: Cosmopolitan in tropical and warm-temperate seas.

Podocerus inconspicuus (Stebbing, 1888)

Podocerus palinurt K. H. Barnard, 1916: 277, pl. 28, fig. 23.

Podocerus inconspicuus: Pirlot, 1938: 356, fig. 160.

Records: PED 16 W (1).

Diagnosis: Head with a low rounded dorsal keel; pereon segments 1-7 and

pleon segments 1 and 2 dorsally carinate; gnathopod 2 male with palm bearing

a flat tooth near the hinge and a small conical one below it.

Distribution: Indian Ocean and west coast of South Africa.

Family Stenothoidae

Stenothoe gallensis Walker, 1904

Stenothoe gallensis: K. H. Barnard, 1916: 1543; 1925: 344. J. L. Barnard, 1955: 3 fig. 1.

Records; JAN 12S (2).

Diagnosis: Gnathopod 2 male with hind margin of article 4 finely crenulate,

palm straight, densely hirsute, a double tooth near base of dactylus; uropod 3

uniramous, ramus bi-articulate and slightly shorter than peduncle, article 2

curved upwards, finely denticulate dorsally.

Distribution: Mediterranean, Caribbean, Hawaii, Indian Ocean.

THE AMPHIPODA OF SOUTHERN AFRICA 299

Stenothoe valida Dana, 1853

Stenothoe affinis: K. H. Barnard, 1925: 345.

“Stenothoe valida: Ledoyer, 1967: 125, fig. 4b. Sivaprakasam, 1967: 373, fig. 2a—b.

Records: JAN 12 Q (1); Cabo da Inhaca (Kalk 1958).

Diagnosis: Gnathopod 2 male with the hind margin of article 4 entire, palm

slightly concave, a large, distally directed tooth and a marked incision near the

base of the dactylus; uropod 3, article 2 of ramus straight, not denticulate.

Distribution: Cosmopolitan in tropical and temperate seas.

Superfamily TALITROIDEA

Family Hyalidae

FHiyale grandicorns Kroyer, 1845

Hyale grandicornis: K. H. Barnard, 1916: 230. Stephensen, 1949: 33, figs 14-15. K. H. Barnard,

1955: 93, fig. 46.

Records: Among seaweeds on the east coast of Inhaca Island (Macnae & Kalk,

1958).

Diagnosis: Eyes large, nearly meeting on top of the head; gnathopod 2 article 2

not lobed, article 3 with a small lobe, palm of male oblique, with a pocket-like

cavity and a double tubercle carrying two spines defining it from a fairly long

hind margin.

Remarks: Hyale grandicornis Kroyer and Hyale novaezealandia (Thompson) were

at one stage separated by differences in spination of uropods 1 and 2 and

pereiopod 4. K. H. Barnard (1916) found that these characters were not

consistently correlated in different populations and united the species.

Apparently, one of the several forms predominates in any one population to the

almost complete exclusion of the other forms. Hurley (1957) suggests that

environmental factors determine the genetic balance achieved by different

populations.

Distribution: Indo-Pacific, Tristan da Cunha, South West Africa, Gough Island.

Parhyale inyacka (Barnard, 1916)

Hyale inyacka K. H. Barnard, 1916: 233, pl. 23, fig. 4.

Parhyale inyacka: J. L. Barnard, 1955: 23, fig. 12. Sivaprakasam, 1969): 562, fig. 6.

Records: IN 159 B (1); MOR 40 Z (18); MOR 75 A (11); MOR 232 D (3).

Diagnosis: Antenna 2 half body length, twice as long as antenna 1; gnathopod 2

male with article 6 elongate-oval, palm oblique, convex; pereiopod 3 with hind

margin of article 2 serrate, a marked indent centrally; article 6 of pereiopods 4

and 5 spinose posteriorly; peduncle of uropod 3 slightly longer than outer

ramus, inner ramus small but distinct.

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks : Shoemaker (1956), in a review of the genus, united P. inyacka (Barnard)

with Hyale hawaiensis (Dana) but this has not been accepted by Bulycheva (1957)

or Sivaprakasam (19690).

Distribution: Cosmopolitan in warm-temperate and tropical seas.

Family Talitridae

Orchestia ancheidos (Barnard, 1916)

Talorchestia ancheidos: K. H. Barnard, 1916: 221, pl. 27, figs 35-36; 1940: 470, fig. 31.

Records: MOR 179 A (5); MOR 180 A (abundant); MOR 193 A (1);

POE 8 A (common); Masiene (near Limpopo River mouth) (Barnard 1940).

Diagnosis: Eyes separated by less than their diameter; coxa 2 with a strong

rounded lobe on upper posterior edge; gnathopod 1 male, article 5 strongly

expanded distally and longer than subtriangular article 6; gnathopod 2 male,

article 6 oval, widest at its midpoint, palm convex, spinose, forming an almost

even curve with hind margin, dactyl strongly curved.

Distribution: Malagasy, Mogambique, South Africa.

Orchestia anomala Chevreux, 1901

Talorchestia malayensis: K. H. Barnard, 1955: 93.

Orchestia anomala: Sivaprakasam, 1969a: 297, fig. 1.

Records: Among Botrychia on intertidal rock faces, west coast of Inhaca and

along drift line, Northern Bay (Macnae & Kalk, 1962).

Diagnosis: Articles 4-6 of gnathopod 1 male with scabrous lobes; dactyl of

gnathopod 2 male with averted point; hind margin of article 2 of pereiopod 5

with numerous serrations, pleonal epimera 2 and 3 with submarginal ridges.

Remarks: K. H. Barnard (1935) united T. malayensis ‘Tattersall with his Orchestra

floresiana, which were synonymized with O. anomala Chevreux by Schellenberg

(1938). Barnard again, however, recorded 7. malayensis as a distinct species in

1955. Ihe synonymy established by Schellenberg is nevertheless generally

accepted since 7. malayensis shows a palm in the female gnathopod 1.

Distribution: Indo-Pacific.

Orchestia notabilis (Barnard, 1935)

Pig.

Parorchestia notabilis K. H. Barnard, 1935: 291, fig. 8.

Records: MOR 85 B (6); MOR 95 C (4).

Diagnosis: Gnathopod 2 male strongly developed, article 6 ovate, the straight

palm separated from the hind margin by a slight step, one conical tooth in the

THE AMPHIPODA OF SOUTHERN AFRICA 301

Fig. 11. Orchestia notabilis (Barnard), male, 9 mm.

A. Antenna 1. B. Antenna 2. C. Maxilla 1. D. Maxilliped. E. Gnathopod 1. F. Gnathopod 2.

G. Uropod 3, lateral view. H. Telson.

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

centre of the palm, a slightly larger one distally and two small rounded pro-

jections between the latter and the hinge; inner margin of dactyl sinuous, tip

not averted; pereiopod 5, article 2 with slight and widely spread setiferous

indents.

Distribution: India, Mocgambique.

Talorchestia australis Barnard, 1916

Talorchestia australis K. H. Barnard, 1916: 220, pl. 27, figs 33-34; 1940: 470, fig. 30.

Records: MOR 250 F (2); MOR 255 F (1); IN 159 F (fairly common).

Diagnosis: Eyes distance apart equal to diameter; coxa 2 not lobed posteriorly;

gnathopod 1 male, article 5 triangular with a prominent apical lobe, longer

than article 6, article 6 not strongly expanded, apically lobed, palm concave;

article 6 of gnathopod 2 male oblong, widest across the defining angle, palm

nearly transverse, slightly convex, a small pellucid lobe at palmar angle,

spinose, dactyl matching palm, evenly curved.

Distribution: Endemic, South West Africa to Mocambique.

Suborder CAPRELLIDEA

Family Aeginellidae

Metaprotella haswelliana (Mayer, 1882)

Metaprotella haswelliana: Sundara Raj, 1927: 126, pl. 16. McCain & Steinberg, 1970: 54.

Records: MOR 212 J (3).

Diagnosis: Last two thoracic segments fused, segment 5 long and slender;

pereiopods 1 and 2 almost as long as branchiae; dorsal surface of head and body

spinose; article 1 of antenna 1 bearing a small tubercle with 1 seta.

Distribution: Indo-Pacific.

Monoliropus falcomanus Mayer, 1904

Monoliropus falcimanus: Sivaprakasam, 1967: 382, fig. 4g—h. McCain & Steinberg, 1970: 56.

Records? PED 7. (PED so (4):

Diagnosis: Hand of gnathopod 2 male long, slender, sickle-shaped, the palmar

edge covered by long setae; basis of gnathopod 2 slender with lateral ridges,

longer than pereon segment 2; branchiae long and slender; pereiopods 1 and 2

very small; penultimate joint of maxillipedal palp produced into a pointed

process; flagellum of antenna 1 nine-articulate in male and eight-articulate in

female.

Distribution: Ceylon, India, Mocgambique. This is the first record from Africa.

THE AMPHIPODA OF SOUTHERN AFRICA 303

Orthoprotella mayert Barnard, 1916

Orthoprotella mayeri K. H. Barnard, 1916: 284; 1925: 372. McCain & Steinberg, 1970: 57.

Records: PED 8 N (4); PED 15 S (3); PED 20 T (5); on the hydroid Lytocarpus

philippinus on Inhaca Island (Macnae & Kalk, 1958).

Diagnosis: Young specimens smooth, but those over 10 mm with lateral spines

on the anterior margins of segment 2 and above the base of gnathopod 2; seg-

ment 3 also with antero-lateral spines and a pair of dorsal tubercles; gnathopod 2

with palm sparingly setose, a single triangular tooth near the finger hinge with

a narrow parallel-sided slit cut in the apex and extending nearly to the basal

line; pereiopods 1 and 2 half as long as branchiae, uniarticulate, apically

setose.

Distribution: Indo-Pacific.

Family Caprellidae

Caprella equilibra Say, 1818

Caprella equilibra: McCain, 1968: 25-30, figs 12-13. McCain & Steinberg, 1970: 19.

Records: JAN 12 L (18).

Diagnosis: Basis of gnathopod 2 less than half the length of pereon segment 2; a

spine between the insertions; palm very oblique, defined by a small tooth and

with a large rectangular tooth distally; large males with very elongate pereon

segment 2 and peduncular articles of antenna 1 enlarged.

Distribution: Cosmopolitan, 0-300 m.

Caprella scaura ‘Templeton, 1836

Caprella scaura: K. H. Barnard, 1925: 371. McCain, 1968: 40-44, figs 17-18. McCain & Stein-

bere,1970: 37.

Records: On Cymodocea, Inhaca Island (Macnae & Kalk, 1958).

Diagnosis: Large, anteriorly-directed cephalic spine; pereon segments 1-2 male

elongate, basis of gnathopod 2 equal to pereon segment 2; gnathopod 2 male

with hand elongate, palm with two teeth and a distal rectangular projection.

Remarks: K. H. Barnard (1925) amalgamated C. laevipes Mayer with C. scaura

but this synonymy has not been followed, since C. laevipes does not bear grasping

spines on the pereiopods.

Distribution: Cosmopolitan.

Hemiaegina minuta Mayer, 1890

Hemiaegina minuta: McCain, 1968: 61-64, figs 29-30. McCain & Steinberg, 1970: 51.

Records: JAN 14 C (1).

Diagnosis: Flagellum of antenna 2 bi-articulate; mandibular palp absent,

molar present; in dorsal view, pereonites centrally expanded; pereiopods 1 and 2

uniarticulate; a pair of ventral spines between the insertions of gnathopod 2.

Distribution: Cosmopolitan in warm and temperate seas.

304. ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Phtisicidae

Phtisica marina (Slabber, 1769)

Phiisica marina: K. H. Barnard, 1916: 283. McCain, 1968: 91-97, fig. 46. McCain & Steinberg

1970: 64.

Records: PED 15 R (1).

Diagnosis: Head anteriorly rounded; gnathopod 1 male, hand subtriangular,

palm very oblique, defined by a projecting lobe armed with several spines;

gnathopod 2 male, carpus shorter than merus, hand widest proximally, palm

defined by two grasping spines, otherwise lacking teeth; pereiopods 1 and 2

six-segmented.

Distribution: Atlantic, extending into the Mediterranean and Black Sea, and

around southern Africa as far as Mocambique.

SUMMARY

A synthesis is presented of the known gammaridean and caprellid amphi-

pod fauna of Mocambique south of 20°S. Material was collected by the Univer-

sity of Cape ‘Town and the South African Museum, reference also being made

to specimens collected by the University of the Witwatersrand. Samples were

taken at Morrumbene estuary, Inhaca Island, Jangamo reef, Maxixe, Lagoa

Poelela, Ponta Zavora and by dredging to depths up to 135 m, a total of 65

species being recovered. One genus Janice, and three species, namely Gitanopsis

mariae, fanice spinidactyla and Gammaropsis inhaca are described as new to science.

Five others, namely Paragrubia vorax Chevreux, Photis kapapa Barnard, Lyssianassa

cinghalensis (Stebbing), Monoliropus falcomanus Mayer and Maera serrata Schellen-

berg, are new records for southern Africa (here defined as Africa south of 20°S).

Brief diagnoses, references and distributions are given for each species.

ACKNOWLEDGEMENTS

I wish to express my thanks to Professor J. H. Day for his advice and

encouragement; also to Dr J. C. McCain for his identification of Monolzropus

falcimanus and to Brian Kensley of the South African Museum for the supply of

specimens from Inhaca and Ponta Zavora.

I am also indebted to J. Laurens Barnard, J. C. McCain, T. E. Sivapraka-

sam, M. Ledoyer, Wim Vader, Margaret Kalk and others for literature and

unpublished information.

Financial support was provided by the South African Council for Scientific

and Industrial Research.

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BARNARD, K. H. 1916. Contributions to the crustacean fauna of South Africa. 5. The Amphipoda.

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additions to the list of Amphipoda. Ann. S. Afr. Mus. 20: 319-380.

BarnarD, K. H. 1935. Report on some Amphipoda, isopoda and Tanaidacea in the collection

of the Indian Museum. Rec. Indian Mus. 37: 279-319.

BARNARD, K. H. 1940. Contributions to the crustacean fauna of South Africa. XII. Further

additions to the Tanaidacea, Isopoda and Amphipoda, together with keys for the identi-

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BARNARD, K. H. 1951. New records and descriptions of new species of isopods and amphipods

from South Africa. Ann. Mag. nat. Hist. (12) 4: 698-709.

BARNARD, K. H. 1955. Additions to the fauna-list of South African Crustacea and Pycnogonida.

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INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins, arranged in the following order:

(1) Heading, consisting of informative but brief title, name(s) of author(s), address(es) of

author(s), number of illustrations (figures, enumerated maps and tables) in the article.

(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-

headings to be used sparingly and enumeration of headings to be avoided. (4) Summary

(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

ILLUSTRATIONS

To be reducible to 12 cm xX 18 cm (19 cm including caption). A metric scale to appear

with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their

proper place in the text), with arabic numbering; items of composite figures to be designated

by capital letters (A, B, C etc.).

REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

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

FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

FiscHEr, P.-H., Duva, M. & Rarry, 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.

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

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn scHULTZE, L.

Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature issued

by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).

The Harvard system of reference to be used in the synonymy lists, with the full references

incorporated in the list at the end of the article, and not given in contracted form in the synonymy

list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

C. L. Griffiths

THE AMPHIPODA OF SOUTHERN AFRICA

PART I

THE GAMMARIDEA AND CAPRELLIDEA OF

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