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ANNALS OF THE ANNALE VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME 86 BAND 86

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 86 BAND

HE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

GAPE TOWN KAAPSTAD

1981-1982

SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY

THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE

D54

List OF CONTENTS

Page

CurRiE, P. J.

The osteology and relationships of Tangasaurus mennelli Haughton (Reptilia,

osuchia) (Published: May 19825)).— sae save 44 oe cis ie Mes owen A eae 247

GosLINER, T. M.

The South African Janolidae (Mollusca, Nudibranchia) with the description of a

new genus and two new species. (Published September 1981.)................ 1

GRINE, F. E.

Description of some juvenile hominid specimens from Swartkrans, Transvaal.

CEnblisneGlOctober 198M.) i sxe Seok ee ce Mig ae ene mre Rp Tae Sin i Set 43

GRINE, F. E.

Occlusal morphology of the mandibular permanent molars of the South African

Negro and the Kalahari San (Bushman). (Published October 1981.) .......... IS y7/

GRInE, F. E.

Relative sizes of the maxillary deciduous canine and central incisor teeth in the

Kalahari San (Bushman) and South African Negro. (Published December

ASUS) rea Rte ee baby trae alae a Sig there io ROS Eee) si fee Re ee ere 229

GRINE, F. E. see VAN DEN HEEVER, J. A.

KENNEDY, W. J., KLINGER, H. C. & SUMMESBERGER, H.

Cretaceous faunas from Zululand and Natal, South Africa. Additional observations

on the ammonite subfamily Texanitinae Collignon, 1948. (Published October

GIL) Ss OUNCE sean Rae elle a ate isk aed ee ee SCE SS, at, is ot OR 115

KLEIN, R. G. see Scott, L.

KLINGER, H. C. see KENNEDY, W. J.

Scorr, L. & KLEIN, R. G.

A hyena-accumulated bone assemblage from Late Holocene deposits at Deelpan,

@ranecibree state: (Published December 19812) 22... seh te gee eee te oe 217

SUMMESBERGER, H. see KENNEDY, W. J.

VAN DEN HEEVER, J. A. & GRINE, F. E.

Dinocephalia type material in the South African Museum (Reptilia, Therapsida).

MiublishedvOctober lO Sila) ete aera cayenne Wea tee Cyr rm ete, Sua tus ee a tae ye 73

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Bonisa Gosliner, 1981

SEPTEMBER 1981 ISSN 0303-2515

PA Mind 7

‘OF THE SOUTH AFRICAN

“MUSEUM

CAPE TOWN

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

FIsCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

Fiscuer, P.-H., DuvAL, M. & RaArFy, 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.

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

und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

bthes

date nt

Lie fil

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 86 Band

September 1981 September

Part 1 Deel

THE SOUTH AFRICAN JANOLIDAE

(MOLLUSCA, NUDIBRANCHIA)

mare THE DESCRIPTION.OF A NEW GENUS

AND TWO NEW SPECIES

TERRENCE M. GOSEINER

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 8000

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

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Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000

OUT OF PRINT/UIT DRUK

iy 20S, 5-8), SGD, 2S, 8, ico), SES, 5, 7-5),

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INGED, 5, 7, tor), 15S), WO), 2, BGS), 2X), 28, 250)

EDITOR/REDAKTRISE

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Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 022 2

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

THE SOUTH AFRICAN JANOLIDAE

(MOLLUSCA, NUDIBRANCHIA)

WITH THE DESCRIPTION OF A NEW GENUS

AND TWO NEW SPECIES

TERRENCE M. GOSLINER

South African Museum, Cape Town

(With 28 figures and 3 tables)

[MS accepted 28 April 1981]

ABSTRACT

Examination of the shallow waters of the Cape Peninsula, South Africa, has revealed the

presence of three sympatric species of Janolidae. The description of Janolus capensis Bergh,

1907 is expanded. Janolus longidentatus sp. nov. is described in detail and, despite its

resemblance to J. capensis, possesses several consistently distinct morphological characteristics.

The two species differ in body shape, position of the gonopore and nephroproct, dentition of

the jaws and radula, arrangement of ganglia within the central nervous system, and in several

aspects of reproductive morphology. The third species differs significantly from all other

members of the Janolidae in that the digestive gland surrounds the stomach rather than

entering the notum and cerata. Bonisa nakaza gen. et sp. nov. is described in detail. The

presence of intermediate morphological characters in several species of janolids necessitates the

synonomy of Janolus Bergh, 1884, and Antiopella Hoyle, 1902.

CONTENTS

PAGE

MNILOMUCHON js tine aes hee teen Ge ate oe 1

Janolus capensis Benen lO07. 5 generac. Aas ae. tan 2

NOCOTS UDGOIGHCHIS S05 Wl aeecbheccsesceaoncses 9

Bonisa nakaza Senn etispsnOVei. 44-25 os eek eee 19

DIScussionvOK ne anolidace sys oe coe eee 26

PNCKAMOWLE GS CMICINS ee hay ficients acca teredire ts Cuenta rae Rennes ot 40

INCIERENCES Men aie heii en ee aay en Nea se 41

INTRODUCTION

The Janolidae are a family of arminacean nudibranchs consisting of 18

species in 5 genera. They are widely distributed in the temperate and tropical

waters of the world. The genus Caldukia Burn & Miller, 1969, and the

monotypic genus Galeojanolus Miller, 1971, are known only from Australia and

New Zealand. Proctonotus Alder & Hancock, 1844, is also monotypic and has

been reported only from Great Britain and the Brittany coast of France

(Thompson & Brown 1976). Antiopella Hoyle, 1902, and Janolus Bergh, 1884,

are much more widely distributed.

Ann. S. Afr. Mus. 86 (1), 1981: 1-42, 28 figs, 3 tables.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Bergh (1907) described Janolus capensis from three specimens dredged

from False Bay, south-western Cape. Intertidal and subtidal collections by

means of S.C.U.B.A. around the Cape Peninsula have yielded specimens of

Janolus capensis as well as two previously unknown species, which are here

described.

Antiopella and Janolus have been considered as distinct genera by most

workers who have studied the Janolidae (Eliot 1906; Pruvot-Fol 1954; Marcus

1958; Burn & Miller 1969; Miller 1971). The South African species possess

morphological features which are intermediate between those utilized to separ-

ate Antiopella from Janolus. For this reason, a review of the generic status of

the Janolidae is provided. To supplement this review, specimens of Janolus

fuscus from California, J. cristatus from England, and J. toyamensis from

_ Hawaii were also examined.

Janolus capensis Bergh, 1907

Figs 1A-B, 2-9

Janolus capensis Bergh, 1907: 90, pl. 7 (figs 6-21). Barnard, 1927: 207, pl. 20 (figs 6-7).

Material examined

10 specimens, 20 m depth off Llandudno (34°01’'S 18°20'E), 23 December

ORE

6 specimens, intertidal, Clovelly, False Bay (34°05’S 18°26’E), 17-18

January 1980

3 specimens, 10 m depth, Rooi Els (34°18’S 18°49’E), 23 January 1980

1 specimen, 10 m depth, Castle Rocks, False Bay (34°18’S 18°29’E), 12

February 1980

5 specimens, 10 m depth, Windmill Beach, False Bay (34°12'S 18°27’E), 9

October 1980

2 specimens, 10 m depth, Rooi Els (34°18’S 18°49’E), 26 October 1980

2 specimens, 5 m depth, Miller’s Point, False Bay (34°14’S 18°29’E), 28

October 1980

Distribution

Atlantic and Indian Ocean coasts of the Cape Peninsula, extending east-

ward to Rooi Els, Cape Hangklip.

External morphology

The living animals (Fig. 1A—B) attain a length of 30 mm. The stout body is

broadest anteriorly, tapering to the acute posterior end of the foot. The

rhinophores (Fig. 2A) are perfoliate with 11 to 16 complete or incomplete

transverse lamellae. Between the rhinophores is the large, convoluted inter-

rhinophoral crest (caruncle) (Fig. 2B). The anus in middorsal, near the poster-

ior limit of the notum. The head (Fig. 2C) is rounded with the mouth situated

SOUTH AFRICAN JANOLIDAE 3

Fig. 2. Janolus capensis Bergh, 1907. A. Rhinophore, scale 1,0 mm. B. Inter-rhinophoral crest,

scale 1,0 mm. C. Ventral view of head, scale 2,0 mm. D. Lateral view, scale 3,0 mm. Figs

2A-B, D are drawn from preserved material, Fig. 2C is from living material.

centrally. Short, blunt dorsoventrally flattened oral tentacles are present on

either side of the head. The foot is rounded anteriorly with a deep transverse

groove. The separate male and female gonopores are situated on the right side

near the middle of the body, while the nephroproct is situated more posteriorly

(Fig. 2D).

The cerata are entirely smooth, somewhat laterally compressed and are

arranged in 5 to 6 closely packed longitudinal rows which are irregularly

arranged. Within each ceras (Fig. 3A) are 2 to 4 irregular lobes of the digestive

gland which branch near the middle of the ceras. The anteriormost cerata lack

extensions of the digestive gland.

The ground colour is translucent white. Opaque white markings may be

present or absent on the rhinophoral lamellae, inter-rhinophoral crest and

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

along the dorsolateral margin of the foot. The cerata are tipped with an apical

band of opaque white guanine crystals which may have a bluish tinge. The

digestive gland within the cerata varies from chocolate brown to brick red or

red-orange.

Digestive system (Fig. 3B)

Near the opening of the mouth, numerous small, simple oral glands are

present on the surface of the outer lips of the buccal mass. The buccal mass is

large and muscular with an ovoid opening. A pair of large, highly dendritic

Fig. 3. Janolus capensis Bergh, 1907. A. Ceras dissected to show branching of digestive gland,

scale 1,0 mm. B. Digestive system, scale 1,0 mm.

SOUTH AFRICAN JANOLIDAE =)

salivary glands extends anteriorly from the stomach and enters the buccal mass

near its anterior limit via glandular ducts. The salivary glands are thick at their

insertion, taper sharply and again expand into the dendritic portion. The

oesophagus is wide throughout its length and enters the highly ridged stomach.

Three major branches of the digestive gland emerge from the stomach. At the

posterior limit of the stomach the intestine curves to the right and continues

posteriorly, terminating at the medial anus. A well-developed anal gland

surrounds the anus.

The buccal mass is large and muscular with an oblong opening (Fig. 4A).

The paired jaws (Fig 4B) are large and strong. The inner masticatory border

(Fig. 5A) is thickened and entirely smooth. There is a raised, arched portion

which strengthens the masticatory edge.

The radula (Figs. 5B, 6) is broad and well developed. There may be from

17 to 21 rows of teeth with 26 to 42 lateral teeth on each side of the narrow,

linear rachidian tooth. The hook-shaped lateral teeth (Fig. 6B) are smooth and

sharply arched. Laterally from the centre of the radula, the lateral teeth

increase in size until about one-third of the breadth of that half of the radula, at

which point they again begin to diminish in size for the remaining outer

two-thirds.

Central nervous system

All the major ganglia of the central nervous system (Fig. 7) are situated

anteriorly in the circumoesophageal nerve ring. The cerebral and pleural

ganglia are almost entirely fused, forming a pair of ganglionic masses separated

by a short commissure. The anteriormost nerves on the dorsal surface of the

Fig. 4. Janolus capensis Bergh, 1907.

A. Buccal mass, scale 1,0 mm.

B. Jaw, scale 1,0 mm.

ANNALS OF THE SOUTH AFRICAN MUSEUM

FAEes

EAW

, scale 300 wm between

h of radula, scale 10 wm

jaw

between squares.

icrograp

is Bergh, 1907. A. Scanning electron

tory border of

lectron mi

ica

ing e

micrograph of mast

squares. B. Scann

Fig. 5. Janolus capens

SOUTH AFRICAN JANOLIDAE

Fig. 6. Janolus capensis Bergh, 1907. A. Scanning electron micro-

graph of central portion of radula. B. Scanning electron micrograph

of lateral tooth from outer portion of radula. Scales 10 wm between

squares.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Janolus capensis Bergh, 1907.

Central nervous system.

Scale 0,5 mm

cerebropleural ganglia bifurcate near the base. The outer branch innervates the

rhinophore and a small rhinophoral ganglion is present near the apex of the

nerve at the base of each rhinophore. The inner branch divides again with both

branches innervating the inter-rhinophoral crest. The eyes are situated on short

nerves which join the cerebropleural ganglia near their juncture with the

somewhat smaller pedal ganglia. At the junction of each eye to the cerebro-

pleural ganglia is a minute optic ganglion. Anterior to each buccal ganglion

(Fig. 3B) is a minute gastro-oesophageal ganglion. The pedal ganglia are

separated by a short commissure.

Reproductive system (Fig. 8)

The ovotestis consists of numerous lobes and gives rise to a slightly

convoluted ampulla which subsequently narrows and bifurcates into the male

wae

| See Eee |

Fig. 8. Janolus capensis Bergh, 1907.

Reproductive system. Scale 4,0 mm.

SOUTH AFRICAN JANOLIDAE 9

and female ducts. The vas deferens is highly convoluted and prostatic through-

out its length. It terminates at an elongate, acutely pointed penis. The

extremely elongate oviduct is muscular and gives rise to a very long receptacu-

lum seminis and continues until it joins the albumen gland of the female gland

mass at the female atrium. /n situ, the oviduct and receptaculum seminis are

surrounded by the two lobes of the voluminous mucous gland. The membrane

and albumen glands are significantly smaller than the mucous gland. Adjoining

the oviduct and female gland mass, at the female atrium, is a small pyriform

bursa copulatrix.

Egg mass

The egg mass (Fig. 9) is highly convoluted and corresponds to type B

(Hurst 1967). There are 38 to 45 eggs per capsule.

Natural history

Janolus capensis is associated with, and feeds upon, several species of

arborescent cheilostomatous ectoprocts, most commonly Menipea triseriata

Busk, 1852, and Onchoporella buskii (Harmer, 1923). Mating individuals and

egg masses are often found on ectoproct colonies as well. Janolus capensis is

found commonly throughout the year along the Atlantic and Indian Ocean

coasts of the Cape Peninsula from the intertidal to at least 40 m in depth.

Janolus longidentatus sp. nov.

Figs 1C, 10-17

Type material

Holotype—SAM-A34883, 1 specimen, 3 m depth, Miller’s Point (34°14’S

18°29’E), 10 December 1980

Paratypes—SAM-A34884, 2 specimens, 10 m depth, Castle Rocks, False

Bay (34°18’S 18°29’E), 1 October, 1980

SAM-—A34885, 1 specimen, 5 m depth, Miller’s Point, False Bay

(34°14'S 18°29'E), 28 October 1980

Other material

3 specimens, intertidal, Clovelly, False Bay (34°05’S 18°26'E), 17 Decem-

ber 1979

2 specimens, 10 m depth, Windmill Beach, False Bay (34°12'S 18°29’E), 11

July 1980

Etymology

The epithet /ongidentatus refers to the elongate cusp of the lateral radular

teeth which characterizes this species.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Janolus capensis Bergh, 1907.

A. Egg mass X 6.

B. Single egg capsule x 50.

SOUTH AFRICAN JANOLIDAE 11

External morphology

Fully mature specimens (Fig. 1C) attain a maximum length of 20 mm. The

slender body is widest anteriorly, tapering to an acute posterior margin of the

foot. The rhinophores (Fig. 10A) are perfoliate with 9 to 12 complete or

incomplete, transverse lamellae. The inter-rhinophoral crest (Fig. 10B) is

highly convoluted, narrowest in the middle. The anus is situated mid-dorsally

near the posterior end of the notum. The mouth is located centrally on the

ventral side of the rounded head (Fig. 10C). A pair of short, blunt oral

tentacles extend from either side of the head. The anteriorly rounded foot

possesses a deep transverse groove at its anterior limit. The separate male and

female gonopores are situated laterally in the anterior third of the right side of

the body, while the nephroproct is located more posteriorly, but within the

anterior half of the body (Fig. 10D).

The entirely smooth cerata are long and slender, arranged in 4 or 5

irregular, longitudinal rows. Within each ceras there are 2 main branches of the

digestive gland which usually branch again above the base of the ceras (Fig.

11). The anteriormost cerata lack extensions of the digestive gland within them.

The general body colour of living animals is translucent white, with a pink

or blue cast. Varying amounts of opaque white pigment are present on the

rhinophores and at the apices of the cerata. The digestive gland within the

cerata is chocolate brown to red-orange.

Digestive system

Near the mouth and surrounding the outer lips of the buccal mass is a

narrow ring of small, simple oral glands. The buccal mass (Fig. 12A) is

somewhat dorsoventrally compressed, muscular, with a circular opening. The

paired salivary glands are very thin anteriorly and expand posteriorly into the

dendritic, glandular portion with numerous ramifications. The oesophagus is a

straight glandular tube which expands into the corrugated, saccate stomach.

From the stomach arise three major branches of the digestive gland: two

branches emanate from the anterodorsal portion of the stomach and give rise to

the left and right anterior digestive branches; the third branch arises at the

ventral portion of the stomach and branches to the right and left posterior

digestive branches. The intestine emerges from the posterior portion of the

stomach and curves to the right, continuing posteriorly to the anus. The anus is

surrounded by a large anal gland.

The jaws (Figs 12B, 13A) are of moderate thickness, angular and termin-

ate posteriorly in a rather acute point. The masticatory border consists of 7 or 8

large, rounded denticles. The radula (Figs 13B, 14) is broad, consisting of 18 to

23 rows of teeth. There are 19 to 26 gradually arched, edenticulate lateral teeth

on each side of the somewhat broad rachidian tooth. In the ten specimens

examined, the rachidian teeth all possess 6 to 10 minute striations on each side

of the base of the elongate central cusp. The lateral teeth have an elongate cusp

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Janolus longidentatus sp. nov. A. Rhinophore, scale 0,5 mm. B. Inter-rhinophoral

crest, scale 0,5 mm. C. Ventral view of head, scale 1,5 mm. D. Lateral view, scale 2,0 mm.

Figs 2A—B, D are drawn from preserved material, Fig. 2C is from living material.

SOUTH AFRICAN JANOLIDAE 13

Fig. 11. Janolus longidentatus sp. nov.

Cerata dissected to show branching of digestive gland.

Scale 1,0 mm.

Fig. 12. Janolus longidentatus sp. nov.

A. Buccal mass, scale 1,0 mm.

B. Jaw, scale 1,0 mm.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Janolus longidentatus sp. nov. A. Scanning electron

micrograph of masticatory border of jaw, scale 300 wm between

squares. B. Scanning electron micrograph of radula, scale 10 wm

between squares.

SOUTH AFRICAN JANOLIDAE

Bee sa ae |

Fig. 14. Janolus longidentatus sp. nov. A. Scanning electron

micrograph of central portion of radula, scale 30 wm. B. Scan-

ning electron micrograph of lateral teeth from outer portion of

radula, scale 30 wm between squares.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

and increase in length towards the outer edges of the radula. Only the

outermost teeth decrease in length.

Central nervous system (Fig. 15)

The major ganglia are situated in the circumoesophageal nerve ring. The

cerebral and pleural ganglia are largely distinct. The anteriormost nerve of the

dorsal surface of the cerebral ganglia bifurcates near its base with the inner

branch innervating the inter-rhinophoral crest and the outer branch innervating

the rhinophores. There is a distinct rhinophoral ganglion near the apex of the

rhinophoral nerve. The eyes are situated at the apex of short optic nerves which

join the central nervous system via small optic ganglia at the junction of the

cerebral and pleural ganglia. Extending from the cerebral ganglia are the buccal

~ nerves which join the round buccal ganglia. Anterior to each buccal ganglion is

a minute gastro-oesophageal ganglion. The pedal ganglia are long with an

elongate commissure between them.

Reproductive system (Fig. 16)

The follicles of the ovotestis are numerous and arranged into distinct lobes.

The thick ampulla has 2 or 3 convolutions and narrows to the bifurcation of the

vas deferens and oviduct. The glandular prostatic vas deferens is short with few

convolutions. It enlarges into a much thicker penis sac which is recurved to an

acutely pointed unarmed penial papilla. The muscular oviduct is embedded

Fig. 15. Janolus longidentatus sp. nov.

Central nervous system.

Scale 1,0 mm.

SOUTH AFRICAN JANOLIDAE i7/

Fig. 16. Janolus longidentatus sp. nov.

Reproductive system.

Scale 1,0 mm.

between two lobes of the mucous gland and branches near the middle of its

length to a short, bulbous receptaculum seminis. The major branch continues

distally to its connection with the albumen gland and female gonopore. The

narrow, linear bursa copulatrix joins the oviduct near the junction of the female

gland mass, at the female atrium.

Egg mass (Fig. 17)

The egg mass is a low, flat spiral consisting of 3 or 4 whorls and

corresponds to type B (Hurst 1967). There are 5 to 9 eggs per capsule.

Natural history

Janolus longidentatus feeds upon the cheilostomatous ectoproct Menipea

triseriata Busk. It is commonly found in False Bay but has not been encoun-

tered on the Atlantic coast of the Cape Peninsula. J. longidentatus appears to

have a seasonal distribution. Specimens have not been found from February to

July and appear to be present largely in the winter and spring months.

Specimens have been found from the intertidal to 30 m depth.

Bonisa gen. nov.

Diagnosis

Body stout, ovoid in outline. Rhinophores perfoliate. Inter-rhinophoral

crest low, triangular with few convolutions. Cerata smooth, caducous, not

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 17. Janolus longidentatus sp. nov.

A. Egg mass X 6.

B. Single egg capsule x 100.

SOUTH AFRICAN JANOLIDAE 19

containing branches of digestive gland. Gonopore near middle of body with

nephroproct in posterior third of body. Oral glands small, simple. Salivary

glands well developed with basal bulb. Digestive gland of three lobes surround-

ing stomach, not extending into notum or cerata. Anal glands absent. Jaws

thick with smooth masticatory border. Rachidian and lateral teeth weakly to

strongly denticulate. Pedal ganglia lateral to well separated cerebral and pleural

ganglia. Optic nerves elongate. Reproductive system hermaphroditic, andro-

diaulic. Vas deferens short, prostatic, expanding into blunt, corrugated penis.

Receptaculum seminis large, muscular and adjacent to gonopore. Bursa copula-

trix small, linear and joining receptaculum seminis at female gonopore.

Type species

Bonisa nakaza sp. nov.

Etymology

Bonisa is named for my wife Bonnie Isabel Julien Gosliner.

Bonisa nakaza sp. nov.

Figs 1D-F, 18-25

Type material

Holotype—SAM-A34886, 20 m depth, off Llandudno (34°01'S 18°20’E),

ZaeDecember 1979.

Paratypes—SAM-A34887, 4 whole specimens, 20 m depth, off Llandudno

(34°01’S 18°20’E), 23 December 1979

Other material

1 specimen, 10 m depth, Oudekraal (33°59'S 18°21’E), 13 January 1980

2 specimens, 10 m depth, Castle Rocks, False Bay (34°18’S 18°29’E), 17

January 1980

Etymology

Nakaza is a Zulu word meaning ‘to adorn with beautiful colours’, and the

name is based on the bright coloration typical of this species.

External morphology

Fully mature specimens may reach a length of 100 mm. The body is broad,

roughly oval in outline. The rhinophores (Fig. 18A) are perfoliate with 14 to 23

transverse lamellae. The slightly convoluted inter-rhinophoral crest (Fig. 18B)

is triangular in shape, widest anteriorly. The mid-dorsal anus is located near the

posterior end of the notum. The broad head (Fig. 18C) is round in shape, with

a secondary, anterior lobe. The paired oral tentacles are moderately elongate

and situated on either side of the head. The foot is transversely grooved at its

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 18. Bonisa nakaza gen. et. sp. nov. A. Rhinophore, scale 1,0 mm. B. Inter-rhinophoral

crest, scale 1,0 mm. C. Ventral view of head, scale 1,0 mm. D. Lateral view, scale 5,0 mm.

Figs 2A—B, D are drawn from preserved material, Fig. 2C is from living material.

SOUTH AFRICAN JANOLIDAE 71)\

anterior limit. The separate male and female gonopores are located near the

middle of the right side of the body (Fig. 18D). The nephroproct is situated

more posteriorly, in the posterior third of the animal. The elongate, readily

caducous cerata are smooth and cylindrical. They are arranged in 16 to 21

diagonal rows per side with 8 to 10 cerata per row. Within the cerata a central

muscular duct is present but this contains no extension of the digestive gland.

At the base of each ceras is a ganglion which bifurcates into two nerves which

extend distally into the ceras (Fig. 19A).

The living animals are variably and brightly coloured. (Fig. 1D-F). The

general body colour ranges from translucent white to yellow or yellow-orange.

Yellow or orange pigment is particularly concentrated on the anterodorsal

surface. The rhinophores possess varying amounts of yellow or orange pigment.

The cerata are strikingly variable in their coloration. On the surface they are

most commonly yellow, basally, with varying amounts and shades of blue

pigment on the more distal portion. Some specimens possess a small subapical

concentration of black or dark-blue pigment within the ceratal ducts. Other

“ae

" Bo FRAN \

y ay, 9

By." Ms aerate aa

LA eri ORO POS OR

‘are NRE Soins

d oon Ke 3a

one SESH EA

AOR

BOIS AS

Wes

—

PNR

r£5

= ~,

Fig. 19. Bonisa nakaza gen. et sp. nov.

A. Ceras, scale 1,0 mm.

B. Digestive system, scale 2,0 mm.

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

specimens, which entirely lack blue or yellow pigment, possess only an apical

band of orange on each ceras and rhinophore. A few juvenile specimens have

cerata which are pinkish with blue pigment and opaque white apices. Since

adult specimens of different colour patterns have been observed copulating,

they are considered to be conspecific.

Digestive system (Fig. 19B)

There is a moderately large ring of oral glands surrounding the outer lips.

The massive buccal mass is highly muscular and occupies the anterior third of

the body cavity. The paired salivary glands are well developed, extending from

the posterior limit of the oesophagus. Near their origin there is a large,

spherical expansion of the salivary gland duct which probably functions as a

vestibule for storage of secretory products. The oesophagus is short and

expands into a thin-walled stomach. On the dorsal surface of the stomach is a

large corrugated portion. Surrounding the stomach are three major lobes of the

highly dendritic digestive gland. A digestive lobe joins the stomach on both the

anterolateral sides of the stomach with a third lobe connecting to the antero-

ventral portion. The intestine emerges from the posteroventral portion of the

stomach and continues posteriorly to the anus. The anus is not surrounded by

anal glands.

The buccal mass (Fig. 20A) has an oblong opening. The jaws (Fig. 20B)

are thick and broad with a smooth masticatory border (Fig. 21) which is

supported by an elevated arch of chitin. The outer borders of the jaws are

sharply indented. The radula (Fig. 22) contains 21 to 46 rows of teeth with 7 to

Fig. 20. Bonisa nakaza gen. et sp. nov.

A. Buccal mass, scale 1,0 mm.

B. Jaw, scale 1,0 mm.

SOUTH AFRICAN JANOLIDAE 23

7 % ; es “

“ 4 a a i thy ie

i pS : 4.

& ass me ame see =e

Fig. 21. Bonisa nakaza gen. et sp. nov. Scanning electron micrograph

of masticatory border of jaw. Scale 100 wm between squares.

33 laterals on each side of the rachidian teeth. The rachidians possess 2 to 5

small or elongate denticles. The lateral teeth possess 2 to 7 denticles on each

side of the elongate central cusp.

Central nervous system (Fig. 23)

The major ganglia of the central nervous system are situated in the

circumoesophageal nerve ring. The cerebral ganglia are closely appressed to

each other without a distinct commissure and are joined posteriorly by the

distinct pleural ganglia. The anteriormost dorsal nerve of the cerebral ganglion

innervates the rhinophores and divides near the middle of its length to give rise

to a very thin branch which innervates the inter-rhinophoral crest. The eyes are

situated at the distal end of the elongate optic nerve which joins the junction of

the cerebral and pleural ganglia via a short optic ganglion. Extending anteriorly

from the cerebral ganglia are the paired buccal nerves which enlarge into the

buccal ganglia on the surface of the buccal mass. Ventral to the oesophagus,

the buccal ganglia are joined by a commissure of moderate length and anter-

iorly each gives rise to a smail gastro-intestinal ganglion. The pedal ganglia are

situated laterally to the cerebral and pleural ganglia rather than posteriorly, and

are connected by an elongate commissure.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 22. Bonisa nakaza gen. et sp. nov. A. Scanning electron

micrograph of central portion of radula. B. Scanning electron

micrograph of lateral teeth from outer portion of radula. Scales 30

um between squares.

SOUTH AFRICAN JANOLIDAE 25

Fig. 23. Bonisa nakaza gen. et sp. nov.

Central nervous system.

Scale 0,5 mm.

Reproductive system (Fig. 24)

The ovotestis consists of numerous follicles which are united into an ovoid

mass. The preampullary duct emerges from the anterior end of the ovotestis

and expands into a highly convoluted, thickened ampulla. The ampulla narrows

anteriorly and divides into the oviduct and vas deferens. The vas deferens is

short, thick, and muscular, and expands into the massive penis sac. The penial

papilla is club-shaped and strongly corrugated. The oviduct is muscular and

joins the large receptaculum seminis near its base. Traversing the base of the

receptaculum seminis is a small, linear bursa copulatrix which joins the recepta-

culum near the female gonopore. At the base of the receptaculum sac is a thick

duct which connects it with the small albumen and membrane portions of the

female gland mass. These glands join the massive mucous gland, which forms

the bulk of the female gland mass.

Egg mass (Fig. 25)

The egg mass is about 20 to 30 mm high, consists of numerous convolu-

tions and corresponds to type B (Hurst 1967). There are one or two eggs per

capsule. The egg mass is frequently deposited on the finger-like projections of

gorgonians.

Natural history

Bonisa nakaza is found in the shallow subtidal, at depths of 3 to 30 m on

both the Atlantic and Indian Ocean coasts of the Cape Peninsula. It is

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 24. Bonisa nakaza gen. et sp. nov.

Reproductive system.

Scale 4,0 mm.

associated with and feeds upon the heavily calcified arborescent cheilostoma-

tous ectoproct, Tubucellaria levinseni Canu & Bassler, 1930.

DISCUSSION OF THE JANOLIDAE

Within the suborder Arminacea, the tribe Pachygnatha is subdivided into

three families, the Madrellidae, Dironidae, and Janolidae (Franc 1968). The

Janolidae have been separated from the other two families by having a broad

rather than narrow radula, a medial rather than lateral anus, and by having

extensions of the digestive gland into the cerata. With the discovery of Bonisa

nakaza in this study, the diagnosis of the Janolidae must be expanded to

include forms with or Without extensions of the digestive gland into the cerata.

The family Janolidae has been previously subdivided into five genera.

Proctonotus Alder & Hancock, 1844, and Caldukia Burn & Miller, 1969, lack

an inter-rhinophoral crest. Caldukia can be separated from Proctonotus by its

distinctively stout rhinophores with transverse lamellae and its significantly

narrower radular ribbon. Of the genera which possess an inter-rhinophoral

crest Galeojanolus Miller, 1971, can be separated from Janolus Bergh, 1884,

and Antiopella Hoyle, 1902, by the helmet-like appendage extending anteriorly

from the head. Bonisa gen. nov. must be added to this group and can be readily

separated from the other janolids by the fact that the digestive gland surrounds

the stomach rather than entering the notum and cerata. In several respects

Bonisa nakaza most closely approaches Galeojanolus ionnae Miller, 1971. Both

species have an ovoid body shape, similar coloration, perfoliate rhinophores, a

low, broad inter-rhinophoral crest with few convolutions, jaws with a smooth

masticatory border, a moderately broad radula with strongly denticulate teeth

and a diaulic reproductive system. Both species lack anal glands. However,

there are several significant differences which distinguish the species. Galeo-

SOUTH AFRICAN JANOLIDAE

Fig. 25. Bonisa nakaza gen. et sp. nov.

A. Egg mass X 3.

B. Several egg capsules x 100.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

janolus ionnae possesses an elongate helmet-like appendage on the anterior end

of the head which may be used in capturing mobile prey (Miller 1971), while

there is a short lobe anterior to the head of Bonisa nakaza. G. ionnae has

papillate, inflated cerata, while they are smooth and cylindrical in B. nakaza.

B. nakaza lacks complex oral glands and is devoid of digestive gland ducts in

the cerata which are both present in G. ionnae. The reproductive s¥stem of G.

ionnae, while not fully mature, has a conical, tapered penis and an elongate vas

deferens, while B. nakaza has a short vas deferens and a blunt, corrugated

penis similar in shape to that described for Janolus cristatus (Schmekel 1970).

Miller described a bursa copulatrix in G. ionnae, while B. nakaza possesses

both a large receptaculum seminis and a small bursa copulatrix situated near

the gonopore. These differences, particularly those of the digestive system,

' warrant generic as well as specific separation.

It is difficult to separate Antiopella and Janolus. Several authors have

united the genera (Thiele 1931; Franc 1968; Thompson & Brown 1976), while

others have maintained them as distinct (Eliot 1906; Pruvot-Fol 1954; Marcus

1958; Burn & Miller 1969; Baba & Abe 1970; Miller 1971). Pruvot-Fol (1954)

stated that Antiopella may be separated from Janolus by virtue of the fact that

the component species possess a denticulate instead of smooth masticatory

border of the jaw, smooth instead of papillate cerata, and branched instead of

undivided ducts of the digestive gland within the cerata. However, several

species of janolids (Table 1) possess intermediate or variable character states or

individual characters. They may also possess a mixture of ‘Antiopella’ and

‘Janolus’ characters. Marcus (1958: 40) noted that the structure of the cerata in

the type species of Janolus, J. australis, is questionable. Several workers who

have suggested the separation of Antiopella from Janolus have stated that jaw

denticulation alone should not warrant generic separation (Pruvot-Fol 1954;

Marcus 1958). The fact that Janolus capensis and J. longidentatus are very

similar in many aspects of their external and internal anatomy (present study)

but differ in jaw denticulation supports the view that this character alone

should not be utilized for generic separation in the Janolidae. The division of

the genital ducts into a diaulic or triaulic configuration varies significantly

within Janolus and Antiopella. The reproductive anatomy is described for eight

of the fifteen species described in both genera. In three species, A. mucloc

(Marcus 1958), Janolus comis (Marcus 1958), and J. toyamensis (present

TABLE |

Antiopella and Janolus characteristics in three species of janolids.

Species Ceratal epithelium Ceratal ducts Jaws

Janolus toyamensis smooth or with branched (A) large

few tubercles (A or I) denticles (1)

Janolus capensis smooth (A) branched (A) smooth (J)

Janolus fuscus smooth (A) branched (J) denticulate (A)

A—Antiopella I—Intermediate J—Janolus

SOUTH AFRICAN JANOLIDAE 29

study), the reproductive system is triaulic. In A. cristata (Schmekel 1970), J.

hyalinus (Schmekel 1970), J. fuscus (present study), J. capensis (present study),

and J. longidentatus (present study) a diaulic configuration is present. The

reproductive system of the type species of Janolus, J. australis, is incompletely

described. There is no correlation between division of the genital ducts and jaw

denticulation. There is, therefore, little basis for separating Antiopella Hoyle,

1902, from Janolus Bergh, 1884, and Antiopella is here regarded as a junior

subjective synonym of Janolus on the basis of priority.

KEY TO THE GENERA OF THE JANOLIDAE

ime izestive sland surrounding Stomach . 22... . i... ite tnt eee ee eee eee Bonisa

Digcstivercland extending into notum and cerata ’. 97 .-05. 03445: oes e sas saves anne 2

PRC -oMInOphoral Crest PLESENE. 25 ow Mee ee ik we Tae eye ee ahaa gemebes 3

Inte roninophnoralccrest:abSemt 2s. Ps e256 eet tee oe oa ele al wes oe hes eda ew dees 4

Sc acawiininelmet-lke EXtenSiOM 2. oan eos Meee ne sek be cee meme eats Galeojanolus

Hela NWAGMOUIREKCENGIOM. 5 oes). nj sesepane ao Heel eyerg ales Bt Ht ge ea bs eo nee Janolus

4 Rhinophoral club stout with transverse lamellae, radula6.1.6................ Caldukia

Rhinophores with inconspicuous lamellae, radula broad.................. Proctonotus

The following species are considered to constitute the family Janolidae

(“indicates type species):

Bonisa gen. nov.

1. *Bonisa nakaza sp. nov.

Caldukia Burn & Miller, 1969

2. *Caldukia affinis (Burn, 1958)

Proctonotus? affinis Burn, 1958: 32, fig. 8, pl. 7 (fig. 15).

Caldukia affinis (Burn, 1958), Burn and Miller, 1969: 23, figs 1-2, pl. 2.

3. Caldukia albolineata Miller, 1970

Caldukia albolineata Miller, 1970: 279, figs 1-10.

4. Caldukia rubiginosa Miller, 1970

Caldukia rubiginosa Miller, 1970: figs 11-21.

Galeojanolus Miller, 1971

5. *Galeojanolus ionnae Miller, 1971

Galeojanolus ionnae Miller, 1971: 491, figs 1-4.

Janolus Bergh, 1884

6. *Janolus australis Bergh, 1884

Janolus australis Bergh, 1884: 19, pl. 8 (figs 15-22), pl. 9 (figs 6-8).

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

7. Janolus barbarensis (Cooper, 1863)

Aeolis barbarensis Cooper, 1863: 59.

Janolus coeruleopictus Cockerell & Eliot, 1905: 48, pl. 8 (figs 12-16). O’Donoghue, 1922: 141.

Janolus barbarensis (Cooper, 1863), O’Donoghue, 1922: 141.

Antiopella aureocincta MacFarland, 1966: 303, pl. 57 (figs 1-5), pl. 63 (figs 13-30), pl. 64 (figs

117) Roller 197053725 in) pant-

8. Janolus capensis Bergh, 1907

Janolus capensis Bergh, 1907: 90, pl. 7 (figs 6-21).

9. Janolus comis Marcus, 1955

Janolus comis Marcus, 1955: 170, pl. 25 (figs 226-236).

10. Janolus cristatus (Chiaje, 1841)

Eolis cristatus Chiaje, 1841: pl. 88 (figs 1-12).

Janus spinolae Verany, 1845: 24, pl. 2 (fig. 9). Pruvot-Fol, 1954: 375.

Antiopa splendida Alder & Hancock, 1848: 190. Alder & Hancock, 1851.

Antiopella cristata (Chiaje, 1841), Hoyle, 1902: 214.

Janolus cristatus (Chiaje, 1841), O’Donoghue, 1924: 1.

11. Janolus flagellatus Eliot, 1906

Janolus flagellatus Eliot, 1906: 374.

(Likely a junior synonym of J. hyalinus (Alder & Hancock, 1854). Eliot

(1910) considered this as an uncertain species. It was distinguished from

J. hyalinus by possession of a flagellar penis which has subsequently been

described in J. hyalinus (Schmekel 1970).)

12. Janolus fuscus O’Donoghue, 1924

Janolus fuscus O’Donoghue, 1924: 16, pl. 2 (figs 18-20).

Antiopella aureocincta Johnson & Snook, 1927: 500. Nomen nudum, Steinberg, 1963: 66.

Antiopella aureotincta MacGinitie & MacGinitie, 1949: 135. Nomen nudum, Steinberg, 1963:

66.

Antiopella aureocincta MacFarland, 1966: 303, pl. 57 (figs 1-5), pl. 63 (figs 13-30), pl. 64 (figs

11-17). Roller, 1970: 372, in part.

13. Janolus hyalinus (Alder & Hancock, 1854)

Antiopa hyalina Alder & Hancock, 1854a: 105.

Janolus hyalinus (Alder & Hancock, 1854), Eliot, 1906: 374.

14. Janolus indicus (Eliot, 1909)

Antiopella indica Eliot, 1909: 143.

Janolus indicus (Eliot, 1909), comb. nov.

15. Janolus longidentatus sp. nov.

16. Janolus mirabilis Baba & Abe, 1970

Janolus mirabilis Baba & Abe, 1970: 65, figs 2-3.

17. Janolus mucloc (Marcus, 1958)

Antiopella mucloc Marcus, 1958: 37, figs 62-71.

Janolus mucloc (Marcus, 1958), comb. nov.

18. Janolus novozealandicus (Eliot, 1907)

Antiopella novozealandica Eliot, 1907: 331.

Janolus novozealandicus (Eliot, 1907), comb. nov.

SOUTH AFRICAN JANOLIDAE Sil

19. Janolus praeclarus (Bouchet, 1975)

Antiopella praeclara Bouchet, 1975: 127, pl. 1 (fig. 3), fig. 5.

Janolus praeclarus (Bouchet, 1975), comb. nov.

20. Janolus toyamensis Baba & Abe, 1970

Janolus toyamensis Baba & Abe, 1970: 63, fig. 1.

Proctonotus Alder & Hancock, 1844

21. *Proctonotus mucroniferus (Alder & Hancock, 1844)

Venilia mucronifera Alder & Hancock, 1844: 163, pl. 2.

Proctonotus mucroniferus (Alder & Hancock, 1844) Alder, 1844: 407.

Zephyrina pilosa Quatrefages, 1844: 130, pl. 3 (fig. 1), pl. 4 (fig. 1), pl. 5 (figs 1-2), pl. 6 (figs 1,

11-12). Pruvot-Fol, 1954: 372.

Morphological variability

Janolus varies in its external and internal morphology. The major morpho-

logical features and their character states in the fifteen described species of

Janolus are summarized in Table 2. Many taxa are incompletely described,

including the type species, but the available morphological data does permit a

review of the variability within the genus. All described species have a simply

rounded head, perfoliate or, exceptionally, papillate (J. comis Marcus, 1955)

rhinophores, an inter-rhinophoral crest, and cerata which contain ducts of the

digestive gland. The inter-rhinophoral crest may be small and rectangular or

may be highly elaborate as in J. cristatus (Alder & Hancock 18546), J. capensis

(present study), J. novozealandicus (Eliot 1907), and J. longidentatus (present

study). The cerata are papillate in J. comis (Marcus 1955), J. hyalinus (Alder &

Hancock 1854b), and J. mirabilis (Baba & Abe 1970), and smooth in the

remainder of the species. The cerata of J. toyamensis are variable, with

scattered, small papillae (Baba & Abe 1970) or with an entirely smooth surface

(present study). The digestive gland may extend only within the basal half of

the cerata in Janolus comis (Marcus 1958), J. hyalinus (Bergh 1904), and

Galeojanolus ionnae (Miller 1971), or extend to the apex of the cerata in the

remaining species of Janolus. Within the cerata the digestive gland may be

branched or unbranched. In species with a branched digestive gland, the

splitting may occur apically in J. cristatus (Alder & Hancock 185456) and

J. barbarensis (MacFarland 1966), near the middle in J. mucloc (Marcus 1958),

J. praeclarus (Bouchet 1975), J. toyamensis (Baba & Abe 1970), and J. capensis

(present study), or basally in J. longidentatus (present study). In most species of

Janolus all cerata contain extensions of digestive gland except J. hyalinus (Eliot

1910), J. comis (Marcus 1958), J. capensis (present study), and J. longidentatus

(present study) where the anteriormost cerata are devoid of digestive gland

tissue.

It appears that in all species of Janolus that have been studied, simple oral

glands and elaborate salivary glands are present and the digestive gland occurs

as three major branches from the stomach. Anal glands appear to be present in

all species except J. comis (Marcus 1955) and J. fuscus (present study).

ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

Comparative morphology of Janolus.

Species of Ceratal Ceratal 5

Janolus Colour epithelium ducts Jaws Radular teeth

J. australis unknown smooth ? unknown smooth rachidian and =

laterals smooth

J. barbarensis . translucent white; cerata smooth branched unknown rachidian smooth, w

with gold and blue laterals: inner 2 denticulate,| |

outer smooth

J. comis light brown with dark- papillate unbranched smooth rachidian denticulate, 4

brown stipples laterals: denticulate, variable} *

J. cristatus cream or brown with smooth branched at 8-12 denticles rachidian and laterals

opaque white cerata apex smooth, rarely denticulate

J. flagellatus unknown unknown unknown smooth rachidian and laterals

smooth

J. fuscus translucent white with smooth unbranched 8-14 denticles rachidian denticulate,

: brown lines, cerata with laterals: inner 2 denticulate,| |

yellow and white outer smooth

J. hyalinus cream with red-brown papillate unbranched smooth rachidian and laterals

blotches; cerata dotted with smooth or finely denticulate

white and brown

J. indicus translucent with red-brown unknown unknown denticulate rachidian and laterals

spots, cerata grey-green smooth

J. mirabilis yellow with brown and papillate unbranched? smooth rachidian, denticulate,

Opaque white; cerata white laterals: inner 3 denticulate,

at apex outer 2 smooth

J. mucloc . transparent white with smooth branched, 8 denticles rachidian denticulate, |

opaque white lines; cerata bifurcate laterals: inner denticulate,

with orange spot and below outers smooth

opaque white lines middle

J. novozealandicus grey with purple stripe and smooth unbranched indistin-t rachidian and laterals

spots; cerata with purple denticulation smooth

spots

J. praeclarus orange with opaque white; smooth branched 6 denticles rachidian and laterals

cerata with yellow bands smooth

and bluish-white apex

J.toyamensis . yellow with white and few minute branched 2 or 3 large rachidian and laterals

brown markings; cerata papillae denticles smooth

with yellow spot and red-

brown apex

J. capensis translucent white with smooth branched smooth rachidian and laterals

Opaque white; cerata red- smooth

brown with opaque white

apex

J. longidentatus translucent white with smooth branched 7-8 denticles rachidian striate, ;

Opaque white; cerata red- basally laterals smooth

brown with opaque white

apex

{

21 x 21.1.21

SOUTH AFRICAN JANOLIDAE

Receptaculum Bursa Prostatic Reproductive

Radular formula seminis copulatrix Penis vas deferens ducts References

24 x 30-54.1.30-54 unknown unknown thicker unknown unknown Bergh 1884

than vas

deferens

6X27 127 unknown unknown unknown unknown unknown Cockerell & Eliot 1905

MacFarland 1966, in part

16 x 12—20.1.12—20 proximal, absent conical, proximalto triaulic Marcus 1955

25 x 41.1.41 serial flagellar penis only Marcus 1958

30 x 40.1.40 proximal, minute, blunt throughout diaulic Alder & Hancock 1851

24 x 33.1.33 semi-serial distal Alder & Hancock 1855

Schmekel 1970

Thompson & Brown 1976

15 x 20.1.20 unknown large, unknown unknown unknown Eliot 1906

elliptical

PAS C22 N22 proximal, distal conical throughout diaulic O’Donoghue 1924

ZO25. 1625 semi-serial MacFarland 1966, in part

present study

15 x 11—13.1.11-13 small, absent elongate, throughout diaulic Alder & Hancock 1854b

proximal, flagellar Alder & Hancock 1855

Bergh 1888, Eliot 1906,

semi-serial Schmekel 1970

Pip sletesil unknown unknown unknown unknown unknown Eliot 1909

DO x5 15 unknown unknown unknown unknown unknown Baba & Abe 1970

18 x 24.1.24 proximal, absent thick throughout triaulic Marcus 1958

semi-serial

within female

gland mass

PX Silos unknown unknown short unknown unknown Eliot 1907

20 x 28.1.28 unknown unknown unknown unknown unknown Bouchet 1975

20 x 10-—20.1.10-20 proximal absent elongate, throughout triaulic Baba & Abe 1970

OX 25S Ae25 flagellar present study

20 x 42.1.42 elongate, distal, elongate, throughout diaulic Bergh 1907

18 x 41.1.41 proximal, minute flagellar Barnard 1927

17 x 26.1.26 semi-serial present study

LETS< AAT a short, distal, conical short diaulic present study

22) 26:1226 proximal, minute

20 x 20.1.20 semi-serial

ANNAL

S OF THE SOUTH AFRICAN MUSEUM

SOUTH AFRICAN JANOLIDAE

TABLE 2

Comparative morphology of Janolus.

Ceratal

Species of pit ducts Jaws Radular teeth Receptaculum Bursa Prostatic Reproductive

Janolus Colour ; aan Radular formula seminis copulatrix Penis vas deferens ducts References

. th ? unknown smoot rachidian and :

J. australis unknown estel) laterals smooth 24 x 30-54.1.30-54 = unknown unknown pueneD unknown unknown Bergh 1884

an vas

deferens

as anched unknown rachidian smooth

J. barbarensis . EransLiceus site ETA smooth brane! NaPeTAIee inner 2 denticul, 16 x 27.1.27 unknown unknown unknown unknown unknown Cockerell & Eliot 1905

with gold an outer smooth y MacFarland 1966, in part

5 5 i th achidi i z

J. comis light brown with dark- papillate unbranched smoo HSE PANE, 16 x 12-20.1.12-20 —_— proximal, absent conical, Proximalto triaulic Marcus 1955

brown stipples » Variable 25x 41.1.41 serial flagellar penis only Marcus 1958

= i th branched at 8-12 denticles rachidian and laterals A A =

J. cristatus cream or brown with sn100, A 30 x 40.1.40 proximal, minute, blunt throughout diaulic Al ancock 185

opaque white cerata pees smooth: rately eaten 24 x 33.1.33 semi-serial distal : ; Ree ee enee

Schmekel 1970

= Thompson & Brown 1976

J. flagellatus . unknown unknown unknown smooth Aechistan and laterals 15 x 20.1.20 unknown Testes unknown unknown unknown Eliot 1906

elliptica’

J. fuscus translucent white with SH Rote ay CCRC eS ag Facclan denteua 21 x 2.1.22 proximal, distal conical throughout diaulic O'Donoghue 1924

brown lines, cerata with Bi nae 26 x 25.1.25 semi-serial MacFarland 1966, in part

yellow and white S present study ;

F ; i branched smooth rachidian and laterals aoa

J. hyalinus cream with red-brown __ papillate wn S 15% 11-13.1.11-13 small, absent elongate, throughout diaul Al ancock 185:

6 blotches; cerata dotted with smooth or finely denticulate proximal, flagellar g) iaulic ALE - Pancake Las

white and brown : f Bergh 1888, Eliot 1906,

semi-serial Schmekel 1970

J. indicus . translucent with red-brown unknown unknown denticulate peehidian and laterals 21 31.1.31 unknown unknown unknown unknown unknown Eliot 1909

spots, cerata grey-green smoot

. mirabilis ellow with brown and papillate unbranched? smooth rachidian, denticulate, ant Ie k k

J. mirabilis seaehe De teoraial White laterals: inner 3 denticulat, 20 x 5.1.5 unknown unknown unknown unknown unknown Baba & Abe 1970

at apex outer 2 smooth

J. mucloc . transparent white with smooth branched, 8 denticles rachidian denticulate, 18 x 24.1.24 proximal, absent thick throughout _ triaulic Marcus 1958

opaque white lines; cerata bifurcate laterals: inner denticulate, semi-serial

with orange spot and below outers smooth within female

opaque white lines middle gland mass

J. novozealandicus grey with purple stripe and smooth unbranched indistin ot. rachidian and laterals 23 %37.1.37 unknown unknown short unknown unknown Eliot 1907

spots; cerata with purple denticulation smooth

spots

J. praeclarus . orange with opaque white; smooth branched 6 denticles rachidian and laterals 20 x 28.1.28 unknown unknown unknown unknown unknown Bouchet 1975

cerata with yellow bands smooth

and bluish-white apex

J. toyamensis . yellow with white and few minute branched 2 or 3 large rachidian and laterals 20 x 10-20.1.10-20 proximal absent elongate, throughout triaulic Baba & Abe 1970

brown markings; cerata papillae denticles smooth 19 x 25.1.25 flagellar present study

with yellow spot and red-

brown apex

J. capensis translucent white with smooth branched smooth rachidian and laterals 20 x 42.1.42 elongate, distal, elongate, throughout diaulic Bergh 1907

opaque white; cerata red- smooth 18x 41.1.41 proximal, minute flagellar Barnard 1927

brown with opaque white 17 x 26.1.26 semi-serial present study

apex

J. longidentatus translucent white with smooth branched 7-8 denticles rachidian striate, 18 x 21.1.21 short, distal, conical short diaulic present study

opaque white; cerata red- basally laterals smooth 22 x 26.1.26 proximal, minute

brown with opaque white 20 x 20.1.20 semi-serial

apex

21x 21.1.21

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

As mentioned in the discussion of the synonomy of Antiopella with Janolus

the elaboration of the masticatory border and shape of the jaw are interspeci-

fically variable (Fig. 26). Previous workers (Pruvot-Fol 1954; Marcus 1958)

have emphasized the distinctness of a smooth versus denticulate masticatory

border. However, Janolus toyamensis (Fig. 26C) possesses three large tubercles

on the masticatory border and J. novozealandicus was described as having

indistinct denticles (Eliot 1907).

The radular teeth are variable within Janolus (Fig. 27) and some intraspe-

cific variations exists. The rachidian teeth are denticulate in J. fuscus (MacFar-

land 1966; present study), J. comis (Marcus 1958), J. mirabilis (Baba & Abe

1970), and J. mucloc (Marcus 1958), striate in J. longidentatus (present study),

and smooth in the remaining species. The rachidian tooth has a broad base in

species with denticulations or striations and is linear in species with smooth

teeth. In J. fuscus (MacFarland 1966; present study), J. mirabilis (Baba & Abe

1970), and J. mucloc (Marcus 1958) the inner 1 to 3 lateral teeth are denticulate

while the outer teeth are entirely smooth. J. cristatus generally has smooth

lateral teeth (Alder & Hancock 1855), but exceptionally (Bergh 1874) they are

denticulate. The presence or absence of denticles varies with age in J. comis

(Marcus 1958) and J. hyalinus (Eliot 1906). Bouchet (1975) noted that the

lateral teeth of J. praeclarus increase in size until the eighteenth row and then

begin to diminish. In J. australis the innermost are largest (Bergh 1884). In

J. cristatus (present study) and J. capensis (present study) the teeth increase in

length on either side of the radula until about one-third of the breadth, and

subsequently diminish. In J. capensis the radular morphology of fifteen speci-

mens of various sizes did not vary significantly. In J. hyalinus (Eliot 1906) and

J. novozealandicus (Eliot 1907), and in at least ten specimens of J. longidenta-

tus (present study) the largest lateral teeth are found at or adjacent to the outer

edge of the radula. The relative length of the base to the cusp of the lateral

teeth varies considerably between species and appears to be useful in the

separation of species.

The central nervous system of Janolus cristatus was described (Alder &

Hancock 1851) as containing distinct cerebral and pleural ganglia. Bergh noted

that these ganglia are fused in Janolus australis (Bergh 1884) and J. capensis

(Bergh 1907). The present study confirms this arrangement in J. capensis, while

J. longidentatus possesses distinct ganglia as in J. cristatus.

The morphology of the reproductive system has been fully described in five

species of Janolus. In J. comis and J. mucloc (Marcus 1958) the arrangement is

triaulic with a short uterine duct connecting the vagina and female gland mass

near the separation of the oviduct and vas deferens from the ampulla. In

J. cristatus and J. hyalinus (Schmekel 1970) the vagina joins the female gland

mass only at the female atrium, producing an androdiaulic configuration. The

present study has shown that J. toyamensis (Fig. 28G) has a triaulic arran-

gement with a proximal receptaculum seminis but with no bursa copulatrix.

J. capensis and J. longidentatus have a diaulic arrangement with a proximal

SOUTH AFRICAN JANOLIDAE a5

Fig. 26. Variation in jaw morphology. A. Janolus fuscus O’Donoghue, 1924. B. Bonisa nakaza

gen. et sp. nov. C. Janolus toyamensis Baba & Abe, 1970. D. Janolus longidentatus sp. nov.

E. Janolus capensis Bergh, 1907. Scale 1,0 mm.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 27. Variation in the rachidian, first, seventh and twentieth lateral teeth in the Janolidae.

A. Janolus capensis Bergh, 1907. B. Janolus longidentatus sp. nov. C. Janolus cristatus (Chiaje,

1841). D. Janolus fuscus O’Donoghue, 1924. E. Janolus toyamensis Baba & Abe, 1970.

F. Bonisa nakaza gen. et sp. nov. Not drawn to scale.

SOUTH AFRICAN JANOLIDAE 37

receptaculum seminis and a minute, distal bursa copulatrix. This configuration

is also present in J. fuscus (Fig. 28F) contrary to that described by MacFarland

(1966, pl. 64 (fig. 11)).

The shape of the penis has been utilized as a characteristic for specific

separation. Janolus flagellatus (Eliot 1906) was described as distinct from J.

hyalinus because its elongate, flagellar penial papilla was thought to be differ-

ent from the blunt papilla depicted by Bergh (1888) for J. hyalinus. Subsequent

work by Schmekel (1970) has shown that the penis of J. hyalinus is flagellar

when fully extended. The shape of the penis varies considerably within species

depending on degree of extension. In comparing penial structure between

species, it is imperative that descriptions be made from specimens with a fully

extended penis.

The egg mass of Janolus cristatus (Alder & Hancock, 1855: pl. 44 (figs

6—7)) is a short, convoluted ribbon of about one and a half whorls, with a single

egg per capsule. A similar egg mass is apparently present in Janolus fuscus

(O’Donoghue 1924). The egg mass of Janolus capensis (present study) is

globose, consisting of many whorls and with 38 to 45 eggs per capsule, while

that of J. Jongidentatus is a flat spiral consisting of 5—9 eggs per capsule.

Janolus longidentatus differs from all other species of Janolus which

possess denticles on the masticatory edge. J. mucloc (Marcus 1958) is triaulic

and therefore differs from the diaulic J. Jongidentatus. J. praeclarus (Bouchet

1975), while much of its anatomy remains unknown, differs from J. longidenta-

tus in its Orange versus translucent white coloration, smaller inter-rhinophoral

crest, branching of the ceratal ducts near the middle rather than basally, more

sharply arched radular teeth that are longest near the middle rather than at the

outer edge of the radula, and jaws with 6 denticles rather than 7 or 8. Janolus

longidentatus differs from J. fuscus in its coloration, more elaborate inter-

rhinophoral crest, smooth inner lateral teeth, and presence of anal glands.

Janolus longidentatus, although similar to J. cristatus in coloration, external and

reproductive anatomy (Schmekel 1970), differs in several significant aspects. In

J. longidentatus the ceratal ducts branch basally while they branch apically in J.

cristatus. The rachidian teeth of J. Jongidentatus are broad with striations versus

the smoothly linear form of J. cristatus. In J. cristatus the radular teeth are

largest at the inner third of the radula, while they are broadest near the outer

margin in J. longidentatus. The lateral teeth are broader and heavier in J.

cristatus. The oviduct is short in J. cristatus and elongate in J. longidentatus. J.

cristatus has a blunt club-shaped penis, while in J. /ongidentatus it is conical.

Despite the difference in the elaboration of the masticatory border of the

jaw, J. longidentatus most closely approaches J. capensis in its morphology. For

this reason a morphological comparison of the three sympatric South African

janolids is provided in Table 3.

The South African species of janolids occur together in the shallow waters

of False Bay. All three species can be found within a few centimetres of each

other. Janolus capensis and J. longidentatus have been observed to feed upon

ANNALS OF THE SOUTH AFRICAN MUSEUM

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SUOTINJOAUOD MOI yum poinjoauos ATYysTY yys1es “y10Ys SUdIOJOp SBA

yeotuos Ieyjosey povesnis09 kat stuod

jouN)sIp posnj youNSsIp eljsues jeinojd pue [eiqo1eo

qjoours ‘dsno a3e3u0]9 YIM yoous ‘podeys-yooy aje[nouep ‘podeys-yooy Y199} [e19}e]

e[NpeI JO UIsIVW 19j}NO IevoU B[Npel JO YIpIM jo ply) 4199} JSOWIOUUI sje1o}e] 1SoSIe]

SUONPII}S YIIM ‘peoiq reoul] ‘YJOOUS sopPluep o31e] YIM ‘peolq YJ00} UvIpPIyoes

Sopoluep g-/ yJOous YJOOUIS Japiog A1ojeonseul

UONIISUI Je Ie[Npueys uOonJesuI je Jode}y IIOAIOSOI [eseq YIM spurs Arearyes

punol ro 0(0) (@[0) Zuojqo sulusdo ynoul

Apoqg Jo d]pprw Iesu Apoq jo o][pprul 0} Jo1193sod Apoq JO pity} 1o11a}sod yooido1ydou

Apoq Jo pity} Jo1ajue Apog Jo 3[pprw edu Apog jo o]pprw re9u a1odouos

aseq Ieau ‘iejnse1 ‘pylupenb J[pplu Ieou ‘Ie[NsoIM “pynnuw juasqe sjonp [eye100

suOINJOAUOS Aue ‘YsTY suolNjoAuos Aueu ‘Yysty SUONNJOAUOD MOJ ‘MOT soo Je1OYdouTYI-19}UI

Jopusys no}s yno}js Apoq

Snjvuap1suo] snjouve sisuadvo snjouve DZDYDU DSIUOg

‘sprjouel uevoLyy ynos jo Asojoydiow saneiedwod

¢ ATaV

SOUTH AFRICAN JANOLIDAE

“Q[2OS 0} UMBIP JON ‘(Apnys yussoid) ‘Aou ‘ds j9 ‘uod DzDynU DSiuog “HY ‘(Apnys yuosoid)

OL6I “9QV w eqeg sisuaupdo] snjouve ‘DH “(Apnys jussoid) p76, ‘onysou0d,.C snosn{ snjouvr ‘4 “(OL6T [OYOWYOS Joye) (TpET ‘oferD) snywjsi9

snjouvr “ZA “(OL6I JPYouyos Joye) (post “YoooueH 2%» Jppy) snuyody snjouve *q “(OLGT [OYewWYIS Joe) Ccoy ‘snoIe] suo snjouve *D *(Apnjs

juasoid) -aou ‘ds snipjuapisuo] snjouvr ‘gq ‘(Apnys juasoid) (06, ‘YsiI9g sisuadvo snjouvs ‘YW ‘snjouvs ut Wa\sks dAONpoOidal Jo UONeLIeA °87 “SI

Pa Su

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40 ANNALS OF THE SOUTH AFRICAN MUSEUM

the same species of ectoproct, while Bonisa nakaza feeds on a more heavily

calcified species. Fdod abundance does not appear to be a limiting factor as

many ectoproct colonies are devoid of janolids. Observation of feeding behav-

iour in Janolus capensis and J. longidentatus has produced no apparent differ-

ences in mode of feeding or handling of prey. No obvious form of resource

partitioning has been observed between the two species, although it may exist.

The hottentot, Pachymetopon blochii, is the most common fish species in

environments where janolids have been observed and is known to be a

generalized omnivore of benthic organisms (Nepgen 1977). Specimens of

Bonisa nakaza, Janolus capensis, and J. longidentatus are readily ingested by

hottentots in the field and even more readily regurgitated in an unharmed state,

suggesting defensive immunity to fish predation. More detailed study of the

three species of janolids is required to determine their competitive population

dynamics, interaction, and possible resource partitioning.

ACKNOWLEDGEMENTS

I should like to thank Dr Roberta Griffiths of the University of Cape Town

for her critical review of the manuscript. Miss Patricia Cook of the British

Museum (Natural History) kindly identified the ectoproct species. The follow-

ing members of staff at the South African Museum provided valuable assistance

in the preparation of the manuscript: Mr William R. Liltved collected many

specimens and made many of the original drawings and all the final inked

figures; Mr Sidney X. Kannemeyer and Mrs Michelle van der Merwe prepared

the final photographic prints; Mrs Gail Strong typed several drafts of the

manuscript, and Mrs Marcelle Scheiner typed the final draft. To them I extend

my appreciation.

SOUTH AFRICAN JANOLIDAE 41

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ALDER, J. & Hancock, A. 1851. Monograph of the British Nudibranchiate Mollusca 5. London:

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ALDER, J. & Hancock, A. 1854a. Notice of some new species of British Nudibranchiata. Ann.

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note on some specimens from Tristan d’ Acunha. Ann. S. Afr. Mus. 25: 171-215.

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395-416.

BERGH, R. 1884. Report on the Nudibranchiata. Report of the Scientific Results of the Voyage of

the H.M.S. Challenger, Zool. 10(26): 1-154.

BERGH, R. 1888. Beitrage zur Kenntniss der Aeolidiaden, 9. Verh. zool-bot. Ges. Wien 38:

673-706.

BeErGH, R. 1904. Malacologische Untersuchungen (9)6(1). Jn: SEMPER, C., ed. Reisen im

Archipel der Philippinen. Wiesbaden: Kreidel’s Verlag.

BERGH, R. 1907. The Opisthobranchiata of South Africa. Trans. S. Afr. phil. Soc. 17(1): 1-44.

BoucueEt, P. 1975. Nudibranches nouveaux des cotes du Sénégal. Vie Milieu 25(1A): 119-131.

Burn, R. 1958. Further Victorian Opisthobranchia. J. malac. Soc. Austr. 2: 20-36.

Burn, R. & Miiier, M. 1969. A new genus, Caldukia, and an extended description of the type

species, Proctonotus? affinis Burn, 1958 (Mollusca Gastropoda: Arminacea, Antiopelli-

dae). J. malac. Soc. Austr. 12: 23-31.

CHIAJE, S. 1841. Descrizione e notomia degli animali invertebrati della Sicilia citeriore osservati

vivi negli anni 1822-1830, Napoli. 7: pls 86-173.

CocKERELL, T. & Exior, C. 1905. Notes on a collection of Californian nudibranchs. J. malac.

12(3): 31-53.

Cooper, J. 1863. On new or rare Mollusca inhabiting the coast of California. 2. Proc. Calif.

Acad. Sci. 3: 56-60.

Euror, C. 1906. Notes on some British nudibranchs. J. mar. biol. Ass. U.K. 7(3): 333-382.

Euior, C. 1907. Nudibranchs from New Zealand and the Falkland Islands. Proc. malac. Soc.

Lond. 7: 327-361.

Eviot, C. 1909. Report on the nudibranchs collected by Mr. James Hornell at Okhamandal in

Kattiawar in 1905-6. In: Report to the government of Baroda on the marine zoology of

Okhamandal in Kattiawar 1: 137-145.

Eviot, C. 1910. A monograph of the British nudibranch Mollusca 8 (suppl.). London: Ray

Society.

Franc, A. 1968. Mollusques gastéropodes et scaphopodes. Jn: Grassk, P., ed. Traité de

Zoologie 5(3): Paris: Masson et Cie.

Hoye, W. 1902. Two points in nomenclature. 1. Cypriniadea versus Cyprina. 2. The genus

Antiopa. J. Conch., Lond. 10(7): 214.

Hurst, A. 1967. The egg masses and veligers of thirty northeast Pacific opisthobranchs. Veliger

9: 255-288.

Jounson, M. & Snook, H. 1927. Seashore animals of the Pacific Coast. New York: Dover.

MACFaRLAND, F. 1966. Studies of opisthobranchiate mollusks of the Pacific coast of North

America. Mem. Calif. Acad. Sci. 6: 1-546.

MacGinitie, G. & MacGinitie, N. 1949. Natural history of marine animals. New York:

McGraw-Hill.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Marcus, E. 1955. Opisthobranchia from Brazil. Bol. Fac. Fil. Cien. Let. Univ. S. Paulo,

Brazil (Zool.) 20: 89-261.

Marcus, E. 1958. On western Atlantic opisthobranchiate gastropods. Amer. Mus. Novit. 1906:

1-82.

MiLLer, M. 1970. Two new species of the genus Caldukia Burn & Miller, 1969 (Mollusca:

Gastropoda: Opisthobranchia) from New Zealand waters. Veliger 12: 279-289.

Miter, M. 1971. A new genus and species of the nudibranch family Janolidae (Mollusca,

Gastropoda, Opisthobranchia) from New Zealand waters. J. nat. Hist. 5: 491-498.

NEPGEN, C. 1977. The biology of the hottentot, Pachymetopon blochii (Val.) and the silverfish

Argyrozona argyrozona (Val.) along the Cape south-west coast. Invest. Rep. Sea Fish.

Branch 105: 1-35.

O’DOoNOGHUE, C. 1922. Notes on the taxonomy of nudibranchiate Mollusca from the Pacific

Coast of North America. Proc. malac. Soc. Lond. 15: 133-150.

O’DonoGHuE, C. 1924. Notes on the nudibranchiate Mollusca from the Vancouver Island

Region. 4. Trans. R. Can. Inst. 15(1): 1-33.

Pruvot-FoL, A. 1954. Mollusques opisthobranches. Faune Fr. 58: 1-460.

QUATREFAGES, J. 1844. Sur les gastéropodes phlébentérés (Phlébentérata Nob.), ordre nouveau

de la classe des gastéropodes, proposé d’aprés l’examen anatomique et physiologique des

genres Zéphyrine (Zephyrina Nob.), Actéon (Acteon Oken), Actéonie (Acteonie Nob.),

Amphorine (Amphorina Nob.), Pavois (Pelta Nob.), Chalide (Chalidis Nob.). Ann. Sci.

Nat., Paris (3) Zool. 1: 129-183.

Ro.ier, R. 1970. A list of recommended nomenclatural changes for MacFarland’s ‘Studies of

Opisthobranchiate Mollusks of the Pacific Coast of North America’. Veliger 12: 371-374.

SCHMEKEL, L. 1970. Anatomie der Genitalorgane von Nudibranchiern (Gastropoda

Euthyneura). Pubbl. Staz. Zool. Napoli 38: 120-217.

STEINBERG, J. 1963. Notes on the opisthobranchs of the West Coast of North America. III.

Further nomenclatural changes in the order Nudibranchia. Veliger 6: 63-67.

THIELE, J. 1931. Handbuch der Systematischen Weichterkunde 1. Jena: Gustav Fischer.

THompson, T. & Brown, G. 1976. British opisthobranch molluscs. Synopses of the British fauna

(New Series) No. 8. London: Academic Press.

VERANY, G. 1845. Janus spinolae. Mag. Zool. (2) 7: 121-122.

ABBREVIATIONS

a anus 1 intestine

ag anal glands m muscle bands

al albumen gland me membrane gland

am ampulla mu mucous gland

b buccal mass n nephroproct

be bursa copulatrix O oral glands

Cc cerebral ganglion Pp penis

cp cerebropleural ganglion pg pedal ganglion

d digestive gland branch pl pleural ganglion

dg digestive gland rc. renopericardial canal

C eye rs receptaculum seminis

f female gland mass S stomach

g gonopores sg Salivary gland

ga _ceratal ganglion vd ___-vas deferens

gl ceratal gland

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature

(particularly Articles 22 and S51).

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Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

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

Nucula largillierti Philippi, i861: 87.

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

Note punctuation in the above example:

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TERRENCE M. GOSLINER

THE SOUTH AFRICAN JANOLIDAE

(MOLLUSCA, NUDIBRANCHIA)

WITH THE DESCRIPTION OF A NEW GENUS

AND TWO NEW SPECIES

OF THE SOUTH AFRICAN

CAPE TOWN _

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

FISCHER, P. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

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

Zool. exp. zen. 74: 627-634.

OH, 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, ee masses and larval development in Conus from the Indian Ocean.

Bull. Bingham oceanogr. Coll. 17 (4):

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

und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 86 Band

October 1981 Oktober

Part 2 Deel

DESCRIPTION OF

SOME JUVENILE HOMINID SPECIMENS

FROM SWARTKRANS, TRANSVAAL

bo Ee GRINE

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 8000

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000

OUT OF PRINT/UIT DRUK

ie ees) 53), LD, 6-59, hatin), ese, ED):

GG, tat), WO). 8, TD, My, MOGES,

GE Sy. I east, 1SCES), LO), 27, SIGE), WIG), 28, 250)

EDITOR/REDAKTRISE

Ione Rudner

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 024 9

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

DESCRIPTION OF SOME JUVENILE HOMINID SPECIMENS

FROM SWARTKRANS, TRANSVAAL

F. E. GRINE

South African Museum, Cape Town

(With 15 figures)

[MS accepted 12 May 1981]

ABSTRACT

Recent work on the hominid fossils from Swartkrans revealed several juvenile specimens

which required cleaning and description. Four such specimens, representing the gnathic and/or

dental remains of three ‘robust’ australopithecine individuals, have been prepared and

described in detail. The specimens, SK 839/852, SK 1595, and SK 2147 have added a number of

deciduous and permanent teeth to the collection from this site.

CONTENTS

PAGE

HA tKOCUCEOM rere Cisse aac wea eee ee, 43

Matcnalandidescriplions (2.2 0 9.2. bees ie 45

SPECIMENISK 839 Wao eh ac Hos ee ak eee 45

WeciduousMdentitions. 4.4) ase ee eee eee 45

Rermanent Gentittion= 2445.) eae eee 50

SPECIMIEMESIKGS 592m eke ate es vd ore PS 53

Deciduousidentition= =. >.e0 ee see ee oo 54

Renmanentdentitionas ole) oe eee 58

Specimens SK 839/SK 852 composite ............ 61

SPECIMMEMYS RGIS OSes ges Reais noe omens a ie 62

Weciduousidentitions. 1.) vo sso ee ee ae 63

Rermanentadentitions oe) eee eee 65

Specimen!" SKQ14 7 ec oy ee see A 67

DISCUSSION Hye eee Oe tae chis es cpg AIOE 69

ANCKNOWIEdSeMIENES = hiss hairs bagels Hoe eee oe 69

FRETENEMCES ee ere ere ee eames eg i ae eee 70

INTRODUCTION

Since 1948, when Robert Broom and J. T. Robinson began working at

Swartkrans, the fossiliferous deposits of this cave system have yielded an

abundance of hominid remains, including a number of juvenile specimens.

Geological and palaeontological investigations have demonstrated that the

Swartkrans deposit consists of older and younger sediment groups (Brain 1958,

1976; Cooke 1963, 1978; Brain, Vrba & Robinson 1974; Hendey 1974; Vrba

1975; Butzer 1976). Butzer (1976) and Brain (1976) have defined the older

breccias as Member 1 and the younger breccias as Member 2 of the Swartkrans

Formation.

Ann. S. Afr. Mus. 86 (2), 1981: 43-71, 15 figs.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

The first hominid specimens recovered from the Swartkrans Formation were

attributed to a new taxon, Paranthropus crassidens, by Broom (1949). The vast

majority of hominid specimens which have been found at Swartkrans (over 95 %

of individuals) have been referred to this taxon under a variety of names, viz.

Paranthropus crassidens, P. robustus, Australopithecus crassidens, A. robustus,

and A. robustus crassidens. Geological studies have shown that these australopi-

thecine remains are derived from the Member 1 sediments (Brain 1976, 1978).

The existence of. individuals of the genus Homo in the Swartkrans deposit

has been well substantiated (Broom & Robinson 1949, 1950, 1952; Robinson

1953a, 1953b, 1961; Tobias 1968, 1978; Clarke, Howell & Brain 1970; Clarke &

Howell 1972; Wallace 1972, 1975; Clarke 1977a, 1977b; Olson 1978). The first

specimen referable to Homo to be found at Swartkrans was the mandible SK 15

(Broom & Robinson 1949, 1950). The jaw was found in a pocket of brownish

breccia surrounded by the typically pink Member 1 sediments, and it was

initially thought that SK 15 was younger than the australopithecine remains

(Broom & Robinson 1949, 1950). Later, however, Robinson (1953b) consid-

ered that the brown breccia encasing the mandible was a pocket of decalcified

primary breccia and that SK 15 and the australopithecine remains were coeval.

More recent work has demonstrated that the brown breccia is, indeed, younger

than the australopithecine-bearing Member 1 sediments (Brain, Vrba & Robin-

son 1974; Brain 1976, 1978).

Three other hominid specimens, a P; (SK 18a), a proximal radius

(SK 18b), and the buccal moiety of a P, (SK 43), were recovered from within

the same pocket of Member 2 breccia as the SK 15 mandible. These four

specimens have been referred to the same taxon (Broom & Robinson 1950,

1952; Robinson 19536; Tobias & Wells 1967).

Several specimens attributed to Homo (e.g. SK 45, SK 847, SK 27, and

SK 2635) have been recovered from the australopithecine-bearing Member 1

breccia. As noted by Clarke (1977a), there is little morphological basis for

assuming that the Homo specimens from Members 1 and 2 belong to the same

specific taxon. On the other hand, it is generally accepted that the australopi-

thecine remains from Swartkrans are representive of a single, species-specific

taxon (Broom & Robinson 1952; Robinson 1956; Tobias 1967; Wallace 1972;

Clarke 1977a; Howell 1978).

Recent work on the hominid fossils from Swartkrans revealed several

juvenile specimens which required cleaning, reconstruction and description.

Four such fossils are dealt with in this paper. The specimens consist generally of

small gnathic fragments with variously preserved deciduous and permanent

teeth. The fossils are catalogued in the Transvaal Museum as SK 839, SK 852,

SK 1595, and SK 2147, and all are derived from Member 1 breccias.

Each specimen will be described as follows: (i) the data presented on the

relevant museum catalogue card, (ii) a brief statement of preservation prior to

cleaning and reconstruction, and (iii) a detailed morphological description of

the specimen after restoration.

‘JUVENILE HOMINID SPECIMENS FROM SWARTKRANS 45

All measurements recorded here were taken by the author, unless stated

otherwise. The dental measurements were taken to 0,1 mm accuracy with a

dial-equipped sliding vernier caliper.

MATERIAL AND DESCRIPTIONS

SPECIMEN SK 839

This specimen was excavated from Member 1 breccia by J. T. Robinson in

1952. The description on the catalogue card reads: ‘Fragmentary juvenile

maxilla containing some deciduous and permanent teeth some of which have

suffered considerable damage.’

Prior to restoration, this specimen consisted of isolated, incompletely

developed and slightly damaged LM' and RM’, isolated and slightly damaged

Rdm* and the distal two-thirds of the Rdm'. In addition, the badly damaged

permanent incisors, the damaged Rdi', the root of the Ldi', and a small bit of

the alveolar portion of the maxilla were preserved in a piece of breccia (Fig. 1).

Fig. 1. Stereoview of the maxillary central deciduous and central and lateral permanent

incisors of SK 839 prior to preparation. Compare with Figures 2-5. Scale in cm.

The crown of the Ldi'! was found in a box with another, unrelated, hominid

specimen from Swartkrans. The two deciduous central incisors and the four

permanent incisors were cleaned, reconstructed and strengthened by the author.

Deciduous dentition (Figs 2-4)

Maxillary central deciduous incisors

The damaged crowns and roots of both incisors are present. The crown of

the Ldi’ is only moderately well preserved; the distolinqual quadrant is missing

and the remaining lingual portion is separated from the buccal part by a narrow

crack. The buccal surface is displaced somewhat by virtue of breaks on either

side of a 0,7 mm wide vertical sliver of enamel. The crown of the Rdi’ is better

preserved, with small pieces of both the mesial and distal edges missing. The

roots of both teeth are poorly preserved; the lingual sides of both are missing

just beyond the cervical lines, and the buccal surfaces are covered by remnants

of the alveolar portion of the maxilla.

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occlusal wear is moderate in degree, with some reduction of crown height;

most of the lingual aspect comprises a lingually sloping and slightly mesiodis-

tally concave dentine exposure. The buccal edge of this surface is rather sharp

and comprises a thin wall of enamel. Lingually, the remaining enamel surface

shows slight faceting. Interproximal attrition is moderate on the mesial face,

with dentine exposed on the incisal aspect of this surface; distally, interproxi-

mal wear appears to be slightly less severe. Neither crown shows any periky-

mata, but on the buccal surface of the Ldi’ there is a rather large hypoplastic

area (MD, 1,0 mm; height, 0,6 mm) surrounded by several smaller pits, whilst

on the Rdi' a considerably smaller, single hypoplastic pit is evident.

The buccal aspect of the crown appears to have been square in outline before

wear. There is no cervical enamel prominence, and this surface is nearly flat

and disposed vertically incisocervically. Wear and damage to the incisolingual

aspect of the crowns have obliterated any morphology that might have been

present.

Fig. 2. Lingual view of the maxillary central deciduous incisors;

and occlusal view of the Rdm!' and Rdm/ of SK 839.

Scale in cm.

Robinson (1956: 122) recorded the existing mesiodistal diameters of both

crowns as 5,7 mm and noted that, because of the appreciable degree of incisal

and interproximal wear, the original dimensions ‘must have been about a

millimetre greater’. The author agrees with Robinson that about 0,5 mm of

dental material has been lost from either side of each crown. The dimensions of

these teeth recorded by the present author are as follows:

MD diameter MD diameter BL diameter

(as measured) (estimated) (estimated)

led. Le ee 5,7 mm 6,7 mm 5,2 mm

Ridit syn eee 5,8 mm 6,8 mm 5,2 mm

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS

Fig. 3. Oblique incisolingual view of the maxillary

deciduous central incisors of SK 839, showing type of

wear. Scale in cm.

Fig. 4. Buccal view of maxillary central deciduous

incisors and the attached alveolar bone of SK 839.

Scale in cm.

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

Maxillary first deciduous molar

The distal two-thirds or more of the right crown is preserved. The portion

of the crown that is missing has been broken away along a line which runs

obliquely buccolingually from a point apparently just behind the tip of the

paracone, through the protocone to the level of the lingual developmental

groove. A slight crack runs distally from the paracone, through the tip of the

metacone where it bifurcates, one limb courses distobuccally, whilst the other

divides the distal marginal ridge. The mesiobuccal root is missing, and the

lingual root has been broken away from the crown with the cervical enamel

margin. The distobuccal root is preserved for what appears to be most of its

length. It is evident that buccally the neck of the radicular system was rather

low, there being less than 1 mm of cementum exposed between the cervical

enamel margin and the point of bifurcation of the two buccal roots. The

distobuccal root, as preserved, slopes away from the crown both distally and

very slightly buccally. The root is straight and the apical end appears to taper

slightly.

It is evident that all four principal cusps—paracone, metacone, protocone

and hypocone—were present. The size of the paracone cannot be determined

accurately, but it seems to have been approximately the same size as, or

perhaps slightly smaller than, the metacone. The protocone was apparently the

largest cusp. The hypocone and metacone are almost the same size. Occlusal

wear is moderate and has produced two bevelled surfaces. Mesially, the

protocone has been worn flat and a moderately large, concave dentine island is

exposed. The protocone is worn slightly below the level of the paracone, which

appears to have shown a very small dentine exposure. The distal portion of the

occlusal surface slopes distolingually; a moderate-sized, concave dentine island

is exposed on the hypocone, while no dentine is shown on the metacone.

Interproximal attrition appears to have been slight to perhaps moderate in

degree. The distal contact facet, which measures approximately 3,1 mm bucco-

lingually by 2,1 mm in height, is ovoid in outline and flattened. It is located, for

the most part, behind the hypocone, while the corresponding facet on the dm?

is situated over the buccal half of its mesial face.

The buccal face, as preserved, shows no cervical prominence or swelling,

and it is faintly convex occlusocervically. There is no trace of a buccal groove.

The cervical margin of the lingual face is not preserved, but this surface—over

the hypocone—is more convex than the buccal surface. The lingual groove is

very shallow and short.

The protocone and paracone appear to have been separated by a very

shallow, narrow groove. The trigon basin is represented by a short and narrow

but oblique groove. The distal trigon crest is narrow but continuous. The distal

marginal ridge is thick and high, extending without any apparent interruption

from behind the tip of the metacone to the hypocone. The talon basin, or fovea

posterior, is represented by a crescent-shaped, relatively deep groove.

It is not possible to record any. crown diameters for this tooth.

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS 49

Maxillary second deciduous molar

The crown of the right tooth is preserved. It is very nearly complete; the

cervical margin has been damaged round the entire periphery save for the

buccal aspect. A large, vertical sliver of enamel has broken away from the

mesial face, and the distobuccal corner of the crown has been displaced slightly.

Several narrow cracks traverse the crown. The radicular system has been

broken away save for a short segment of the mesiobuccal root.

Viewed from the occlusal aspect, the crown is square in outline. All four

principal cusps are present and well developed. The protocone is by far the

largest cusp, followed by the metacone and the paracone, the latter being

slightly smaller. The hypocone is well developed and is approximately the same

size as the metacone. Occlusal wear is slight, and has reduced the cuspal tips to

nearly the same height. Wear is heaviest on the protocone, which shows large

enamel facets but only a tiny pit of exposed dentine. Small dentine exposures

are displayed also by the metacone and paracone. Generally, the mesial portion

of the crown has been worn more heavily than the distal part. Interproximal

contact with the dm’ appears to have been slight, while no distal contact facet is

present.

The mesial marginal ridge is thick and well developed and, though worn, it

appears to have coursed without interruption from the front of the paracone to

the protocone. Distal to the mesial marginal ridge, the fovea anterior is

represented by a short, narrow groove which is completely enclosed distally by

a short and thin but continuous epicrista. The epicrista runs parallel to the

mesial marginal ridge. The trigon basin is broad and rather deep, its floor being

represented by a triradiate groove. The distal trigon crest is moderately well

developed but is incised and thinned in its middle by a narrow but deep groove.

The distal marginal ridge is moderately thick and high and runs without

interruption from the metacone to the hypocone. It is slightly lower in its

middle, with two faint grooves on its mesial aspect; it runs continuously, high

up on to the metacone. The talon basin is represented by a deep, broad groove

which runs obliquely between the hypocone and protocone to incise the lingual

surface of the crown.

The buccal surface is rather flat and vertically disposed over the upper

third of its height, and is slightly swollen and convex over the cervical

two-thirds. There is no distinct cervical enamel prominence, and the cervical

enamel line courses, for a short distance, towards the bifurcation of the two

buccal roots. The buccal groove is rather weakly expressed; it is broad but very

shallow towards its occlusal end, and courses cervically for less than half of the

height of the crown where it terminates gradually. A number of very small

hypoplastic pits cover the buccal face around both the occlusal and cervical

extremities of the buccal groove.

The lingual surface is more convex than the buccal, and curves in a

continuous arc from the occlusal margin to the cervical line. The cervical

enamel margin is missing, but it appears that no pronounced prominence was

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

present. The lingual groove is deep and broad over most of its course, but

terminates gradually approximately half-way towards the cervical margin.

There is no trace of a Carabelli trait. The protocone is ringed on its mesial and

lingual aspects by a thin band of numerous, tiny, hypoplastic pits. This band is

situated near the occlusal surface.

The radicular system is represented solely by a short segment of the

mesiobuccal root. In so far as it is preserved, it courses slightly mesialward and

is considerably broader in its buccolingual than in its mesiodistal dimension. It

appears that the radicular neck was low.

Robinson (1956: 128) recorded the mesiodistal and buccolingual diameters

of this crown as 10,5 mm and 11,4 mm respectively. The measurements

obtained by the present author are as follows:

MD diameter MDdiameter BL diameter

(as measured) (estimated) (as measured)

RGIS cet oe eee 10,7 mm 10,8 mm 11,5 mm

Permanent dentition (Fig. 5)

Maxillary central permanent incisors

The left and right maxillary central incisors are represented. Both teeth

consist of rather badly damaged crowns only. They are unerupted. A root is not

present.

The crowns were badly crushed and broken in preservation, and both have

been reconstructed from several different pieces. The left crown is crushed and

the central and distal regions of the lingual surface have been displaced. The

mesial, buccal and distal surfaces have suffered considerable loss of enamel

about the cervical margin. The mesiobuccal quadrant of the left crown is

missing, and the mesial end of the lingual surface has suffered slight crushing.

The mesial end of the incisal edge is slightly rounded, while the distal

extremity of this edge shows a broad curvature. The incisal edge comprises

large mesial and distal mammelons and a considerably smaller and lower

median mammelon. Viewed from the buccal aspect, the crown is judged to

have had a tapered outline, the cervical region having been narrower than the

incisal portion. The buccal surface is gently convex incisocervically. Neither

tooth exhibits either perikymatous or hypoplastic enamel.

Lingually, there is a moderate cervical enamel swelling which appears to

have been symmetrically disposed. The mesial marginal ridge is much more

pronounced, and a relatively narrow, low median ridge projects vertically from

the cervical enamel prominence towards the incisal edge. The lingual surface is

slightly concave over the upper half of the crown, both incisocervically and

mesiodistally.

MD diameter MDdiameter BL diameter

(as measured) (corrected) (corrected)

TEL | ee oN sie 8,2 mm 8,6 mm =

Ribaie. once 8,6 mm 8,6 mm 6,8 mm

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS al

Maxillary lateral permanent incisors

The crown of the right tooth is well preserved, with only slight damage to

parts of the cervical enamel margin. The crown of the left tooth is severely

crushed and distorted and, because the right crown is so well preserved, it was

deemed unnecessary to attempt a reconstruction of its antimere.

The tooth is unerupted. There is no root present. The crown appears to be

completely developed.

The mesial end of the incisal edge is gently rounded, while the distal

extremity is slightly lower than the mesial and is more broadly rounded. The

incisal edge is gently rounded and rather smooth; there is no mammelon

development.

Viewed from the buccal aspect, the crown has a tapered outline, with the

cervical region narrower than the incisal portion. The buccal surface is slightly

convex incisocervically, with a faint, broad, flattening in the centre of this

surface over the upper half. This area is bounded both mesially and distally by

faint ridges that course from the cervical region to the respective corners of the

incisal edge. The distal side of this face displays several faint perikymatous

ridges; there is no evidence of hypoplastic enamel.

Lingually, the cervical region shows a slight swelling. The mesial marginal

ridge is slightly developed, the distal marginal ridge is moderately well deve-

loped and there is no median ridge present. The mesial and distal marginal

ridges converge cervically to blend into the cervical swelling. The lingual

surface is slightly concave, especially mesiodistally.

MD diameter BL diameter Height

(as measured) (as measured) (as measured)

Rug hee). 6,0 mm 5,5 mm 8,7 mm

Maxillary first permanent molars

The unerupted crowns of both the left and right maxillary first permanent

molars are present. Both crowns have suffered from damage to the cervical

enamel; generally, the right tooth is better preserved. The buccal side of the

left crown is separated and displaced from the rest of the tooth by a crack

which runs through the tips of the paracone and metacone. Nothing of the

radicular system is present.

The two crowns are nearly identical in morphological detail (Fig. 5).

Viewed from the occlusal aspect, the crown is nearly square in outline, with all

four principal cusps present and well developed. The protocone is the largest

cusp, followed by the paracone and metacone, which are nearly equal in size.

The hypocone is the smallest cusp. The mesial marginal ridge is moderately

thick; it is higher and thicker near the protocone where a moderate-sized

protoconule is present. The protoconule is delineated on either side by shallow

grooves. A short epicrista projects from the mesial end of the paracone and,

although this crest is delineated on either side by deep, narrow grooves, there is

SZ. ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Lingual view of the permanent incisors and occlusal view of

the first permanent molars of SK 839. Scale in cm.

no anterior fovea present. The trigon basin is deep. The distal trigon crest is

moderately thick, but is incised midway between the protocone and metacone

by a relatively deep pit which is continuous with the deep, but narrow, groove

between the protocone and hypocone. On the lingual surface this groove

continues vertically for approximately half of the crown height, where it ends

abruptly.

The lingual surface projects from the occlusal to the cervical margin; the

cervical half of this face is slightly convex. The mesiolingual corner of the

protocone presents two short, vertical grooves which are separated by approxi-

mately 2,7 mm of enamel. However, there is no alteration of the general

curvature of this face. The Carabelli feature is represented by grooves. This

feature is only slightly expressed on the left crown.

The buccal surface is less expanded than the lingual and is only faintly

convex occlusocervically. The buccal groove is vertically deep and narrow and

extends only over the upper third of the crown. It ends abruptly. Neither the

buccal nor the lingual surface shows any indication of hypoplastic enamel.

Robinson (1956: 81) considered these crowns to be incompletely developed

and recorded estimated, complete mesiodistal and buccolingual dimensions of

13,2 mm and 13,9 mm respectively. The present author agrees with Robinson

that the crowns are immature, but he is unable to judge accurately how much,

if any, additional increase in size would have been attained. The measurements

of the crowns as recorded are as follows:

MD diameter BL diameter

(as measured) (as measured)

|)" eaten os il eR pc 12,5 mm ==

RIVER ire el 12,8 mm 14,0 mm

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS Se)

SPECIMEN SK 852

This specimen was excavated from Member 1 breccia by J. T. Robinson in

1952. The description on the catalogue card reads: ‘Poor specimen of juvenile

mandible with left dm, (broken); dm,, moderately worn; right dm, appreciably

worn; dm, moderately worn and incomplete; and fragment of M, erupting.’

Prior to cleaning and restoration, this specimen consisted of an isolated

and nearly complete Rd, as well as the isolated, complete crowns of the LI,, LI,

and LC. The major part of this specimen consisted of the poorly preserved

mandibular corpora with the roots of the Ldm,, the damaged Rdm,, Ldm,,

Rdm, and the metaconid of the RM,, and the matrix-covered crown of the LM,

(Fig. 6). The mandibular corpora were preserved in misalignment, and wedged

Fig. 6. Stereoview of the mandibular corpora and dentition of SK 852 prior to restoration.

Compare with Figure 8. Scale in cm.

between them was a large piece of unidentified bone. The region of the RM,

was covered in plaster. The two corpora were removed from the matrix and

cleaned. In this process the hypoconid and hypoconulid of the RM, were

discovered beneath the plaster; this piece of crown was reconstructed in correct

anatomical position. The corpora and their contained teeth were cleaned and,

where necessary, strengthened with plaster.

The mandibular corpora are poorly preserved, somewhat distorted and

very incomplete. Meaningful descriptions or measurements for either of these

are not possible (see Figs 8 and 10).

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

Deciduous dentition (Figs 7-9)

Mandibular deciduous canine

The right tooth only is present. It is reasonably well preserved with rather

severe damage to the mesial portion of the crown, but the root is nearly

complete with only the apical tip missing.

Enamel from the mesial and mesiolingual parts of the crown has been

broken away. The tip is damaged also and a moderately wide vertical crack

courses across the middle of the buccal surface.

Viewed from the buccal aspect, the crown is rather ‘mitten-shaped’ with a

large primary cusp situated mesially and a small distal cuspulid. The buccal

surface is faintly convex incisocervically over the cervical third of the crown’s

height, and a cervical prominence is not present. The distal cuspulid is worn,

but there remains the trace of a short, faint vertical groove between it and the

main cusp.

Fig. 7. Stereoview of the lingual aspect of the Rd. of SK 852.

Scale in cm.

Lingually, there is a slight cervical prominence which is skewed distally.

There are no tubercles supported by this prominence. The cervical enamel line

courses downwards from mesial to distal. The mesial third of this surface has

been damaged. A moderately deep, triangular-shaped groove separates the

main cusp from the distal cuspulid. A moderate enamel ridge courses from the

tip of the distal cuspulid downwards to blend into the cervical enamel pro-

minence.

Occlusal wear is slight. A well-developed enamel facet is present along the

distal slope of the main cusp and on the tip of the distal cuspulid. This facet is

narrow but highly polished, and it is continuous along the main cusp to a

horizontal disposition on the distal cuspulid. Dentine is not exposed. Interprox-

imal attrition with the dm, was mild, with only a small, slightly flattened,

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS 5)5)

circular contact facet (0,7 mm diameter) near the top of the distal cuspulid. The

dimensions of the crown are as follows:

MD diameter MD diameter BL diameter Height

(as measured) (corrected) (as measured) (estimated)

RG La 5,7 mm 5,9 mm 4,6 mm 6,5 mm

The root of this tooth is single. It has a flattened ovoid outline in

cross-section, with the longer axis (5,1 mm) running from mesiobuccal to

distolingual and the shorter axis (3,8 mm) running from mesiolingual to disto-

buccal. The root is rather straight and long. It is estimated that the original

length was approximately 13,0 mm (measured length, 11,4 mm).

Mandibular first deciduous molars

The obliquely shorn-off roots of the left tooth and the somewhat damaged

crown and roots of the right tooth are present. The right crown has suffered

enamel loss along the entire distal surface, and the occlusal surface has a wide

crack over the top of the hypoconid and metaconid. Two narrow, vertical

cracks course across the buccal surfaces of the protoconid and hypoconid.

Viewed from the occlusal aspect, the crown has a somewhat irregular

rectangular outline. The mesial end is slightly narrowed and projects anteriorly,

while the distal end of the crown is broad and appears to have been flattened.

The protoconid, hypoconid, metaconid, and entoconid are preserved. It

appears that the hypoconulid was absent, or if it was present originally it is

evident that it would have been very small. Wear and damage do not permit an

accurate assessment of the relative cuspal sizes.

Occlusal wear is moderate, with considerable reduction and flattening of

cuspal height. The occlusal surface has been worn so that a broad, rather

flattened plane has been produced; this plane slopes slightly downward distally

from the tips of the metaconid and protoconid. It appears that a small island of

dentine was exposed on the top of the protoconid, while the metaconid shows a

somewhat larger exposure. The top of the hypoconid is damaged, but it is

evident that at least a moderately large dentine island was present. The

entoconid is covered by a large, concave dentine basin. A second facet is

present in front of the protoconid tip. This facet slopes downward mesially as a

well-defined, polished enamel surface along the mesial marginal ridge.

Interproximal wear is judged to have been mild both mesially, as discussed

above, and distally. Although the distal surface of the dm, is missing, the

mesial surface of the dm, presents a moderate-sized, rectangular and somewhat

flattened contact facet. On the Ldm, this facet measures 3,5 mm in breadth and

2,3 mm in height.

The mesial marginal ridge is relatively thick and courses round from in

front of the protoconid to the mesial end of the metaconid. It is thicker buccally

than lingually, where, mesial to the metaconid, it is considerably thinner.

Nevertheless, the mesial marginal ridge is continuous around the front of the

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

tooth; it is not incised. Though the region is worn, it appears that a small

plesioconulid might have been present mesial of the tip of the protoconid. The

fovea anterior is represented by a short, relatively deep, transverse groove

which is enclosed completely by the mesial marginal ridge and a continuous,

high and moderately thick crest between the metaconid and protoconid. A

broad contact is present between the metaconid and hypoconid, and the buccal

groove is more mesial than the lingual groove.

The lingual surface is slightly convex occlusocervically with a faint cervical

enamel prominence present. The lingual groove is faint and fades imperceptibly

approximately half-way down this face.

The buccal surface over the hypoconid is rather bulbous in appearance and

is moderately to markedly convex occlusocervically. A slight cervical enamel

prominence is present. This face of the protoconid is convex over the cervical

half, the occlusal half is somewhat flattened and slopes outwards cervically from

the protoconid tip. A faintly developed tuberculum molare is present; the

cervical portion of the protoconid surface projects slightly laterally and the

cervical enamel line dips below the level of this margin on the hypoconid. The

buccal groove is rather narrow, but relatively deep. It courses vertically to end

abruptly about half-way down this surface.

A small area of hypoplastic enamel is present on the buccal surface of the

protoconid, just mesial to the buccal groove.

The principal dimensions of the crown are as follows:

MD diameter MD diameter BLtrigonid — BL talonid

(as measured) (corrected) (as measured) (as measured)

RGN eek) 4 ode 9,4 mm 10,0 mm 73) shoe 8,1 mm

The radicular system comprises broad mesial and distal root plates and a

very low neck. The two plates are directed almost vertically downward,

expanding buccolingually towards their apical ends, and each has two separate

radicular canals. The apical third of each root plate is bifid.

The apical ends of the mesial and distal radiculae of the left tooth are

separated on the buccal side by some 7,3 mm. The mesial root plate of the left

tooth measures approximately 6,5 mm buccolingually at the cervical margin

and 9,2 mm at the apical end. The distance from the cervical line to the buccal

apex of the mesial plate is about 8,8 mm, and the lingual side of the plate

measures some 9,3 mm in length.

Mandibular second deciduous molars

The left and right teeth are present. The left crown and root system are

more complete than those of the right. The cervical half of the buccal surface

and the upper part of the root of the left tooth are broken away.

On the right tooth most of the protoconid and hypoconid have been

broken away, together with much of the distal root plate. Both crowns show

several fine cracks which course across the surfaces in various directions; the

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS 57

right tooth has been affected by this cracking more than the left. In so far as

they are preserved, the crowns are nearly identical in morphological detail.

Viewed from the occlusal aspect, the crown is rectangular in outline. The

five principal cusps are present and well developed. The metaconid is by far the

largest cusp with the hypoconid, protoconid and entoconid nearly equal in size.

The hypoconulid is, by a slim margin, the smallest cusp. In general, the cusps

are rather bulbous in appearance, with narrow and nearly vertically sided

grooves separating them.

Occlusal wear is slight, with all cusps showing enamel wear, though this is

slightest on the entoconid. The other cusps have been reduced in height

somewhat and they show flattened occlusal surfaces. The buccal cusps have

been worn slightly more heavily than the lingual cusps. The tops of the

protoconid, hypoconid, and hypoconulid are worn to a nearly flat, horizontal

plane. Except for a small pit of dentine on the protoconid, there is no dentine

exposure despite the reduction in cuspal height. Interproximal attrition with the

dm,, as noted above, is slight, and there is no contact distally with the M,.

The mesial marginal ridge of the dm, is very thick and, though worn, it

appears to have been relatively high. It continues uninterrupted from in front

of the metaconid across to the protoconid. Behind this ridge the fovea anterior

is represented by an H-shaped groove. The mesial transverse groove is narrow

and relatively shallow, and it is incompletely enclosed behind by short, access-

ory transverse crests from the metaconid and protoconid. A short, narrow

longitudinal groove separates these two crests. The distal transverse groove is

somewhat longer, broader and deeper than the mesial groove, and it is

completely enclosed behind by a very thick, continuous ridge between the back

of the protoconid and the principal crest of the metaconid. The metaconid

displays a broad contact with the hypoconid; the two buccal grooves, which

surround the hypoconid, and the lingual groove, between the metaconid and

entoconid, are arranged in a symmetrical Y-shaped pattern. The mesiobuccal

developmental groove is thus situated mesiad of the level of the lingual groove.

The distal marginal ridge is relatively thick and is uninterrupted in its course.

This ridge is slightly worn but it is apparent that it supported a tiny accessory

cuspulid (the tuberculum sextum) which is represented solely by a faint convex-

ity of the distal crown surface. The cuspulid, despite its small size on the left

tooth, is even smaller and more weakly expressed on the right crown. The

fovea posterior is represented by a relatively deep but narrow crescent-shaped

groove which is continuous with the talonid basin via a deep, narrow groove

between the hypoconulid and entoconid.

The buccal surface has suffered damage. However, it is evident that the

mesiobuccal groove was better developed than the distobuccal groove. The

distal groove is relatively deep but narrow, and extends vertically for about

one-third of the crown height where it terminates rather abruptly.

The lingual surface is somewhat inflated in appearance. It is slightly convex

occlusocervically with a slight cervical enamel prominence at the base of the

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

metaconid and a rather marked prominence at the base of the entoconid. The

enamel line continues round underneath the crown for a short distance between

the mesial and distal radiculae. The lingual groove is of moderate depth but

narrow, and continues vertically to the cervical margin. Over the lower third of

its course it is bifid, with a faint accessory groove running parallel to it mesially.

The distal half of the metaconid surface displays a number of minute hypoplas-

tic pits, and these are more numerous and marked near the lingual groove.

MD diameter BL trigonid BL talonid

(as measured) (as measured) (as measured)

Rai. 8h es 12,2 mm 9,7+ mm 9,.8+ mm

RRC ier aces 12,4 mm _ —

The buccolingual diameters recorded here are minimum values; because of

the damage the buccal surface has suffered, it is not possible to estimate

accurately the original dimensions of the trigonid and talonid portions of the

crown.

The root system of the second deciduous molar comprises broad mesial

and distal radicular plates and a very low neck. Because of damage to the

cervical region, the breadth of the plates cannot be measured accurately. The

plates appear to diverge slightly as they course downward. The length of the

buccal side of the mesial plate of the left tooth measures at least 10,5 mm.

Permanent dentition (Figs 8-9)

Mandibular central permanent incisor

The isolated crown of the left central incisor is present. The crown is

complete and well preserved. A very short segment of the developing root is

represented on the mesial and distal aspects of the tooth. The tooth had not yet

erupted at the time of death.

The mesial and distal corners of the incisal edge are slightly rounded. A

large, high mammelon is supported at either extremity, with a much lower

tubercle between the mesial and distal mammelons.

Viewed from the buccal aspect, the crown has a high, tapered outline; the

mesiodistal diameter is considerably greater incisally than cervically. The buccal

surface is slightly convex incisocervically and a cervical enamel prominence is

not present. Numerous faint, horizontal, perikymatous lines cover this face

entirely.

Lingually, a moderate basal prominence is present. A tubercular structure

is not supported by this swollen base. The basal prominence is symmetrically

orientated. The mesial marginal ridge is thin and very faintly expressed; the

distal marginal ridge is slightly better developed. The lingual surface is

flattened.

MD diameter BL diameter Height

(as measured) (as measured) (as measured)

Bee oeeeut 5,1 mm 5,7 mm 10,0 mm

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS 59

Mandibular lateral permanent incisor

The isolated crown oi the leit tooth is present. The crown is well preserved

and nearly complete, with only slight enamel loss to the buccal cervical

and somewhat more severe damage to the base of the lingual side. The

developing root is represented by short segments on the mesial and distal

aspects of the tooth. The tooth had not yet erupted at the time of death.

The crown is somewhat more robust in appearance than that of the central

incisor. The mesial and distal comers of the incisal edge appear rather

angulated with the mammelons at these extremities. The incisal edge is, mn

general, horizontal, and it supports five small mammelons.

Viewed from the buccal aspect, the crown is somewhat rectangular

outline. The mesiodistal breadth across the incisal portion is greater than that

Fis. 9. Lingual view of the LC, LL, LI, and Rd, (from left to nght) of SK 852.

Scale m cm.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

across the cervical region of the crown, but the relative difference between

these measurements is not so marked as in the central incisor. The buccal

surface is shghtly convex incisocervically. Very faint perikymatous lines and a

broad but shallow groove are present on this face.

Although the cervical extremity of the lingual surface has been damaged, it

is evident that a shght to moderate basal prominence was present. The basal

swelling does not show tubercles. The mesial marginal ridge is slightly devel-

oped; the distal marginal ridge is more faintly expressed than the mesial.

Between these mdges, the lingual surface is faintly concave both mesiodistally

and incisocervically.

MD diameter BL diameter Height

(as measured) (corrected) (as measured)

| 3) SER es See nr 5,6 mm 6.4 mm 9.9 mm

Mandibular permanent canine

The isolated crown of the left mandibular canine is well preserved and is

nearly complete. The lingual cervical enamel margin has suffered only shghtly

from damage. Mesially, a short, thin sheet of the developing root is present,

but this is not represented on any other part of the tooth. The tooth had not yet

erupted at the time of death.

Viewed from the buccal aspect, the crown is nearly square in appearance;

the mesial corner of the tip is shghtly rounded, while the distal portion slopes

downward for about half of the crown’s height from just behind the centre of

the crown. The tip of the tooth is obtuse; mesially this surface is nearly

horizontal. The buccal surtace is slightly convex occlusocervically as well as

mesiodistally. No evidence of enamel pathology is shown on this face.

Lingually, the cervical prominence is moderate and symmetrical. Lingual

tubercules are not present. The mesial marginal mdge is faintly expressed; it

appears as a low, thin band coursing round from the occlusal surface of the

crown for approximately half of the crown height, where it blends impercept-

ibly with the basal swelling. The distal marginal mdge is moderate; it originates

occlusally approximately in the middle of the distal slope of the crown. This

ridge becomes thickened towards the cervical prominence. A thin, low, vertical

enamel ridge is situated mesiad of the distal marginal mdge, and this vertical

crest originates at the cervical edge and continues cervically for just less than

half of the crown height. The ‘median’ and distal marginal mdges are separated

by a depression which is broader and relatively shallow occlusally, but which

becomes increasingly narrower and deeper as it passes cervically, where it

partially separates the distal mdge from the cervical swelling. The lingual

surface is, for the most part, rather flat.

MD diameter BL diameter Height

(as measured) (as measured) (as measured)

LC 7,3 mm 8.1 mm 9.0 mm

ID SPSOMENS FROM SWARTIERANS 61

ees li ss stwated m &S opt

and_ except for the occlusal surface. oaly the distal surface and the posterior

half of the buccal face are exposed. A narrow crack courses across the tp of the

hhypocomsuial and conimucs veriically over the desta] surface: sheht enamel

dsplaccment has ocomred along tims cack. The root & represented by 2

relatively shori shoei below the cervical marem. and @ s&s evident that the mesial

amd dssial rooi plaics had begun development at ai the time of death. The mght

tooth i represented by a portion of the metacomd and a second picce of the

the nehi Gown iS misses.

Viewed irom the occlusal aspect. the Gown appears to have a rectansular

outime. All five prmopal cusps ate presemi and are well developed The

mctacomd scoms 10 be the largest cusp. Fe i svc by

the protocomd. entocond, hypocomsd. and hypoconulad.

— io have

been moderately well developed. thick and comimuous. Distal to 1, the fowea

amfcnmor i represenicad by a relatively shori_ bul Geep_ tamsverse sroove. The

fowea SS dramed disially by a very narrow, but deep, stoove between the

protocom”d and metacom. An “accessory amiemor fowea, 2s presented by the

dm... t noi shown by tus tooth The prmopal cesi of the metacomd has a

shorn. drstally deflected mdse (a Geiiecime wonkle) which contacts the hypoco-

ma. Dhe meiacomd s more Gsially cloneaie than the protocomd. The imezual

developmental sroove is situated Gistad to the mesiobuccal sroove—and thus

the pumiary occlusal fissure paiiem ss symmetmally Y-shaped. The dsl

maremal midec rs represenied by a small cospuhd (ihe tobercolum sexium)

Winch ss demarcated from the hypoconulhd and eniocomd by shallow. narrow

grooves. The fovea posicnor ts m the fomm of a small bat relatively Geep pit A

Ralrow sroowe beiween the cntocomd and hypocomulkd commects the fovea

postenor with the disiobuoccal sroove.

The postenor part of the buccal surface ss shehily comvex occlesocerwacally-

amd the dssiobuccal sroove s deep bat short. Ii courses vertically for siehily

more than a third of the cown heieht, where i ends m a pi.

The Gamensions of this cown cannot be deiemmmed accurately.

sPnomens SK 8§39/SK 852 comPosiiz

The niehi first and sccond mauillary Geadsous molars of SK $59 and the

aie st. ne ee

lanonships beiween bla ts fcat

eked ac ivcry mood (Fie: 10). and they are very smmular

m them sencral state of wear. Casts were made of the upper and lower scoond

by Gnine (1977). The occlusal surfaces of the casis were exammed by scammme

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Lingual view of the occlusal relationship between the Rdm?

of SK 839 and the Rdm, of SK 852. Scale in cm.

electron microscopy. The types, degree, and orientation of the abrasive scrat-

ches on the two teeth were found to be very similar.

Also, the permanent teeth of the two specimens display degrees of calci-

fication that are compatible with their having belonged to the same individual.

The states of preservation of these fossils, for example the manganese staining

patterns of the teeth, are also similar.

These findings support Wallace’s (1972, 1973, 1978) suggestion that SK 839

and SK 852 represent the maxillary and mandibular remains of a single juvenile

individual.

SPECIMEN SK 1595

This specimen was recovered from the Member 1 breccia during the

1948-52 operations at Swartkrans. It was discovered in 1966 amongst a number

of supposedly non-hominid faunal remains by C. K. Brain, who performed

some preliminary preparation on it. The description on the catalogue card

reads, ‘Fragment of maxilla with a slightly worn M' left and an erupting

incisor?’

Prior to cleaning and restoration, this specimen consisted of a damaged left

upper molar, the tip of a permanent lateral incisor and a small piece of what

appeared to be the incisal edge of an incisor (Fig. 11). Also, a narrow, thin

sheet of maxillary alveolar bone was present round the incisors, and at the back

of the specimen the outline of what appeared to be part of an upper molar

crown was visible in the matrix.

Preparation revealed a very badly damaged and fragmented LI' (Figs

12-13), much of the crown of the LI? (Fig. 12), the mesial part of what is here

considered to be the LM! (Fig. 14), and the distal moiety of what is here

believed to be a slightly worn Ldm? (Fig. 14).

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS 63

Fig. 11. Stereoview of SK 1595 prior to preparation. Compare with Figures 12-14.

Scale in cm.

= all

Fig. 12. Lingual view of LI’ and LI* of SK 1595. Scale in cm.

Deciduous dentition (Fig. 14)

Maxillary second deciduous molar

The distal half or more of the crown of the left second deciduous molar is

present. The mesial portion of the crown has been broken away along an

uneven transverse line which courses through the middle of the protocone

lingually, across the distal edge of the paracone buccally. Nothing of the

radicular system remains.

It is evident that all four principal cusps were present originally, and it is

probable that all were well developed. The metacone and hypocone are both

well developed and they are nearly equal in size.

Occlusal wear is slight on the preserved part of the crown. The protocone

is the most heavily worn cusp; it has been reduced in height more than the two

distal cusps, and it shows a large, faintly convex enamel facet. The hypocone

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Buccal view of LI’ of SK 1595.

Scale in cm.

Fig. 14. Occlusal view of the Ldm? and the mesial

portion of the LM’ of SK 1595. Scale in cm.

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS 65

displays a large, flattened enamel facet which slopes downwards mesiobuccally.

The metacone shows the lightest wear of the three cusps, with only a small

enamel facet. Dentine is not exposed. An interproximal attrition facet is not

present on the distal surface of the crown.

Occlusally, the distal trigon crest is moderately thick but it is incised on its

mesial and distal surfaces by a shallow groove. The distal marginal ridge is thick

and complete. It is somewhat lower in its middle but is continuous high on to

the distal aspect of the metacone. Four small cuspules are supported by this

ridge; each cuspule is only faintly demarcated. The talon basin is relatively

large and deep. A deep, narrow, longitudinal groove drains this basin near the

middle of its mesial aspect, and this groove is continuous with the deep, narrow

lingual groove between the protocone and hypocone.

The lingual surface over the hypocone is moderately convex occlusocervi-

cally and, although the cervical margin is damaged, it appears that a cervical

enamel prominence was present. The lingual groove is relatively deep but

narrow, and it courses vertically to the cervical margin. Approximately two-

thirds of the way down the crown the groove is interrupted by a narrow

(1,5 mm thick), horizontal enamel band.

Buccally, the surface is more swollen in appearance than the lingual. The

face of the metacone is moderately convex occlusocervically. The buccal groove

is rather deep and somewhat broader than the lingual. It continues vertically

for about half of the crown height, where it ends abruptly in a deep pit. The pit

is surrounded buccally by a thin, continuous enamel crest from the paratone to

the metacone. Near the distal side of the metacone a short, shallow and

narrow, vertical groove courses from the occlusal margin. This groove partially

demarcates the most buccal of the distal cuspule (= distostyle) from the main

body of the metacone. A few minute hypoplastic pits are shown on the

mesiobuccal surface of the metacone, near the occlusal margin of this surface.

Some hypoplastic mottling is present also on the distal surface of the crown.

The mesiodistal diameter of this crown is estimated to have been between

11,2 and 11,4 mm. The buccolingual diameter measured across the distal parts

of paracone and protocone is some 12,5 mm, the original dimension is judged

to have been perhaps another 0,2 mm greater.

MD diameter BL diameter BL diameter

(estimated) (as measured) (estimated)

WdmMe res fa es. 11,3 mm 12,5 mm 12,7 mm

Permanent dentition (Figs 12-14)

Maxillary central permanent incisor

The incomplete, isolated crown of the left maxillary incisor is represented.

The crown, as preserved, was reconstructed from seven different fragments.

Most of the lingual surface is missing, save for the mesial marginal ridge and a

narrow bit of the incisodistal edge. The mesial surface is nearly complete, but

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

the cervical region is damaged. The buccal face is only slightly more complete

than the lingual, where the mesial and incisal parts are present.

The crown is unworn and probably had not yet erupted—at least it had not

yet reached occlusion—at the time of death.

The mesial corner of the incisal edge is squared off, with the incisal and

mesial edges meeting at just slightly more than 90°. The distal corner is

considerably more rounded than the mesial. The incisal edge appears gently

curved mesiodistally. This edge is faintly scalloped, and the mesial part sup-

ports two broad, low mammelons.

Viewed from the buccal aspect, the crown appears to have been rather

square in outline; its mesiodistal diameter is judged to have been only slightly

less cervically than incisally. The buccal face is slightly convex incisocervically.

Fine perikymatous lines are visible over much of the preserved surface.

Lingually, the mesial marginal ridge is only slightly developed. It is

extremely thin and low incisally, and expands somewhat as it courses cervically.

MD diameter BL diameter Height

(as measured)

|S) a See Poet oes 9.0 mm = —

Maxillary lateral permanent incisor

The left maxillary lateral incisor is represented by an isolated, somewhat

damaged crown. The cervical region round the entire periphery of the crown

has suffered enamel loss, this being heaviest distobuccally and lingually. The

crown appears to be fully, or nearly fully, developed, and it is relatively small,

especially when compared to the I’.

The tip of the crown is surmounted by two small mammelon-like tubercles.

Mesial to the tip, the occlusal edge courses only slightly downward to the

mesial corner, which is rather strongly curved. The edge distal to the tubercles

slopes cervically much more strongly, and the distal ‘corner’ is broadly rounded

and lower than the mesial. i

Viewed from the buccal aspect, the crown is almost square in general

outline, but with the distal occlusal ‘corner’ reduced. The buccal surface is

faintly convex occlusocervically and there is no evidence of either perikymatous

or hypoplastic enamel.

Lingually, the cervical region is damaged; the presence, or otherwise, of

the basal prominence cannot be determined. There is no evidence of the

existence of lingual basal tubercles. Both the mesial and distal marginal ridges

are faintly developed; they course round the periphery of the lingual face from

the tip as thin, low enamel ridges. Towards the cervical margin, however, these

ridges become slightly more prominent. The lingual surface is flat.

MD diameter BL diameter Height

(as measured)

[es eae Ae 6,2 mm _ a

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS 67

Maxillary first permanent molar

The left molar is represented by the damaged mesial portion of the crown.

This piece consists of much of the mesial surface, the lingual part of the

paracone, the entire mesial marginal ridge, and the mesial part of the paracone.

The preserved part of the crown is unworn and the tooth is judged to have been

unerupted at the time of death.

The mesial marginal ridge is moderately thick and high. It is continuous,

but is incised by two shallow grooves at the base of the tip of the paracone. A

small cuspule (= epiconule) is demarcated by these grooves. The crest is

expanded distally in its middle, and a small pit is situated atop it in this

position. The fovea anterior is represented by an elongate, irregular and

narrow transverse groove; it is broadest in front of the paracone. The proto-

cone and paracone are separated by a relatively deep, broad groove by which

the fovea anterior appears to be continuous with the trigon basin.

Measurements are not possible.

SPECIMEN SK 2147 (Fig. 15)

There is no catalogue card for this specimen. It was discovered in the

collection, labelled as an isolated lower premolar; it is, in fact, an isolated,

incomplete Ldm'.

Prior to preparation, the crown sat atop, and was partially covered by, a

small piece of typical ‘Pink’, or Member 1, Swartkrans breccia.

The specimen consists of the mesial two-thirds of a moderately worn

crown, with much of the mesiobuccal root present. The distal part of the crown

has been broken away along an irregular transverse line which cuts through the

distal edge of the protocone, and through the lingual extent of the metacone

where it turns sharply mesially to the posterior extent of the paracone. The

crack extends directly lingually from this point along the plane occupied by the

lingual developmental groove. Several narrow cracks traverse the protocone

and the paracone. The cervical portion of the buccal surface is cracked, with

considerable displacement of two rather large pieces of enamel. The mesial face

of the crown has suffered slight enamel loss and cracking. A narrow horizontal

crack traverses the mesial surface of the mesiobuccal root, and the tip of the

lingual part of this root has been damaged near its apex.

The occlusal outline of the crown cannot be determined. The protocone is

a very large cusp; the paracone is well developed also, but considerably smaller

than the protocone. Mesial to the paracone a large mesiostyle (= parastyle or

paraconule) is present. The mesiostyle is separated from the paracone by a

relatively deep, broad groove.

Occlusal wear is moderate. The protocone has been reduced considerably

in height, with a large, nearly flat and slightly lingually sloping enamel facet. A

large, elongate and ovoid-shaped, concave dentine exposure is present in the

middle of the protocone. The buccal side of the crown is less heavily worn; the

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

paracone, mesiostyle and mesial marginal ridge all show enamel wear. A small

island of dentine is exposed on the tip of the paracone.

Interproximal attrition, at least mesially, appears to have been moderate.

A large, slightly concave facet for the d° is present at the buccal extremity of

the mesial surface, it measures 2,8 mm buccolingually and 2,5 mm in height.

Occlusally (Fig. 15), the mesial marginal ridge is well developed. It is thick

and relatively high, and extends continuously from the mesiostyle to the

mesiolingual aspect of the protocone. The fovea anterior is represented by a

Fig. 15. Occlusal view of SK 2147 Ldm’.

Scale in cm.

relatively deep, broad groove which is continuous with the incision between the

paracone and mesiostyle. The fovea is enclosed distally by a high, moderately

thick enamel crest between the anterior part of the paracone and the proto-

cone. The trigon basin is represented by a relatively deep and broad \-shaped

groove. The tines of the A encompass part of the buccal aspect of the protocone

between this cusp and the paracone and metacone. The stem of the \ has been

damaged, but it is evident that it represented the buccal developmental groove

between the paracone and metacone.

The lingual surface of the protocone is slightly convex occlusocervically

and a slight cervical enamel prominence is shown.

The cervical portion of the buccal surface has been damaged, but this

portion was seemingly less convex than the lingual face, with a slight cervical

enamel prominence. The occlusal two-thirds of the buccal face is relatively flat.

The buccal groove is missing. A faint groove courses vertically for a short

distance from the occlusal incision between the paracone and the mesiostyle. It

is apparent that there was no tuberculum molare over the mesiocervical aspect

of the buccal surface.

JUVENILE HOMINID SPECIMENS FROM SWARTKRANS 69

MD diameter BL diameter BL diameter

(as measured) (corrected)

Lich eee see sss 9.8 mm 10,0 mm

The mesiobuccal root has a flattened, ovoid outline in cross-section; it is

expanded buccolingually and compressed mesiodistally. The long axis of the

root, in cross-section, runs slightly mesiobuccally-distolingually. The bucco-

lingual diameter measures some 5,3 mm and the mesiodistal some 3,9 mm at

the cervical margin. The root is relatively straight, and it courses rather

markedly mesially from the cervical line to its tip. It diverges buccally some-

what. This root is estimated to have been approximately 8,5 mm long.

DISCUSSION

Four specimens representing the jaws and/or teeth of some three juvenile

hominid individuals from Swartkrans have been prepared and described in

detail. The remains are all derived from Member 1 breccia. These specimens

have added a number of deciduous and permanent teeth to the already sizeable

collection from this site.

Robinson (1956) referred one of these specimens (SK 839) to Paranthro-

pus robustus. Wallace (1972) suggested that SK 839 and SK 852 represent a

single individual, and he included this composite in his hypodigm of Australopi-

thecus robustus from Swartkrans. The results of the present study support

Wallace’s suggestion that SK 839 and SK 852 belonged to a single individual.

SK 1595 was also referred to A. robustus by Wallace (1972) on the basis of the

occlusal wear pattern exhibited by the dm’.

The fourth specimen, SK 2147, was catalogued as an isolated permanent

premolar. The author considers this tooth to be a left dm’, and as such it

represents, to date, the second specimen of this tooth found at Swartkrans. The

other dm’, SK 91, has been referred by both Robinson (1956) and Wallace

(1972) to the ‘robust’ australopithecine taxon. The morphology shown by

SK 2147 is similar to that evinced by SK 91, and the dm’ of SK 2147 is also

referred to the Swartkrans ‘robust’ australopithecine taxon.

An analysis of the morphological and metrical data available for the

collection of deciduous and permanent teeth from Swartkrans is beyond the

scope of the present paper and will be presented elsewhere.

ACKNOWLEDGEMENTS

I am grateful to Drs C. K. Brain and E. S. Vrba (Transvaal Museum) for

permission to prepare and describe specimens in their care. I thank Prof. P. V.

Tobias and Dr T. D. White for reading and constructively criticizing this paper.

Mr A. R. Hughes and Mr H. Thackwray assisted with photography. This work

was supported by a grant from the Senate Research Committee, University of

the Witwatersrand.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

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6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature

(particularly Articles 22 and 51).

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

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

nov., syn. nov., etc.