279 ’ Sac ut . eee . a= & 2 oe eee Mam,
Te tek See eee ee ee ee est ee ee ee ee a en ee eed re ee oe ee ee ee te ee ree ae ae ee a a es

HARVARD UNIVERSITY

eB

LIBRARY
OF THE

Museum of Comparative Zoology

~

> ew Se

TH AFRICAN MUSEUM

VOLUME 48

ee at

ANNALS

GF THE

SOUTH AFRICAN MUSEUM

VOLUME 48

PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM
1964-1967

MUS. COMP. ZOOL.
LIBRARY

wae G1? ele lm
yi te os Gio BEC 7 19?

Twi
¥ eA Mike e polit) _ HARVARD
ia4") UNIVERSITY

| : ROTI at %
: y HAP ule » : j # rn i, ; Oe) Ae ie, hoe S ?

@ PRINTED IN SOUTH AFRICA BY’

Eliot OF CONTENTS

BARNARD, K. H.
Isopoda and Amphipoda collected by the Gough Island scientific survey (published
September 1965)
Barry, T. H.
On the epipterygoid—alisphenoid transition in Therapsida (published November
1965)
Bonk, E. L. & SINGER, R.
Hipparion from Langebaanweg, Cape Province and a revision of the genus in Africa
(published November 1965)
Boonstra, L. D.
The girdles and limbs of the Gorgonopsia of the Tapinocephalus zone (published
November 1965)
BoonstrA, L. D.
The girdles and limbs of the pristerognathid Therocephalia (published April 1964)
BoonstrA, L. D.
The Russian dinocephalian Deuterosaurus (published September 1965)
Boonstra, L. D.
The skull of Struthiocephalus kitchingi (published November 1965)
way, J. H.
A review of the family Ampharetidae (Polychaeta) (published April 1964)
Gess, F. W.
Contribution to the knowledge of the South African species of the genus Ceramius
Latreille (Hymenoptera: Masaridae) (published September 1965)
Haropine, J. P. & Situ, W. A.
Some South African fresh water Copepoda (published April 1967)
Hu. ey, P. A.
The validity of Raja rhizacanthus Regan and Raja pullopunctata Smith, based on a
study of the clasper (published November 1966)
Juss, R. A.
A new palaeoniscid fish from the Witteberg Series (Lower Carboniferous) of South
Africa (published November 1965)
KulrrEr, J. G. J.
Contributions to the knowledge of the South African species of the genus Pisidium
(Lamellibranchiata) (published April 1964)
LAWRENCE, R. F.
New cavernicolous spiders of South Africa (published April 1964)
Mixiarp, N. A. H.
Hydroids of the Vema Seamount (published November 1966)
Mi.yiarp, N. A. H.
The Hydrozoa of the south and west coasts of South Africa. Part II. The Lafoeidae,
Syntheciidae and Sertulariidae (published February 1964)
Miiarp, N. A. H.
The Hydrozoa of the south and west coasts of South Africa. Part III. The Gymno-
blastea and small families of Caluptoblastea (published June 1966)
PEnriTH, M. J.
The fishes of Tristan da Cunha, Gough Island and the Vema Seamount (published
March 1967)
PenriTH, M. J.
A new species of flatfish, Mancopsetta milfordi, from South Africa, with notes on the
genus Mancopsetta (published September 1965)
PEenriTH, M.-L.
Studies on the South African Clinidae. I. Description of a new species of Pavoclinus,
and redescription of Gynutoclinus rotundifrons (Barnard) (published September

1965)

Page

195

399

273

237

I2!I

233

251
97

219

515

497

267

523

181

2er

LIST OF CONTENTS

PenriTH, M.-L.
See TALBOT, F. H.
SINGER, R.
See Bonk, E. L.
SmirH, W. A.
See Harvine, J. P.
TALsor,(F. EH.
See WAPENAAR, M.-L.
TAxsoT, F. H. & PenritH, M.-L.
Ctenogobius cloatus Smith, 1960, a synonym of Ctenogobius saldanha (Barnard, 1927) 189
(published September 1965)
WAPENAAR, M.-L. & TALBoT, F. H.
Note on the rigidity of the pectoral fin of Makaira indica (Cuvier) (published April 167
1964)

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Bicorona Millard, 1966 (Corynidae), 440
Mentzichthys Jubb, 1965 (Palaeoniscidae), 270
Speleoderces Lawrence, 1964 (Leptonetidae), 62

Kinnosaurus Boonstra, 1964 (Pristerognathidae), 143

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to A f ~C. MUS. COMP. ZOOL

LIBRARY

MAY 29 1964

HARVARD
UNIVERSITY,

THE HYDROZOA OF THE SOUTH
AND WEST COASTS OF SOUTH AFRICA
PART II. THE LAFOEIDAE, SYNTHECIIDAE
AND SERTULARIIDAE

N. A. H. MILLARD

February 1964 Februarie
Volume 48 Band
Peru) © )\ eel

ANNALS OF THE SOUTH AFRICAN MUSEUM

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MUS. COMP. ZOOL
LIBRARY

MAY 29 1964

THE HYDROZOA OF THE SOUTH AND WEST COASTS OFARVARD

SOUTH AFRICA UNIVERSITY
PART II. THE LAFOEIDAE, SYNTHECIIDAE AND SERTULARIIDAE
By

N. A. H. MiILtarp

Koology Department, University of Cape Town
(With 16 figures in the text)

CONTENTS
PAGE
Introduction . ae peat re
DIAMOnAst sy. 8
EASE OF SPECIES Oye. G
Bam. Gafoeidae . . §.. 7
Fam. Syntheciidae . . 22
Fam. Sertulariidae . . 25
DMRIMATV tt se? sy. SY. 154.
Etereneessreya = fre 2. 55
INTRODUCTION

This paper represents the second part of a systematic account of the
hydroids of the south and west coasts of South Africa. The first part, dealing
with the Plumulariidae, appeared in these Annals, vol. 46, 1962. The scope of
the work and the source of the material were detailed in the introduction to
_ Part I, and need not be repeated here.

As before the details of the collecting stations are given in the station list
which follows, and only the catalogue numbers are quoted in the systematic
account. Occasional records which do not come from the south or west coast,
but which have been mentioned for some special reason, have been put between
brackets.

The author wishes to acknowledge the help of all those bodies or individuals
who have helped materially or financially to build up the very extensive
collection now present in the University of Cape Town, and also the South
African Museum for access to the material dredged by the s.s. Pieter Faure.
Acknowledgements are also due to the British Museum of Natural History for
_ permission to examine material housed there, and to the Munich Museum for
the loan of mounted slides.

Type specimens of new species have been deposited in the South African
Museum and have been given a Museum registered number in addition to the
University catalogue number.

The Trustees of the South African Museum acknowledge with thanks
receipt of grants from the University of Cape Town and the Council for
Scientific and Industrial Research towards the cost of publication.

I

Ann. S. Afr. Mus. 48 (1), 1964: 1-56, 16 figs.

ANNALS OF THE SOUTH AFRICAN MUSEUM

STATION LIST

A. Littoral material from Oudekraal on the west coast of the Cape Peninsula.
Position: 33°58°5'S/18°22-2’E.

Date
A 123 153-34
A 381 25.38.34
A 382 13.5.34

A 384 25.8.34
AFR. Material dredged by the government research vessel, r.s. Africana.

Date Position Depth (m.) Bottom
AFR 736 17.8.47 30°42°4’S/15°59°2’E 201 co gn S, Sh
AFR 743 21.8.47 30°2'S/15 2°E 364 gn S
AFR 835 20.11.47 ?35°9/S/19°2’E 188
AFR 866 9.1.48 34°36-8’S/19°16-4’E 38 S,R
AFR 945 19.3.48 36°25’S/21°8’E 177 S,R
AFR 1028.0 15.5.48 28°28’S/32°25:8’E 27 fS,R

B, BB. Littoral material from Lambert’s Bay on the west coast. Position:
32°5/S/18°14’E.
Date
B 105 28:7.38
B 114 29.7.98

B.1g7 || 4.7.38
BB 13 18.1.57

BMR. Bushman’s River Estuary, on sand and muddy banks. Date: September,
1950. Position: 33°41'S/26°40’E. Depth: 2-44m.
BRE. Breede River Estuary, littoral. Date: 7.7.51. Position: 34°25’'S/20°51°5’E.

CP. Littoral material from various localities on the west coast of the Cape
Peninsula.

Date Locality Position
CP 336 12.5.49 Oudekraal 33°58°5'S/18°22-2’E
CP 379 15.4.53 Sea Point 33°55°2'S/18°22-6’E

CP 650 1.2.61 Bakoven 93°57 8) lo 227 a7e
CPR. Material from various localities in the Cape Province.
Date Locality Position Depth (m.)
CPR 7 15.1.50 The Haven 32°15’S/28°57’E littoral
CPR 9 30.4.50 Glentana Strand 34.°4’S/22°20’E littoral
CPR 46 20.6.59 Umgazi Bay 31°43’S/29°26’E a |

E. Littoral material from Port Elizabeth on the south coast. Date: 9.7.36
Position: 33°56'S/25°36’E.
KNY. Knysna Estuary, on the south coast. Date: July, 1947. Position:
34.°5'S/23°4'E (average).
Depth (m.) Bottom
M

KNY 22 1-4

KNY 30 5-7 S,M
KNY 57 II‘5 R
KNY 70 2-6 S

KNY 71 7, Sh, S, M

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 3

L. Littoral material from East London, on the south coast. Date: 10.7.37.
Position: 33°1'S/27°54’E.
LAM. Dredged in Lambert’s Bay, west coast.

Date Position Depth (m.) Bottom
LAM 2 16.1.57 32°4°5/S/18°18-3’E 17 S, R
LAM 7 18.1.57 32°5'9/18°17-9’E 23 R, Sh, $
LAM 9 17.0657, 92 4:7 0/18 17-7 EB 23 S, Sh
LAM 13 19.1.57 32°4'5/18°18-1’E 18 R
LAM 14 18.1.57 32°5'5/18°17'7'E 17 5, oh, R
LAM 18 18.05.57 °32°4°6°5/18'17°8’E 17 R
LAM 23 T7057 92°4°t'S/18°18-6/E 15 S, Sh
LAM 30 19.1.57 32°5°1'S/18°17°7’E 20 R
LAM 35 19.1.57 32°5°5'9/18°17°7’E 27°5 R, Sh
LAM 40 19.1.57 32°5°5/9/18°17-6’E 28 S, Sh
LAM 41 200.57), 92° R ola 19°) 20 S, Sh
LAM 45 STs 1 eo RSG too" 2, 8 S, R
LAM 46 22.1.57 32°4°49/18°17-7’ E 23 R
LAM 59 23.1.57 32°9'S/18°18’E 16 i, ats
LIZ. Dredged in Algoa Bay, Port Elizabeth, south coast.
Date Position Depth (m.) Bottom
LIZ 2 5-4-54 33°55°7'S/25°37°2'E Or5a RE |
Rie -7 6.4.54 33°58-1’S/25°38-9’E 9 St, R
LIZ 11 6.4.54 33°57°2'S/25°38’E 9°5 R, Clay
LIZ 13 6.4.54 33°58°2’S/25°38-8’E 75 S
LIZ 16 7.4.54 33°58°4'S/25°40°5’E 14 St
LIZ 27-40 11.4.54 34°0°8'S/25°42-4’E 6 R
MB. Dredged in Mossel Bay, south coast.
Date Position Depth (m.) Bottom
MB 8-12 12.1.56 934°4°3/S/22°13-0’E 19 R
MB 15 13.1.56 34°r1:1’S/22°10-1’E 16 5
MB 24 190.50 34 Til 'S/22°9°9°E 19 R
MB 26 13.1.50° 34°11'1'S/22°10-1’E 21 S
MB 39 16.1.56 34°10°1’S/22°8-0’E 9 R
MB 47 Mot 5oy -S4- rig 3/22 10-0'E, 10 R
MB 52 7.1.50 34°11°0'5/22°9'9'E 14 R,S
MB 58 18.1.56 34°4°3’S/22°13°5’E 12°5 R
MB 59 18.1.56 34°4°1'S/22°13°9/E a." R
MB 64 18.1.56 34°4°8’S/22°13°1’E 26 - co S, Sh, R
MB 69 19.1.56 34°8-6'S/22°7:3’E 13°5 S,R
MB 72 19.1.56 34°9°1'S/22°7-2’E 12 R, S, Sh
MB 84 21.1.56 34°11-4/S/22°10-1’E 29 R
MB 88 18.1.56 34°4°8'S/22°13°1’E 26 co S, Sh, R
NAD. Dredged off Natal, east coast.
Date Position Depth (m.) Bottom
NAD 1 17.5.58 30°47°1’S/30°29°1’E 44 St
NAD 22 12.8.58 29°58’S/31°2’E 49

PP. Littoral material from Paternoster, west coast. Date: 24.9.57. Position:
32°43'S/17°55’E.
S. Littoral material from Still Bay, south coast. Date: 10.1.32. Position:

34°23'S/21°26’E.

4. ANNALS OF THE SOUTH AFRICAN MUSEUM

SAMH. Material from the South African Museum. Specimens 146-350 and
374-382 were dredged by the s.s. Pieter Faure. Their positions were given
in the original records as compass bearings off salient points on the coast,
and were probably not very accurate. These have been converted into
latitude and longitude and are given to the nearest minute.

Date Position Depth (m.) Bottom
SAMH 146 23.6.1898 South of Mossel Bay
SAMH 151-156 15.7-1898 34°8’S/22°16’E St
SAMH 171-172 11.11.1898 33°49’S/25°56’E
SAMH 173-178 19.11.1898 33°45'S/26°44’E 73-78°5 M
SAMH 181 22.12.1898 32°52’S/28°12’E
SAMH 182-192 28.12.1898 33°9’S/28°3’E 86 S, Sh, R
SAMH 204-209 7.3.1899 33°59'S/25°51'E 24-27
SAMH 213 24.3.1899 33°50'S/26°35’E gi M
SAMH 216-220 19.6.1899 34°26’S/21°42’E f{S
SAMH 228-232 20.9.1899 34°15'S/22°10°5’E M
SAMH 234-237 5-7-1900 34°27’S/20°58’E 51 Crl
SAMH 246 11.10.1900 34°8’S/22°59°5’E 73 S, Sh, Crl
SAMH 251-252 15.7.190I 33°13°5'S/27°58’E 89 brk Sh
SAMH 256-272 17.7.1901 33°7'S/27°47°5E £S
SAMH 281-282 25.7.190I1 32°50'S/28°18-5’E 86 brk Sh
SAMH 288-294 13.8.1901 32°45’S/28°26’E 53 brk Sh, St
SAMH 295 13.8.1901 32°47'S/28°28’E 82 brk Sh
SAMH go1 15.8.1901 32°42’S/28°26’E 31 R
SAMH 310-314 10.9.1901 33°54’S/26°51’E 120 brk Sh, St
SAMH 316-318 23.9.1901 34°5'S/26°34’E 115 S, Sh, bk Spks
SAMH 321 19.2.1902 34°32'S/24°27-5’E 137 S, Sh, R
SAMH 325 22.9.1904 34°12’S/22°15°5’E 51 £S
SAMH 334 4.10.1904. 34°12’S/22°15°5’E 51 £S
SAMH 348 19.10.1904 34°15°5’S/22°14’E 64 M
SAMH 350 22.8.1905 33°52’S/26°9’E M
SAMH 358-359 19.6.1914 33°55’S/18°27’E
SAMH 374-382 15.3.1899 33°47'S/26°19’E 18-29 S, Sh, St
SAMH 403 —.4.1962 28°37’S/16°25’E 10-20
SAMH 409 —.10.1962 28°37’S/16°25’E 10-20
SB. Saldanha Bay, west coast.

Date Position Depth (m.) Bottom
SB 150-168 —.9.57 33°2°5/S/18°2’E littoral R
SB 194 1.5.59 33°3°5'9/17°59°2'E 20 R,S
SB 253 22.4.62 33°3'S/17°56-6’E 35 fkhS

SCD. Dredged off the south coast.

Date Position Depth (m.) Bottom
SCD 3 18.4.58 34°30’S/24°40’E 102 R
SCD 5 19.4.58 34°15'S/25°5’E II R, Sh
SCD 29 22.6.58 33°38-6’S/26°54-7’E 56 R
SCD 33 21.5.58 35°3'S/27°56-2’E 65 S, Sh
SCD 36-37 19.5.58 32°15°2'S/28°57-7'E 49°5 R
SCD 50 18.5.58 31°38-8’S/29°34°4’E 33 R
SCD 52 20.8.58 34°1'S/25°45°5’E 46 R
SCD 56 19.8.58 33°37'S/26°56-6’E 46
SCD 60 16.8.58 33°2’S/27°56-2’E 46
SCD 61 15.8.58 32°17°7'S/28°54°5’E 49
SCD 75 16.7.59 32°33’S/28°38’E 55 ,M
SCD 79 16.7.59 32°43'S/28°28’E 58 St, Sh

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA

SCD 82
SCD 84-85
SCD 94
SCD 96
SCD 1o1
SCD 103
SCD 104
SCD 106
SCD 108
SCD 112
SCD 113
SCD 114
SCD 115
SCD 117-118
SCD 119
SCD 122
SCD 126
SCD 138
SCD 141
SCD 145
SCD 153-154
SCD 169
SCD 175
SCD 179
SCD 184
SCD 191
SCD 206
SCD 219-239
SCD 250
SCD 254
SCD 258
SCD 265
SCD 276
SCD 284
SCD 290
SCD 296-297
SCD 301
SCD 304-305
SCD 320-322
SCD 324
SCD 330
SCD 345
SCD 354

Date
17-7-59
177-59
20.7.59
20.7.59
21.7.59
22.7.59
237-59
237-59
23-7-59
20.7.59
26.11.59
26.11.59
26.11.59

14.2.60

f 14.2.60

' 14.2.60
3.6.60
28.8.60
28.8.60
28.8.60
25.11.60
24.11.60
30.11.60
24.11.60
25.11.60
29.11.60
30.11.60
29.11.60
30.11.60
16.7.61
TAs 7.0%
14.7.61
14.7.61
6.2.62
6.2.62
6.2.62
6.2.62
8.2.62
9.2.62
9.2.62
1.2.62
12.2;62
11.10.62

Position
33°9°7'S/27°54°7'E
33°3'S/27°55'E
33°55°5/9/25°51'E
34°21'S/25°41’E
34°33'S/24°1’E
35°7'S/22°15’E
34°33'S/21°28’E
34°35'S/21°10’E
34°35'S/21°11’E
33°55°5'9/25°51'E
34°24’S/21°45’E
34°29'S/21°49°5’E
34.°54°49/22°12°2’E
34°24'S/21°46’E
34°33'S/21°52’E
34°40°5'S/22°0’E
34°26°5'S/21°48’E
34°35/S/21°56’E
34°46’S/22°5’E
34°46’S/22°5’E
34°3'S/25°59’E
33°58-9/S/25°41°4’E
34.°20'S/23°31’E
33°58°9'S/25°41°4’E
34°23’S/26°1’E
34°4°3'S/23°25°8’E
34°51'S/23°41’E
34°28 /23°28-4’E
34°48'S/23°39’E
33°7°3'S/28°1E
33°53°8'S/25°42°5’E
33°48'S/25°47'E
33°53 °8'S/25°42°5’E
33°1'S/27°55E
33°4S/27°57’E
33°9'S/28°2’E
33°39'S/27°15’E
34°0'S/25°53’E
34°15/S/25°50°5’E
34°27'S/25°57'E
34°3°5'S/23°23’E
34.°16'S/22°17’E
32°8’S/29°12’E

Depth (m.)
51

TB. Material dredged from Table Bay, Cape Town.

Bt
TB 10
Pea DT
TB 12
TB a1

TRA. Material collected by commercial trawlers.

TRA 23
TRA 33
TRA 35

Date

11.2.47
11.2.47
25.10.46
11.2.4.7
15.12.57

Date
Q.11.47

Position
33°4.7°5'S/18°24°3’E
33°50°5'S/18°25°8’E
33°52'S/18°28’E
33°47°5'S/18°24’E
33°48-6’S/18°24-6’E

Position
34°49’S/20°21°5’E

20.7.49 34°55’S/21°10’E
21.1.50 34°34’S/20°50’E

Depth (m.)
19-20

27

LB
19-20

15

Depth (m.)

C. 91°5
c. gO

70

Bottom
br S, Sh
R
bk M, S
Sh
R
S

co S, brk Sh

co: 8, Sh, St
bk M,S
S, 7 Sh
gr-gn M
co S, Sh
R

kh §

kh §

bk M
co&fS
kh §

kh §

R

R
R,khS
R

co S, brk Sh

en M
kh S
S»R, M

Bottom
5, nb, St
R
S
Sh, St

St, brk Sh, S

Bottom

S,R
M, S

ANNALS OF THE SOUTH AFRICAN MUSEUM

Date Position
TRA 97 —.7.50 34°30'S/20°50’E
TRA 38 -—.7.50 34°30'S/20°56’E
TRA 42 —.7.51 34°30'S/20°55’E
TRA 56 28.11.52 34°40'S/21°35’E
TRA 59 26.11.52 34°28’S/21°45’E
TRA. 92 —.1.54 35°3’S/21°50’E
TRA 151 6.3.58 34°51'S/19°55’E
TRA 156 15.10.58 34°12’S/18°22’E
TRA 159 6.7.58 33°56'S/25°36’E

WCD. Dredged off west coast.

Date Position
WCD 1 25.2.59 34°9°8’S/18°16°5’E
WCD 7 24.3:59 34°9°3'S/18°17°5’E
WCD 12 24.3.59 34°9°4’S/18°16°5’E
WCD 18 29-4.59 33°5°6'S/17°54°5'E
WCD 30 15.12.59 34°10°5'S/18°14-3’E
WCD 34 15.12.59 34°11°2’S/18°20°2’E
WCD 56 21.9.60 32°4°6’S/18°18’E
WCD 81 15-9.49 34°5'S/18°21’E
WCD 100 2.7.61 32°5°5'S/18°17-3'E

Acryptolaria conferta (Allman, 1877)
Filellum antarcticum (Hartlaub, 1904)

Hebella furax Millard, 1957
Hebella scandens (Bale, 1888)
Hebella urceolata n. sp.

Lafoea fruticosa M. Sars, 1851

1786)

Salacia articulata (Pallas, 1766)

Salacia disjuncta n. sp.

Sertularella africana Stechow, 1919

LIST OF SPECIES

Family Lafoeidae

Depth (m.)
73

&
S$
§

DAP Ree
PLlrnMAYD

“aS
aes
Q

Bottom

POD BREW Rw
Li = ae

Scandia mutabilis (Richie, 1907)
Kygophylax africana Stechow, 1923

K.ygophylax armata (Ritchie, 1907)
Kygophylax cornucopia Millard, 1955

Kygophylax enigmatica n. sp.
Kygophylax sibogae Billard, 1918

Family Synthecitidae
Fiincksella cylindrica (Bale, 1888)
Synthecium dentigerum Jarvis, 1922

Family Sertulariidae

Amphisbetia bidens (Bale, 1884)
Amphisbetia minima (Thompson, 1879)
Amphisbetia operculata (Linn., 1758)
Crateritheca acanthostoma (Bale, 1882)
Dictyocladium coactum Stechow, 1923
Diphasia tetraglochina Billard, 1907
Dynamena cornicina McCrady, 1858
Dynamena cristoides Lamx., 1824
Dynamena quadridentata (Ell. & Sol.,

Synthectum ?elegans Allman, 1872
Synthecium hians Millard, 1957

Sertularella agulhensis n. sp.
Sertularella arbuscula (Lamx., 1816)
Sertularella capensis Millard, 1957
Sertularella congregata n. sp.
Sertularella dubia Billard, 1907
Sertularella falsa Millard, 1957
Sertularella flabellum (Allman, 1886)
Sertularella fusiformis (Hincks, 1861)
Sertularella gilchristi n. sp.

Sertularella goliathus Stechow, 1923

Sertularella mediterranea Hartlaub, 1901

Sertularella megista Stechow, 1923

Sertularella polyzonias (Linn., 1758)

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA y

Sertularella pulchra Stechow, 1923 Sertularia turbinata (Lamx., 1816)
Sertularella striata Stechow, 1923 Symplectoscyphus arboriformis (Markt.,
Sertularella xantha Stechow,: 1923 1890)

Sertularia distans (Lamx., 1816) Symplectoscyphus macrogonus (‘Treb., 1928)
Sertularia marginata (Kirch., 1864) Thyroscyphus aequalis Warren, 1908

Family Lafoeidae
Acryptolaria conferta conferta (Allman, 1877)
Fig. 1 A-C, E

Cryptolaria conferta Allman, 1877: 17, pl 12 (figs. 6-10).
Acryptolaria conferta: Totton, 1930: fig. 19a. Leloup, 1937: 29, fig. 19.

Records. West coast: AFR 736Y. South Coast: SCD 101G.

Description. Colonies reaching a maximum height of 3:9 cm. Stem fascicled
except for the terminal branches, but slender and flexible; branching in a
roughly alternate manner and roughly in one plane, but on the whole very
irregular in appearance.

Hydrotheca adnate to stem or branch for over half height; adcauline
wall approximately straight and parallel with axis of stem in lower part of
adnate section, curving outwards in upper part of adnate section and in free
section; abcauline wall usually slightly convex opposite base of adcauline wall,
curving evenly outwards beyond this; margin slightly everted. Base of adcauline
wall of hydrotheca always above level of top of adnate part of the one below.
Diameter at margin approximately 14 to 24 times that at base.

Free, solitary hydrothecae also present, arising separately from hydrorhiza
or from epizootic stolons creeping over the surface of mature stems. These
hydrothecae erect, at right angles to stolon, widening to margin which is
slightly everted, quite symmetrical or (more often) somewhat irregular in
shape.

No coppiniae.

Measurements. See under subspecies australis.

Remarks. Both these colonies are growing on the surface of worm-tubes, and
include stems in various stages of development. Certain observations on the
method of growth were thus possible.

The hydrorhiza forms a branching reticulum giving off solitary hydrothecae
and upright stems. In a young upright stem the first hydrotheca and the
hydrocaulus arise separately and side by side from the hydrorhiza (fig. 1C).
It appears thus that in the development of the colony a solitary hydrotheca
is produced first, and that this is followed by the growth of a separate tube of
the hydrorhiza past it, and in contact with it, to become the hydrocaulus of
the stem. The development of the rest of the stem proceeds by a method of

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

RR Ren Ieee ee ae BEN Se ems Weed Yay | 4 |

i
Y

Va
an,

VO

E G
F

Fic. 1. Acryptolaria conferta (Allman).
. Part of the stem from AFR 736Y, to show branching. Portions of the peripheral tubes in
position.
Solitary hydrothecae from AFR 736Y.
. Solitary hydrotheca and young stem from AFR 736Y, to show origin.
. Subsp. australis, solitary hydrothecae from SCD 175N.
E.-G. A few hydrothecae from SCD 101G, SCD 101F and SCD 175N respectively. The last
two are included in subsp. australis.

GOW >

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 9

sympodial growth. At a later stage the superficial tubes are developed as
branches from the hydrocaulus arising opposite the top of the adnate part of
the hydrothecae, and remaining in communication with it by a series of
connexions in this position. Branches arise in the same position as the accessory
tubes and are in cytoplasmic continuity with both the coenosarc of the stem
and that of the accessory tube.

The free hydrothecae which are often present on the surface of older
stems appear to belong mostly to young epizootic colonies growing on the
surface of older ones.

These solitary hydrothecae are similar to those produced by various
species of Lafoea (e.g. L. dumosa (Fleming), L. gracillima (Alder) and L. fruticosa
M. Sars) and are almost identical with them. They also resemble hydrothecae
of the creeping L. tenellula Allman, which is included by some authorities in
L. dumosa. It is evident, thus, that extreme care should be taken in assigning
such stolonic colonies to a particular species unless the branching form is
present as well. For example, the L. tenellula described by Stechow in 1925
(p. 453, fig. 23) was growing on a colony of Acryptolaria humilis Allman and is
probably the creeping form of that species.

The general appearance of the colonies described above resembles Allman’s
figure (pl. 12, fig. 6), and the shape of the hydrotheca resembles that illustrated
by Totton (fig. 19a) from Madeira and by Leloup (fig. 19B) from French
Indo-China. The measurements are in range of those quoted by most authors.

This is the first record of the species from South Africa.

Acryptolaria conferta australis Ritchie, 1911
Pig. 1D, FG

Cryptolaria conferta var. australis Ritchie, 1911: 826, pl. 84 (fig. 2), pl. 87 (fig. 1).
Acryptolaria conferta var. australis: Totton, 1930: 163, fig. 19 c-e. Ralph, 1958: 315, fig. 4 a-g.

Records. South coast: AFR 835E. SCD ror1F, 103F, 175N.

Description. Several colonies, the largest reaching a height of 11 cm. Stem
fascicled, stiff and rather woody; branching alternate and always in one plane;
branches variable in length, generally arising next to every third hydrotheca of
the stem and often rebranching, those on the same side being separated by a
distance of 3-4 mm. Branches often anastomosing.

Hydrotheca adnate to stem or branch for over half height; adcauline
wall convex throughout; abcauline wall curved gracefully outwards, the
curvature being more marked in the distal half. Hydrothecae overlapping,
with base of adcauline wall always below level of top of adnate part of the
hydrotheca below. Diameter at margin approximately 14 to 2} times that at
base.

Free hydrothecae also present in SCD 175N, arising from epizootic
stolons.

No coppiniae.

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm.)

A. conferta australis

A. conferta conferta

AFR 835E SCD175N AFR 736Y SCD 101G
Hydrotheca, length adcauline, adnate
part We oe 9 0°75-0°95 0°67-0°85 0°37-0'49 0°33-0°45
*length adcauline, free part 0'20-0'72 0°27-0°55 0°24-0°55 O*12-0°75
diameter at base .. O'1lI-O0'13 O'II-O'14 0°065-0°10 0:06-0-09
diameter at margin O'19-0'23 0°23-0°26 0°135-0°16 =0°13-0°15,
diameter, margin/base 1*54-1°92 1°64-2°18 1°40-2°29 = 167-233,
Solitary hydrotheca,+ length .. O°71-1°27 0°43-0°82 0°57=0°61
diameter at margin 0°16-0°20. -O*II5—O°15, =O" 12-015

*Including rejuvenated margins.

+Without rejuvenated margins.

Remarks. This material shows the distinctive characters of subspecies australis,
namely the overlapping hydrothecae and the characteristic thecal shape. It
differs markedly from the material which has been assigned to the nominal
subspecies in its growth-form, which is stiffer and more regular, and in the
measurements of its component parts, which are larger, though the proportions
are similar. Neither of the last two characters are necessarily of systematic
value, but it is difficult to assess the variability until more material is available.

Filellum antarcticum (Hartlaub, 1904)

Filellum ?antarcticum: Millard, 1958: 175.
Records. South coast: PAFR 835Z. MB 69C. ?SCD 175R.

Description. The second sample, growing on a polyzoan, bears the remains of a
coppinia in a somewhat dilapidated condition. However, the accessory tubes
are clearly visible; they are usually forked at the end (in one case twice),
occasionally simply truncated, but never curved as in F. serpens. The other
samples bear no coppiniae.

Measurements (mm., in MB 690(C).

Hydrotheca, length of free part, without reduplications ..
diameter at margin

0-18—0-30
~ ~ O-10—0°12
Remarks. The structure of the accessory coppinial tubes in MB 60C establishes
with certainty the presence of this species in South Africa for the first time.
The identity of unfertile samples is doubtful. Stechow (1925, p. 458) has
reported the presence of the closely related F. serpens in South Africa, but
since his specimens were unfertile, these records are also subject to doubt.

Hebella furax Millard, 1957
Fig. 2B—D
Hebella furax Millard, 1957: 200, fig. 8.

Records. South coast: MB 15B, 24T, 26E, 39W, 47K, 72G. SAMH 256, gor.
SCD 37U, 84Z, 117R, 179B.

Description. Colonies epizootic on Lytocarpus filamentosus (Lamarck), Thecocarpus

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA II

formosus (Busk) and T. flexuosus solidus Millard. Very few parasitic hydrothecae
present among these samples.

Hydrothecae similar to those previously described, though a few are a
little shorter and thus the proportion of length/diameter is less.

Gonothecae (described for the first time) borne on hydrorhiza, not sharply
demarcated from pedicel, widening towards distal end, often curved towards
one side, with smooth or slightly corrugated walls. Pedicel with 2-5 spiral
annulations. Containing several medusa-buds in various stages of development.
The young gonotheca is closed distally by a slightly convex operculum, which
tends to crumple in microscopic preparations, but has no valves. In the ripe
gonotheca the operculum is absent and the margin usually everted.

Oldest medusa about 0-4 mm. deep and 0:25 mm. wide, with at least
2 long, spirally coiled, marginal tentacles and a 4-lipped mouth.

Length of gonotheca, including pedicel a ie .. 1°64-2°52 mm.
maximum diameter ne es vie sis .. 0°50-0°81 mm.

Remarks. The examination of numerous samples of this species shows that the
epizootic form is far more common than the parasitic one. It is possible that
H. furax is conspecific with H. parasitica (Ciamician), but the gonophores of
the latter have not so far been described.

Hebella scandens (Bale, 1888)

Hebella scandens: Millard, 1957: 202. Millard, 1958: 176. Vervoort, 1959: 237, fig. 12.

Hebella calcarata: Ralph, 1958: 306, fig. 1 a-s.

Records. West coast: WCD 18V, 34K. South coast: MB 8M, 64M. SAMH 183,
Seine 2og, 402. oC) 377, 5271, 75F; 79Q, 84Y, 108H, 118L, 154J, 175M,
i7on. TRA 92R, 151H.

Hebella urceolata n. sp.
Fig, 2A

Holotype: SCD 154H (South African Museum registered number, SAMH 410).

Description. Colony epizootic on Halecium beani (Johnston). Hydrorhiza unseg-
mented, giving rise to hydrothecal pedicels at irregular intervals.

Hydrotheca about three times as long as wide, distinctly demarcated
from pedicel, gibbous below and narrowing above to just below margin.
Margin smooth, everted, straight or slightly oblique, sometimes rejuvenated.
Pedicel short, not annulated, widening distally. Hydrotheca separated from
pedicel by an annular thickening of the wall, to the distal region of which is
affixed a delicate diaphragm. :

Most hydrothecae show a tendency to be asymmetrical with the margin
more strongly everted and the basal region more gibbous on the surface
directed towards the distal end of the host colony, while the annular thecal
thickening is more strongly developed on the opposite side.

Gonophores absent.

L2 ANNALS OF THE SOUTH AFRICAN MUSEUM

nd

Fic. 2. Hebella spp.
A. Hebella urceolata n. sp. Various hydrothecae from the holotype.
B.-D. Hebella furax Millard. B and C, gonothecae containing young medusae from SCD 84Z.
D, a young gonotheca and a hydrotheca from MB 72G. .

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 13

Measurements (mm..)

Hydrorhiza, diameter Hy wi ms “4 uy ie 0*07-0'10
Pedicel, height, to diaphragm e ie 3 ts 4 O‘II-O'17
Hydrotheca, height, from diaphragm __... i 'p og 0:81-0'93
diameter, near base _.. se ee ns 7 2 0*31-0°35
diameter, at margin .. Li 4 bes a5 a 0'25-0'33

Lafoea fruticosa M. Sars, 1851
Fig. 3

Lafoea fruticosa: Allman, 1888: 34, pl. 16 (fig. 2, 2a). Broch, 1918: 12. Stechow, 1925: 456,
fig. 24B. Totton, 1930: 157, fig. 13. Fraser, 1944: 223, pl. 46 (fig. 206). Vervoort, 1946: 201,
fig. 83 c, d.

Records. West coast: WCD 1L. South coast: SOD 175Q.

Description. Branching colonies reaching a maximum height of 9:0 cm. Stems
fascicled, branching mainly in one plane but growing together in clusters and
anastomosing with each other to produce a shrubby effect.

Hydrothecae arising from all sides of the stem and branches, forming an
angle of about 40-60° with branch. Asymmetrical in shape, generally with a
double curvature on the adcauline side and a more or less straight abcauline
wall. Margin slightly everted. Pedicel short and slightly twisted.

An epizootic colony present on the surface of WCD IL, giving rise to
upright stems and solitary hydrothecae. The latter generally more slender and
with thinner perisarc than those borne by upright stems.

Coppinia present in WCD IL, consisting of closely packed, more-or-less
hexagonal gonothecae surmounted by acrocysts, which are loosely attached
and come away easily on handling. Gonotheca approximately 0-4 mm. in
height and 0-15 mm. in diameter at the shoulder, with a short mouth-funnel of
approximately 0-05 mm. in height and 0-05 mm. in diameter at the everted
margin. Tubular hydrothecae long (well over 3 mm.) and much coiled.

Measurements (mm.) WCD 1L SCD 175Q
Pedicel, height . . 5. rie & ne .. O'I0-O'IQ O'14-0°22
Hydrotheca, height .. oi 2s .. 0'42-0°66 0°34-0°59
Pedicel + hydrotheca, heey - .. 0°52-0:77 0:46—0°77
Hydrotheca, diameter at margin... a .. OtI4-O'19 O°14-0°16

Height of hydrotheca + pedicel

diameter at margin eos lo SAS aa

Hydrotheca height/pedicel height .. +) .. 2:63-6:00 1°55-3°28
Solitary hydrotheca + pedicel, eae: ty ~« .O*41-0"73
diameter at margin * 2 .: | O*I0-O°14

Remarks. It was difficult to assign this material to a species, as it has features
which resemble both L. fruticosa and L. dumosa. It has been assigned to the former
partly because of the appearance of the coppinia, which resembles that illus-

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

0-S5 mm.

B+C

Fic. 3. Lafoea fruticosa M. Sars.

A. Part of a fascicled stem, to show branching.

B and C. Hydrothecae.

D. An epizootic colony with solitary hydrothecae and an upright stem growing on the surface
of an older colony.

E. Part of a section through a coppinia, showing gonothecae (one with an acrocyst) and tubular
hydrothecae. _

F. Surface view of part of a coppinia.

(A and B from SCD 175Q, C-F from WCD 1L.)

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA I5

trated by Fraser and Vervoort, and partly because of the shape of the
hydrotheca, which resembles more that illustrated by Stechow, who has
described both species from South Africa. The shape and proportions of the
hydrotheca are intermediate between those described and illustrated by Totton
for L. dumosa and L. fruticosa.

Since the branches arise from the accessory tubes in this species it is
often difficult to distinguish epizootic colonies from the accessory tubes and
branches. However, the accessory tubes always run strictly parallel to the
axial tube and are in cytoplasmic connexion with the coenosarc of the latter,
whereas the epizootic hydrorhiza wanders at will over the surface of the stem,
with which it has no communication, and gives off both upright stems and
solitary hydrothecae.

Scandia mutabilis (Ritchie, 1907)
Scandia mutabilis: Millard, 1957: 202. Millard, 1958: 176.
Records. South coast: KNY 30U.

Kygophylax africana Stechow, 1923
Fig. 4A—F
Kygophylax africana: Stechow, 1925: 445, fig. 18.
Records. West coast: AFR 743H. South Coast: SAMH 321.

Description. Hydrorhiza growing over the surface of worm-tubes, bearing
nematothecae, solitary hydrothecae and upright stems in all stages of develop-
ment, the latter reaching a maximum height of 9-5 cm. Stem and main branches
thickly fascicled with some of the accompanying tubes extending on to the
basal part of practically all the smaller branches. Stem bearing alternate
hydrothecae and alternate branches given off at the base of every third and
fourth hydrotheca. Branches often rebranching in the same manner, or
according to a different scheme in which subbranches arise at the base of every
third hydrotheca alternately to the right and the left. The hydrotheca at the
origin of each branch is not strictly in the axil but shifted slightly onto the
branch itself. All branches in one plane, many of the larger ones anastomosing
with other parts of the colony.

Smaller branches (Stechow’s ‘cladia’) unsegmented, but often with one
or two corrugations close to the base; bearing alternate hydrothecae, of which
the two rows are in one plane.

Hydrothecal pedicel short (and covered by the accompanying tubes on
the thick part of the stem), sometimes witha distinct indentation on the adcauline
side. Hydrotheca long, tubular, curved away from branch, of equal diameter
throughout from just above the level of the diaphragm (SAMH 321, fig. 4F), or
widening slightly towards margin (AFR 743H, fig. 4E). Margin slightly
everted. Hydrotheca set at a varying angle to stem or branch—the angle may
be as large as 50° (more common in AFR 743H), or very small so that the

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

adcauline wall is almost in contact with the axial tube (more common in
SAMH 321). Diaphragm in form of annular thickening, set obliquely, with
adcauline side lower than abcauline.

Nematotheca tubular and of equal diameter throughout, borne on a very
short and narrow pedicel, and separated from it by a delicate diaphragm.
Nematothecae borne irregularly on the peripheral tubes of the stem, and one
on the base of each hydrothecal pedicel. The nematotheca of the first hydro-
theca of a branch, however, is situated on the branch itself immediately
beyond the hydrotheca instead of on its pedicel. Both hydrothecae and nemato-
thecae often with reduplicated margins.

Coppiniae present around the stem and larger branches, and also borne
on the substratum by the hydrorhiza, reaching about 10 mm. in length and
4 mm. in diameter. Consisting of adpressed gonothecae (about 9-12 visible
in cross-section), and numerous branching nematophores. Gonotheca penta-
gonal or hexagonal in surface view; in lateral view widening from base to
top of adpressed part, bearing a free tubular neck surmounted by 2 divergent
sharply pointed horns. Each with 2 apertures situated on opposite sides of the
distal end of the neck immediately below the horns. Male and female gonothecae
borne in separate coppiniae, but exactly the same in appearance, female
containing a cluster of planula larvae. Nematothecae borne on branching
perisarcal tubes which arise from the peripheral tubes of the stem and penetrate
between the gonothecae, continuing beyond them to reach a total height of
14 to 24 mm.

Solitary hydrothecae borne by hydrorhiza and also by young epizootic
colonies growing on the surface of the larger ones. Usually smaller than normal
hydrothecae and with longer pedicels. Shape variable; quite symmetrical and
straight, irregular, or curved as in the normal type.

Measurements (ram., without reduplications) AFR 743H SAMH 321
Final branches, distance between two hydrothecae .. 0°34—-0-48 0-21—0°38
diameter, above hydrotheca at s .. 0'05-0°0Q 0-05—0-07
Pedicel, length adcauline va ee Me .. 0'03-0°095 0:02-0:05
Hydrotheca, length adcauline .. the oe .. 0:'26-0°33 0°24-0°30
length abcauline .. Si ue we .. 0°23-0°90) Q71G—0-25
diameter at mouth Bs ae bi .. 0*'09-0°10 0:07-0:08
diameter across diaphragm bs a .. 0'04-0:06 0:05-0:07
Nematotheca, length, including pedicel fi .. 0°07-O'12 0:0Q—0°17
diameter at margin of es Mm .. 0'025—-0°04 0°02—0°025

Remarks. Although there are slight differences in the appearance of the hydrothe-
cae in these two colonies (e.g. the hydrothecae in SAMH 321 have narrower
mouths and shorter pedicels and are more closely set than those in AFR 743H)
I can see no reason for specific distinction. Both bear coppiniae (male in
SAMH 321, female in AFR 743H) which are exactly alike in construction.
Although the branching in general conforms to the scheme outlined above,

i

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA

E

Fic. 4. Zygophylax spp.

AF. <ygophylax africana Stechow. A, part of a fascicled stem to show branching (branches
cut off short). B, part of a section through a female coppinia, showing gonothecae and
branching nematothecae. C, surface view of part of coppinia (horns of gonothecae
displaced by pressure of coverslip). D, solitary hydrothecae from epizootic colony.
E and F, portions of branches, to show hydrothecae and nematothecae.

G. <ygophylax armata (Ritchie). Part of a branch.

(A and F from SAMH ga21, B-E from AFR 743H, G from SCD 254P.)

17

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

there are many irregularities, and the branches differ greatly in thickness and
in the degree to which they subdivide. ‘The general effect is far from regular.

The species has only once been reported, by Stechow in 1923b, off Cape
Town. Stechow described a smaller sterile colony reaching only 1-8 cm.
in height, and less heavily fascicled. I have seen a slide of Stechow’s material
loaned by the Munich Museum, and there is no doubt that it is the same species.
The coppiniae are described here for the first time.

The coppiniae of this species are strikingly similar to those of Cryptolaria
pectinata (Allman) (cf. Stechow, 1925, fig. 20). The latter, however, are dioecious,
and the female gonotheca has a relatively shorter neck and horns. The two
species are also similar in the general appearance of the colony and in the
subopposite method of branching. This emphasizes the close relationship
between the genus Cryptolaria and certain species of <ygophylax. In fact, Crypto-
laria, by the presence and nature of its diaphragm, its nematothecae, subopposite
branching and coppinial structure, is more closely related to certain species
of <ygophylax than the various species of <ygophylax are to one another.
Cryptolaria differs only in the adherent hydrothecae, which as Stechow has
shown (1925, p. 450) is a matter of degree only and does not occur in the
young stems. I cannot help but feel that the adherent nature of the hydrotheca
is not a character of high systematic value, and that it will be necessary in the
future to unite Cryptolaria with <ygophylax, and possibly Acryptolaria with
Lafoea. I hesitate to do so at this stage as my knowledge of the variability of the
species in these genera is insufficient.

Kygophylax armata (Ritchie, 1907)
Fig. 4G
Brucella armata Ritchie, 1907: 533, pl. 2 (fig. 2-2c).
Records. South coast: SCD 254P, 297X.

Description. Two colonies, the larger reaching a height of 3-0 cm., with stem
and larger branches fascicled. Branching at the base rather irregular, but
larger branches and their subdivisions in one plane. The 2 rows of hydrothecae
not always in one plane, but with a tendency to shift on to the anterior surface
_ of a branch. Further details of hydrothecae and nematothecae as described by
Ritchie, except that the individual measurements are somewhat smaller. —

Solitary hydrothecae present on hydrorhiza and in epizootic colonies,
with the same proportions as those on branched stem, but less curved, and
sometimes completely symmetrical.

Coppiniae absent.

Measurements (mm., without reduplications).

SCD 254P

Hydrocladium, diameter above hydrotheca Be A .. 0'07—0:09
distance between 2 consecutive hydrothecae .. oe .. 0°23-0°35
Hydrothecal pedicel, length adcauline si bi .. 0°03-0-06

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 19

Hydrotheca, length adcauline ve Hit AN a .. 'O*19—0°26
length abcauline. . ae yf is -# oh .. 0°18-0:26
diameter at mouth i we ar oe a -s OWT-O'RS
diameter at level of diaphragm aie y ms .. 0*06-0:075

Nematotheca, length, including pedicel .. ‘- ee .. 0°05-0'07
diameter at mouth y 4¢ Ag Ap es .. 0°03-0'04

Remarks. This species has only been recorded once before, from off Gough
Island in 100 fathoms, and is a new record from South Africa. It can be
distinguished from Z. biarmata Billard by the nature of the coppinia; and as
far as I can determine, the trophosomes of the two species can also be distin-
guished by the closer approximation of the hydrothecae in <. armata and their
more pronounced asymmetry.

Kygophylax cornucopia Millard, 1955
Kygophylax cornucopia Millard, 1955: 219, fig. 3. Millard, 1957: 203.

Records. West coast: TB 1B (reported by Millard, 1955). TRA 156F. South
coast: CPR 46K. LIZ 27L, MB 59U. SAMH 192. SCD 36V, 153P.

Description. Colonies epizootic on Antennella africana Broch, A. secundaria
(Gmelin) and Monostaechas natalensis Millard.

Kygophylax engmatica n. sp.
Fig. 5A—-F

Holotype: WCD 12E (South African Museum registered number, SAMH 411).

Description. Colony epizootic on Nemertesia ramosa Lamx. and reaching a height
of 2-3 mm.

Hydrorhiza adherent to stem of host, unsegmented, branching irregularly.

Stem upright and pinnate, divided by straight nodes into long internodes,
each of which gives off a hydrocladial apophysis near the distal end. Apophysis
with a large mamelon on the upper surface. The first few internodes of the
stem sometimes much shorter than the others, and occasionally without
hydrocladia. An internodal ridge sometimes present near base of stem internode.

Hydrocladia alternate, the two rows in the same plane. Hydrocladium
forming an angle of about 30°—40° with stem. Consisting of one basal internode,
followed by a pedicel and hydrotheca which form one unit and are not exter-
nally demarcated from one another. Pedicel and hydrotheca expanding evenly
towards margin. Thecal margin even, not everted. Pedicel and hydrotheca
separated internally by a diaphragm. Diaphragm bilaterally symmetrical,
with aperture close to adcauline side, and abcauline section sloping obliquely
towards distal end of hydrocladium. Pedicel sometimes with a low internodal
ridge near base, and sometimes showing evidence of regeneration.

One nematotheca on each hydrothecal pedicel, seated about midway
along adcauline surface, 2-chambered, moveable.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

a eT

(i
se?

G

a 05 mm.

H

Fic. 5. <ygophylax spp.
A.-F. Zygophylax enigmatica, n. sp., from the holotype. A, an upright stem. B, two hydrocladia
on a larger scale. C-F, gonothecae in various stages of development.
G.-H. Zygophylax sibogae Billard, from SCD 301H. G, ventral view of part of a branch. H,
anterior view.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA a |

Hydranth completely retractable into hydrotheca, hypostome conical
when contracted but capable of great distension.

Gonothecae borne on hydrocladial apophyses, very large in comparison
with size of hydrothecae. Each gonotheca laterally compressed in a plane at
right angles to the axis of the stem, flask-shaped, and tapering to a slender neck.
Neck sometimes curved to one side, with margin obliquely set. Containing
about 8-10 eggs which develop zn situ to planula larvae.

Measurements (mm.)

Hydrorhiza, diameter ne ie ‘is a6 .. 0'04-0:08
Stem internode, length (not the ee 5 ‘3 Me ay ». 0'20—0'34
diameter .. : .3 ae 7 .. 0°04-0:07
Hydrocladium, basal ES anout. fect is ae 4.3 .. 0°03-0'07
pedicel, length to diaphragm .. ia ig: a .. O*I10—0°20
hydrotheca, height from diaphragm .. a ie .. 0°07-0'10
diameter at margin - a! se i 3 ». 0°095-0°11
Nematotheca, length ae Ls: ie 5 i .. 0°035-0°05
Gonotheca, length .. a ie Aa se 3 .. 0'60-0°71
diameter .. a ae a * om sic wx; 0'29-0-92

Remarks. This minute species is obviously closely related to <. cornucopia Millard,
particularly in the structure and proportions of the hydrotheca and its pedicel
and in the absence of a coppinia, but differs from it in the presence of an upright,
pinnate stem.

Both species show tendencies towards the Plumulariidae, but cannot be
included in that family since the hydrothecae are not sessile, but borne on a
pedicel from which they are separated by a diaphragm. The continuity of
pedicel and hydrotheca is characteristic of other species of < ygophylax.

Kygophylax sibogae Billard, 1918
Fig. 5G-H
KIgophylax stbogae Billard, 1918: 21, fig. 1. Totton, 1930: 167, fig. 21. Ralph, 1958: 311, fig. 2,
e-i.

Records. South coast: SCD gorH.

Description. A number of fascicled stems reaching a maximum height of 3:1 cm.,
branching pinnately and in one plane. Branches generally arising below every
third and fourth hydrotheca, the lower ones fascicled at base and often
rebranching in a similar manner.

Hydrothecae borne in two longitudinal rows on the anterior surface of stem
and branches, with a sharp angle between the rows, and with the members of
one row twisted slightly away from those of the other row. Hydrotheca strongly
curved towards distal end of branch, widening to circular aperture which faces
towards branch and slightly to one side. Borne on long pedicel of approximately
the same length. Diaphragm well developed, often funnel-shaped, Pedicel
usually separated from stem apophysis by distinct node,

[5.49 ANNALS OF THE SOUTH AFRICAN MUSEUM

Nematothecae borne on peripheral tubes and one on the lateral surface of
each hydrothecal apophysis, tubular and narrowing slightly to margin, normally
about 0:07 mm. in length, but often abnormally lengthened by regeneration.

Gonophores absent.

Measurements (mm.)

Hydrocladium, distance between 2 pedicels 4) r¢ -. 0°28-0°37
diameter .. a A i a ae A .. O*05-0°07
Pedicel, length (without apophysis) oi ay i ». 0*20—0°26
Hydrotheca, length adcauline Be x ne Bs .. 0°16—-0-20
maximum length De Be oy ds ah .. 0°26—0°30
diameter at mouth “. ae Ne A x. .. O*tII-O'13
diameter at diaphragm at oe Lay bes ». 0°045-0°05
Nematotheca, length 1 we oe a ee .. 0°07—0°32
maximum diameter... i i if 5 . » 0°025—-0'035
diameter at mouth J os iy AP ni .. O*°015—0°03

Remarks. The hydrothecae in this colony have a beautifully regular arrangement,
more so than those figured by Ralph, and in appearance are exactly like
Billard’s diagram. The measurements of hydrothecal length and pedicel length
are slightly less than those given by Totton and Ralph, but there is no doubt
about the identification of the material.

This species is closely related to <. infundibulum Millard, 1958, differing
from it in the greater length of the pedicel and the more marked curvature of
the hydrotheca.

XK. sibogae is known from the East Indies and New Zealand. This is the
first record from South Africa.

Family Syntheciidae

Hincksella cylindrica pusilla Ritchie, 1910
Fig. 6A-D
Sertularella cylindrica var. pusilla Ritchie, 1910: 817, pl. 77 (fig. 9).

Hincksella cylindrica var. pusilla: Billard, 1925: 124. Vervoort, 1959: 247, fig. 19 b, c.
Synthecium cylindricum var. pusilla: Leloup, 1935: 31, fig. 14.

Records. South coast: SCD 297Y.

Description. A single colony of about 6 unfascicled and unbranched stems reaching

a maximum height of 0-7 cm. Internodes of irregular length, separated by

slightly oblique nodes, and each bearing a hydrotheca near the distal end.
Hydrothecae very delicate, and most of them crumpled, adnate for about

+ adcauline side, then bent outwards. Margin slightly everted, often regenerated.
Gonothecae absent.

Measurements (mm., without reduplications)

Internode length tn ie ae RY MF ie .. 0'29-0°54.
diameter at node hye ‘es oy a as .. 0°05—-0°09

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 23

2
ss
x +r
Bee a Ee 0-5 mm

y

Fic. 6. Syntheciidae.
A.-D. Hincksella cylindrica pusilla Ritchie. Hydrothecae from SCD 297Y.
E.—J. Synthecium dentigerum Jarvis, from SCD 84G. E and F, portions of hydrocladia. G and H,
2 ae a ates viewed from the distal end (adcauline surface indicated by x). J, gonotheca
in side view.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hydrotheca, total length in centre .. We bb, iG .. 0°49-0°60
length adcauline, adnate part Pe te ee -» O*°15-0°18
length adcauline, free part A ie 2% A 1. O°44-0°47

adnate part/adcauline length a, i 1s if .. 0°24-0°29
diameter at margin Mi ne ‘e if M2 .. O*°I4-0°17

Remarks. 'This is the first record of the species from South Africa. Subspecies
pusilla has previously been recorded from the Mergui Archipelago, East
Indies, Japan, tropical West Africa and the West Indies.

Synthecium dentigerum Jarvis, 1922
Fig. 6E-J

Synthecium dentigerum Jarvis, 1922: 344, pl. 25 (fig. 15 a, b). Totton, 1930: 172.
Records. South coast: SAMH 375. SCD 52G, 84G, 112F. TRA 151G.

Description. A number of colonies reaching a maximum height of 4-0 cm. Stem
unfascicled, pinnate, divided into internodes by straight nodes, which are
sometimes obscure. Arrangement on internodes variable, common arrangements
including 2 pairs of hydrothecae with a pair of opposite hydrocladia arising
between them, and one pair of opposite hydrocladia followed by one pair of
hydrothecae.

Hydrotheca as described and figured by Jarvis, with one large, adcauline,
internal tooth, but the presence of this tooth is by no means constant and is
sometimes found on only a few hydrothecae of a colony. Occasional hydrothecae
with a small triangular thickening in the centre of the abcauline wall.

Gonothecae (not previously described) borne on stem or hydrocladia,
arising within hydrothecae; generally pentagonal when viewed from above,
though sometimes two of the angles are smoothed out resulting in a rather
flattened triangle; with 5-6 transverse folds on the flat surfaces, which may
peter out on the angles or continue over them, but do not produce a definite
zigzag line of junction; tapering distally to a small circular opening. Sex not
determinable.

Measurements (mm.)

Hydrocladium, internode length .. if oe e .. 0*58-0-76
Hydrotheca, length abcauline ae &. es if .. 0°34-0°41
length adcauline, adnate part .. re ie he .. 0°38-0-48
length adcauline, free part... ay = ne .. 0*08-0°17
adnate part/adcauline length .. Bs! a = .. 0°69-0°85
diameter near base ‘ s Ne As see .. O°14-0'17
diameter at margin B ms ue ie .. 0°16-0-20
Gonotheca, length .. a y a Ne es .. I‘OI-I-40
maximum diameter ses aie He Ms oe .. 0°56—0-77

Remarks. The dimensions of this material are slightly less than those given by
Totton for S. dentigerum, but are closer than to S. carinatum Totton, a closely

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 25

related species with internal teeth. The gonotheca of the latter is also
different.

S. dentigerum has only been reported once, from Chagos in the Indian
Ocean by Jarvis. This is the first record from South Africa.

Synthecium elegans Allman, 1872
Synthecium ?elegans: Millard, 1957: 203, fig. gD. Millard, 1958: 182.
Records. South coast: MB 8U, 64N.

Remarks. As the gonothecae of this species have still not been found in South
Africa, the identification must remain uncertain.

Synthecium hians Millard, 1957
Synthectum hians Millard, 1957: 204, fig. 9 A—C.

Records. South coast: MB 12Y, 64L. SAMH 151, 188, 281, 374. SCD 37L,
52F, 84H, 119N, 250K, 254N, 296F. TRA 38G.

Family Sertulariidae
Amphisbetia bidens (Bale, 1884)

Thuiaria bidens: Day, Millard & Harrison, 1952: 404 (listed).
Amphisbetia bidens: Millard, 1957: 220. Millard, 1958: 182.

Records. South coast: BMR 23M. KNY 30K, 70D (recorded by Day et ai.
1952). LIZ 7P, 11H. MB 15F, 24N, 47L, 88M. SAMH 261, 378. SCD 60A,
84K, 153V, 330E.

Amphisbetia minima (Thompson, 1879)
Amphisbetia minima: Millard, 1957: 221. Millard, 1958: 183. Ralph, 1961: 774, figs. 8 a—h.

Records. West coast: B 114F. CP 650A. TRA 156J. WCD 34], 81F. South
coast: MB 24Q.

Description. Stems reaching a maximum height of 0:5 cm., and bearing up to
15 pairs of hydrothecae. Male and female gonothecae present, similar in
structure, usually arising below the first pair of hydrothecae, but occasionally
below the second or third pair.

Amphisbetia operculata (Linn., 1758)

?Sertularia aperta Allman, 1886: 138, pl. 19 (figs. 1, 2).

Sertularia operculata: Day, Millard & Harrison, 1952: 404 (listed).

Amphisbetia operculata: Millard, 1957: 221. Millard, 1958: 183. Millard, 1961: 204. Ralph, 1961:

775, fig. 8 i-k.

Records. West coast: B 137. LAM 14T, 45Y. SAMH 403. SB 194M. TB a21E.
TRA 156H. WCD 34G, 81C. South coast: BMR 9M. CPR 9C. KNY 22G,
30L, 57H, 70A (recorded by Day et al. 1952). LIZ 2F, 7Q, 11G. MB 24P,
52A. SAMH 232, 358, 376. SCD 84L, 284A, 304H.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks. Many of these colonies show a form of branching approaching that of
A. fasciculata (Kirch.), and similar to that figured by Allman, 1886, pl. 14, for
Sertularia crins (considered by Billard, 1910, who examined the types, to
be conspecific with A. operculata), with an elongated ‘main stem’ and subsidiary
branches which may subdivide once or twice. There is sometimes, but not
always, a difference in internode length and thickness between the smaller
branches and ‘main stem’ (where the length may be almost twice as great), but
the difference is never so marked as described by Ralph, 1961, for A. fasciculata.
Since this and the normal form may occur in the same colony, it is not possible
to distinguish two species. ‘The height of the colony never exceeds about 14 cm.

Sertularia aperta Allman, 1886, from the Cape of Good Hope is almost
certainly a growth-form of the same species.

Crateritheca acanthostoma (Bale, 1882)
Fig. 7

?Dynamena pluridentata Kirchenpauer, 1864: 14, fig. 10.

Sertularia acanthostoma: Bale, 1884: 85, pl. 4 (figs. 7-8). Billard, 1907: 352. Warren, 1908: 303,
pl. 46 (figs. 23-26), fig. 7. Bale, 1913: 131.

Stereotheca acanthostoma: Millard, 1958: 199.

Crateritheca acanthostoma: Ralph, 1961: 756, fig. 2c.

Records. South coast: SCD 50A.

Description. Several pinnate stems reaching a maximum height of 2-4 cm.
Arrangement of hydrocladia, internodes and hydrothecae as in previous
descriptions.

Hydrotheca adnate for a little over half adcauline length, with no external

ridges or furrows, but with 3 intrathecal septa. These include (fig. 7A, C, D):

(a) an adcauline septum about half-way up the hydrotheca in the form
of a horizontal shelf bearing 2 or 3 minute denticles on the free inner
edge, )

(b) a lower abcauline septum about a quarter of the way up, in the form
of a horizontal shelf behind which is an opening for the passage of a
strand of ectoderm, and below which is situated the abcauline blind
pouch of the hydranth,

(c) an upper abcauline septum about three-quarters of the way up in
the form of a narrow horizontal shelf forming the base of a longitudinal
trough communicating with the cavity of the hydrotheca and
extending to the margin. This trough contains a cluster of large
nematocysts embedded in ectoderm which is continuous with the
strand passing behind septum b.

Hydropore surrounded by a raised funnel-shaped ridge. Operculum of a single
membranous plate filling complete orifice. Marginal teeth and soft parts as
described by Warren.

Gonothecae absent.

Remarks. The genus Crateritheca is recognized here on the basis of remarks by

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA ar,

Totton, 1930, p. 207, and Ralph, 1961, although the diagnosis of the latter
author will need modification to include C. acanthostoma.

Stechow in 1919 included Warren’s material from Natal and Billard’s
material from Madagascar in Kirchenpauer’s species ‘Dynamena’ pluridentata
from the Cape of Good Hope, although in 1925 he kept them separate. As
Kirchenpauer’s species is inadequately described and illustrated, and might
from the diagrams equally well be Stereotheca elongata, no certainty can be
reached on this matter.

The South African material of the species differs in certain respects from
the Australian material as shown by dissections of hydrothecae made under a
high-power dissecting microscope. Thus, the hydrotheca has a smaller propor-

0:2 mm.

C

Fic. 7. Crateritheca acanthostoma (Bale).

A. A single hydrotheca in side view. Adcauline surface on right.

B.—-D. Free-hand diagrams drawn from dissections of hydrothecae. B, distal view of complete
hydrotheca with operculum in position. C, distal view of cross-section through abcauline
wall taken near distal end. D, distal view of cross-section through hydrotheca at a slightly
deeper level.

a, the adcauline septum; b, the lower abcauline septum; c, the upper abcauline septum.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

tion of the adcauline wall adnate, the adcauline intrathecal ridge is not
continued round the complete diameter of the hydrotheca, and the lower
abcauline intrathecal ridge is perforated. These differences, however, are not
considered sufficient basis for the separation of another species, although they
may prove to be of subspecific value. Further material from both countries is
needed to show the degree of variation possible.

Dictyocladium coactum Stechow, 1923.
Dictyocladium coactum: Stechow, 1925: 466, fig. 27. Millard, 1957: 206.

Records. South coast: AFR 866.0.A. SAMH 264, 314. SCD 3C, 36Z, 52P,
85R. TRA 151K.

Diphasia tetraglochina Billard, 1907
Fig,
Diphasia tetraglochina Billard, 1907: 358, fig. 7. Billard, 1925: 139.
Records. South coast: SAMH 267.

Description. Small unbranched stems growing on algae and reaching a maximum
height of 0-25 cm. Hydrorhiza with internal thickenings of perisarc. Nodes
markedly oblique, resembling hinge-joints. Each internode bearing one pair
of opposite hydrothecae.

Hydrothecae asymmetrical, with both members of a pair twisted and
tilted towards one surface of the stem (here termed the anterior surface),
and sometimes contiguous with one another on the opposite surface (in the
distal region of some stems only). See figure 8B. Hydrotheca adnate for 3 to #
of adcauline length, bent outwards, widening towards margin. Perisarc usually
thickened in centre part of abcauline wall and often around margin as well.
Margin with 4 small, pointed teeth: 2 latero-adcauline and 2 latero-abcauline,
but displaced towards the anterior surface of the stem by the twisting of the
hydrotheca. Operculum of 1 adcauline valve. eae with no abcauline
blind pouch.

Gonotheca (not previously described) borne on anterior surface of stem
immediately below first pair of hydrothecae, resembling a prickly pear in
appearance, with a variable number of small spines in distal half and a small
terminal aperture at the end of a short neck.

Measurements (mm.)

Internode length Me Se ae e: se .. 0°30—-0°54.
Hydrotheca, length ae hae a m a a .. 0°27-O0°41
length adcauline, contiguous part .. Se ne .. 0°00—-0°17
length adcauline, adnate part re 213 ne .. 0°24-0°31
length adcauline, free part iM ue ay. ae .. 0*°09-0°15
adnate part/adcauline length .. fe a be .. 0°65-0°74
diameter at margin ma "- ag us sks .+ @1@-e25
Gonotheca, length .. ns nit Ue st Ww .. "OF 70-93

maximum diameter am aio uA S: bie .. 0'38-0°45

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 29

ig
) D

E

A, C-E

ge eee ee Siri.

Fic. 8. Diphasia tetraglochina Billard.
. Astem with 3 pairs of hydrothecae.
Sketch of distal view of a pair of hydrothecae to show asymmetry. Operculum in place in
right hydrotheca.
. The distal pair of hydrothecae from a stem with 5 pairs.
. The proximal pair from another stem.
. The gonotheca.

HOO wp

Remarks. This rare species has only been recorded once before, from Madagascar.
It was not possible to determine the sex of the gonothecae.

Dynamena cornicina McCrady, 1858

Fig. 9
Sertularia densa Stechow, 1919: 93, fig. J}.
Sertularia cornicina: Jarvis, 1922: 338.
Dynamena cornicina: Billard, 1925: 188, pl. 7 (fig. 23), fig. 40. Broch, 1933: 86, fig. 36. Vervoort,
1941: 206, fig. 3.

Records. South coast: LIZ 16C, 4oJ.

Description. Two colonies, one very rich, growing on coralline algae. Only
simple stems present, reaching a maximum height of o-g cm. Stem with short,
proximal, athecate region terminated by a hinge-joint, and a long thecate
region divided into regular internodes by constricted nodes. Each internode
bearing a pair of opposite hydrothecae. Consecutive pairs of hydrothecae close,

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

separated by a distance less than the height of a hydrotheca. Members of a
pair of hydrothecae contiguous with one another on anterior surface in distal
region of stem, but completely separate in proximal region.

Hydrotheca tubular, adnate for over half adcauline length, smoothly
bent out in distal half. Margin with two large, but delicate, lateral teeth and a
short median adcauline tooth. Abcauline wall thickened just below margin.
No internal teeth. Operculum of 2 valves: one large abcauline, and one small
adcauline divided into 2 by a median line.

D haya GRY

Fic. 9. Dynamena cornicina McCrady.

A. and B. The upper 3, and the lower 3, pairs of hydrothecae respectively from a stem bearing
12 pairs in all (anterior view).
C. The margin of a hydrotheca showing the operculum. (Abcauline valve in thick broken line.)
D. The gonotheca.
(All from LIZ 16C.)

Gonothecae abundant, borne on hydrorhiza, ovoid, distinctly annulated
throughout, with a broad, operculate aperture.

Measurements (mm.)

Internode length ss) ry di a ar ss, .. 0754-0765
diameter at node By ae i % By: .. 0°07-O0°14
Hydrotheca, length abcauline va Kon oe si .. 0*29-0°45

length adcauline, contiguous part... Ns si .. 0:00-0:26

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 31

length adcauline, adnate part .. =, re . .. 0°35-0°46
length adcauline, free part... ‘f af i .. O*°14-0'25
adnate part/adcauline length .. * re ag .. 0*59-0°76
diameter at margin Sis - ae 4 - .. Of13-O'175
Gonotheca, length .. re ut a ie ae .. 1:06-1°38
Mmaxmim diameter ..° . re ne .. 0:76-0:88

Remarks. In this material the pairs of hydrothecae are ined set on the stem,
thus resembling that illustrated by Stechow, 1919.

The species has been reported by Jarvis, 1922, from tropical East Africa,
but this is the first record from South Africa.

Dynamena crisioides crisioides Lamouroux, 1824
Dynamena crisioides: Millard, 1958: 183. Vervoort, 1959: 260, fig. 27 a, b.
Records. South coast: CPR 7A.

Dynamena quadridentata nodosa Hargitt, 1908

Dynamena quadridentata var. nodosa: Billard, 1925: 197, fig. 43D. Millard, 1958: 186, fig. 6B.
Pasythea quadridentata: Warren, 1908: 312, fig. 11.
_Dynamena gibbosa Billard, 1925: 199, fig. 45.

Records. South coast: SCD 50P.

Remarks. The South African material of this species appears to be intermediate
between D. quadridentata nodosa and D. gibbosa, which is included here as a
synonym.
Salacia articulata (Pallas, 1766)
Fig. 10G

Salacia articulata: Millard, 1957: 208 (synonymy). Millard, 1958: 186. Millard, 1961: 205.

Records. West coast: LAM 7N, 30N. SB 153R, 168E. WCD 12C, 81E. South
coast: L 452B. MB 47X, 84B. SAMH 152, 262, 295, 377. SCD 36X, 52N,
56R, 84M, 138J, 153Q, 296G, 330D. TRA o2L.

Salacia disjuncta nov. sp.
Fig. 10A—F

Types. Holotype: SCD 37K (South African Museum registered number,
SAMH 412). Paratype: SCD 296H. (Both from South coast.)

Description of Holotype. A single rooted stem 1-4 cm. in height. Stem upright,
moderately stiff, not zigzag, pinnate, divided by straight nodes into distinct
internodes each of which bears 3 pairs of hydrothecae and 1 pair of opposite
hydrocladia arising between the first and second pairs of hydrothecae. First
internode of stem with 2 pairs of hydrothecae only. Stem constricted at nodes.
Members of a pair of hydrothecae opposite or subopposite, not in contact with
one another. The two rows of hydrothecae in 1 plane and on opposite sides of the
stem. Consecutive pairs of hydrothecae on an internode separated by a short

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

AL Rae Ese on:

Fic. 10. Salacia spp.

A.-F. Salacia disjuncta n. sp. A, whole colony. B, 2 hydrothecae of a group. C-E, various views
of the margin, C showing the operculum (in broken line). F, part of stem and
hydrocladia. : she

G. Salacia articulata (Pallas). Part of stem and hydrocladium as a comparison with S. disjuncta.

(A-C, F from SCD 37K; D-E from SCD 296H; G from F 254.)

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 33

interval or slightly overlapping; the group on one internode well separated
from that on the next. Hydrocladia arising from the stem at right angles, the
2 rows in one plane.

Hydrocladium borne on a stem apophysis of variable length, divided into
distinct internodes of variable length by constricted nodes. Each internode
consisting of a slender proximal region without hydrothecae occupying up to
half its length, and a wider distal region bearing 2 or 3 pairs of hydrothecae
arranged in a close and overlapping group reminiscent of the genus Dynamena.
Members of a pair of hydrothecae opposite or subopposite, not in contact with
each other; the 2 rows in one plane.

Hydrotheca tubular, narrowing slightly to margin, slightly turbinate in
proximal region, smoothly bent outwards in distal region, adnate for most of
adcauline length, but with at least 1/10 free. Margin more or less parallel to
axis of stem in cauline hydrothecae, but variable in hydrocladial hydrothecae
(facing slightly downwards in the proximal pair of a group and slightly upwards
in the distal pair). Aperture widened transversely, with no marginal teeth or
occasionally with 2 low, rounded, lateral lobes. Operculum of 1 large abcauline
valve. Hydranth with no abcauline blind pouch.

Gonothecae absent.

Measurements (mm., exclusive of regenerations). Holotype Paratype

Stem, normal internode, length i es .. 1°86-2:16 2:06-2:34
diameter at node .. se ce Pe »» 0'22-0°30 0°15-0°25
Hydrocladium, internode length af ae -» O°QI-1°72 1°20-1°78
diameter at node .. os tf ME .. 0*°09-0°12 0-*09-O'II
Hydrotheca, length abcauline (cauline only) .. 0°29-0°36 0°30-0°42
*length adcauline, adnate part .. i .. 0°30-0:46 0:30-0°47
*length adcauline, free part se oh .. 0°08-0:25 0:08-0:21
*adnate part/adcauline length .. ne .. 0°55-0°85 0-61-0°85
diameter at margin be a “ .. O*'IO-O'14 O*10—-0'12

*Including cauline hydrothecae and those on the distal ends of hydrocladial groups.

Remarks. This species is very distinct and easily recognized by its strictly
opposite hydrocladia and marked grouping of the hydrothecae on the hydro-
cladia. Although quite different in appearance from S. articulata, it resembles this
species in many details of construction and is closely related to it. It can be
distinguished by:
(a) the position of the hydrocladium which forms a right angle with
the stem,
(b) the more slender stem internodes,
(c) the nature of the hydrocladial internodes, with their slender proximal
region and marked bunching of hydrothecae in the distal region,
(d) the hydrothecae, which are more bent outwards and are not
completely adnate.

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

The paratype consists of 2 stems, the larger 1-9 cm. in height, in which
certain irregularities occur; thus 2 of the stem internodes bear but one
hydrotheca each, and there is a tendency towards stolon formation from the
tips of the hydrocladia.

Genus SERTULARELLA
Remarks on the diagnosis of species
As has been indicated by Picard (1956) and Millard (1958), the shape of

the hydrotheca is sometimes used as a basis for specific differentiation in
Sertularella. Thus, Picard distinguishes between species in which the margin is
tilted towards the distal end of the colony, and those in which the margin is
tilted towards the base.

However, it has been found in practice that the category is not always
obvious, due to variations in the curvature of the walls and elongation of one
or other marginal tooth, and if such categories are to be used it is necessary
that they should be more accurately defined. It is proposed, therefore, to
distinguish three hydrothecal types, which are defined as follows (fig. 11):

Fic. 11. Sertularella.

The three categories of hydrothecal shape. Types a and c could be converted to b by
the enlargement of the abcauline marginal tooth (dotted lines). See text for description.

(a) Hydrothecal margin perpendicular to axis. Forms with a hydrotheca which
is obviously symmetrical in lateral view, and in which a line drawn
at right angles to the margin and passing through the lateral marginal
tooth, will bisect the perisarcal thickening at the base of the adcauline
wall, e.g. typical examples of S. fusiformis and S. capensis.

(b) Hydrothecal margin tilted towards adcauline side. Forms in which a line
drawn at right angles to the margin and passing through the lateral
marginal tooth will pass through the abcauline wall outside the

— -

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 35

base of the hydrotheca (e.g. S. mediterranea), or in less extreme examples
through the hydropore (e.g. S. africana).

(c) Hydrothecal margin tilted towards abcauline side. Forms in which a line
drawn at right angles to the margin and passing through the lateral
marginal tooth will pass through the adcauline wall (distal to the
perisarcal thickening at its base), e.g. S. arbuscula, S. xantha.

In the following descriptions the categories will be used in this sense. It
will be seen that some species belong strictly to one particular category, but
that others can vary from one to another.

Measurements

Measurements of adcauline and abcauline hydrothecal lengths are taken
from the base of the hydrotheca to the tip of the respective marginal tooth
across any curvature which may be present. Adcauline measurements include
the perisarcal thickening at the base of the adcauline wall. All measurements are
exclusive of marginal regenerations. The diameter of the internode is measured
diagonally across the node.

Sertularella africana Stechow, 1919

Sertularella fusiformis: Warren, 1908: 295, fig. 5C, D.

Sertularella tenella: Stephenson, Stephenson & du Toit, 1937: 374 (listed).

Sertularella africana: Mallard, 1957: 207, figs. rol, 11F.

Records. West coast: A 382. LAM 2H, 9U. PP 1T. SB 168F. South coast:
CPR 7B. E134. S 65D (recorded by Stephenson ef al. 1937). SAMH 208, 218.
TRA 159C.

Remarks. Examination of Warren’s material from Park Rynie, Natal, in the
British Museum (reg. no. 22.3.6.20) confirms the identity of the above material.

Sertularella agulhensis, nov. sp.
Oop aeN

Holotype: TRA 151F, from south coast, (South African Museum registered
number, SAMH 413).

Description. Stem stiff, fascicled, reaching a maximum height of 3:2 cm.,
branching (and often rebranching) in a pinnate fashion, normally with 3
hydrothecae between the origins of successive branches. All branches in one
plane. Nodes oblique and distinct in terminal regions, often indistinct in older
parts of colony. Hydrothecae fairly closely set with the margin of one usually
overlapping the base of the next, one to each internode, the two rows in one
plane. Perisarc thick.

Hydrotheca adnate for about half adcauline length, distinctly annulated
for most of length (about 6 annulations), with convex adcauline wall and
straight or slightly concave abcauline wall. Abcauline side of margin produced
more than the rest. Margin perpendicular to axis of hydrotheca or tilted slightly

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 12. Sertularella spp.
A. Sertularella agulhensis n. sp.
B.-D. Sertularella capensis delicata n. subsp. B and C, solitary hydrothecae. D, an upright stem -
and a solitary hydrotheca.
F. Sertularella capensis capensis Millard, gonotheca.
E, G, H. Sertularella gilchristi n. sp.
(A from TRA 151F, B from SAMH 294, C from SCD 50Q, D from NAD 22F, E and G from
the holotype, F from SCD 37G, H from SCD 153X.)

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 37

towards adcauline side. 3 internal teeth: 1 abcauline, 2 latero-adcauline, the
latter often quite small.
Gonothecae absent.

Measurements (mm.)

Internode length = - ee ify - ak .. 0*40—0'59
diameter across node .. ie 23 is se .. 0'24-0'33
Hydrotheca, length abcauline a: #3 ns me .. 0°52-0°67
length adcauline, adnate part ie we pes + ,O°30-0'°97
length adcauline, free part Hes “ sis ft. .. 0°34-0°44
adnate part/adcauline length .. i ae 5 .. O°4I-0°51
diameter at mouth 34 ah bas ies te .. 0'23-0°28
maximum diameter... ee a: me Ae . + (O"9 1-087

Remarks. This species resembles S. mediterranea in the shape of the hydrotheca
and in the presence of 3 internal teeth, but differs from it in the fascicled stem
and annulated hydrotheca.

The structure of the hydrotheca strongly resembles that of S. richardsoni
Ralph, 1961, but S. agulhensis differs from it in the smaller size of the parts
and in the fascicled stem.

Sertularella arbuscula (Lamx., 1816)

Sertularia polyzonias: Busk, 1851: 118 (pp.)

Sertularia arbuscula?: Busk, 1851: 118.

Sertularella tumida Warren, 1908: 297, fig. 6A—C. Jarvis, 1922: 342.

Sertularella arbuscula var. quinquelaminata Leloup, 1934: 1, figs. 1-3.

Sertularella arbuscula: Millard, 1957: 208, figs. 1oB, 11C. Millard, 1958: 188. Millard, 1961: 204.

Records. West coast: A 381. SB 194K. WCD 34H, 81D. South coast: KNY 30N,
71E (reported by Day ef al. 1952). LIZ 16D. MB 8L, 15C, 64H. SAMH 155,
182, 204, 213, 216, 228, 234, 288, 318. SCD 37C, 61B, 75D, 85D, 117H, 153U,
169Z, 175H, 184P, 354E. TRA 35F, 38F, 42F, 92S, 156G.

Remarks. I have examined specimens of Jarvis’s material of S. twmzda from the
Indian Ocean and Warren’s material of S. tumida from Algoa Bay in the British
Museum of Natural History, and can confirm their identity as typical S.
arbuscula. Unfortunately Warren’s material from Park Rynie, Natal, (Warren,
1908, fig. 6B) was not available, but I feel that Stechow (1925, p. 485) was
wrong in assigning it to S. pulchra, which species is easily distinguished by the
adcauline annulations and 4 internal teeth. I have seen unmistakable material
of S$. arbuscula in which some of the hydrothecae are very little bent out as in
Warren’s diagram, and I have also seen gonothecae which are corrugated in
the distal region.

The presence of 2 extra internal teeth (var. quinquelaminata of Leloup) is a
common variation (found also in S. mediterranea), but usually occurs only in
some hydrothecae of a colony.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sertularella capensis capensis Millard, 1957
Fig. .12F

Sertularella capensis Millard, 1957: 210, fig. 10H.
Records. South coast: SAMH 272. SCD 37G, 79K.

Description. ‘Three colonies similar to holotype.

Female gonothecae (not previously described) borne on front of stem,
each arising just below the base of a hydrotheca. Spindle-shaped, distinctly
annulated throughout, with 4 (or occasionally more) marginal spines. No
external marsupium and eggs released within the gonotheca.

Remarks. See under subspecies delicata.

Sertularella capensis delicata nov. subsp.
Fig. 12B—D

Sertularella tenella: Hartlaub, 1901: 64, pl. 5 (fig. 24), (material from Algoa Bay).

Types and Records. Holotype: (NAD 22F) (South African Museum registered
number, SAMH 414). Other records: SAMH 294, SCD 50Q.

Description of holotype. Stem short, slender and geniculate; unbranched; bearing
a small number of hydrothecae (up to 8 observed) and sometimes only one;
usually annulated at base and in the region of nodes. |
Hydrotheca similar in shape and appearance to the nominate subspecies,
but with a smaller proportion of the adcauline wall adnate (less than half);
with one or two distinct ridged annulations; mouth rounded in section.
Gonophores absent.

Measurements (mm.) Holotype SAMH 294 -

Internode length ea Hs 4 46 .. 0°49-0°97
diameter across node vd ae te .. O*10—-0°13

Hydrotheca, length abcauline .. 0°34-0°45 0°40-0°45
length adcauline, adnate part 0°20—0°24. 0°20—0°22
length adcauline, free part 0'25-0°34 0°34-0°35
adnate part/adcauline length 0°37-0°46 0:36—-0-°39
diameter at mouth 0°25-0°29 0:°28—-0°30
maximum diameter 0°30—-0°37 0°41-0°43

Remarks. The samples other than the holotype consist almost entirely of solitary
hydrothecae, and stems bearing more than one hydrotheca are rare. Such
specimens are similar to the material from Algoa Bay described and figured
by Hartlaub in 1go1 and ascribed by him to S. tenella. With this diagnosis
I do not agree, as the hydrotheca of S. tenella has more annulations, a smaller
proportion of the adcauline wall adnate to the stem and a narrower, squared
mouth. The proportion between the diameter at the mouth and the abcauline
length is 0:59-0:79 mm. in the holotype of S. capensis delicata, as against 0-39
mm. in typical S. tenella (from Stechow’s measurements, 1923c, p. 186).

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 39

That the solitary hydrotheca is merely a young stage of an upright stem
is shown by the presence of both on the same hydrorhiza. Moreover, in the
solitary hydrotheca there is an extension of the internode along one wall
containing a bud of coenosarc, which will obviously form the continuation of
the stem at a later stage. This extension is visible in Hartlaub’s diagram, and
also in Leloup’s diagram of Thyroscyphus intermedius f. peculiaris (1935, fig. 15)
indicating that the latter is merely a young Sertularella. Sertularella campanulata
Warren, 1908, however, has no such extension and can justifiably be included
in a separate genus, i.e. Calamphora.

The subspecies delicata differs from the young stems and branches of the
nominate subspecies in the more slender stem with annulations and a hydrotheca
which has a smaller proportion adnate to the internode.

S. capensis shows some resemblances to S. gayi var. annulata Allman, 1888,
as revised by Billard, rg10, p. 10, fig. 3, but differs from it in the growth-form.

Sertularella congregata nov. sp.
Fig. 13A—D

Types. Holotype: SCD 254Q (South African Museum registered number,
SAMH 415). Paratype: SAMH 185. Both from South coast.

Description. Colonies reaching a maximum height of 6—7 cm. Stem stiff, thick
and fascicled, giving off alternate hydrocladia, which are all in one plane.
Long, fascicled branches, similar to the stem in structure, often replacing
hydrocladia. Segmentation not visible in fascicled part of stem and branches,
but oblique nodes usually visible in the distal, unfascicled parts (faint in
holotype, distinct in paratype). One hydrotheca to each internode and one
hydrocladium normally arising at the base of every third hydrotheca. Hydrothe-
cae on stem moderately closely set, the margin of each just overlapping the
base of the next on the opposite side.

Hydrocladia with nodes invisible in distal regions. Hydrothecae crowded,
the margin of one often reaching half-way up the length of the next on the
opposite side, and sometimes even to the base of the next on the same side. The
two rows of hydrothecae in the same plane. The hydrocladium usually starts
with several well-spaced hydrothecae separated by oblique nodes, and the
' crowding and disappearance of nodes becomes more pronounced towards the
distal end.

Hydrotheca adnate for half or more of adcauline length, then bent smoothly
outwards with margin tilted towards abcauline side. The sides parallel for
almost entire length, though usually narrowing very slightly near margin.
Marginal teeth low. No internal teeth. A pronounced perisarcal thickening
present on abcauline wall below margin, and continued as an annular ridge
for about half-way round hydrotheca. Margin often regenerated.

Gonotheca borne on hydrocladium and flattened against its surface, with
about 8 low annulations in distal region, and with 5 short, conical, marginal
spines.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

(/-

‘\

—

i

i \
®
B

F

Fic. 13. Sertularella spp.
A.-D. Sertularella congregata n. sp. A, gonotheca. B, portion of stem and hydrocladium. C and
D, a few hydrothecae from hydrocladium.
E.-G. Sertularella pulchra Stechow.
(A-C from holotype, D from SAMH 185, E-F from SCD 37H, G from SAMH 219.)

3

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 41
Measurements (mm.)
Holotype Paratype

Internode length (stem) , 0'40-0°42 0°35-0°46
Hydrotheca, length abcauline .. 0°39-0°46 0:37-0°45

length adcauline, adnate part 0°35-0'41 0°38-0'43

length adcauline, free part 0:28-0°36 0:26-0'33

adnate part/adcauline length 0°49-0'58 0:54-0°62

diameter at mouth O°19-O'2I 0*20-0°22

maximum diameter _" a .. O'24-0':27 0°25-0°27
Gonotheca (2 only), length .. om 2. ‘* 2°88

maximum diameter Ae ae uh .. 1'16-1-28

Remarks. This species shows a superficial resemblance to S. diaphana, S. lata,
S. quadridens and others, where the nodes on the hydrocladia occur at irregular
intervals, and where the nodes on the stem occur after every third hydrotheca.
The fundamental difference in S. congregata is that the nodes, when visible,
occur after every bydrotheca, both on the stem and hydrocladia, but on the
distal parts of the hydrocladia the nodes seem to be eliminated by the crowding
of the hydrothecae. S. congregata also differs from S. diaphana in the nature of
the gonotheca.

Sertularella dubia magna Millard, 1958
Fig. 14A-F

Sertularella dubia var. magna Millard, 1958: 189, fig. 7A.

Records. West coast: ?>LAM g30L (young colony). South coast: SAMH 186,
252, 282. SCD 79J, 82L, 85C, 108G, 112C, 320K.

Description. A number of colonies, all (except LAM 30L, which is unbranched)
with stiff, fascicled stems branching in one plane and in a pinnate manner.

Hydrothecae rather variable in structure and appearance—some are
comparatively short and fat with well-marked abcauline thickenings as
illustrated by Billard, 1907, fig. 3 and Millard, 1958, fig. 7A; others are more
slender with distinct corrugations on the adcauline wall and poorly developed
abcauline thickenings; and still others show a tendency for an elongation of
the abcauline marginal tooth so that the margin becomes tilted towards the
adcauline side. Internal bydrothecal teeth never present.

Gonotheca (not previously described) distinctly annulated in distal
region, with a short terminal neck bearing 2—7 small spines (usually 5-6).
Arising opposite the base of a hydrotheca. All female where the sex could be
definitely ascertained. One colony (part of SCD 112C) with practically smooth
gonothecae (possibly male).

Measurements (mm., excluding LAM 30L, which is a doubtful record)

Internode length + as is ee ny ss .. 0*40-0°72
diameter across node .. 18 - ae i .. 0*20-0°39

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hydrotheca, length abcauline ie Ae * a .. 0°48-0°65
length adcauline, adnate part .. ye Be a ». 0°32-0°53
length adcauline, free part th a T ». O°31-0°46
adnate part/adcauline length .. ie) ue ee »» 0°45-0°58
diameter at mouth a ch pa mp te .. 0'22—0°30
maximum diameter... a Me Pa ae .. O'31-0°43

Gonotheca, length .. se side es, My its .. 2°67-3°94
maximum diameter... i oh ref 14 -. 1°06—1-48

Remarks. The variability of the shape of the hydrotheca in this species has been
the cause of some hesitation in identification, and there appear to be 3 common
forms: (i) the short, fat hydrotheca with the margin tilted towards the abcauline
side and a well-developed abcauline thickening, as originally, described (fig..
14A, B), (ii) a longer hydrotheca with thinner perisarc and distinct adcauline
corrugations (fig. 14C), and (iii) a hydrotheca with the margin tilted towards
the adcauline side due to elongation of the abcauline marginal tooth (fig. 14F).
These forms cannot be separated as distinct species or subspecies, as inter-
grading forms occur, and specimens in the same sample may vary from form (i)
to form (ii), or from form (ii) to form (i). The macroscopic appearance of the
colony and the structure of the gonotheca is identical in all cases.

All the specimens described have greater dimensions than those quoted
by Billard, 1907, for the nominate subspecies, and can thus be included in
the subspecies magna. So far no overlap has been observed.

The form with adcauline thecal corrugations shows resemblances to
S. gayi var. robusta Allman, 1874, but the gonothecae are different (those of
S. gayt possessing a bilabiate aperture) and the measurements are not so great
as those quoted by Billard in 1906 (p. 185) for var. robusta.

Some of the specimens also show resemblances to S. crassicaulis (Heller),
but the latter species has a dichotomously branched stem.

Sertularella falsa Millard, 1957

Sertularella ?tumida: Day, Millard & Harrison, 1952: 404 (listed).
Sertularella falsa Millard, 1957: 211, figs. 1oF, 11D.

Records. South coast: KNY 70F (reported by Day e¢ al. 1952).

Sertularella flabellum (Allman, 1886)
Sertularella flabellum: Millard, 1957: 212, figs. 1oG, 11G. Millard, 1958: 190.

Records. West coast: TB 21D. WCD 12B. South coast: MB 47U. SAMH 171,
178, 205, 217, 263, 312. SCD 5B, 29M, 36Y, 52Q, 56S, 85E, 106P, 112B,
1535, 175J, 206N, 265K, 2908, 320J. TRA 595, 92P, 151J.

Sertularella fusiformis (Hincks, 1861)

Sertularella ellisii f. ellisti: Picard, 1956: 264, fig. 3d, e.
Sertularella fusiformis: Millard, 1957: 213, figs. 1oC—D, 11E.

a?

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 43

Fic. 14. Sertularella dubia magna Millard.
A.-D., F. Hydrothecae from various colonies to show variation in shape

E. Gonotheca.
(A from PF 12308H (False Bay), B from SAMH 252, C and E from SAMH 282, D from SCD
112C, F from SCD 85C.)

se

a

a ee eek PE

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Records. West coast: B 105, 114B. CP 336B. PP 1S, 4M. South coast: MB 24W.
SAMH 231, 271. SCD 37F, 61E, 75E.

Remarks. Picard in 1956 has included S. fusiformis in SS. ellisiz, but does not seem
to have considered the question of the number and position of internal hydro-
thecal teeth. In his diagrams he shows internal teeth situated immediately
below the marginal teeth, which is certainly not the case in South African
material allocated to S. fusiformis, where internal teeth alternate with the
marginal teeth. Stechow, 1923c, described S. ellisiz as possessing 3 small internal
teeth (thus differing from S. mediterranea where the teeth are large), and until
more work has been done on the variability of this character. I prefer to retain
S. fusiformis as a separate species. The number of internal teeth in the latter is
known to be variable, but as has been pointed out before (Millard, 1957, p. 214;
1958, p. 187) the position in South African material at any rate is constant,
and differs from that in S. ellzszz.

Sertularella gilchristt nov. sp.
Fig, 1ak, (Gib
Types and Records. Holotype: SCD 85J (South African Museum registered
number SAMH 416). Other records: SCD 153X. Both from south coast.

Description of holotype. Stem thick, fascicled, reaching 5-0 cm. in height, branch-
ing and rebranching profusely in an irregular pinnate fashion, but not necessarily
in one plane. General effect bushy. ‘Terminal branches not geniculate, with
distinct oblique nodes and sometimes an annulation above each node. Inter-
nodes of variable length, each with one hydrotheca.

Hydrotheca swollen below and narrowing very markedly to margin;
adnate for a little less than half adcauline height; symmetrical, or (more
commonly) bent outwards with margin tilted towards abcauline side; annu-
lated, with 3 or more annulations passing all round hydrotheca or (less
commonly) with annulations indistinct or confined to adcauline surface. Margin
with 4 teeth, of which the abcauline one may project slightly more than the
others. 3 well-developed internal teeth, 1 abcauline and 2 latero-adcauline.

Gonotheca fusiform, annulated in distal region, with 3 or 4 spines at the
distal end of the slender terminal region.

Measurements (mm.) Holotype SCD 153X
Internode length ne pa at ate .. 0°28-0°45 0°39-0°53
diameter across node A ee =i .. 0719-05225 eres
Hydrotheca, length abcauline .. - sie .. 0°33-0°41 0°36-0-41
length adcauline, adnate part... sa .. 0°22-0-27) 0-22-37
length adcauline, free part mt ay .. 0'24-0'34 0:26-0°35
adnate part/adcauline length .. oh .. 0'40-0:48 0°40—0°49
diameter at mouth ae sis be .. O'II-O'14 O°12-0°13
maximum diameter Ec, “fh ee .. Q°2I-0'24. 0°22-0°25
Gonotheca, length oo a ay oN .. 1°84-1-86

maximum diameter aa a he Hd 0:97

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 45

Remarks. This species shows strong resemblances to S. robusta Coughtrey, 1875
(see Trebilcock, 1928: 16, and Ralph, 1961: 824), but differs from it in the
fascicled and freely branching stem, and also to a lesser extent in the shape
of the hydrotheca which has a decidedly narrower mouth.

Sertularella goliathus Stechow, 1923
Sertularella goliathus: Millard, 1957: 215, figs. 10A, 11A.
Records. West coast: WCD 7P. South coast: SCD 106Q. TRA 37N.

Sertularella mediterranea mediterranea Hartlaub, 1901

Sertularella mediterranea: Millard, 1957: 215, figs. 1oE, 11B. Hamond, 1957: 316, fig. 24. Millard,
1958: 190. Vervoort, 1959: 272, fig. 33a.

Sertularella ?gaudichaudi: Day, Millard & Harrison, 1952: 404 (listed).

Sertularella ellisii f. mediterranea: Picard, 1956: 264, fig. 3b.

Records. South coast: A 123, 384B. KNY 57J (recorded by Day et al. 1952).

SAMH 206.

Sertularella mediterranea asymmetrica Millard, 1958

Sertularella mediterranea Hartlaub var. asymmetrica Millard, 1958: 191, fig. 7B.
Records. South coast: CPR gD. SAMH 270. SCD 60Y.

Sertularella megista Stechow, 1923

Sertularella polyzonias f. robusta Kirchenpauer, 1884: 38.

Sertularella polyzonias var. robusta: Hartlaub, 1901: 88, pl. 5 (fig. 1). Stechow, 1925: 479.
Sertularella megista: Stechow, 1925: 480, fig. 36. Millard, 1957: 217, figs. 10L, 11J.

Records. West coast: WCD 12A, 56L. South coast: AFR 945M. MB 47V, 52H,
64J. SAMH 181, 246, 310, 316. SCD 5D, 37E, 52R, 85F, 96C, 108E, 153T,
175K, 239C, 254R, 265C, 290R, 320].

Description. Numerous colonies from all round the coast, the tallest reaching a
height of 10:3 cm. Most colonies are unfascicled and sparsely branched, and
individual stems may reach a length of over 7 cm. without branching. Only the
sturdiest colonies are fascicled, and these only to a small extent and in the basal
region. The stems, thus, usually have a flexuous appearance, surprisingly so
for a species with so thick a perisarc.

With the abundant material available it is apparent that there is great
variation in length of internode and size of hydrotheca. New measurements
are thus included to illustrate the range of variation in the species.

The hydrotheca is characterized by a sharp bend in the adcauline wall
at the point where it separates from the stem, but the angle of the margin is
influenced by the amount of bending and the length of the abcauline marginal
tooth, which may be produced. Normally the margin is perpendicular to the
axis, but when the bend is marked (and the angle within the hydrotheca lies
between 110° and 120°) it is tilted towards the abcauline side, and when the

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

bend is less marked (with the internal angle over 125°) and the abcauline tooth
produced it is tilted towards the adcauline side.

Measurements (mm., complete range, including False Bay material described
in 1957).

Internode length by ue a om ws x ». O°7I-1°52
diameter across node .. My 4 si if: .. 0*40-0°82
Hydrotheca, length abcauline rae ay ws ea .. 0°80-1-28
length adcauline, adnate part ie + 4 .. 0°67-1-08
length adcauline,'free‘part — . . am ay: a .. 0°34-0°72
adnate part/adcauline length .. ae i My ». O°55-0°71
diameter at mouth ys iy ae Be i! .. 0°33-0'52
maximum diameter bay ie ne bs Pe .. 0°47-0°72
Mature gonotheca, length .. et Mg S a ve, | Rees
maximum diameter... “te = ie Rs .. 0°85-1-90

Remarks. 1 have included in the synonymy of this species Kirchenpauer’s
material of S. polyzonias f. robusta from the Cape of Good Hope. Kirchenpauer’s
type material unfortunately no longer exists, but was examined and illustrated
by Hartlaub in 1901, and I agree with the latter author that it should be
removed from S. polyzonias. The hydrotheca has the typical angle in the
adcauline wall characteristic of S. megista, and the size is within range.

Sertularella polyzonias (Linn., 1758)
Sertularella polyzonias: Millard, 1957: 217, figs. 10J, 11H. Millard, 1958: 191. Millard, 1961: 205.

Records. West coast: TB 10, 12A, 21G. South Coast: MB 47W. SAMH 175.
SCD 37J, 85G, 112D, 153W, 175L.

Sertularella pulchra Stechow, 1923
Fig. 13E-G

Sertularella pulchra Stechow, 1923b: 113. Stechow, 1925: 485, fig. 39.
Records. South coast: SAMH 219, 313. SCD 37H, 154L, 296].

Description. Fascicled stems reaching a maximum height of 4-8 cm. and bearing
alternate hydrocladia, which are usually very regular in arrangement, one
arising below the base of every third hydrotheca. The 2 rows of hydrocladia
in one plane.

Hydrothecae generally as described by Stechow, though in most colonies
a few of the hydrothecae are bent gently away from the stem, with the adcauline
wall slightly convex and the abcauline wall slightly concave (fig. 13 F). Asa
result the margin is perpendicular to the axis or tilted towards the abcauline
side. Adcauline striations very obvious, and only rarely absent. 4 internal teeth,
alternating with the marginal teeth, sometimes incomplete in young parts of
the colony.

Gonothecae of 2 types. Empty ones, presumably male, as described by

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA

Stechow, with narrow distal end and 4 marginal spines. Female similar, but

with wider distal end and 5-6 short marginal spines (fig. 13 E).

Stechow’s

material from

Measurements (mm.) Stmon’s Bay
Internode length .. 0°43-0°82 0°49—-0°63
diameter across node 0°20-0°42 0:16-0:34
Hydrotheca, length abcauline 0:58-0°78 0:5 8-0°72
length adcauline, adnate part 0:32-0°46 0:35-0'44
length adcauline, free part .. 0°36-0°46 0°39-0°56
adnate part/adcauline length 0°44-0°55 0'40-0°53
diameter at mouth .. 0°21-0°28 =: 0:21-0:28
maximum diameter 0:29-0'38 0°31-0:36
Gonotheca, length 2°95-3°42 2°33-29 84
maximum diameter 0-99-1°63 =: 0'93-1 31

Remarks. 1 have seen two mounted slides of Stechow’s material from Simon’s
Bay (loaned by the Munich Museum) which confirm the identity of this
material. Measurements taken from Stechow’s material are included above for
comparison.

Stechow includes in S. pulchra part of Warren’s material described under
the name of S. tumida (material from Park Rynie, 1908 fig. 6B). With this I
do not agree, as Warren shows no striations on the hydrothecae and only 3
internal teeth. His material can probably be included in S. arbuscula.

Sertularella striata Stechow, 1923
Fig. 15

Sertularella striata Stechow, 1923: 10. Stechow, 1925: 470, fig. 30.
Records. South coast: LIZ 7Z. MB64K. SCD 85H.

Description. Stems reaching a maximum height of 1-1 cm., usually unbranched,
but occasionally giving off 1 or 2 small branches which arise immediately
below a hydrotheca. Details of structure and measurements as described by
Stechow.

_ Gonotheca (not previously described) arising from stem opposite the base
of a hydrotheca, annulated throughout, or with annulations becoming indistinct
in basal third. Margin with 3 or 4 minute spines. Female with external
marsupium.

Present Stechow’s

Measurements (mm.) material — material

Internode length hs : is AN .. 0°30-0:76 0°45-0°77
diameter across node wi sm oF .. O°10-0'16 O-10—-0°15

Hydrotheca, length abcauline .. on Me .. 0°36-0°48 0°41-0°47

length adcauline, adnate part .. ah .. 0*20-0'26 0:21-0:26

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

length adcauline, free part oe SPARES .. 0°18-0'31 0°25-0°34

adnate part/adcauline length Se .. 0°39-0°56 0°40-0°49

diameter at mouth 3% ve i .. O*15-O°21 O*17-0'23

maximum diameter oe A hy .. 0°23-0°30 0:23-0°30
Gonotheca, length A ay Ab Ry) .. 1°35-1°79

maximum diameter As S As .. 0°86-0°95

Remarks. The identity of this material was confirmed by comparison with a
mounted slide of Stechow’s material from the Agulhas Bank, kindly loaned by
the Munich Museum. The measurements of the latter are included above for
comparison.

fpr eee a TNT

Fic. 15. Sertularella striata Stechow.
A. and C. Hydrothecae from different colonies.
B. The gonotheca.
- (A and B from SCD 85H, C from LIZ 7Z.)

The species shows variability in the length of internode, which may be
shorter or longer than that illustrated by Stechow; in the degree of annulation
on the hydrothecal walls, which may be less marked; in the proportion of the
adcauline thecal wall adnate to the stem; and in the angle of the thecal margin.
The latter may be perpendicular to the axis as illustrated by Stechow, but
may also be tilted towards the adcauline side. This is the case in some of the
hydrothecae on Stechow’s slide, and in most of the hydrothecae in the present
material.

This species is closely related to S. africana, differing from it in the greater
number of more distinct annulations which extend over the full length of the
hydrotheca.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 49

Sertularella xantha Stechow, 1923
Sertularella xantha: Millard, 1957: 218, figs. 1oK, 111.

Records. West coast: TB 12B. WCD 30U. South coast: SAMH 146, 154, 220,
325, 334, 348. SCD 33C, 103E, 104E, 108F, 113F, 114F, 115N, 122W, 126K,
141J, 145B, 184Q, 191R, 219W, 239D, 258P, 276T, 290Q, 322G, 324], 345A.
TRA 33L, 35A, 38B, 42E, 56V, 92T.

Sertularia distans gracilis Hassall, 1848

Sertularia heterodonta Ritchie, 1909: 79, fig. 4. Jarvis, 1922: 339.

Sertularia distans var. gracilis: Billard, 1925: 175, fig. 33. Leloup, 1935: 47, figs. 28, 29. Millard,
1957: 221, fig. 12. Millard, 1958: 193. Pennycuik, 1959: 197.

Records. South coast: BRE 60R. CPR 7C. LIZ 13E. MB 24R. SAMH 266.

SCD 37Y, 50M, 85Q, 112E.

Description. A number of colonies, none taller than 0-8 cm., all with unbranched
stems, except in a few instances where side-branches arise from within hydrothe-
cae. Distance between consecutive pairs of hydrothecae very variable, and
internodes sometimes very long and slender, as illustrated by Ritchie, 1909, for
S. heterodonta. Interna) pegs of perisarc in lower part of hydrotheca very charac-
teristic, and internal teeth in distal region below margin also common—I
abcauline (fairly common), or 1 abcauline and 2 latero-adcauline (more rare
and in occasional hydrothecae only).

Gonothecae present in 2 colonies, as illustrated by Leloup, 1935; female
with external marsupium.

Remarks. Examination of Ritchie’s type material of S. heterodonta (on loan to
the British Museum) showed that this species is distinguished from the common
form of S. distans gracilis only by the presence of 3 internal hydrothecal teeth,
longer and more slender internodes, and slightly smaller hydrothecae. The last
2 characters can be attributed to variations in growth-form which are paralleled
in the South African material. The presence of a single abcauline internal
tooth has been reported in S. distans gracilis by a number of authors, and the
presence of 2 extra latero-adcauline internal teeth (which were not present
in all the hydrothecae of the type material) is here reported in South Africa.
I therefore include S. heterodonta Ritchie as a synonym for S. distans gracilis.

Sertularia marginata (Kirch., 1864)
Sertularia marginata: Millard, 1957: 224, fig. 13. Ralph, 1961: 785, fig. 12a-g (synonymy).
Records. South coast: SCD 305D.

Description. A young colony consisting of 5 stems reaching a maximum height
of 0-6 cm., of which one bears 1 branch, another 2, and the rest are simple.

Sertularia turbinata (Lamx., 1816)

Sertularia loculosa: Bale, 1884: pl. 4 (figs. 5, 6), pl. 9 (fig. 12), pl. 19 (fig. 9). Bale, 1913: 121,
pl. 12 (figs. 7, 8). Warren, 1908: 306, fig. 8. Jarvis, 1922: 340.

50 | ANNALS OF THE SOUTH AFRICAN MUSEUM

Sertularia brevicyathus : Nutting, 1904: 60, pl. 6 (figs. 1, 2). Jarvis, 1920: 338, pl. 24 (fig. 6).
Tridentata acuta Stechow, 1921: 231. Stechow, 1923c: 207.
Sertularia turbinata: Billard, 1925: 177, fig. 34. Millard, 1958: 197, fig. 8B. Vervoort, 1959: 275;

figs. 35, 36.
Sertularia restricta Totton, 1930: 205.

Sertularia acuta: Millard, 1958: 192, figs. 8A, F.

Records. South coast: MB 58E. ‘SCD 85S.

Description. Two small colonies reaching a maximum height of 0-7 cm. One of
them (MB 58E) is an old colony with thick perisarc and smoothly worn hydro-
thecal margins; the whole surface is thickly overgrown with epiphytes. In both
colonies the nodes tend to be indistinct, particularly near the proximal end of a
stem.

Kemarks. Up to the present time S. acuta (Stechow) has been held to be separate
from §. turbinata (Lamx.), although Billard (1925: 178), was dubious about
the matter, remarking that the former species differs from the latter only in
the shorter internodes, and in the shorter hydrothecae which narrow more
abruptly to the margin.

The two samples in this collection are intermediate between the material
of S. acuta and S. turbinata previously described (Millard, 1958) in internode
length and abcauline thecal length, and the 4 samples together form a perfect
gradation from one extreme to the other. It is thus no longer possible to keep
the two species separate, and S. acuta must be considered as a somewhat stunted
growth-form of S. turbinata. Selected measurements (in mm.) of the 4 samples
are included for comparison:

Fydrotheca, Hydrotheca,
Internode length diameter at

length abcauline margin
RHB 52C 0°39-0'54. 0°16—-0:21 0°09—-0'12
SCD 85S 0°52—0:62 O°19Q—0:22 0°0Q—-0'II
MB 58E 0°55-0°76 0°22-0'32 0°13—0°16
PZ 13B 0:66-0:85 0'25-0°31 O°12—-0-17

In all 4 samples the hydrotheca is very similar in appearance, and grades
evenly from the form illustrated for S. acuta (Millard, 1958: fig. 8A) to that of
S. turbinata (Millard, 1958: fig. 8B).

The gonothecae of S$. turbinata have now been described by Vervoort
(1959: fig. 36 b, c) and are exactly similar to those of S. acuta (Millard, 1958:
fig. 8F; Warren, 1908: fig. 8D), except for the presence of 2 minute distal
spines in the former.

It was previously thought (Millard, 1958: 198) that the nature of the
nodes was a distinguishing character between S. acuta (straight) and S. turbinata
(oblique). But the new materia] has shown that this is a variable character
and the nodes may be straight, oblique or invisible, and further that both
straight and oblique nodes may occur on the same stem. This is discounting the
hinge-joints, which always form the termination of an extra athecate internode
and which occur quite irregularly.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA pal

S. turbinata is closely related to S. ligulata Thornely, and the difference
between them is largely one of shape, which is obvious to the eye, but which is
not shown up at all clearly by the measurements, due mainly to the tendency
for variation between the proximal and distal regions of a stem in both species.
In S. ligulata the members of a pair of hydrothecae are more erect and tend
to diverge at a higher level than in S. turbinata. As a result the hydrothecae
of the former are in general contiguous and adnate for a greater length, and
the maximum diameter across a pair tends to be less. S. ligulata also has a
slightly wider thecal mouth and lower marginal teeth (Millard, 1958: figs.
8B, 9A, B). If these 2 species are to be kept separate, the presence of a ligula
in S. ligulata cannot be used as a specific character, as it is also clearly evident in
a mounted specimen of S. turbinata (MB 58E). As a rule specimens are not
sufficiently well preserved to determine the presence or absence of this character.

Symplectoscyphus arboriformis (Markt., 1890)

Sertularella arboriformis Marktanner-Turneretscher, 1890: 228, pl. 4 (fig. 5). Stechow, 1912: 358,
fig. C.

Records. West coast: SAMH 409. WCD 81L. South coast: SAMH 156, 172, 184,

Se7eett, 17. oGD 5C, 37D, 61C, 75C, 138K, 290T. TRA 92Q.

Description. Colonies reaching a maximum height of 11-0 cm., some with

gonothecae. Structure agreeing perfectly with Marktanner’s description.

Stechow’ s

Measurements (mm.) material
Internode length 6 As, ti oe .. 0:60-0:80 0:°51-0°78
diameter across node Ry ae Wf .. O°17-0°33 O*1Q-0°42
Hydrotheca, length abcauline .. ¥ - .. 0'34-0'47 0:°46-0-63
length adcauline, adnate part... fis .. O°38I-0°40 0-28-0°43
length adcauline, free part ne ie .. 0°38-0°46 0°35-0°50
adnate part/adcauline length .... aN .. 0°42-0°51 0°36-0°53
diameter at mouth a ie be .. 0°28-0'38 0°25-0°35
Gonotheca, length Ne Sah we i, .. 1°35-1°92 1°46-1°75
maximum. diameter acs is ve .. 0°71-0:96 0-81—0-96

Remarks. ‘Two slides of Stechow’s material from Algoa Bay were examined by
courtesy of the Munich Museum. The measurements are included above for
comparison.

Symplectoscyphus macrogonus (Treb., 1928)

Sertularella macrogona Trebilcock, 1928: 11, pl. 1 (fig. 4).

Symplectoscyphus macrogonus: Millard, 1957: 219. Ralph, 1961: 708, fig. 14 a, b.

Records. West coast: BB 13P. CP 379, 650B. LAM eG, 7M, oT, 13], 145, 18L,
23M, 30K, 35K, 40P, 41G, 46M, 59B. PP 1U. SB 150D, 161Y, 194L, 253D.
TB 11, 21F. WCD 81J, tooT. South coast: BMR 23N. SAMH 265. SCD
179C.

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

Thyroscyphus aequalis Warren, 1908
Fig. 16
Thyroscyphus ramosus: Billard, 1907: 342.
Thyroscyphus aequalis Warren, 1908: 344, pl. 48 (figs. 38-40), fig. 23. Jarvis, 1922: 337. Millard,
1958: 199.
Thyroscyphus regularis : Ritchie, 1910: 811, pl. 77 (fig. 7). Jaderholm, 1923: 5. Stechow, 1925: 463.
Cnidoscyphus aequalis: Splettstésser, 1929: 82, 124, figs. 78-82. Kramp, 1947: 13.
kecords. South coast: SAMH 173, 251, 268, 350, 379. SCD 2oL, 37A, 61D,
79L, 84J, 94F, 117J, 153R, 169X, 2548, 265D. TRA 23B, 35D, 38D, 56T,
g2A.

Description. Numerous samples, some consisting of a few fragments only, the
largest reaching 40 cm. in height. Stem weakly fascicled at base in larger
colonies only, branching in an irregularly alternate manner. Nodes distinct or
only faintly indicated. Hydrotheca-bearing apophysis without basal septum.
Hydrothecal pedicel about half width of apophysis, spirally annulated to a
varying degree, often regenerated.

Hydrotheca of rather variable shape and size, usually expanding to margin
and practically symmetrical, but often with narrower mouth and slightly
bulging adcauline wall. Margin with 4 teeth and perisarcal thickening below
edge. Diaphragm in form of annular thickening of perisarc, which is better
developed on adcauline side and often invisible on abcauline side. An internal
ridge of perisarc sometimes present about one-third of height of hydrotheca for
attachment of annular fold of hydranth. Two batteries of large rod-shaped
nematocysts present in ectodermal lining of hydrotheca in adcauline and
abcauline position respectively.

Gonotheca (male) elongated, widening towards distal extremity, which is
obliquely truncated, smooth or roughly corrugated. Containing one large
gonophore, which is extruded into an external marsupium when mature.
Measurements of hydrothecae (mm.). Samples in which typical nematocyst batteries

could be recognized indicated by *.
Height Diameter

Diameter
from at ———.
diaphragm mouth height
TRA 35D* a My ay -. I*O05-1°2I 0°90-0-99 0°74-0:92
TRA 36D": z% ee a -. I-05-1°21 0°76—0-99 0:72-0°82
(AFR 1028.0.A __.. a ei .. 0°78-1:01 0:65-0:77 0-69-0°86)
TRA 23B* .. A 4 a, .. 1:07-1:28 0-80-1-13 0-68-0-98
SAMH 173* Ef ie ie -. I‘OI-1:28 0:83-0:93 0:68-0:92
SAMH 350* Me oy: ha -. 0°92-1'25 0:76-0:90 0-62-0:93
(NAD 1W* ve - y -. I*00-1'02 0-61-0-74 6°61-0-73)
SCD 2oL* .. 0*79-I°00 0°55-0°75 0:58-0:93
SCD 37B 0:83-1:05 0:44-0:64 0:47-0:64
From Jarvis. . 0°85-0°9 0°75 0-83-0°88
From Jaderholm I*I-1°2 0-9 0+75-0°82

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 53

M

rae eich een eerie armen oo a

Fic. 16. Thyroscyphus aequalis Warren.

A.-K. Hydrothecae and pedicels from various colonies to show variation.

L.—M. Male gonothecae, M with an external marsupium.

(A from TRA 35D; B and C from AFR 1028.0.A (Natal); D and E from TRA 23B; F and L
from SAMH 173; G from SAMH 350; H, I and M from SCD 29L; J and K from SCD 37A.)

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

From Splettstésser . . ny S .. O*°8-I'I5 0*75-0-92

From Warren ae zis He Be, 1-18 0°79 0°67
From Ritchie Ws os Ht ie 0-96 0°63 0-66
From Stechow Hp ite AF Np 0:96 0:60 0-62

Remarks. T. aequalis is closely related to T. torresi (Busk) and according to
Splettstésser and Kramp the latter is distinguished from the former by (i) the
presence of a perisarcal septum at the base of the hydrothecal apophysis, (ii) the
presence of a single battery of large nematocysts situated in the ‘Deckelplatte’
rather than in the ectodermal lining of the hydrotheca, (iii) the shape of the
hydrotheca which has a narrower mouth and somewhat protruberant adcauline
wall, and (iv) the hydrothecal pedicel which is not annulated.

In none of the South African material is there a septum at the base of the
apophysis, and in all specimens where nematocysts are preserved the batteries
are arranged as described by Warren, apparently indicating that all the material
is conspecific. Yet there is so much variation in the shape and size of the
hydrotheca and in the nature of the pedicel that it appears that characters
(ii) and (iv) are of less systematic value than previously supposed (see fig. 16).

The shape of the hydrotheca varies from a large form with a wide mouth
and practically symmetrical sides to a smaller form with a narrower mouth
and protruberant adcauline wall (see diameter/height ratios above), but all
intermediate stages are present and nowhere can a definite dividing line be
placed between the two forms. Sometimes both forms occur in the same sample.
A similar variation is evident in the literature among records assigned by
Splettstdsser to Cnidoscyphus aequalis; thus, the material of Jarvis and Jaderholm
belongs near the top of the series and that of Stechow near the bottom.

The hydrothecal pedicel in T. aequalis is said to be ‘annulated’. This
annulation, when well developed, is due to the presence of a groove running
spirally around the pedicel and completing a maximum of 2} turns, usually
with half a turn more on the abcauline than on the adcauline side. However,
the development of this groove varies considerably (and often within the same
colony), and many pedicels would come into the category of ‘not annulated’
(fig. 16 C). The amount of annulation bears no relation to the size and shape
of the hydrotheca, but may possibly be related to the age, as there is a tendency
for more distinct annulation in older parts of the colony. In old colonies, how-
ever, the spiral grooving is usually obscured by successive regenerations of
the pedicel which are marked by deep transverse septa across its width.

The gonotheca, of which only the male has been observed, is similar to
that described by Ritchie, and is unique among male hydroids for the extrusion
of the sexual products into an external marsupium.

SUMMARY

In the three families of hydroids considered here, a total of 50 species is
recorded from the south and west coasts of South Africa. Of these 12 belong
to the family Lafoeidae, 4 to the Syntheciidae and 34. to the Sertulariidae.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 55

6 new species are described, namely Hebella urceolata, <ygophylax enigmatica,
Salacia disjuncta, Sertularella agulhensis, Sertularella congregata and Sertularella
gilchristi, and one new subspecies, namely Sertularella capensis delicata. 7 of the
other records are new to South Africa.

REFERENCES

Auiman, G. J. 1874. Report on the Hydroida collected during the expeditions of H.m.s. Porcupine.
Trans. Zool. Soc. Lond. 8: 469-481.

Auitman, G. J. 1877. Report on the Hydroida collected during the exploration of the Gulf
Stream by L. F. De Pourtalés, assistant United States Coast Survey. Mem. Harv. Mus.
comp. Kool. 5: 1-66.

ALLMAN, G. J. 1886. Description of Australian, Cape and other Hydroida, mostly new, from the
collection of Miss H. Gatty. 7. Linn. Soc. (Xool.) 19: 132-161.

Autman, G. J. 1888. Report on the Hydroida dredged by u.m.s. Challenger during the years
1873-76. Part I1.—The Tubularinae, Corymorphinae, Campanularinae, Sertularinae and
Thalamophora. Rep. Voy. Challenger 1873-76. 23: 1-90.

Bae, W. M. 1884. Catalogue of the Australian hydroid zoophytes. Sydney: Australian Museum.

Bate, W. M. 1913. Further notes on Australian Hydroids, II. Proc. roy. Soc. Victoria (n.s.) 26:
114-147.

Bittarp, A. 1906. Hydroides. Expéd. sci. ‘Travailleur’ et du ‘Talisman’ 8: 153-244.

BILLARD, A. 1907. Hydroides de Madagascar et du Sud-Est de l'Afrique. Arch. Zool. exp. gén. (4)
7? 335-396.

BILLARD, A. 1910. Revision d’une partie de la collection des Hydroides du British Museum.
Ann. Sci. nat. zool. (9) 11: 1-67.

BittarpD, A. 1918. Notes sur quelques espéces d’hydroides de l’expédition du ‘Siboga’. Arch.
Kool. exp. gén. 573 21-27.

BILLARD, A. 1925. Les Hydroides de l’expédition du Siboga. II. Synthecidae et Sertularidae.
Siboga Exped. monogr. 7b: 117-232.

Brocn, H. 1918. Hydroida, part II. Danish Ingolf-Exped. 5 (7): 1-205.

Brocu, H. 1933. Zur Kenntnis der adriatischen Hydroidenfauna von Split. Skr. norske Vidensk-
Akad., Mat-natur. Kl. 1933 (4): 1-115.

Busk, G. 1851. A list of Sertularian Zoophytes and Polyzoa from Port Natal, Algoa Bay, and
Table Bay, in South Africa; with remarks on their geographical distribution, and obser-
vations on the genera Plumularia and Catenicella. Rep. Brit. Ass. 1850: 118-120.

Day, J. H., Mittarp, N. A. H. & Harrison, A. D. 1952. The ecology of South African estuaries.
Part III. Knysna: a clear open estuary. Trans. roy. Soc. S. Afr. 33: 367-413.

Fraser, C. McL. 1944. Hydroids of the Atlantic coast of North America. Toronto: University Press.

Hammon, R. 1957. Notes on the Hydrozoa of the Norfolk coast. 7. Linn. Soc. (Zool). 432 294-324.

Hart avs, C. 1901. Revision der Sertularella-Arten. Abh. Naturw. Hamburg 16: 1-143.

JADERHOLM, E. 1923. Hydroids from West and South Africa. Medd. Géteborgs Mus. Zool. 26: 1-7.

Jarvis, F. E. 1922. The Hydroids from the Chagos, Seychelles and other islands and from the
coasts of British East Africa and Zanzibar. Trans. Linn. Soc. Lond. Zool. 18: 331-360.

KIRCHENPAUER, G. H. 1864. Neue Sertulariden aus verschiedenen Hamburgischen Sammlungen,
nebst allgemeinen Bemerkungen tiber Lamouroux’s Gattung Dynamena. Verh. K. Leopold-
Carol. deuts. Akad. Naturf. Dresden 31 (3): 1-16.

KIRCHENPAUER, G. H. 1884. Nordische Gattungen und Arten von Sertulariden. Abh. Naturw.
Hamburg 8 (3): 1-54.

Kramp, P. L. 1947. Hydroids collected by the ‘Skagerak’ Expedition in the Eastern Atlantic
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Letovp, E. 1934. Trois hydropolypes de la Baie de la Table, Afrique Australe. Bull. Mus. Hist.
nat. Belg. 10 (19): 1-8

Letoup, E. 1935. Hydraires Calyptoblastiques des Indes Occidentales. Mem. Mus. Hist. nat.
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Letoup, E. 1937. Hydropolypes et Scyphopolypes recueillis par C. Dawydoff sur les cétes de
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MARKTANNER-TURNERETSCHER, G. 1890. Die Hydroiden des k.k. naturhistorischen Hofmuseums.
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56 ANNALS OF THE SOUTH AFRICAN MUSEUM

Minzarp, N. A. H. 1955. New species of Hydrozoa from South Africa. Ann. S. Afr. Mus. 41:
215-222.

Mitiarp, N. A. H. 1957. The Hydrozoa of False Bay, South Africa. Ann. S. Afr. Mus. 43:
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Miuiarp, N. A. H. 1958. Hydrozoa from the coasts of Natal and Portuguese East Africa. Part I.
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Mixuarp, N. A. H. 1961. A report on Busk’s collection of South African Hydroids. Ann. Mag.
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The Plumulariidae. Ann. S. Afr. Mus. 46: 261-319.

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269-355.

4

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TABLE oF CoNnTENTs and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 42 in. X 7 in. (74 in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (black type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

Smiru, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication ; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmirH, C. D. 1954. South African plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).
Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:

Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.

When another species has been called by the same name:

[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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MUS. COMP. ZOOL
LIBRARY

R. F. LAWRENCE JUN 2 4 1964

HARVARD
UNIVERSITY,

NEW CAVERNICOLOUS SPIDERS
OF SOUTH AFRICA

April 1964 April
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JUN 2 4 1964

HARVARD
NEW CAVERNICOLOUS SPIDERS FROM SOUTH NMOS py

OOL

By
R. F. LAwRENCE

Natal Museum, Pietermaritzburg

(With 24 text-figures)

CONTENTS

PAGE
Introduction .. bie ais Se GT
Descriptions of Sees. . 58

A list of the cavernicolous Rraetinicts rie Morapods
of South Africa .. as d's ois 7,
Summary ag as ae fs ead
eee iedeement + ¢3 ap fi soe Ms
References nie * os ats ar ee ey

INTRODUCTION

At the end of the last century only three species of cave arachnids had been
described from South Africa, all of them spiders, by E. Simon (1893, 1894,
1896).

The number of Arachnida and Myriopoda known at the present time,
including those described in the present paper, is twenty-two, seventeen of them
being Araneae. Most of these have been taken from the Table Mountain and
Kalk Bay caves in the Cape Peninsula, and the Cango Caves at Oudtshoorn.
Specimens have also been collected in: Skeleton Cave, Oudtshoorn; Guano
Cave, Hotpot and Onmeetbarediepgat, Bredasdorp; two small caves in Natal, at
Champagne Castle in the Drakensberg and at Noodsberg in the New Hanover
district. In the Transvaal Simon collected a number of arthropods at a small
cave in the suburbs of Pretoria on the banks of the Apies River, and at the
Makapan Cave near Potgietersrus. It is worthy of note that in the cave at
Pretoria, Simon mentions (1894: 63) finding a number of specimens of an
undescribed pseudoscorpion (Chilifer, sic) which he sent to M. Balzan for
identification. In the Makapan Cave he discovered a new species of tenebrionid
beetle, Eurychora simon, and observed, but apparently did not collect, Myria-
poda. Many of these caves are large, forming long horizontal galleries or
extending deep below the surface, and almost all the specimens were collected
in totally dark parts of the caves.

It is of interest to compare the numbers of cave Arachnida found in South
Africa with those of neighbouring regions of the African continent. In 1931
Fage published a comprehensive list of cave spiders from all parts of the world,
omitting however the three South African species already described by Simon.

a)
Ann. S. Afr. Mus. 48 (2), 1964, 57-75, 24 text-figures.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

According to this list no less than twenty-six species had been described from
East Africa by Berland (1914) and Simon and Fage (1922) as compared with
seventeen from South Africa up to 1963 and eight from the (Belgian) Congo
(Leleup, 1956). It is possible that collecting in South African caves has not
been as extensive as in East Africa.

The Congo has a much larger cavernicolous myriapod fauna than South
Africa, Leleup (1956) listing nine species of Diplopoda and three of Chilopoda,
while only one diplopod species has been found in the whole South African
region and no Chilopoda. On the other hand only eight species of spiders have
been recorded from the caves of the (Belgian) Congo.

A number of immature or incomplete specimens, which it has been impos-
sible to describe and include in the South African faunal list, indicate that a
fairly large number of South African cave spiders still await description. This
would especially apply to the Bredasdorp caves, the Oudtshoorn caves other
than the Cango Caves, and various caves in Natal and the Transvaal. It might
also be interesting to explore for comparison some of the long tunnels which
have recently been excavated for the South African railways, such as the
3-mile-long tunnel between Pietermaritzburg and Howick in Natal.

The large number of specimens placed at my disposal by Mr. J. R.
Grindley of the South African Museum were collected by himself and other
staff members of this Institution at various times between 1929 and 1961, and
by the South African Spelaeological Association.

DESCRIPTIONS OF SPECIES
Family Dictynidae
Genus HAEMILLA E. Simon
Haemilla grindleyi n. sp.
(Fig. 1)

Holotype: 1 2 (S.A.M. 10004), Wynberg Caves, Table Mountain, collected
J. R. Grindley, August 1956.

Colour: Carapace light reddish brown, its anterior margin a little darker,
sternum reddish brown, mouthparts and chelicerae dark reddish brown;
legs (including coxae) yellow brown, the apical segments reddish; pedipalps
yellow brown, tibia and tarsus reddish brown. Abdomen dark olive green
dorsally, with a series of indistinct lighter chevron markings, some intermixed
lighter spots and stripes laterally, ventral surface olive green with a pair of
widely separated parallel longitudinal lighter stripes.

Eyes: Anterior row with lower margins forming a straight line, medians
less than a diameter apart, their own diameter from the laterals; posterior row
distinctly procurved, medians a little more than their diameter apart, two
diameters from the laterals; posterior medians distinctly larger than anterior
medians, median quadrangle considerably wider behind than in front (by a

NEW CAVERNICOLOUS SPIDERS FROM SOUTH AFRICA 59

little less than the diameter of an anterior median), as long as or a very little
longer than posteriorly wide; laterals subequal, subcontiguous, anterior medians
3-4 times their diameter from edge of clypeus.

Chelicera with 7-8 teeth on superior margin, 4—6 on inferior margin not
much smaller than the superior ones.

Fic. 1. Haemilla grindleyi n. sp. 9. Vulva.
Fic. 2. Haemilla profundissima n. sp. 2. Vulva.
Fics. 3, 4. Loxosceles valida n. sp. 3. 3, dorsal pattern of abdomen; 4, pedipalp from
inner side.

Vulva as in fig. 1, resembling in its basic pattern those of tuberculata Lawr.
and tanganensis Simon & Fage.

Calamistrum on metatarsus IV with a row of 33 modified hairs occupying
only one-eighth of its total length; the segment slightly sinuous in the region
of the calamistrum.

Legs: Patellae III and IV with 1 posterior spine, anterior patellae without;
tibiae I and II with 2 pairs of inferior spines, 2 lateral pairs; metatarsi I and II
with 4 lateral spines on each side, I with 1 small inferior spine at apex, II with
2 pairs of inferior spines. All tarsi with a ventral brush-like scopula, anterior
legs with a metatarsal scopula in addition.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pedipalp: ‘Tibia and tarsus with a dense brush-like scopula, tarsus with 5-6
long spines in addition.

Dimensions: Length of carapace 5:1, abdomen 6 mm.

Further material: 1 immature 9, Wynberg Caves (S.A.M. Broo1o), collected
R. F. Lawrence, March 1931: 1 9, Kalk Bay Caves (S.A.M. B7894), collected
R. F. Lawrence, July 1932; 3 9° (2 fragmentary) from Wynberg Caves (S.A.M.
B10007, B10008, B10009), collected J. R. Grindley, August 1956; 1 immature 9,
Bats Cave (S.A.M. Brooo6), Table Mountain, collected J. R. Grindley,
September 1960; 1 juvenile 9, (S.A.M. B 10006), Oread Halls, Kalk Bay Caves
(S.A.M. B1o005), collected J. R. Grindley, June 1956.

The species seems to be nearest tuberculata Lawrence from Natal. It differs
from all other species of the genus in the form of the vulva and in the small
extent of the fourth metatarsus occupied by the calamistrum, this being only
one-eigth (two-fifths in tuberculata) of its length. Only one other species, caverni-
cola Lawrence from Natal, appears to have been recorded from caves.

Haemilla profundissima n. sp.

(Fig. 2)

Holotype: 1 Q (S.A.M. Bioo11), Onmeetbarediepgat, Bredasdorp, Cape
Province, collected C. Gow, July 1961.

Colour: Carapace light reddish brown; legs light reddish brown, the basal
segments with an olive tinge, the mouthparts all dark reddish brown, sternum
reddish brown, chelicerae blackish brown. Abdomen dorsally olive green with
very numerous small light dots, a median stripe in anterior two-thirds with
chevron markings and a row of ill-defined spots on each side, lighter; lateral
surfaces with mixed spots and stripes, venter with two widely separated parallel
white stripes.

Eyes: Anterior row from in front slightly procurved, subequal or the
laterals a little larger, medians less than their diameter apart and a diameter
from the laterals; posterior row from in front distinctly procurved, the
medians distinctly smaller than anterior medians, subequal to the laterals,
13—2 times their own diameter apart, 2-3 diameters from the laterals; laterals on
each side the radius of a posterior lateral apart, the posterior distinctly smaller
than the anterior; median quadrangle as long as posteriorly wide, wider behind
than in front by the diameter of a posterior median eye, anterior medians the
length of median quadrangle or 2—3 times their own diameter, from the edge of
clypeus.

Mouthparts: Labium a little longer than in tuberculata Lawr. (cf. Lawrence
1939: 270, fig. 1b), reaching almost to the apices of the maxillae or more than
three-fourths of their length.

Chelicerae: Inferior margin with 7-8 teeth, the distal 3 very small, superior
margin with 7.

Vulva as in figure 2, differing remarkably from all other species in its basic
pattern.

NEW CAVERNICOLOUS SPIDERS FROM SOUTH AFRICA 61

Legs: Tibia I and II with 3 inferior and 3 lateral pairs of spines, metatarsi
apparently similar but the spines obscured by a thick brush of hairs on the
under side of metatarsi and tarsi, and also on tibia I. Metatarsus IV with a
slight sigmoid curve seen from above, the calamistrum very distinct, composed
of about 45 stout modified hairs and occupying a little less than a third of its
total length.

Dimensions: Length of carapace 5:3, abdomen 5:6 mm.

Family Sicartidae
Genus LOXOSCELES Lowe

Loxosceles valida n. sp.

(Figs. 3, 4)

Holotype: 1 3 (S.A.M. Broo12), Echo Halt Caves, Table Mountain, Cape
Town, collected J. R. Grindley, April 1954 (labelled No. 10).

Colour: Carapace in general rich reddish brown, cephalic portion with its
lateral margins and 4 narrow parallel stripes behind the median eyes, thoracic
portion with some large ill-defined radial markings from the foveal depression
outwards, all a little darker than the background; chelicerae reddish brown;
sternum light yellow brown, the margins narrowly reddish brown, an irregular
cluster of blackish olivaceous spots in the middle; coxae infuscated in apical
fourth, remainder yellow; labium reddish brown, maxillae a little lighter, the
apices of both with narrow white border. Abdomen dorsally with a brown
pattern on light yellow background (fig. 3), ventral surface mostly brown,
ventral spinners with a light transverse band in basal half. All legs reddish-
brown, femora I and II a little darker, metatarsi and tarsi I-IV a little lighter.

Carapace with a group of fairly coarse black bristles at anterolateral angle
and behind the lateral eyes, cephalic portion with 7 distinct longitudinal rows
of forwardly directed similar bristles, 3 behind the median, 2 behind the lateral
eyes; thoracic portion with 3 weak rows of bristles on each side, the two posterior
ones reaching the lateral margin, the anterior row abbreviated; sides of carapace
with a regular marginal row of bristles but otherwise smooth, shiny.

Eyes of both the median and lateral pairs contiguous, a line joining the
anterior laterals touching the posterior margins of the medians; median pair
separated from the laterals by a little more than their combined width, from
anterior margin of clypeus by 14~2 times the long diameter of a median eye.

Chelicerae with a band of black bristles on the lateral half of its anterior
surface; apex of chelicera ending in a large black triangular tooth on the
inferior margin.

Legs without spines, very long and, as far as patellae, very strong; femora
almost entirely smooth but the ventral surfaces of the anterior ones with numer-
ous soft fine hairs in basal half. Tibiae and distal segments with regular rows of
large spine-like black bristles increasing in length and slenderness distally,
anterior tarsi with a weak scopula ventrally, the posterior ones with a distinctly

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

denser scopula and at the apices of metatarsi in addition. Legs I, II, IV, III,
II only a little shorter and not weaker than I.

Pedipalp as in figure 4 seen from inner side; tarsus and tibia with coarse
bristles, more numerous on inner than outer surfaces, more dense on tarsus than
tibia; tarsus bluntly triangular, tibia much inflated, almost twice as deep as
patella seen from the side (its depth two-thirds its greatest length), ovoid, much
wider than the remaining segments seen from above.

Dimensions: Length of carapace 5°5, width 4-6; length of abdomen 7-3 mm.
Leg I: femur 15, patella-tibia 22, metatarsus 21, tarsus 2°8 mm.

Further material: All the remaining material consists of females as follows:
1 9 (S.A.M. B10018), Powder Room Cave, Table Mountain, collected South
African Spelaeological Association, March 1956. 1 9 (S.A.M. B10016), Wynberg
Caves, Table Mountain, collected South African Spelaeological Association,
February 1956. 1 2 (S.A.M. B1ioo15), Devil’s Pit, Kalk Bay, collected J. R.
Grindley, June 1954. 1 immature 2 (S.A.M. B1o0013), Tartarus Cave, Kalk Bay
Mountains, collected J. R. Grindley, July 1961. 1 immature 2 (S.A.M. Bioor7),
Giant’s Workshop, ‘Table Mountain Caves, collected J. R. Grindley, July 1956.
I immature 2 (S.A.M. B7892), Wynberg Caves, Table Mountain, collected
R. F. Lawrence, March 1931. 1 2 (S.A.M. B1ioo14), Bats Cave, Table
Mountain, collected J. R. Grindley, September 1960.

These specimens are in general smaller, with shorter legs than the dg,
leg I only about 3 times the body length. The distinctive pattern of the dorsum
of abdomen is almost identical with that of the g except that the transverse
bars or chevrons are in some cases more numerous, 7-8 in number. Total length
of largest 9 (S.A.M. Broor4 from Bats Cave), 14:5, leg I 43 mm.

Remarks: Five other species of Loxosceles are known from southern Africa:
two from South West Africa, bergerz Strand and szmillima Lawrence, two from
the Cape Province, pilosa Purcell and spinulosa Purcell, and one from the
Transvaal, speluncarum Simon. All these species are very much smaller, with
shorter legs, the first being only 3 times or less the total length of body while
in valida it is more than 6 times. The only other known cavernicolous species

of the genus, L. speluncarum, was found in a cave in calcareous deposits of the —

Apies River valley near Pretoria, while Simon and Fage recorded an unidenti-
fied juvenile from Haitajwa Cave on Zanzibar Island (1922: 528). The new
species has in all the specimens a strong, very clearly defined colour pattern,
without the least sign of the depigmentation characteristic of cave animals.

Family Leptonetidae
Subfamily Ochyroceratinae
SPELEODERCES new genus

Carapace and abdomen well chitinized, the latter with large oval dorsal
scute but no ventral scute. Eyes consisting of three widely separated pairs,
two lateral and one median, as in Cangoderces Harison, but the median pair

3

NEW CAVERNICOLOUS SPIDERS FROM SOUTH AFRICA 63

situated much farther back. Chelicerae with 4 large teeth on superior margin;
labium much wider than long; maxillae wide, fairly short, not meeting distally,
their external and internal apices angular. Legs very long and slender, femora
subparallel throughout. Patella of male pedipalp much longer than tibia and
strongly toothed, remaining segments normal. T'ype-species of genus: Speleo-
derces scutatus n. sp.

Speleoderces scutatus n. sp.
(Figs. 5-8)

Holotype: 1 3, (S.A.M. Bioorg), Wynberg Caves, Table Mountain,
collected South African Spelaeological Association, February 1956.

Colour: Carapace ventrally and dorsally yellow with an orange tinge,
abdomen yellow white, dorsal scute a little darker, legs yellow white.

Carapace narrowing a little anteriorly, the anterior margin truncate but
rounded at the antero-lateral angles.

Eyes as in figure 5, separated from anterior margin of carapace by an
indistinct groove, the laterals on each side occupying a low indistinct rounded
tubercle; position of posterior medians unusual in being well back, more or less
in a line with the posterior of the two laterals; posterior medians well separated
by about their own diameter from each other, far removed from the laterals
on each side which are contiguous.

Mouthparts as in figure 6, labium very short, much wider than long, maxillae
wide, their inner margins converging, the distal margins quite straight with
both the inner and outer angles distinct though rounded.

Chelicerae: Claw very wide at base but narrowing abruptly in distal third,
superior margin with 4 teeth, the basal one largest and well separated from
the remaining three which are subgeminate; basal half of the claw with minute
but very regular and distinct saw teeth (fig. 7).

Legs very long and slender, the anterior longer than posterior pair, the
femora not incrassate basally; all patellae at dorsal apex with 1, tibiae dorsally
with 2 very long erect setae, legs clothed otherwise with series of fairly numerous,
much shorter setae as in Cangoderces lewisi Harison (1951, p. 83, fig. 1).

Pedipalp as in figure 8 seen from outer side, patella much longer than tibia,
subparallel, with a strong hooked tooth at its base directed dorsally and a
little to the outer side, another large claw-like tooth at its apex.

Abdomen: Except on the scute, clothed with long slender setae, its dorsal
surface with a large oval scute, not strongly chitinized but distinct, covering
all except the posterior fifth or sixth of dorsal surface.

Dimensions: Length of carapace 0-65, of abdomen 0-85 mm.

Female: A single 2 specimen (S.A.M. Broog20) from Bats Cave, Table
Mountain, collected by the South African Spelaeological Association, February
1956, is undoubtedly the female of this species. It differs from the male only in
the somewhat shorter legs, more rounded abdomen and the total absence of a
dorsal scute on the latter.

64. ANNALS OF THE SOUTH AFRICAN MUSEUM

Fics. 5-8. Speleoderces scutatus n. sp. 3. 5, eyes from above; 6, sternum and mouthparts; 7,
chelicera from below; 8, pedipalp from outer side.
Fics. 9-11. Spermophora peninsulae n. sp. 3. 9, apex of chelicera; 10, pedipalp from outer side;
11, chelicerae from in front.

NEW CAVERNICOLOUS SPIDERS FROM SOUTH AFRICA 65

Dimensions: Total length 1-1 mm.

Remarks: The new genus appears to differ from all other genera of the
family Leptonetidae interpreted in its widest sense to include the Ochyrocer-
atidae. These differences lie in the arrangement of the eyes, the structure of the
chelicerae, labium and maxillae, while in possessing a dorsal abdominal scute
it seems to approach the Oonopidae. When themale of Cangoderces Harison has
been discovered it may be found to be most nearly related to this genus.

Family Pholcidae
Genus SPERMOPHORA Henz

Spermophora peninsulae n. sp.

(Figs. 9-14)

Holotype, 1 3, paratype, 1 9 (S.A.M. B7897), Kalk Bay Caves, Cape Penin-
sula, collected R. F. Lawrence, September 1932.

Male (holotype)

Colour: ‘The specimen completely bleached, the chitinous structures of the
mouthparts and pedipalp a little darker; carapace dorsally and ventrally
yellow, a little darker than the abdomen which is quite pale, legs pale.

Chelicerae: The fang stout and short, a very conspicuous triangular tooth
on its margin proximal to the fang (fig. 9); anterior surface with two pairs of
broad chitinous teeth near its apex (fig. 11), the more distal pair directed
inwards, the second pair downwards; base of chelicera near clypeal margin
with an oblique projecting ridge behind which is a depression extending almost
the whole width of the segment.

Abdomen longish, cylindrical, rounded, considerably longer than that of 9.

Pedipalp as in figure 10 seen from the outer side, tibia and basal part
of tarsus with the sockets of numerous spines which in most cases have been
lost.

foc LV, IL, TIT.

Dimensions : Total length 2-6 mm.

Female (paratype)

Colouring similar to that of the ¢; chelicerae unmodified; the chitinized
parts of the epigastric area as in figure 14; pedipalp with long, slender, pointed
tarsus; legs very long, the femora long and stout, almost twice as wide at base
as at apex.

Dimensions: Length of carapace 1-1, length of abdomen 1-9 mm.

Remarks: Another 3 from the Kalk Bay Cave system (Oread Halls,
collected J. R. Grindley, June 1954, S.A.M. Broog21) has a jet-black sternum
and indistinct markings on the abdomen, but the pedipalp is structurally
similar to that of the holotype ¢. Four tubes of female Spermophora from the
Wynberg and Powder Room caves I assume to be the same species as the types,
although in the types all traces of any original colour pattern have been lost
after more than 30 years of immersion in alcohol.

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

21

Fics. 12-14. Spermophora peninsulae n. sp. 2. 12, dorsal colour pattern; 13, the same of another 9
from the side; 14, epigastric region.
Fics. 15-17. Teutana fagei n. sp. 9. 15, mouthparts and anterior margin of sternum; 16, carapace
seen in profile; 17, vulva.
Fics. 18, 19. Theridion proxima n. sp. 2. 18, mouthparts and sternum; 19, vulva.
Fics. 20-22. Lephthyphantes rimicola n. sp. 20, chelicera of 9; 21, mouthparts of 9; 22, pedipalp of g
seen from the side.

CD Gee a)

NEW CAVERNICOLOUS SPIDERS FROM SOUTH AFRICA 67

The colouring of the female from Powder Room Cave (S.A.M. B10023) is
as follows: :

Colour: Carapace with dark but not black markings, as in figure 12, a
fairly wide and distinct though slightly crenulated blackish stripe passing forward
from each ocular group to edge of clypeus, these diverging slightly anteriorly;
chelicerae and mouthparts brown, sternum blackish brown, a little lighter in the
middle, contrasting strongly with the yellow coxae. Abdomen with black
markings dorsally as in figure 12, a cuneiform white marking at posterior apex
just above the spinners, seen from the side as in figure 13 (drawn from another
®, Wynberg Caves, S.A.M. Broo24, collected August 1956); ventral surface
with a wide median black band widening to include the epigastric area which is
brown and separated on each side by a fairly narrow sinuous white stripe from
the blackish sides of the abdomen. Legs yellow with a reddish tinge, tibiae
with a narrow, lighter apical annulation.

Further material: 1 9 (S.A.M. B10023) with about 24 eggs, Powder Room
Cave, collected Spelaeological Association, March 1956; 2 99 (S.A.M. Broo22
and B10024), Wynberg Caves, collected J. R. Grindley, August, 1956; 1 4,
2 99 (S.A.M. Bioo21), Oread Halls, collected J. R. Grindley, June 1954;
1 9 (S.A.M. Broo2r), Bats Cave, Table Mountain, collected South African
Spelaeological Association, February 1956; 4 99 (S.A.M. B7896), Wynberg
Caves, Table Mountain, collected R. F. Lawrence, September 1932; 1 g
(S.A.M. B7895), Wynberg Caves, Table Mountain, collected R. F. Lawrence,
March 1931.

Simon (1892-1903: 471) writes of two types of colouring, pale species
occurring in caves and houses, strongly pigmented ones under stones; all the
four East African species, ensifera, globosa, minotaura and nigrescens, appear to
belong to the latter group. Unfortunately no adult males are present among the
Wynberg Caves material and in their absence I have assumed these specimens
to be identical with the Kalk Bay Caves species, the absence of pigmentation
in the type of the latter being due to prolonged immersion in alcohol.

The only species of this genus hitherto found in caves is S. minotaura
Berland from the Campbell Cave, Kenya, though it has also been found in the
forests of Kenya at high and low altitudes.

Genus SMERINGOPUS Simon
Smeringopus pallidus (Blackwall)

One immature 9, (S.A.M. B1o0025), Skeleton Cave, Oudtshoorn, collected
J. R. Grindley, September 1961. |

Although immature the colouring of the single specimen resembles that
of the female of pallidus (= elongatus) as redescribed by Kraus (1957: 220, fig. 6).
No species of the genus appears to have been described from caves but speci-
mens of Smeringopus are often found near the entrance of many caves and should
be regarded as troglophiles rather than troglobionts.

68 : ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Theridiidae
Genus TEUTANA Simon

Teutana faget n. sp.

(Figs. 15-17)

Holotype: 1 9 (S.A.M. B10026), Skeleton Cave, Oudtshoorn, Cape Province,
collected by J. R. Grindley, September 1961.

Colour: Garapace, sternum and mouthparts brown with a slight reddish
tinge, the radiations from the thoracic fovea a little darker, otherwise without
markings. Legs, coxae to tarsi uniform brown with a faint olive tinge; abdomen
above reddish violet, a recurved white transverse stripe at anterior apex with
a short backwardly projecting stripe in the middle, posterior to this two pairs
of ill-defined elongate white markings, above the spinners a short median
elongate marking followed by a transverse series of 3-4 very fine white lines;
ventral surface with a large comma-shaped white marking laterally to each
operculum, posterior to this two elongate, inwardly directed white stripes, the
second considerably smaller than the first; between the spinners and epigastric
furrow, a large vase-shaped, ill-defined lighter marking (somewhat similar to
the well-defined red marking on the venter of Latrodectus geometricus), vulva and
opercula blackish brown.

Carapace seen from the side level but slightly depressed in the region of the
thoracic fovea, which is deep, transversely arcuate and slightly recurved,
clypeus strongly rounded and projecting, with a well-marked transverse groove
just below the eyes separating them from the rest of the clypeus (fig. 16).

Eyes: Anterior row from above slightly recurved, from in front straight,
medians subequal to laterals or a very little smaller, less than their diameter
apart and about the same distance from the laterals; posterior row slightly
recurved seen from above, medians a little more than their own radius apart,
about a diameter from the laterals which are a little larger. Median quadrangle
longer than posteriorly, wide, a little wider behind than in front; clypeus about
14 times length of median quadrangle.

Chelicera: Inferior margin with 2 or 3 small, indistinctly geminate teeth.

Mouthparts: Labium at least twice as wide as long, the anterior apex not
triquetrous, more or less truncate, straight in the middle, somewhat rounded at
the sides, its apex falling well below the middle point of the maxillae (fig. 15).

Vulva as in figure 17, the median septum very strongly chitinized, blackish,
ill-defined.

Legs unspined, with rows of weak setae and soft hairs.

Dimensions: Carapace 3°3, length of abdomen 7, width 5:3 mm.

Further material: 1 smaller 9 (S.A.M. Broo27) from Onmeetbarediepgat,
Bredasdorp, Cape Province, collected C. Gow, July 1961.

The species differs from the two species of Teutana described by O. P.
Cambridge from the Cape Peninsula, lepida and connexa, at least in its larger size
and detailed differences in the pattern of the vulva, while agreeing with them in

NEW CAVERNICOLOUS SPIDERS FROM SOUTH AFRICA 69

the colouring of the ventral surface. It differs from T. albovittatus Lawrence
from the Umfolosi River, Zululand, in its considerably larger size, colour
pattern of abdomen and structure of the vulva. Only one other species of the
genus, 7. grossa, appears to have been recorded from caves (in Europe only).

Genus THERIDION Walck

Theridion proxima n. sp.
(Pigs. 16,, 19.)

Holotype: 1 9 (S.A.M. Broo28), Skeleton Cave, Oudtshoorn, Cape
Province, collected J. R. Grindley, September 1961.

Colour: Carapace yellow with a very fine brown margination, sale
portion and thoracic fovea brown, eyes surrounded by blackish brown rings,
chelicerae light olive brown contrasting with the yellow clypeus; sternum yellow
brown with narrow blackish brown margin, mouthparts light reddish brown.
Legs yellow, the femora with 2 subapical, tibiae with 4 blackish annulations,
metatarsi with smaller basal middle and apical annulations, the middle one
faint or obsolete. Abdomen dorsally as in Tullgren’s description of kibonotense
and O. P. Cambridge’s of purcelli, the black area surrounding the light median
longitudinal band diffuse, a large white marking on each side in posterior
third near the lateral margin; ventral surface with spinners surrounded by a
blackish ring which is continuous with the median blackish area of dorsal
surface, a medium-sized white spot below the epigastric fold.

Eyes seen from above with anterior row slightly recurved, medians a
little larger than laterals, about their diameter apart and a little less from the
laterals; posterior row well procurved, medians distinctly larger than the
laterals, three-fourths their diameter apart, their diameter or a little more from
the laterals; laterals touching on each side, the posterior distinctly smaller
than anterior; median quadrangle a little wider in front than behind, a little
wider than long; clypeus about 14 times the Jength of median quadrangle.

Mouthparts: Labium much wider than long, its anterior margin truncate,
almost straight, with a row of 6 long black setae, the segment reaching to about
a third the length of the perpendicular maxillae (fig. 18).

Legs: 1, 1V, II, 111; femora with 2 inferior rows of setae mounted on small
round tubercles, those of leg II much more regular than the others (the setae
of femur I largely missing), each row with 12-15 setae; patella with a long,
strong, black seta at dorsal apex; anterior tibiae with 1-3 similar dorsal setae,
IV with 4; metatarsi with a ventral row of setae, those of II stronger and more
regular than the others; tarsus IV with a row of 6 long, slender modified hairs
diminishing progressively in length distally.

Vulva as in figure 19 resembling that of T. kibonotense in its general pattern.

Dimensions: Length of carapace 2, length of abdomen 3-6 mm.

Remarks: The species obviously resembles Tullgren’s kibonotense (1910) and
O. P. Cambridge’s purcellii (1903) both in colouring and the basic pattern of

7O ANNALS OF THE SOUTH AFRICAN MUSEUM

the vulva, but of the two is probably more closely related to kibonotense. About
17 species of this large and widespread genus are known from southern Africa,
including an almost cosmopolitan form often found in greenhouses, T. tepidario-
rum C. L. Koch; Thertdion rufipes Lucas has been recorded from caves in Spain.

Family Linyphiidae
Genus LEPHTHYPHANTES Menge
Lephthyphantes rimicola n. sp.
(Figs. 20-23)
Holotype, 1 2, paratypes, 1 3, 2 92 (S.A.M. B7893), Wynberg Caves, Table
Mountain, collected R. F. Lawrence, March 1931.
Female (holotype)

Colour: Carapace and legs yellow to white without markings, the eyes
surrounded by black rings, posterior medians connected with anterior medians

Fic. 23. Lephthyphantes rimicola n. sp. 2. Vulva.
Fic. 24. Phanotea gowi n. sp. 9. Vulva.

by a subtriangular black marking spanning the space between them; abdomen
much faded but darker than remainder of body, light terracotta witha number ~
of large round indistinct blotches. ;
Eyes: Posterior row slightly procurved, the medians distinctly the largest
of all the eyes; anterior row straight to very slightly recurved, the medians
much the smallest of all the eyes, half the diameter of an anterior lateral and
close to each other; laterals contiguous, the anterior a little smaller than the
posterior; median quadrangle as long as posteriorly wide, much wider behind
than in front; clypeus a little less than the length of median quadrangle.
Mouthparts as in figure 21, maxillae very wide, subparallel, labrum semi-
circular, much wider than long; chelicerae as in figure 20, fang long and
robust, superior margin with 3 large teeth, the basal smallest, inferior margin
with 5 distinctly smaller sharp teeth, the basal separated from the others which

NEW CAVERNICOLOUS SPIDERS FROM SOUTH AFRICA 7h

are subcontiguous. Chelicerae subparallel, not narrowing much distally;
sternum cordiform, its anterior margin very wide and almost straight.

Vulva as in figure 23.

Legs I-IV long and very slender, the femora ventrally with a more or less
regular series of long, fine setae; remaining segments with long, slender, erect
dorsal setae as follows (the seta on the patella much longer than any others) :
patella at apex with 1 almost as long as the segment, tibia with 2 or 3 in distal
half, these about a third the length of the segment; metatarsus with 1 subbasal
seta, tarsus without; legs otherwise clothed with regular rows of numerous
short, weak setae.

Pedipalp tarsus long, slender, longer than tibia, its apex with a long edentate
claw, not much thicker than the adjacent setae.

Male (paratype)

Not differing from the female except that the chelicerae are somewhat
more divergent and more tapered apically, the legs relatively longer and more
slender. Carapace not modified, in general similar to that of the 9.

Pedipalp as in figure 22 seen from the side, very complex, the embolus very
long and sinuously curved.

Dimensions: 9: Length of carapace 0-9, of abdomen 1:2 mm. ¢: Total
length 1-9 mm.

Remarks: ‘The material on which the new species is based is very old and
fragile but it will be possible to check the description and figures from fresh
material which should not be difficult to obtain as the spiders are fairly common,
spinning small sheet webs 25-35 mm. in diameter across the fissures in the
vertical rock walls of the cave.

The Linyphyiidae are very little known in South Africa and I have
considerable doubt as to whether this cave form really is a species of Leph-
thyphantes or whether a new genus is required for it. In some respects it agrees
with Linyphia (in the appearance of the male pedipalp, with the British species
Linyphia (Agyneta) cauta O. P. Cambridge). No members of this genus, however,
appear to inhabit caves. In the considerable reduction of the anterior median
eyes it agrees with various genera from different parts of the world, such as
Porrhomma, Troglohyphantes and Asthenargus, the latter being represented by the
species A. inermis in East African grottoes; from all of these, however, it also
differs in at least one important character.

Family Agelenidae
Subfamily Cybaeinae
Genus PHANOTEA Simon
Phanotea gowi n. sp.
(Fig. 24)
Holotype: 1 2 (S.A.M. Broo29), Guano Caves, Bredasdorp, Cape Province,
collected C. Gow, July 1961.

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

Colour: Legs and carapace yellow to orange, carapace becoming progres-
sively more reddish anteriorly ; sternum yellow, the margins orange, mouthparts
reddish brown, chelicerae dark reddish brown.

Eyes: Anterior row slightly recurved, medians distinctly smaller than the
laterals, less than their own diameter apart, about a diameter from the laterals;
posterior row a little procurved, medians a little smaller than anterior medians,
laterals smaller than anterior laterals, medians their own diameter apart, 2
diameters from the laterals; median quadrangle distinctly longer than
posteriorly wide, a little wider behind than in front; anterior medians a little
more than their own diameter from the edge of the clypeus.

Chelicerae very strong, inflated and rounded in front, a deep anterior
constriction near their bases below the clypeus; lower margin with only 2
small subequal teeth, differing in this respect from all other species of the genus;
superior margin with 3 teeth, the middle one large.

Pedipalp with thick scopula ventrally on the distal two-thirds of tarsus.

Vulva as in figure 24, with a large median plano-convex cordiform plate
overlying two posterior sclerites which almost meet in the middle line below it.

Legs: Patellae unspined; tibia I and II with 4 inferior pairs of spines,
metatarsus I and II with 3 inferior pairs, I with 2 anterior lateral spines in
addition, II without lateral spines, tarsus and most of metatarsus I with a
distinct scopula, II with a distinct scopula only on tarsus; tibia III and IV with
3 inferior irregularly paired spines, 2 pairs of lateral spines, metatarsus III and
IV with 3 inferior pairs, 3 lateral pairs and 1 or 2 superior spines in addition;
tarsi III and IV but not metatarsi with distinct scopula.

Dimensions: Length of carapace 6-4, total length 15:3 mm. (including
chelicerae).

Further material: One specimen, Hotpot, Bredasdorp, Cape Province,
collected South African Spelaeological Association, May 1960.

The genus consists mostly of cave-living species, natalensis, simoni and
peringueyt ; it appears to be most closely related to peringuey2 in the eye disposition
and spine formula of the legs but can be easily distinguished from it by having
only 2 instead of 3 teeth on the inferior margin of chelicera and in the pattern
of the vulva.

Subfamily Hahniinae

A single, very fragile and much bleached subadult male from the Wynberg
Caves (S.A.M. B7896), collected R. F. Lawrence, September 1932, is evidently
a member of this subfamily, several species of which have been recorded from
caves.

The spinners are long and slender, arranged in a more or less transverse
row with the superior pair especially long, the apical segment being subequal
to the basal in length. There are, however, only six eyes, all of them large with
the anterior medians absent, the general arrangement similar to that of Bigozs
Simon from the Philippine Islands, in which, however, the anterior median

NEW CAVERNICOLOUS SPIDERS FROM SOUTH AFRICA 73

eyes, though minute, are present. It does not seem to be closely related to
Scotussa zodarioides Simon, the only member of the subfamily recorded from the
Cape Peninsula, and a new genus may have to be created for its reception.

A LIST OF THE CAVERNICOLOUS ARACHNIDA AND MYRIAPODA OF SOUTH AFRICA

IO.

tI.

es

16.

ni.

Species

. Haemilla cavernicola

Lawrence, 1939

. Haemilla grindley

Lawrence, 1964

. Haemilla profundissima

Lawrence, 1964

. Phyxelia makapanensis

EK. Simon, 1894

. Loxoscles speluncarum

E. Simon, 1893

. Loxosceles valida

Lawrence, 1964

. Speleoderces scutatus

Lawrence, 1964

. Cangoderces lewisi

Harison, 1951

. Spermophora peninsulae

Lawrence, 1964

Smeringopus pallidus
(Blackwall), 1858

Teutana fagei
Lawrence, 1964

Theridion proxima
Lawrence, 1964

. Lephthyphantes rimicola

Lawrence, 1964

. Phanotea peringueyr

E. Simon, 1896

. Phanotea natalensis

Lawrence, 1951
Phanotea simoni

Lawrence, 1951
Phanotea gowt

Lawrence, 1964

ARANEAE

Family
Dictynidae

bP)
Sicariudae
39

Leptonetidae

393

Pholcidae

bP)

Theridiidae

Linyphiidae

Agelenidae

Locality
Noodsberg Caves, Natal.

Wynberg Caves, Table Mt.,
and Kalk Bay Caves,
Cape.

Onmeetbarediepgat, Bredas-
dorp, Cape.

Makapan Cave, Transvaal.

Apies River Cave, Pretoria,
Transvaal.

Wynberg Caves, Table Mt.,
Cape.

Wynberg Caves, Table Mt.,
Cape.

Cango Caves, Oudtshoorn,
Cape.

Wynberg Caves, Table Mt.,
and Kalk Bay Caves,

Cape.

Skeleton Cave, Oudtshoorn,
Cape.

Skeleton Cave, Oudtshoorn,
Cape.

Skeleton Cave, Oudtshoorn,
Cape.

Wynberg Caves, Table Mt.,
Cape.

Cango Caves, Oudtshoorn,
Cape.

Noodsberg Caves, Natal.

Champagne Castle Cave,
Drakensberg, Natal.

Guano Cave, Bredasdorp,
Cape.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

OPILIONES
Species Family Locality
18. Speleosiro argasiformis Sironidae Wynberg Caves, Table Mt.,
Lawrence, 1931 Cape.
19. Speleomontia cavernicola. ‘Triaenonychidae § Wynberg Caves, Table Mt.,
Lawrence, 1931 Cape.
20. Larifuga sp. (1964) bg Wynberg Caves, Table Mt.,
Cape.
PSEUDOSCORPIONES
21. Chthoniella cavernicola Chthoniidae Wynberg Caves, Table Mt.,
Lawrence, 1935 Cape.
DIPLOPODA
22. Harpethrix caeca Sphaerotrichopidae Wynberg Caves, Table Mt.,
Lawrence, 1962 Cape.
SUMMARY

Nine new species of cavernicolous spiders from South Africa are described,
one of which is included in a new genus. The new species are: Haemilla grindleyi,
Haemilla profundissima, Loxosceles valida, Speleoderces scutatus (new genus), Spermo-
phora peninsulae, Teutana fagei, Theridion proxima, Lephthyphantes rimicola and
Phanotea gowi. A list is given of the twenty-two species of Arachnida and
Myriapoda now known from South Africa.

ACKNOWLEDGEMENT

The Trustees of the South African Museum gratefully acknowledge a grant
from the South African Council for Scientific and Industrial Research for the
publication of this paper.

REFERENCES

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NEW CAVERNICOLOUS SPIDERS FROM SOUTH AFRICA 75

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141-165.

Leeup, N. 1956. La faune cavernicole du Congo belge et considérations sur les coléoptéres
reliques d’Afrique intertropicale. Ann. Mus. Congo belge 8vo Sci. zool. 46: 1-171.

Simon, E. 1893. Histoire naturelle des araignées. 1: 273. Paris: Roret.

Simon, E. 1894. Note sur les arthropodes cavernicoles du Transvaal. Ann. Soc. ent. Fr. 63: 63-67.

Simon, E. 1896. Description d’un arachnide cavernicole de |’Afrique australe. Bull. Soc. ent. Fr.
1896: 285-286.

Simon, E. 1897. Histoire naturelle des araignées. 2: 244. Paris: Roret.

Sion, E. & Face, L. 1922. Araneae des grottes de l’Afrique orientale. Arch. Zool. exp. gén. 60:

523-555:

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PNSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including Taste or Contents and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. x 7 in. (7$ in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (black type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

Smitu, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

Smit, C. D. 1954. South African plonias. Jn Brown, X. Y. Marine faunas. and ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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MUS. COMP. 70
eo Me KUIPER OL
J J LIBRARY

JUN 2 4 1964
“CONTRIBUTION TO THE KNOWLEBDGEaOF
| YE TY
PHE SOUTH AFRICAN SPECIES OF THE GENUS
PISIDIUM (LAMELLIBRANCHIATA)

April 1964 April
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MUS. COMP. ZOOL
LIBRARY

JUN 2 4 1964
CONTRIBUTION TO THE KNOWLEDGE OF THE SANE RSA
SPECIES OF THE GENUS PISIDIUM (LAMELLI
By

jd. G. J. JSuiper

c.o. Institut Néerlandais, Paris

(With 32 figures in the text)

CONTENTS
PAGE
Introduction . : - : : eM iig
Abbreviations : 78
Key to the South aia species of Pisidium 78
Descriptions of species. ; ; Ry ;°)
Summary. : , ; : . 94
References’ _. i : : : = 194
INTRODUCTION

The fauna of South Africa includes, as far as we know, seven species of
the genus Pisidium: pirothi Jickeli (syn. lepus Kuiper), harrisoni n.sp., costulosum
Connolly, Jangleyanum Melvill & Ponsonby, viridarium Kuiper, ovampicum
Ancey, casertanum (Poli).

Pisidium langleyanum, P. costulosum and P. harrisoni seem to be South African
endemics. The others are also known from Central Africa, while P. pirothi
and P. casertanum also occur in North Africa. Another species, P. artifex Kuiper,
which has been described from specimens taken in high mountain tarns in
Kenya, has perhaps to be added to the South African fauna.

The Pisidium fauna of South Africa is probably richer in species than the
present paper suggests. A number of dubious forms are known but not yet
described. Publication should wait until their specific identity has been cleared
up. To this end, much more collecting has to be done. In the first place large
series from each locality, preserved in alcohol (not in formalin which destroys
the shell), are needed. This is necessary for a study of their little-known anatomy
and in order to establish the subgeneric position of the species.

The identification of the African pisidia is generally difficult, largely
because of the high degree of discontinuity in their distribution and the specific
poverty of the Piszdium associations. Whereas in palearctic regions ten species
of Pisidium often occur in the same habitat, most of the African localities seem
to be inhabited by only one or two species, rarely three. Quantitatively I have
the impression that pisidia are not rare in South African inland waters. I have
seen many series from widespread localities, a great many more than are

(a
Ann. S. Afr. Mus. 48 (3), 1964, 77-95, 32 text-figures.

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

recorded in this paper, which were, however, all spoiled by formalin and so
could not be identified.

The present study is entirely based on material examined by myself.
This would not have been possible without the aid of many malacologists
who kindly put at my disposal the specimens and documents I needed. I am
specially grateful to Dr. A. D. Harrison, at the time principal research officer of
the National Institute for Water Research, Pretoria, who has furnished me, during
many years, with valuable samples of Pzsidium collected in South African rivers.

My grateful thanks are also due to the South African Council for Scientific
and Industrial Research for its assistance in the publication of this paper.

ABBREVIATIONS
~The following abbreviations are employed in the lists of distribution:
BML = British Museum (Natural History), London.

CSIR = Catalogue of the South African Council for Scientific and Industrial
Research, Pretoria.

IZRP = Institute for Zoological Research, Potchefstroom University.

KINB = Koninklijk Belgisch Instituut voor Natuurwetenschappen (Institut
Royal des Sciences Naturelles de Belgique), Brussels.

LMP = Laboratoire de Malacologie du Museum d’ Histoire Naturelle, Paris.

MHNG = Musée d’Histoire Naturelle, Geneva.

MMAT = Koninklijk Museum voor Midden Afrika (Musée Royal de l’ Afrique
Centrale) (formerly: Congo Museum), Tervuren, Belgium.

NIWR = National Institute for Water Research, Pretoria.

NMP = Natal Museum, Pietermaritzburg.

RML = Riyksmuseum van Natuurlike Historie, Leiden, Netherlands.

SAM = South African Museum, Cape Town.

SMF = Natur-Museum und Forschungs-Institut Senckenberg, Frankfurt,
Germany.

SSAE = Swedish South Africa Expedition 1950-1.

ZIUL = Zoological Institute, Lund University, Sweden.

ZMA = Zoologisch Museum, Amsterdam University, Netherlands.
ZMB = Zoologisches Museum, Alexander Humboldt University, Berlin.

Key TO THE SouTH AFRICAN sPECIES OF PISTIDIUM

1. Ligament external,* projecting outside shell; ligament-pit long and narrow; shell inequi-

lateral; sculpture regularly and finely striate ste bi , Se is pirotht
eaten internal, not visible externally .. ae oe an : é aie 2
2. Umbo far back; shell outline oblong ovate, anterior part very ee ee rather
regularly striate 3 ih ak harrisont
Umbo median or submedian; shell aarline pare or peninenale equidaiceal or inequilateral;
sculpture costulate or finely striate .. we ahs ae am oi as eee

* Kuiper (1962) distinguishes three positions of the ligament-pit: 1, introverted position
(ligament internal); 2, extroverted position (ligament external); 3, enclosed or normal position
(ligament internal, may be slightly visible externally, but does not project outside the shell).

S.A. SPECIES OF GENUS PISIDIUM (LAMELLIBRANCHIATA) 79

g. Sculpture regularly costulate; beaks submedian .. .. costulosum
Sculpture regularly and densely striate; nepionic shell often nocueled by 4-7 sharp, dis-
tinctive striae; shell outline subtrigonal .. . langleyanum
Sculpture P ilatly and densely striate; shell Te Seas seals: calli haretened at the end
of its inner slope ae .. viridarium
Sculpture irregularly striate, ie oa fs shell patina ae or subtrees. Ae ON

4. Sculpture irregular, very fine; shell swollen, diameter being often more than Boene beaks
tumid and extremely broad, ee ok at artifex
Sculpture irregular, very fine; shell normally olen Tees: ede half height; posterior
and anterior end nearly sols rounded; ventral margin of the shell much more curved
than the dorsal margin which is nearly straight; beaks median; hinge very narrow. . ovampicum
Sculpture irregular, fine; shell normally swollen, anterior end more pointed than posterior
end; inequilateral; dorsal margin more curved than ventral margin; beaks submedian;
central part of hinge rather broad .. a te fe oe ss .. casertanum

DESCRIPTIONS OF SPECIES
Pisidium pirothi Jickeli
(figs. 1, 9, 30)

Pisidium pirothi Jickeli, 1881: 340.

Pisidium (Fossarina) casertanum var. alexandrina Pallary, 1909: 75, pl. 4, fig. 34.
Pisidium cf. clarkeanum Gardner, 1932: 86, pl. 8, figs. 15-18.

Pisidium costulosum Haas, 1936: 43.

Pisidium clarkeanum var. exile Stelfox, Favre, 1943: 6-11, figs. 1-3.

Pisidium lepus Kuiper, 1957: 85, figs. 1-4.

Pisidium lepus var. dampfi Kuiper, 1957: 88, figs. 5, 6.

Pisidium (Afropisidium) pirothi Kuiper, 1962.

[Non] Pisidium costulosum Connolly, 1931.

Remarks

Pisidium pirotht is known from several localities in Egypt, south Sudan
and the Central African Republic (formerly French Sudan). It has been
collected in South Africa only in the Great Letaba River near Letaba Camp in
the Kruger National Park by F. Haas in 1931 (Haas, 1936, p. 43, s.n. P.
costulosum; Kuiper, 1957, p. 85, s.n. P. lepus n.sp.). These specimens are pre-
served in the Senckenberg Museum, Frankfurt, Germany (holotype SMF
155627, paratypes SMF 152628 and SMF 152886). Paratypes of P. lepus
have also been deposited in the Zoological Museum, Amsterdam (ZMA/K
4577), the South African Museum, Cape Town (SAM Aa2g771), and the
Natural History Museum, Geneva.

For eighty years P. pirotht remained an enigmatic species, owing to the
fact that the original description was not accompanied by figures and, secondly,
that the type-series seemed to be lost. Recently, however, Jickeli’s original
specimens were found (ZMB 37447, 1 lectotype and 6 paratype valves). ‘This,
in addition to the discovery of several other localities in Africa, made it possible
to define the range of what I some years ago called Pisidium lepus n.sp.

Pisidium pirotht can be easily distinguished from all other South African
species of this genus by its prominent external ligament and, consequently, the
peculiar situation of its ligament-pit. I drew attention to this fact in my paper
on P. lepus (Kuiper, 1957, p. 87). Its long, very narrow ligament-pit is bordered

1

ANNALS OF THE SOUTH AFRICAN MUSEUM

80

S.A. SPECIES OF GENUS PISIDIUM (LAMELLIBRANCHIATA) 81

ventrally by a kind of pliciform callosity which has its analogy in the genus
Eupera (Bourguignat, 1854; Klappenbach, 1960).

A similar ligament has been found in the endemic species of Lake Tan-
ganyika, Pisidium giraudi Bourguignat.

The shell of P. pirothi is thin, subtransparent, and inequilateral, with a
more or less pronounced, rather regular sculpture of concentric fine striae
(about 10 striae on $ mm. in the middle of the shell).

Details of the hinge structure are figured by Kuiper (1957, figs. 1 and 2).
See also figure 9 of this paper.

Pisidium pirothi is closely related to the Indian P. clarkeanum Nevill, the
Javanese P. javanum Benthem Jutting and the South American P. sterkianum
Pilsbry. All these species have an external ligament and belong to the sub-
genus Afropisidium (Kuiper, 1962, p. 55). The Indian species is considerably
larger and has a heavier shell than both the African and Indonesian species.
J. Favre (1943, p. 11) classified the Egyptian specimens provisionally as a
variety of the Indian species. I prefer to consider P. pirothi a distinct African
species. I shall come back to this matter in a separate paper on the Central
African pisidia.

Pisidium lepus is the type of the subgenus Afropisidium.

Pisidium harrisoni n.sp.

(figs. 2; 12.99)
Description

Shell very inequilateral, elongate ovate, thin, beaks low, scarcely promi-
nent, placed far back at one-quarter of the shell length. Dorsal margin short,
half the shell length, slightly curved. Ventral margin broadly rounded. Anterior
end very prolongate, obtusely pointed, upper side flattened. Posterior end
faintly curved, perpendicular. Sculpture consisting of irregular fine concentric
striae, 10-15 on 4 mm. in the middle of the shell. Ligament internal. The
holotype is a closed shell. Its dimensions are: L. 4:1 mm., H. 3:1 mm., D. 2°1
mm.

Type locality

Vaal River between Morgenzon and Amersfoort, Transvaal (CSIR Val.
443 AA), leg. A. D. Harrison, 15.1X.1958.

“=

Fic. 1. Pisidium pirothi Jickeli, Kruger National Park, Transvaal; L = ligament visible externally.

Fic. 2. P. harrisoni n.sp., holotype, Vaal River, Transvaal. Fic. 3. P. costulosum Connolly, Klein

Berg River, Cape Province. Fic. 4. P. langleyanum Melvill & Ponsonby, Lake Chrissie, Transvaal.

Fic. 5. P. viridarium Kuiper, Maseru, Basutoland. Fic. 6. P. ovampicum Ancey (paratype of

P. georgeanum Kuiper), Gwyang River, Cape Province. Fic. 7. P. casertanum (Poli), Mokhotlong,

Basutoland. Fic. 8. P. artifex Kuiper, paratype, Mt. Kenya. Enlargements: 1-5 and 7, X15; 6
and 8, x20.

82 _ ANNALS OF THE SOUTH AFRICAN MUSEUM

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Mages oe en
F cy Y make aa
Ym See! Aline aoe
Le

S.A. SPECIES OF GENUS PISIDIUM (LAMELLIBRANCHIATA) 83

Material

Holotype in the South African Museum, Cape Town (SAM A29773);
paratypes in the Zoological Museum of the Amsterdam University (ZMA/K
4979, 4980, 4982, 4983).

Pisidium harrison was also collected in the Vaal River near Vereeniging
(CSIR Val. 210 B) and at the Barrage (CSIR Val. 141 B), 1956 and 1958,
by Dr. A. D. Harrison. These specimens are all smaller than the type. Dimen-
sions of some paratypes:

L. 3-40 mm. H. 2-80 mm. D. 1-90 mm. (CSIR Val. 141 B).
L. 2°80 mm. H. 2-30 mm. D. 1:50 mm. (CSIR Val. 443 AA).
L. 2°35 mm. H. 1-85 mm. D. 1:10 mm. (CSIR Val. 210 B).

Description of the hinge of one of the paratypes (ZMA/K 4983): Hinge
plate narrow, in the left valve nearly as broad below the umbo as laterally, in
the right valve very narrow under the umbo; length (distance between the
cusps of AI and Pi) half the shell length. Cardinal teeth C2 and C4 weak and
short, C4 slightly posterior; C3 thin, thickened posteriorly. Cardinals nearer
to anterior laterals than to posterior laterals. Lateral teeth AI and PI long and
narrow, AIII and PIII very short; AII and PII shorter than AI and PI.
Ligament-pit interior short, one-sixth of the shell length (fig. 12).

Remarks

I dedicate this species to Dr. A. D. Harrison, presently senior research
fellow in the zoological department of the University College of Rhodesia and
Nyasaland, Salisbury, Southern Rhodesia.

Pisidium harrisoni may easily be distinguished from all other African species
hitherto known by its particular oblique shape. In this respect it somewhat
resembles the palearctic species P. subtruncatum Malm which extends as far
south as Algeria and Tangier.

Pisidium costulosum Connolly

(figs. 3, 11, 31)
Pisidium costulosum Connolly, 19314: 310, pl. 13, figs. 24-27; 1939: 626, fig. 57.
[Von.] Pisidium costulosum Haas, 1936-43.

co

Fics. 9-14. Hinges of Pisidium. Fic. g. P. pirothi Jickeli, Kruger National Park, Transvaal; left
valve at the top; AI, AII, AIII, anterior lateral teeth; PI, PII, PIII, posterior lateral teeth;
C2, Cg, C4, cardinal teeth; L, ligament-pit. Fic. 10. P. viridarium Kuiper, Chindamora Reserve
near Salisbury, Southern Rhodesia; left valve at the top; PI, PII, PIII, posterior lateral teeth;
C, callosity at the inner end of PIII. Fic. 11. P. costulosum Connolly, Klein Berg River, Cape
Province; right valve at the top. Fic. 12. P. harrisoni n.sp., paratype, Vaal River, Transvaal;
left valve at the top. Fic. 13. P. artifex Kuiper, paratype, Hall Tarns, Mt. Kenya; left valve at
the top. Fic. 14. P. casertanum (Poli), Mokhotlong, Basutoland, left valve at the top. Enlargement:
see scale in fig. 14.

84. ANNALS OF THE SOUTH AFRICAN MUSEUM

Fics. 15-19. Some forms of Pisidium langleyanum Melvill & Ponsonby. Fic. 15. Lectotype, left
valve at the top, C4 reduced. Fic. 16. Paratype, left valve, nearly oval. Fic. 17. Trigonal shel
with prominent beaks, right valve at the top, C4 reduced; specimen from Jonkershoek neat
Stellenbosch. Fic. 18. Left valve from Lake Chrissie; AS, anterior adductor scar; PS, posterior
adductor scar. Fic. 19. Subtrigonal specimen from Potchefstroom, Transvaal. Enlargements:

figs. 15, 16, 17 and 19, see scale in fig. 17; fig. 18, see scale in figure.

S.A. SPECIES OF GENUS PISIDIUM (LAMELLIBRANCHIATA) 85

Distribution (fig. 31)
Cape Province
Berg River near Cape Town, at three different places (CSIR Gbg. 303 D;
Gbg. 400 K; Gbg. 455 N), leg. A. D. Harrison, 1951 (ZMA/K 4189-4191).
Klein Berg River, dam in Tulbagh Kloof (CSIR Gbg. 732 L), leg. A. D.
Harrison, 1953 (SAM A29775; ZMA/K 4187, 4188).

Orange Free State

Rustfontein, Rhenoster River (near Bloemfontein), type locality (Connolly)
(BML 1937.12.30.9066, lectotype, and 9067-9, three paratypes).

| Transvaal

Vaal River, barrage near Vereeniging (CSIR Val. 141 B, Val. 155), leg.
A. D. Harrison, 1956 (ZMA/K 4978, 4991, 4994).

Klip River, headwater near Muller’s Pass, between Memel and Newcastle
(CSIR Val. 620 E), leg. A. D. Harrison, 1959 (ZMA/K 4977).

Klein Jukskei River near Linden, Johannesburg (CSIR Cro. 111 A and
130 E), leg..B. R. Allanson, 1956 (ZMA/K 4972, 4975).

Klein Jukskei River, Johannesburg (CSIR Gen. 44 A), leg. A. D. Harrison,
1954 (ZMA/K 4192).

Braamfontein Stream near Northcliff, Johannesburg (CSIR Cro. 48 B
and 118 F), leg. B. R. Allanson, 1956 (ZMA/K 4973, 4976).

Zandfontein Stream, confluence with Jukskei Stream, near Leeukop
Prison Farm, Pretoria district (CSIR Cro. 109 A), leg. B. R. Allanson, 1956
(ZMA/K 4974).

Vaal River near Standerton (CSIR Val. 29 J), 7.XI1I.1955 (ZMA/K 4997).

Description

Pisidium costulosum may easily be distinguished by its regular sculpture
of concentric narrow ribs. Its shape does not vary greatly. The characteristic
sculpture, however, shows some variability. On the lower part of the shell the
ribs are relatively widely spaced (about 5 ribs on } mm.), the spaces between
the ribs being at least twice as wide as the ribs themselves. Towards the beaks
the ribs become narrower and closer (up to 15 ribs on } mm.). Sometimes the
smooth nepionic shell is surrounded by some strong, more widely spaced ribs.

The shell is regularly porous, the distribution of the pores not being
limited to the interstices of the ribs. The ligament is long and internal. The
largest known specimen of P. costulosum measures: L. 3:4 mm., H. 2:8 mm.,
D. 2:1 mm. This was collected in the Klein Berg River, Cape Province (ZMA/K
4187).

Remarks

Apart from the Berg River, P. costulosum is also known from the Vaal and
Orange River systems. It seems to be a typical South African species. It has
only been collected in rivers and streams, so far never in brooks. P. costulosum
is ofcen associated with P. langleyanum.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fics. 20-24. Lectotype of Pisidium ovampicum Ancey. Fic. 20. Right valve. Fic. 21. Dorsal view
of both valves, right valve at the top; for abbreviations see explanation to fig. 9. Fic. 22. Ventral
view on hinge, right valve at the top; for abbreviations see explanation to fig. 9. Fic. 23. View
on hinge, left valve at the top; AS, anterior adductor scar; PS, posterior adductor scar. Fic. 24.
Profile view. Fic. 25. P. georgeanum (= P. ovampicum), paratype as figured by Kuiper, 1952;
a, left valve; b, ventral view on hinge of left valve; c, right valve; d, ventral view on right valve;
e, outline of juvenile specimen; f, profile view of single valve. Enlargement: figs 20-25, see scale
above fig. 24.

S.A. SPECIES OF GENUS PISIDIUM (LAMELLIBRANCHIATA) 87

The population recorded by Haas (1936, p. 43) in the Kruger National
Park, Transvaal, does not belong to P. costulosum, but to P. lepus (Kuiper 1957),
which proved to be a synonym of P. pirothi Jickeli.

Pisidium langleyanum Melvill & Ponsonby

(figs. 4, 15-19, 27)
Pisidium langleyanum Melvill & Ponsonby, 1891: 237.
Pilsbry & Bequaert, 1927: 351.
Connolly, 1939: 625.
Kuiper, 1960: 67, figs. 18-23.
[?] Pistdium cf. langleyanum: Boettger, 1910: 455, pl. 28, fig. 19 a—b.
[Non] Pisidium langleyanum: Melvill & Ponsonby, 1892, pl. 5, fig. 7.
Distribution (fig. 27)
Cape Province
Port Elizabeth, leg. J. H. Ponsonby (Melvill & Ponsonby, 1891, p. 237;
BML 1902.7.30.40-42, lectotype and 2 paratypes; SMF 152884, 1 paratype,
leg. Ponsonby, 1889, ex coll. O. Boettger; Kuiper, 1960, figs. 18-23).
Jonkershoek near Stellenbosch, fish-ponds (ZMA/K 4185; BML 15 sp.).
Hogsback near Alice, small stream at the edge of the plateau at a height of
about 4,000 to 4,500 feet (CSIR Misc. 10 and Misc. 62 B), leg. A. D. Harrison,
1950 and 1953 (ZMA/K 3762, ZMA/K 4177).
Kimberley, leg. J. A. van Eeden, 1.X.1958 (IZRP; ZMA/K 13481).
Grahamstown, leg. J. A. van Eeden, 13.III.1959 (IZRP; ZMA/K 13485).
Uniondale, leg. J. A. van Eeden, 24.III.1959 (IZRP; ZMA/K 13487).

Orange Free State

Rustfontein, Rhenoster River (near Bloemfontein) (Connolly, 1939, p. 625;
BML/Mus. Cuming 1937.12.30.9073, I specimen).

Natal

Durban, leg. J. A. van Eeden, 30.VII.1959 (IZRP; SAM A29769;

ZMA/K 13488).
Transvaal

Klein Vaal (NIWR Val. 733 LL), about 9 miles from its confluence with
the Vaal River (ZMA/K 13489).

Headwater stream of Vaal River between Breyton and Lake Chrissie
(CSIR Val. 743 DD), 22.VII.1959 (ZMA/K 4980).

Lake Chrissie, leg. F. G. Gawston (NMP).

Potchefstroom, leg. M. I. Livingston (NMP).

Description

The shell of P. langleyanum is characterized by its subtrigonal shape and
its regular sculpture of fine concentric striae. Its nepionic shell is sometimes
bordered by some (4-7) sharp striae. The shell is densely porous. Its dorsal
margin is more strongly rounded than its ventral one. The shell of the type is

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

ae eK eK

1000 MILES

PISJIDIUM OVAMPIcUM ANCEY

26

ree

CO ee

PIS, PIROTH!) JICKELI

216)

PIS. COSTULOSUM CONN,

31

Pils. HARRISONI N-SP.

32

Fics. 26-32. Distribution of Pisidia in South Africa. Pisidium ovampicum Ancey and P. casertanum
(Poli) are also known from Madagascar.

= Terw?

S.A. SPECIES OF GENUS PISIDIUM (LAMELLIBRANCHIATA) 89

rather thin and has a narrow hinge (figs. 15, 16). The posterior laterals are
parallel and do not converge proximally. A detailed description of the type is
given by Connolly (1939, p. 625), who calls it ‘a somewhat immature shell
with hinge system poorly developed’ (fig. 15). In heavier shells, such as those
from Hogsback, Cape Province, the outer posterior lateral (PIII) bends
proximally slightly towards the inner posterior lateral (PI).

The outline of the shell is more or less ovate when the beaks are not
prominent, and trigonal when they are prominent. Both forms may occur in
the same population (Jonkershoek, Cape Province).

The ligament is internal. It is, however, partially visible externally,
resulting from a slight gaping of the valves behind the beaks, a feature which is
rather common in the genus Prsidium.

Dimensions of the lectotype: L. 2-9 mm., H. 2:5 mm., D. 1-8 mm. Dimen-
sions of a number of other specimens from several localities:

fe omm, H.3-4mm. D.2-6mm. Kimberley.

L.38mm. H.33mm. D.2:8mm. Lake Chrissie, swollen shell (fig. 18).
fieaamm. HH. 32mm. D.2-4mm. Jonkershoek, trigonal shell (fig. 17).
ho32mm. H.26mm. D.2:3mm. Port Elizabeth.

L.3-omm. H.26mm. D.2:2mm. Hogsback.

L.2-9mm. H.2mm. D.2:0mm. Jonkershoek.

26mm. H.2:4mm. D.1t-9mm. Jonkershock.

L.26mm. H.2:3mm. D.1-8mm. Hogsback.

me24mm, H.otmm. D. 1-5 mm.” Hogsback.

be2-2mm. H.2o0mm. D.1:5 mm. Potchefstroom (fig. 19).

Details of the hinge structure are figured by Kuiper (1960, figs. 18-20, 23).

Remarks

The specimen figured by Melvill & Ponsonby (1892, pl. V, fig. 7) is not
Pisidium langleyanum, but another species (Kuiper, 1961, p. 67).

O. Boettger (1910, p. 455) recorded 6 subfossil valves from Witkop,
Bechuanaland, under the name ‘Pisidium cf. langleyanum’. Connolly (1939,
p- 625) has placed this form into the synonymy of P. ovampicum Ancey. In my
opinion this is not correct. Boettger’s figures (1910, pl. 28, fig. 19 a—b) prove
that it is not the ovate P. ovampicum, but a more trigonal species, perhaps indeed
P. langleyanum. One of these specimens, a very small left valve, which I had the
opportunity to examine (SMF 152885), did not allow me to decide with cer-
tainty about its specific identity. I have not seen the other valves.

Pisidium langleyanum is known only from southern Africa. It is closely allied
to P. kenianum which is distributed in Central Africa. Perhaps they will prove
to be identical. Both species are related to the Indian P. nevillianum Theobald.
P. kenianum has a trigonal outline and is heavier. It can be considerably larger
than P. langleyanum (up to 4:6 mm.). The distance between the northernmost
known localities of P. langleyanum (Transvaal) and the southernmost ones of
P. kenianum (in Tanganyika) is about 1,500 miles.

gO ANNALS OF THE SOUTH AFRICAN MUSEUM

Pisidium viridartum Kuiper

(figs. 5, 10, 29)
Pisidium viridarium Kuiper, 1956: 61-63, figs. 1-8.
Distribution (fig. 29)

Cape Province

Gwyang River near George (CSIR Frw. 88), leg. A. D. Harrison, 4.V.1950
(ZMA/K 4178).
Basutoland

Maseru, Roma Mission Station, 5.[V.1958, leg. J. A. van Eeden (IZRP;
SAM A29772; ZMA/K 13483).

Remarks

Pisidium viridarium was described from specimens collected in Kenya.
This species is also known from Southern Rhodesia, Ethiopia, Uganda, Ruanda
and Congo.

Pisidium viridarium has a thin, ovate shell with submedian beaks. Young
specimens somewhat resemble P. ovampicum. The regular sculpture of fine,
equidistant concentric striae is the same as in P. kenianum and P. langleyanum.
Its PIII is proximally thickened (fig. 10) as is often found in P. kenianum. The
shell is densely porous. The affinities between P. viridarium, P. kenianum and
P. langleyanum have not as yet been cleared up. The resemblance to the Euro-
pean P. personatum which, incidentally, has been collected in Uganda, is
probably phenotypical. P. vzrzdarium attains larger dimensions than P. personatum,
one of the paratypes being nearly 5 mm. long. 7

Details of the hinge structure are figured by Kuiper (1956, figs. 1, 4, 5, 6).

Ecologically P. viridarium seems to prefer marshy biotopes. It also lives in
the mud out of the current in rivers where the aquatic vegetation is dense.
It has been found associated with P. kenianum and P. ovampicum.

Pisidium ovampicum Ancey
(figs. 6, 20-26)
Pisidium ovampicum Ancey, 1890: 162.
Pisidium georgeanum Kuiper, 1952: 46-48, figs. a—/.
[Non] Pisidium ovampicum Connolly, 1939: 625.
Distribution (fig. 26)
Cape Province
Gwyang River near George, type locality of P. georgeanum Kuiper (CSIR
Frw. 88), leg. A. D. Harrison, 4.V.1950 (holotype of P. georgeanum in RML,
paratypes in LMP, MHNG, SAM A29774, ZMA/K 3604).
Jonkershoek near Stellenbosch, fish-ponds, leg. A. C. Harrison, 1947
(ZMA/K 4184).

S.A. SPECIES OF GENUS PISIDIUM (LAMELLIBRANCHIATA) QI

South West Africa

Ovamboland (Ancey: Ovampoland = Damaraland), Omanbondé, type
locality of P. ovampicum, leg. viatores Andersson and Chapman (KINB I.G.1059,
lectotype of P. ovampicum).

Remarks

Pisidium ovampicum has been an enigmatic species for more than half a
century. The original description is fairly good, but it is impossible to interpret
because details of the hinge structure and figures were not given, and the type
material was apparently lost.

Connolly (1931a, p. 325), who therefore considered Ancey’s species as
being ‘null and void, a just fate for such a slovenly description’,* has tried to
give a new identity to P. ovampicum. He (1939, p. 626) described the hinge of
specimens which he thought specifically identical with P. ovampicum Ancey.
He too, however, failed to publish illustrations.

P. ovampicum thus remained an obscure species until recently when an
authentic specimen of this species was found in the Dautzenberg collection
(KIN, Brussels). I now designate this specimen the lectotype of P. ovampicum
Ancey. It was, as supposed by Connolly (1939, p. 626), ‘unopened’. I opened it
by soaking the shell in water for some days.

The specimens from Lake Chrissie, on which Connolly based his redescrip-
tion ‘with the purely tentative suggestion that they may represent Ancey’s lost
species’ (Connolly, 1939, p. 625), do not belong to P. ovampicum but to P.
casertanum.

Description

Description of the lectotype P. ovampicum. Shell thin, ovate, nearly equi-
lateral, opaque, no pores visible, glossy. Beaks smooth, median, not prominent,
scarcely rising above the dorsal shell margin. Sculpture consisting of very fine
irregular striae and two slight growth-lines. Greatest shell length in the hori-
zontal median plane, greatest shell height in the vertical median plane. Ventral
margin more rounded than dorsal margin. Anterior and posterior margins
nearly equally rounded. Ligament internal, dark brown. Hinge plate narrow.
In the left valve the lateral parts are scarcely broader than the central part;
in the right valve the lateral parts are a little broader than the central part. The
lateral teeth, PI, PII, AI and AII are narrow and have pointed, distal cusps.
PIII and AIII are short, small teeth. Cardinal teeth thin, short and straight,
C3 being the longest of all. C2 is a short tooth on the ventral margin of the
hinge-plate; C4 as long as C2 and parallel to it. The distance between the
cusps of AII and PII is 1-5 mm.; between those of AII and Ca, 0-6 mm.;
between PII and C2, 0o-g mm. Ligament-pit 0-5 mm., narrow. Posterior
adductor a little larger and situated lower than the anterior adductor. Dimen-

* It is curious that Connolly (1939, p. 626), quoting Ancey’s original description, omitted
its beginning: “Testa fragilis, pallide cornea... .

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

sions: L. 2-6 mm., H. 1-9 mm., D. 1-5 mm. (in the original description L. 22
mm,, H..27 mm.) D.14, mm.)

The differences between the paratype of P. georgeanum and the lectotype of
P. ovampicum are slight. Shape and sculpture are nearly the same. Equally large
specimens of P. georgeanum have slightly more prominent beaks and conse-
quently are a little higher. The junction between the dorsal and the anterior
as well as the posterior margins of P. georgeanum are slightly more angulate.
In both forms the hinge-plate is narrow, the laterals are thin and sharp, the
cardinals small. In the lectotype of P. ovampicum, C4 lies slightly more posteriorly
with respect to C2 than in the paratypes of P. georgeanum. In both forms the
ventral margin is more curved than the dorsal margin. Pisidium georgeanum
is porous, whereas the lectotype of P. ovampicum has no pores at all.

Details of the hinge structure of P. georgeanum are figured by Kuiper

(1952, figs. a—d).

Remarks

The form georgeanum is widely distributed in South and Central Africa,
as well as on the isle of Madagascar. I have seen series from Uganda, Ethiopia,
Congo and Madagascar.

The following records turned out not to be P. ovampicum:

(a) ‘A single weathered valve from Damara or Ovamboland’ (Connolly,
1939, p. 625; BML 1937-12.30.9070).

(b) Two ‘better grown and preserved specimens from Lake Chrissie’ (Connolly,
1930, p. 625; BML 1937.13.30.9971—9972), originally published by Hutchinson
(1932, Pp. 49).

(c) The subfossil valves from Witkop Bechuanaland (SMF 152885), mentioned
by O. Boettger (1910) as P. cf. langleyanum and placed by Connolly in the
synonymy of P. ovampicum.

I am not yet certain of the identity of the specimens mentioned under
(a) and (c). I believe that (6) belongs to Pisidium casertanum (Poli). As to the
habitat of these two specimens Hutchinson (1932, p. 49) wrote: ‘... taken ona
rock surface under very wet soil at one of the springs bordering the lake.’ This
agrees well with the ecology of P. casertanum.

Dr. A. D. Harrison wrote me the following details on the first-mentioned
locality: “The Gwyang, where these specimens were collected, is a small stream
with a muddy bottom covered with Chara or Nitella; they were not very nume-
rous and I had to sort through three or four sieve-fulls of bottom before we
obtained the specimens in the tube. The water is acid with a PH of about 5;
it is brown or peat stained.’

As to the second locality, the fish-ponds near Jonkershoek, Dr. Harrison
noted: ‘The specimens were originally found fastened on to the noses of young
trout.’ This observation is very interesting in connexion with the passive dis-

S.A. SPECIES OF GENUS PISIDIUM (LAMELLIBRANCHIATA) 93

persal of these small clams by means of other animals, a subject about which
very little is known.*

Pisidium artifex Kuiper
(figs. 8, 13)

Pisidium langleyanum (non M. & P., 1891), Melvill & Ponsonby, 1892: 94, pl. 5, fig. 7.
Pisidium artifex Kuiper, 1960: 68-74, figs. 1-17.

Holotype in BML. Paratypes: SMF 162841-2; SAM A29770 (paratype
Nr. 17); Coryndon Museum, Nairobi; ZMA/K 4909-4925, 4927.

Remarks

This species is known only from some high mountain tarns in Kenya. It is
easily recognizable by its extraordinary swollen shape, its diameter being the
same or even more than the height of the shell, and by its very broad and tumid
beaks.

Details of the hinge structure are figured by Kuiper (1960, figs. 1, 2, 3, 6
and 9).

In the original description of P. artifex, I drew attention to the fact that the
specimen figured by Melvill & Ponsonby (1802, pl. 5, fig. 7) cannot be P.
langleyanum but probably belongs to P. artifex. If this should prove to be true,
P. artifex has to be added to the South African fauna. Melvill & Ponsonby
(1892), however, do not mention any locality.

Pisidium casertanum (Poli)
(figs. 7, 14, 28)

Cardium casertanum Poli, 1791: 65, pl. 16, fig. 1.

Pisidium ruwenzoriense Germain, 191 1a: 135.

Pisidium ovampicum (non Ancey) Connolly, 1939: 625.

Pisidium edouardi Kuiper, 1953: 26-28. |

There are several specimens in the South African Museum (A29767 and

A29768) from Ethiopia and Cyrenaica.

Distribution (fig. 28)
Cape Province
Krom River, Stellenbosch (CSIR Ers. 6 L), leg. A. D. Harrison,
22.VIII.1952 (ZMA/K 4757).
Transvaal
A spring bordering Lake Chrissie (Hutchinson, 1931, p. 49, s.n. Pzsidium
sp. juv.; Connolly, 1939, p. 625, s.n. P. ovampicum; BML 1937.12.30.9971—72).

Basutoland
Makkeka Mountains, 15 miles E.N.E. Mokhotlong, alt. 9,500 ft. In slow-
flowing part of the stony high mountain stream, leg. Per Brinck (SSAE),
8.1V.1951 (ZIUL 2931; ZMA/K 4414).

* See the classic work of H. Wallis Kew: The dispersal of shells. London, Kegan Paul & Co.,
1893, pp. 61 sqq. [Ed.].

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

South West Africa

Grootfontein, farm Urupupa, leg. Thomson, 1912 (SMF 152883, one
right valve ex coll. O. Boettger).

Remarks

Pisidium casertanum (Poli) is the commonest and most variable species in
the holarctic region. In the Southern hemisphere this species is less variable.
In northern Africa it lives in the coastal region from Morocco to Cyrenaica.
It has not yet been found in Egypt. P. casertanum (Poli) var. alexandrina Pallary
belongs not to this species but to P. pirothi Jickeli.

In equatorial Africa it is known from many high mountain localities in
Ethiopia, Uganda and Tanganyika. It has also been collected in Rhodesia
(Herrington, 1962, p. 34). P. casertanum has been described from Madagascar
under the name P. edouardi. The Austrian Madagascar Expedition 1958
collected this species commonly in central Madagascar. The specimens from
the Makkeka Mountains, Basutoland, collected by the Swedish South Africa
Expedition 1951, are identical with the types of P. edouard: from Madagascar

(Kuiper, 1953, pl. 1, figs. 1-5). The dimensions of the five specimens taken in

the Makkeka Mountains are:

L,3:8 mum. SE 3-2 2 ame.
L: 9-6 mm, » iT: 970:mim., |) Desa:
Le 9:4 mum.) Ele eon) er -omima:
L.12-o7°mm, He 2:4 moms moe
L. 28 mm. E2-3.mm. ° D455 aa

The East African mountain range seems to provide a climatic bridge from
the palearctic region to the southern African area for this species. The dis-
persion involved is perhaps due to the migration of aquatic birds.

The identification of this variable species is often difficult. I have based
my conclusions on conchological characters only, as these are more reliable

than anatomical characters for specific identification in the genus Pisidium.

SUMMARY

The seven, or perhaps eight, species of the fresh-water lamellibranch
genus Pisidium which are found in South Africa are described, with one
new species, P. harrisoni. Taxonomic notes and figures and remarks on distribu-
tion are included.

REFERENCES

Ancey, C. F. 1906. Description of two new Cleopatra and a new Pisidium. Nautilus 20: 45-46.

BoETTGER, O. 1910. Die Binnenkonchylien von Deutsch-Siidwest-afrika und ihre Beziehungen
zur Molluskenfauna des Kaplandes. Abh. senckenb. naturf. Ges. 32: 431-456.

BourcuicenaT, J. R. 1854. Du genre Pisidium. Aménités malacologiques 1: 20-68.

Conno.iy, M. 1931a. Contributions to a knowledge of the fauna of South West Africa. IX.
The non-marine Mollusca of South West Africa. Ann. S. Afr. Mus. 29: 277-333.

Ry A vate ences

ii ta ee, A ll

S.A. SPECIES OF GENUS PISIDIUM (LAMELLIBRANCHIATA) 95

Conno ty, M. 19315. Descriptions of new non-marine Mollusca from North, South and Central
Africa, with notes on other species. Ann. Mag. nat. Hist. (10) 8: 305-321.

Conno ty, M. 1939. A monographic survey of South African non-marine Mollusca. Ann. S. Afr.
Mus. 33: 1-660.

Favre, J. 1943. Revision des espéces de Pisidium de la Collection Bourguignat du Muséum
d’Histoire Naturelle de Genéve. Rev. suisse Zool. 50, fasc. suppl.: 1-64.

GarpneR, E. W. 1932. Some lacustrine Mollusca from the Faiyum Depression. A study in
variation. Mém. Inst. égypt. (Egypte) 18: 1-119.

Germain, L. 1911a. Contributions a la faune malacologique de l’Afrique équatoriale. XXIV.
Mollusques nouveaux de la région du Tchad et de l’Est africain. Bull. Mus. Hist. nat., Paris
17: 133-136.

GERMAIN, L. 19115. Etude sur les mollusques terrestres et fluviatiles recueillis au cours de la
mission de délimitation du Niger-Tchad. Docum. sci. Miss. Tilho 2: 165-245.

Haas, F. 1936. Binnen-Mollusken aus Inner-Afrika, hauptsachlich gesammelt von Dr. F. Haas
wahrend der Schomburgk-Expedition in den Jahren 1931-32. Abh. senckenb. naturf. Ges.
431: 1-156.

HeErrincTon, H. B. 1962. A revision of the Sphaeriidae of North America. Misc. Publ. Mus. Zool.
Univ. Mich. 118: 1-74.

Hutcuinson, G. E., Pickrorp, G. E. & ScHuuRMAN, J. F. M. 1932. A contribution to the
hydrobiology of pans and other inland waters of South Africa. Arch. Hydrobiol. (Plankt.)
24: I-154.

Jickxeui, J. F. 1881. Land- und Siisswasser Conchylien Nordost-Afrika’s, gesammelt durch J.
Piroth. 7b. dtsch. malakozool. Ges. 8: 336-340.

KLAPPENBACH, Miguel A. 1960. Ueber die Gattungen Byssanodonta und Eupera. Arch. Mollusk.
89: 141-143.

Kuiper, J. G. J. 1952. Pisidium georgeanum, une nouvelle espéce africaine. Basteria 16: 46-48.

Kurrer, J. G. J. 1953. Description de trois nouvelles espéces de Pisidium de Madagascar. 7.
Conchyliol. g: 26-31.

Kurrrr, J. G. J. 1956. Pisidium viridarium, eine neue Art aus Ost-Afrika. Arch. Mollusk. 85: 61-63.

Kurrrr, J. G. J. 1957. Pisidium lepus, eine neue Art aus Afrika. Arch. Mollusk. 86: 85-90.

Kurrrr, J. G. J. 1960. Pisidium artifex, eine neue Art aus Kenya. Arch. Mollusk. 89: 67-74.

Kuiper, J. G. J. 1962. Note sur la systématique des Pisidies. 7. Conchyliol. 102: 53-57.

Me tvit1, J. C. & Ponsonsy, J. H. 1891. Descriptions of nine new terrestrial and fluviatile
mollusks from South Africa. Ann. Mag. nat. Hist. (6) 8: 237-240.

_ Me vit, J. C. & Ponsonsy, J. H. 1892. Descriptions of seventeen new terrestrial mollusks from
South or Central Africa, in the collection of Edgar L. Layard Esq. Ann. Mag. nat. Hist. (6)
9: 87-97.

Nevitt, G. & NeEvitx, H. 1871. Description of new Mollusca from the Eastern regions. 7. Asiat.
Soc. Beng. 40, 2: I-11.

PaLLary, P. 1909. Catalogue de la faune malacologique de l’Egypte. Mém. Inst. égypte (Egypte)
6, 1: I-91. .
Pirssry, H. A. & BEQuaAERT, J. 1927. The aquatic mollusks of the Belgian Congo. Bull. Amer.

Mus. nat. Hist. 53: 69-659.
Pout, J. X. 1791. Testacea utriusque Siciliae ... 1. Parmae.

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PENS TRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TABLE OF CONTENTS and SumMARyY. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 42 in. x 7 in. (7 in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (black type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

Smiru, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

Smity, C. D. 1954. South African plonias. In Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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MUS. COMP. ZOOL
LIBRARY

MAY 29 1964

HARVARD
UNIVERSITY,

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA)

By
J. H. Day
Koology Department, University of Cape Town

CONTENTS

PAGE

Introduction ae ile 2 wee OF
Discussion of diagnostic characters ah). 9S
Division into sub-families and genera .. 103
Sub-family Melinninae.. a Jen Tob
Table of type-species .. me TOA:
Key to genera Bs se 22, LOG
Generic definitions .. ane ste ey LOO
Sub-family Ampharetinae ne LO”
Table of type-species .. fe LOU
Key to genera a ate ie ee:
Generic definitions .. Se a by Ae:
Summary .. o sa a acer IG
References .. = es aa Je, REDO

INTRODUCTION

The most complete account of the family Ampharetidae is that of Hessle
(1917). He reviews earlier work including that of Malmgren (1865) and Fauvel
(1897) on family and generic characters, Nilsson (1912) on the nervous system,
Wiren (1885) on the alimentary canal, and Meyers (1887) on the nephridia.
After giving further evidence from his own researches on the nephridia and
structure of the stomach he goes on to discuss generic characters. While he
follows the main lines laid down by Malmeren he finds that the genera were so
narrowly defined by the latter worker that they seldom include more than one
or two species, so that broader generic definitions are necessary. He criticizes
Fauvel’s work on the enumeration of the anterior segments and his genera as
being based partly on internal anatomical characters. Surprisingly enough
Hessle’s own generic divisions may be criticized on the same lines. Although
he stresses that the number of nephridia may vary from species to species in
the same genus he has grouped the genera largely on the relative sizes and
arrangement of these internal structures.

Since the publication of Hessle’s work many new genera have been erected,
particularly by Augener (1918), Chamberlin (1919), Benham (1921),
Annenkova (1930), Caullery (1944) and Eliason (1955). In general these later
workers focused attention on external features, particularly the development

97
Ann. S. Afr. Mus. 48 (4), 1964, 97-120.

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the paleae and the number of thoracic setigers, and they do not describe the
nature of the nephridia nor the morphology of the stomach. As a result it is
difficult to correlate the genera they describe with those defined in Hessle’s
monograph. There is also another source of confusion. Nilsson and Hessle
recognize only a single two-ringed buccal segment in front of the paleal or
first branchiferous segment, so that the latter becomes segment II. This inter-
pretation is followed by Annenkova (1930), Eliason (1955), and Uschakov
(1957). On the other hand Malmgren (1865), Fauvel (1897 and 1927) and
Caullery (1944) recognize two segments in front of the paleae, so that the latter
which also bears the first pair of branchiae becomes segment III. At first sight
it would seem of little taxonomic importance whether the first branchiferous
segment is labelled segment II or III, but as many workers do not state which
system of numbering they are using, statements such as ‘setae present on seg-
ment III’ or ‘nephridia absent from segment V’ or ‘notosetae of segment XIII
modified’ lead to confusion.

To date about 150 species have been described, distributed through about
49 genera. Fifty-six species are ascribed to the three genera Amphicteis, Amage,
and Ampharete, and the remaining 94 are distributed through the remaining
46 genera, many of which are monotypic. The need for revision is obvious.

In the account that follows the various diagnostic characters are discussed,
the grouping of genera is considered, and tables and keys are provided for the
whole family.

I wish to thank Dr. Olga Hartman for constructive criticism during the
preparation of this paper, and both Mr. R. Sims of the British Museum and
Mr. J. B. Kirkegaard of the Copenhagen Museum for allowing me to examine
material housed in their respective institutions. The Trustees of the South
Afritan Museum are grateful to the South African Council for Scientific and
Industrial Research and the University of Cape Town for grantis in aid of
publication.

DiscussION OF DIAGNOSTIC CHARACTERS

The head and buccal tentacles. The Ampharetidae are deposit feeders and
the food particles are collected by mobile projections that arise from the roof
of the buccal cavity. The most common type are buccal tentacles which are
either smooth with a longitudinal groove on one side as in the Terebellidae or
papillose; often these papillae are pinnately arranged, but not always, and ©
buccal tentacles of this type are best described under the more general term—
‘papillose’. Apart from tentacles, folded or probosciform feeding membranes
have been described in Amythas membranifera Benham and Pabzts deroderus
Chamberlin. An examination of Isolda whydahensis Augener showed a long
ribbon-shaped structure protruding from the mouth with the distal end split
into a number of short tentacles.

As stated, these feeding organs arise from the roof of the buccal cavity and ©
in some cases the mouth cavity leads straight back into the pharynx; in other
cases, however, an internal shelf or horizontal septum separates the tentacular

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 99

cavity above from the pharynx below. According to Hessle this septum
corresponds to the dorsal lip of the 'Terebellidae. ,

Above the mouth there is a hood-shaped lobe which Hessle refers to as the
*“Tentakelmembran’ and Fauvel as the prostomium. Since it bears the eyes and
nuchal grooves and contains the cerebral ganglia the latter term will be used
here. In some genera it also bears a pair of glandular ridges which diverge
anteriorly and project from the antero-lateral margins as a pair of blunt
processes.

When the buccal tentacles are retracted the prostomium may appear
bluntly pointed with obvious lateral grooves, so that the whole prostomium
(or tentacular membrane) has been described as trilobed. When the buccal
tentacles are fully extended, however, the whole head is stretched, the anterior
margin is straightened, and the lateral grooves disappear. This mobility of the
head means that the exact shape of the prostomium is of little systematic value.
The glandular ridges, however, are always recognizable and the eye-spots can
usually be found.

The segmentation of the head region. As described earlier (Day 1961) the region
behind the prostomium is telescoped and the segments are distorted. ‘The
branchiae are often grouped on a transverse branchial ridge which partially
overhangs the segments immediately behind the prostomium. The ventral
surface in Melinna and related genera is distorted by the formation of a pair of
lateral folds which slope back from the mouth region towards the dorsum of
segment VI where they unite to form a transverse dorsal crest. On the basis of
dissection and examination of external features I have accepted the interpreta-
tion of Malmgren, Fauvel, and others that there are two achaetous segments
preceding the branchiae in all genera. The first branchiferous segment, which
bears the paleae in some genera, thus becomes segment III.

The branchiferous region (segments III-VI). In the more primitive genera,
including Phyllocomus, Melinna, Isolda, Amphicteis and Ampharete, there are four
pairs of gills. Dissection of the blood-vessels of Amphicteis gunneri, described by
Day (1961), showed that the four gills are supplied by four blood-vessels
corresponding to segments III, IV, V and VI. With the telescoping of the
anterior segments, however, there is a tendency for the four pairs of gills to
be grouped on the dorsal surface of segments III and IV. In Phyllocomus the
first two pairs are on segment III and the last two pairs are segmentally
arranged. The same arrangement has been reported in Amage auriculata and
Amphicteis posterobranchiata. In other genera the arrangement may be 2 : 2 or
3 : 1 where three pairs of gills are arranged in a transverse row across segment.
III with the fourth pair immediately behind.

In many genera the primitive number of four pairs of gills has been
reduced to three and in Auchenoplax Eblers and its synonym Melinnoides Benham
only two pairs have been reported.

The gills may be separate from one another or united by a basal web.)

| ele) ANNALS OF THE SOUTH AFRICAN MUSEUM

Commonly this web unites only the first three pairs of gills, leaving the fourth
posterior pair free. The gills may be smoothly cylindrical in shape or may
develop lateral papillae or flanges or even a series of lateral lamellae. Commonly
the branchial papillae are arranged in two lateral rows, and such branchiae
have been termed bipinnate, but this is not always the case. In Pterampharete
luderitzt, for example, numerous irregularly arranged papillae arise from the
anterior surface of the branchial axis. In generic descriptions such gills are
best described by the broader term papillose. Similarly the lateral lamellae of
such forms as Phyllamphicteis collaribranchis Augener and Phyllocomus (olim
Schistocomus) hilton (Chamberlin) may be arranged in one or more rows.

The telescoping of the branchiferous region often results in the fusion of
segments and the loss of setae. In Jsolda and Melinna of the sub-family Melinninae
the neurosetae of the four segments III-VI are present though those of segment
VI may be lacking in some species. Segment III never has notosetae, and
segment IV has notosetae in the form of stout hooks behind the gills, but the
notosetae of segments V and VI are either small or absent. In Melinnopsis —
McIntosh the hooks of segment IV are lacking, but notopodial capillaries are ~
present on segments V and VI.

In the sub-family Ampharetinae neurosetae are never developed on segments
III to VI. In a few cases all four segments may be distinct, but usually segments
III and IV are fused and occasionally segments III, IV and V. The notosetae
are variously developed. In primitive forms such as Amphicteis the notosetae of
segment III are enlarged to form stout paleae and those of segments I[V—VI are
all present as normal capillaries. Often, however, the capillary notosetae of 1V
and V are small or absent. Similarly the notosetae of segment III may remain
small and not form paleae, or may be entirely lacking. In Sabellides, for example,
the notosetae of segment III are either absent or represented by small capillaries,
segment IV is fused to segment III and lacks setae, and the first normal capil-
laries are on segment V. In Neosabellides elongatus there are no setae on segments
III and IV, and in Paramage madurensis Caullery reports that the first bundle of
notosetae appears on segment VI though earlier achaetous notopodia remain.
In view of the many variations in the setation of the branchiferous segments
III to VI and the difficulty of deciding whether small setae present on the fused
third and fourth segments represent small paleae of segment III or small
capillaries of segment IV, it would seem undesirable to base generic divisions
on the presence or absence of paleae as has been done in the past.

Internal characters: diaphragm, nephridia, stomach. Apart from a few minor
details no new researches are reported here under this heading. The information
concerning the type-species of each genus has been extracted from Hessle
(1917), supplemented by additional information from Annenkova (1930), and
has been summarized in tables I and II. In several genera no information is
available concerning the internal anatomy.

The coelomic cavity of the first few segments is separated from the rest of
the thorax by a relatively stout septum or diaphragm. According to Hessle this

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) aio

always lies between segments IV and V, but according to the enumeration of
segments adopted here it lies between segments V and VI. Following Hessle’s
terminology the nephridium in segment V (Hessle’s segment IV) is termed the
anterior nephridium, and all later ones are posterior nephridia.

Never more than five pairs of nephridia have been reported; in some cases
there are only three and in exceptional cases only two. At first sight the number
of nephridia would appear to provide a reasonable basis for classification, but
as Hessle himself has pointed out, their distribution is not constant. In some cases
four pairs may be present in segments V to VIII, in others the nephridium in VI
is missing and the four pairs are in V, VII, VIII and [X; again V may be missing
and the four pairs are in segments VI-IX. The same variability occurs in those
species in which there are only three pairs of nephridia. They are located in
segments V, VI and VII in Sosanopsis wirent, Parhypania brevispina and Hypania
invalida, and in segments V, VII and VIII in Sabellides octocirrata. Even when the
number is reduced to two pairs there are differences in arrangement. In
Phyllocomus crocea and Schistocomus (=Phyllocomus) hiltont the nephridia are in
segments VII and VIII, and in Lysippides fragilis they are in segments V and
VII.

Hessle further reports that the number of pairs of nephridia is not constant
within the single genus Ampharete but may vary from two pairs situated in
segments V and VII for A. acutifrons and A. arctica to four pairs situated in
segments V, VII, VIII and IX for A. lindstromi and A. kerguelensis. The num-
bering of the segments given here is of course one higher than that given by
Hessle.

In view of this variability of the nephridia and the labour involved it would
seem undesirable to base generic divisions on these structures as has been done
by Hessle. None the less the nephridial papillae of the anterior nephridia in
segment V are occasionally visible. In Sabellides and Pterampharete the ducts are
elongated dorsally and two nephridial papillae may be found on the branchial
ridge between the two groups of gills. In Anobothrus, Sosane and Asabellides the
ducts open on a single nephridial papilla in the same position. These characters,
while not in themselves sufficient for generic distinction, provide useful
confirmatory evidence. ,

The alimentary canal is normally a fairly straight tube divided into
pharynx, oesophagus, stomach and intestine. At the anterior end of the stomach
where it joins the oesophagus a pair of lateral pouches project forward in
Ampharete, Microsamytha and Anobothrus, but not in Melinna nor fourteen other
genera which have been investigated by Wiren (1885), Hessle (1917) or
Annenkova (1930). There is also a curious invagination of the ventral wall of
the stomach of some genera, which is referred to by Hessle as an ‘innere Blind-
sack’ and is here termed an internal diverticulum. It has been found in Amage,
Amphicteis, Amphisamytha and Hypania, but not in fourteen other genera which
have been examined. Since such characters demand dissection they are of
doubtful value in practical classification.

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

The posterior thorax. As stated earlier, the first branchiferous (or paleal)
segment is reckoned here as segment III and the branchiferous region as
extending over segments III to VI. The beginning of the posterior thorax on
segment VII is marked by the first appearance of uncini in the neuropodia in

all genera. It is the most important and most easily recognizable landmark on ~
the body. Segments anterior to it may be distorted or fused and individual ©

segments may lack notosetae or nephridia, but segment VII and the posterior
thoracic segments are always well defined and fully developed. The maximum
number is 14 and the minimum number is 11 in all genera except Mugga,
where there are 9. It is suggested therefore that the numbers of such uncigerous
thoracic segments provide better generic characters than the total number of
thoracic setigers which have led to such confusion.

The notopodia of the posterior thorax are conical projections which in
Amphictets and several other genera bear a terminal papilla or ‘notopodial
cirrus’. ‘The neuropodia are short projecting pinnules and in a few genera such
as Phyllocomus they bear a superior papilla or cirrus above the row of uncini.

The notosetae are winged capillaries which are very uniform in shape
with few exceptions. In Anobothrus the 8th uncigerous segment has its notopo-
dium elevated and the tips of the notosetae are minutely spinulose. In Mugga
the same phenomenon occurs on uncigerous segment 9, and in Sosane and
Sosanopsis it occurs on uncigerous segment 12.

The uncini are flattened tooth plates of various shapes but are all roughly
triangular to quadrangular, with one or more series of teeth above the base.
In the Melinninae there is always a single series of teeth, but in the Ampharetinae
the teeth of the thoracic uncini may be arranged in 1, 2 or as many as 5 vertical
rows. Abdominal uncini often have more rows of teeth. The base of the uncinus
may be long and well separated from the tooth rows, or it may be short and
curve up towards the lowest tooth to form a bluntly rounded prow. Hessle and
others have quoted such differences as generic distinctions, but to me they
appear to be no more than specific characters, for the shapes vary very consider-
ably. The number of tooth rows seems to be more important when there are
only one or two vertical series, but when there are three or more they lose their
value.

The abdomen. There is seldom any abrupt narrowing between thorax and ~

abdomen, merely a change in the parapodia. This, however, is well marked.
The notosetae disappear and the notopodia are either reduced or absent. In
Melinna, Amphicteis, Amage, Phyllocomus and many other genera, rudimentary
notopodia persist on abdominal segments, and this is regarded as the more
primitive condition. In Ampharete, Sabellides and others, notopodia are entirely
lacking. Where notopodial cirri are present on the thorax they tend to persist on

the rudimentary notopodia of the abdomen, as may be seen in Amphicteis and ~

Amage. Similarly, where neuropodial cirri are present on the thorax these also
are continued on the abdomen. Actually the uncigerous pinnules very often
have an obscure superior papilla and this tends to become better marked towards

all. omar jet!

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 103

the end of the abdomen. This is particularly well shown by Sabellides octocirrata
where the superior papillae become long cirriform projections.

The whole abdomen consists of a variable number of segments. In the
Melinninae there is always a large number of segments, the extremes being a
minimum of 20 in Melinnexis antarctica and a maximum of go in Irana hetero-
branchia. In a few primitive genera of the Ampharetinae there are also large
numbers of abdominal segments. Thus Phyllocomus crocea has 45 but in the great
majority of genera the number is much less, 12-18 being the usual range. It
is doubtful whether the exact number is constant for a species, and certainly
it is not of generic importance.

The abdominal uncini are more or less similar to those of the thorax but
usually there are more teeth arranged in more vertical rows or transverse arcs.

The pygidium is terminal and may be encircled by a number of low
indistinct papillae or may bear a reduced number of longer anal cirri.

_DIVISION INTO SUB-FAMILIES AND GENERA

Hessle (1917) does not divide the Ampharetidae into sub-families, although
his remarks on p. 90 show that he is inclined to separate Melinna and Isolda
from other genera in this way. Chamberlin (1919) proposed three sub-families,
namely the Melinninae with dorsal hooks behind the gills but no paleae, the
Ampharetinae with paleae but no hooks, and the Samythinae with neither paleae
nor hooks.

Since the publication of Chamberlin’s monograph many new genera
have been described. The diagnostic characters of the various type-species are
set out in tables I and II, and it will be immediately obvious that there are
several genera related to Melinna. Not all of these possess dorsal hooks behind
the gills, but they all possess fine acicular neurosetae in segments III, IV, V
and often VI. I agree with Hessle in regarding this as an important and primi-
tive feature which links the Amphaetidae to Terebellides and other genera of the
Trichobranchidae and through them to the Terebellidae. I have therefore enlarged
Chamberlin’s diagnosis of the Melinninae to include all Ampharetidae with
acicular neurosetae in segments III-VI whether they possess dorsal hooks or
not. ‘he amended characters of the sub-family are given later.

Chamberlin’s sub-families Ampharetinae and Samythinae are distinguished
by the presence or absence of paleae. The variability of these setae has been
described earlier; as the accompanying tables show, the setation of the whole
branchial region is not sufficiently reliable for the distinction between genera,
let alone sub-families. Several other possible groupings of genera have been
tried on the basis of the number of branchiae, the number of uncigerous thoracic
segments and the number of tooth rows on the thoracic uncini. None of them
were satisfactory in that they distinguished groups of genera with many charac-
ters in common. It was concluded that all the Ampharetidae apart from the
Melinninae must be included within the sub-family Ampharetinae and that
Chamberlin’s sub-family Samythinae must be sunk.

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA)

104

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‘| Fav,

ANNALS OF THE SOUTH AFRICAN MUSEUM 105

The reliability of the various characters which may be used to distinguish
genera has been discussed earlier. To be of practical value in a taxonomic key
such characters must not only be constant within a group of species which are
obviously similar in other ways, but they must also be easily seen when sorting a
sample. It is suggested that among such characters are the number of gills, and
the number of uncigerous thoracic segments, the presence of glandular ridges
on the prostomium and the possession of notopodial cirri. These have therefore
been used as the main basis for grouping genera with the addition of several
other features in specialized cases. It is tempting to use the obvious ornamenta-
tion of the gills as well as their number, but after careful consideration this
was omitted for it would have increased the number of monotypic genera
without breaking up the three large genera Amphicteis, Amage and Ampharete. In
any case the gills are often missing so that their ornamentation cannot be
decided, and only the scars remain to determine their number.

As stated earlier, 49 genera are currently recognized as valid. In the
accompanying key these have been reduced to 33 of which 6 are assigned to the
Melinninae and 27 to the Ampharetinae. This means that 16 genera have been
sunk, and reference to the tables will show that 11 of these were monotypic and
the other 5 contained 2 species each. One new genus has been erected in the
sub-family Melinninae.

It is hoped that this revision will make the recognition of genera much
simpler, but inevitably it will be found that many species have been assigned
to the wrong genus. This is certainly true of two species assigned to Lysippe by
myself and one referred with hesitation to Neosabellides.

Key to sub-families
1. Segments III—V (or III—-VI) with fine acicular neurosetae. No paleae.
Post-branchial hooks sometimes present .. : “ins .. Melinninae

Segments III—VI without neurosetae. No post- ties hooks. Paleae
sometimes present .. ta ie is bs ie .. Ampharetinae

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

SuUB-FAMILY Melinninae Chamberlin 1919 (characters amended)

Buccal tentacles never pennate or papillose, usually smooth with a groove
along one side. Paleae absent. One or two pairs of stout notopodial hooks may
be present behind the gills. Small acicular neurosetae embedded in segments
III, [TV and often V and VI. Uncini are from segment VII and always have a
single series of teeth. Numerous (20-90) abdominal segments.

Key to genera of Melinninae

1. Stout notopodial hooks behind the gills .. me Ue ai am 2
No notopodial hooks ae , Bh ae 5
2. Notopodial hooks on two seopanee: Prosiouinin ‘Gloasarea Ab .. Moyanus
(doubtful)
Notopodial hooks on segment IV only. Prostomium short .. ae 3
3. 4 pairs of gills be ae in 4
: pairs of gills, including both conecih al papilese ae by .. Jrana
2 pairs of papillose gills .. aA ag 28 ae, aa .. Odvorpata
(doubtful)
4. Allgillssmooth .. or + Ag so .. Melinna
Some gills smooth, some aamhese te Me ae hie .. Isolda
5. 4 pairs of smooth gills a, ao as oi a a .. Melinnopsis
3 pairs of smooth gills My ite ey nF Ar oy .. Melinnopsides

MOYANUS Chamberlin 1919

Prostomium elongated and probosciform. Buccal tentacles mounted on
long tongue-shaped projection. Four pairs of smooth gills. A dorsal crest on
segment VI. Segments [II-VI with notopodial hooks on segments IV and V
and fine acicular neurosetae on segments III, IV and V. Twelve uncigerous
thoracic segments and about 65 abdominal ones. Thoracic uncini with a single
penies (et tert Type-species Moyanus explorans Chamberlin 1919.

MELINNA Malmeren 1866

Buccal tentacles smooth with a groove on one side. Four pairs of smooth
gills. A dorsal crest across segment VI. Segments III—-VI with notopodial hooks
on segment IV and notopodial capillaries on segments V and VI. Fine acicular
neurosetae on segments III—V and sometimes on VI as well. Fourteen uncigerous
thoracic segments and 30-50 abdominal ones. Thoracic uncini with a single
series feo Type-species Sabellides cristata Sars 1851.

IsOLDA Miiller 1858

Buccal tentacles smooth with a groove on one side. Four pairs of gills of
which 2 are smooth and 2 are papillose. A dorsal crest across segment VI.
Segments III-VI with notopodial hooks on segment IV and notopodial
capillaries on segments V and VI. Fine acicular neurosetae on segments III-V
and sometimes on VI as well. Twelve to 13 uncigerous thoracic segments and
25-36 abdominal ones. Thoracic uncini with a single series of teeth.

Type-species Isolda pulchella Miiller 1858.

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 107

IRANA Wesenberg-Lund 1949

Buccal tentacles smooth. Three pairs of gills of which 1 is smooth and 2 are
papillose. A dorsal crest across segment VI. Segments III-IV with notopodial
hooks on segment IV and fine acicular neurosetae on segments III-VI. Twelve
uncigerous thoracic segments and about 90 abdominal ones. Thoracic uncini
with a single series of teeth.

Type-species Lrana heterobranchia Wesenberg-Lund 1949.

OEORPATA Kinberg 1867

An incompletely described genus with two pairs of pennate gills; noto-
podial hooks on segment IV; many abdominal segments; uncini with a

single series of teeth. Type species Oeorpata armata Kinberg 1867.

MELINNOPSIS McIntosh 1885
Synonyms Melinnexis Annenkova 1930 and Melinnides Wesenberg-Lund 1950.

Buccal tentacles smooth with a groove along one side. Four pairs of smooth
gills. Dorsal crest on segment VI present or absent. Segments III-VI without
notopodial hooks on segment IV but with notopodial capillaries on segments V
and VI. Fine acicular neurosetae on segments III—-V and sometimes on VI as
well. Ten to 14 uncigerous thoracic segments and 25-50 abdominal ones.
Thoracic uncini with a single series of teeth.

Type-species Melinnopsis atlantica McIntosh 1885.

MELINNOPSIDES gen. Nov.

Buccal tentacles smooth with a groove along one side. Three pairs of
smooth gills. No dorsal crest on segment VI. Segments III—VI without noto-
podial hooks on segment IV but with notopodial capillaries on segments V and
VI. Fine acicular neurosetae on segments IIJ-V. Ten uncigerous thoracic
segments and about 30 abdominal ones. Thoracic uncini with a single series of
teeth.

Type-species Melinnopsis capensis Day 1955.

SUB-FAMILY Ampharetinae Chamberlin 1919 (characters amended)
(including Samythinae Chamberlin 1919)

Buccal tentacles either smooth with a groove along one side or papillose.
Paleae present or absent. No notopodial hooks behind the gills. Neurosetae
absent from segments III to VI. Neuropodial uncini start on segment VII
and may have one or more series of teeth. Few (8) to many (60) abdominal
segments. !

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109

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA)

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ANNALS OF THE SOUTH AFRICAN MUSEUM

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II!

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA)

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I12

ANNALS OF THE SOUTH AFRICAN MUSEUM

Key to genera of Ampharetinae

I.

10.

Il.

12.

13,

4 pairs of gills
3 pairs of gills
2 pairs of gills; first row fe uncini ite ae

Glandular ridges on prostomium. Buccal tentacles always smooth and
grooved along one side

No glandular ridges on prostomium. Buccal Buen citer sical or
papillose

Notopodial cirri present
No notopodial cirri

14 uncigerous thoracic segments ..
II uncigerous thoracic segments ..

14 uncigerous thoracic segments
13 uncigerous thoracic segments
II uncigerous thoracic segments

14 uncigerous thoracic segments
13 uncigerous thoracic segments
12 uncigerous thoracic segments
II uncigerous thoracic segments

Notopodial cirri present
Notopodial cirri absent

Tentacles papillose. Abdominal notopodia seldom present
Tentacles smooth and grooved. Abdominal notopodia always oe

No nephridial papilla on the branchial ridge
One nephridial papilla on the branchial ridge

No specialized posterior notosetae or elevated notopodia. Over 30
abdominal segments .. .

Specialized notosetae present in some Senos pocieniae ee
Less than 20 abdominal segments

One nephridial papilla on the branchial ridge ..
No nephridial papilla on the branchial ridge

Tentacles smooth with a longitudinal groove. No nephridial papillae on
the branchial ridge
Tentacles papillose. 2 nephridial elles on the prance ee

14, uncigerous thoracic segments ..

13 uncigerous thoracic segments ..

12 uncigerous thoracic segments ..

II uncigerous thoracic segments .. ..

g uncigerous thoracic segments. Last avin ado plewied aad tee
modified notosetae

Glandular ridges on prostomium
No glandular ridges on prostomium

Abdominal notopodia present. Tentacles numerous and smooth. Paleae
usually absent : Bt ni a ae fa Exe
Abdominal notopodia absent. Tentacles represented by a folded
membrane. Paleae usually absent : oe
(With paleae, other characters unknown)

Tentacles papillose
Tentacles smooth

. Adorsal ridge between ae Vi and VII. Paleae present on segment

Ill a,
No dorsal ridge. No setae on Seernene IIT

. Tentacular lobe prolonged

No elongate tentacular lobe

Amphicteis
Amage
Parphypania
Hypania
Grubianella

7
Lysippe
8
12
Phyllamphicteis
Lysippides
9
10

Ampharete
Asabellides

Phyllocomus

II

Sosane
Sosanopsis

Paramage
Sabellides

14
Microsamytha

16
Glyphanostomum

Mugga
Samythopsis
15

Samytha

Ampythas
Aryandes

17
18

Melinnampharete
Neosabellides

Pabits
Samythella

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 113

AMPHICTEIS Grube 1851

Synonyms Crossostoma Gosse 1855, ?Rytocephalus Quatrefages 1865, and Paramphicteis Caullery

1944.

Prostomium with a pair of glandular ridges. Buccal tentacles smooth with
a groove along one side. Four pairs of gilis. Segments ITI-VI without neuro-
setae. Segment III often with paleae; segments IV—VI with notopodial capil-
laries. Fourteen uncigerous thoracic segments. Notopodial cirri present.
Thoracic uncini with a single vertical series of teeth. Thirteen to 19 abdominal
segments sometimes with rudimentary notopodia as well as uncigerous pinnules.

Type-species Amphitrite gunneri Sars 1835.

AMAGE Malmeren 1866

Prostomium with glandular ridges. Buccal tentacles smooth with a groove
along one side. Four pairs of gills. Segments III-VI without neurosetae.
Segments IV—VI usually with notopodial capillaries. Eleven uncigerous
thoracic segments. Notopodial cirri present. Thoracic uncini with 1 or 2 series
of teeth. Eight to 15 abdominal segments usually with rudimentary notopodia
as well as uncigerous pinnules.

Type-species Amage auriculata Malmgren 1866.

PARHYPANIA Annenkova 1928

Prostomium with glandular ridges. Buccal tentacles smooth. Four pairs
of gills. Segments III-VI without neurosetae. Segment III often with paleae,
segments IV—VI usually with notopodial capillaries. Fourteen uncigerous
thoracic segments. Notopodial cirri absent. About 16-20 abdominal segments
sometimes with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Amphicteis brevispinus Grube 1860.

HYPANIA Ostrooumov 1897

Prostomium with glandular ridges. Buccal tentacles smooth. Four pairs
of gills. Segments III-VI without neurosetae. Segment III often with paleae,
segment IV may have notosetae, segments V and VI always with notopodial
capillaries. Thirteen uncigerous thoracic segments. Notopodial cirri absent.
About 15-30 abdominal segments sometimes with rudimentary notopodia as
well as uncigerous pinnules.

Type-species: Amphicteis invalida Grube 1860.

GRUBIANELLA McIntosh 1885

Prostomium with glandular ridges. Buccal tentacles smooth. Four pairs
of gills. Segments III-VI without neurosetae. Notopodial capillaries present
on segments V and VI and sometimes on segment IV. Eleven uncigerous
thoracic segments. Thoracic uncini usually with 2 series of teeth. Notopodial
cirri doubtful. About 25 abdominal segments; the last few are reported to be
swollen in the type but this is probably an abnormality.

Type-species: Grubianella antarctica McIntosh 1885.

II4 | ANNALS OF THE SOUTH AFRICAN MUSEUM

LYSIPPIDES Hessle 1917
Synonyms Amphisamytha Hessle 1917 and Hypaniola Annenkova 1928.

Prostomium without glandular ridges. Buccal tentacles smooth with a
groove along one side. Four pairs of gills. Segments III-VI without neurosetae.
Notosetae present on segments V and VI and usually III and IV as well.
Fourteen uncigerous thoracic segments. Notopodial cirri absent. Thoracic
uncini with 1-3 series of teeth. Between 8 and 23 abdominal segments usually
with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Amphicters fragilis Wollebaek 1912.

PHYLLAMPHICTEIS Augener 1918

Synonym Paiwa Chamberlin 1919.
Prostomium without glandular ridges. Buccal tentacles smooth. Four pairs

of gills. Segments ITI-VI without neurosetae. Notosetae present on segments V
and VI and usually on III and IV as well. Fourteen uncigerous thoracic
segments. Notopodial cirri present. Thoracic uncini with 1-3 series of teeth.
_ Between 15 and 20 abdominal segments sometimes with rudimentary notopodia ~
as well as uncigerous pinnules.

Type-species: Phyllamphicteis collaribranchis Augener 1918.

LysIpPE Malmgren 1866
Synonym Pierolysippe Augener 1918.

Prostomium without glandular ridges. Buccal tentacles smooth. Four —
pairs of gills. Segments IITI-VI without neurosetae. Small notosetae often
present on segments III and IV and notosetae always present on V and VI.
Thirteen uncigerous thoracic segments. Thoracic uncini with 2-3 series of
teeth. Notopodial cirri absent. About 12-15 abdominal segments sometimes
with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Lysippe labiata Malmgren 1866.

AMPHARETE Malmgren 1866
Synonym Branchiosabella Claparéde 1863.

Prostomium without glandular ridges. Buccal tentacles papillose. Four —
pairs of gills. No nephridial papilla on the branchial ridge. Segments II-VI
without neurosetae. Notosetae of segment III usually enlarged to form paleae;
notosetae of segment IV usually absent; notosetae of segments V and VI
always present. Twelve uncigerous thoracic segments. Notopodial cirri absent.
Thoracic uncini usually with 2 series of teeth. About 12-15 abdominal segments
which have uncigerous pinnules but usually lack rudimentary notopodia.

Type-species: Amphicteis acutifrons Grube 1860.

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 115

ASABELLIDES Annenkova 1929
Synonym Pseudosabellides Berkely and Berkely 1943.

Prostomium without glandular ridges. Buccal tentacles papillose. Four
pairs of gills. One nephridial papilla on the branchial ridge between the two
groups of gills. Segments III—VI without neurosetae. Notopodial capillaries
present on segments V and VI but usually absent from III and IV. Twelve
uncigerous thoracic segments. Notopodial cirri absent. Thoracic uncini usually
with 2 series of teeth. About 21 abdominal segments which have unererreus
pinnules but lack rudimentary notopodia.

Type-species: Sabellides sibirica Wiren 1883.

PHYLLOCOMuUS Grube 1878
Synonym Schistocomus Chamberlin 1919.

Prostomium without glandular ridges. Buccal tentacles smooth with a
groove along one side. Four pairs of gills. Segments III-VI without neurosetae.
Notopodial capillaries present on segments IV, V and VI. Twelve uncigerous
thoracic segments. No specialized notosetae. Notopodial cirri present or absent.
Thoracic uncini with a single series of teeth. Numerous (over 30) abdominal

segments. Type-species: Phyllocomus crocea Grube 1878.

SOSANE Malmeren 1866
Synonym Anobothrus Levinsen 1883.

Prostomium without glandular ridges. Buccal tentacles smooth. Four
pairs of gills. One nephridial papilla on the branchial ridge between the two
groups of gills. Segments III-VI without neurosetae. Notosetae usually present
on all four segments ITI to VI and often enlarged to form paleae on segment III.
Twelve uncigerous thoracic segments with one or more of the posterior noto-
podia elevated and bearing specialized notosetae. Notopodial cirri absent.
Thoracic uncini with 1-2 series of teeth. About 13 abdominal segments usually
with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Sosane sulcata Malmgren 1866.

SOSANOPSIS Hessle 1917

Prostomium without glandular ridges. Buccal tentacles smooth. Four
pairs of gills. No nephridial papillae on the branchial ridge. Segments ITI—-VI
without neurosetae. Segments V and VI with notopodial capillaries and usually
segment IV as well. Twelve uncigerous thoracic segments with the last noto-
podium elevated and bearing specialized notosetae. Notopodial cirri absent.
Thoracic uncini with 2-3 series of teeth. About 11 abdominal segments usually
with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Sosanopsis wirent Hessle 1917.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

PARAMAGE Caullery 1944

Prostomium without glandular ridges. Buccal tentacles smooth. Four pairs
of gills. No nephridial papillae on the branchial ridge. Segments III-VI without
neurosetae. Segment VI with notopodial capillaries but notosetae often absent
from more anterior segments. Eleven uncigerous thoracic segments. No
specialized posterior notosetae. Notopodial cirri absent. Thoracic uncini usually
with a single series of teeth. About 11 abdominal segments with rudimentary
notopodia as well as uncigerous pinnules.

Type-species: Paramage madurensis Caullery 1944.

SABELLIDES Milne-Edwards 1838

Synonyms Heterobranchus Wagner 1885 and Pterampharete Augener 1918.

Prostomium without glandular ridges. Buccal tentacles papillose. Four
pairs of gills. A pair of nephridial papillae on the branchial ridge between the
two groups of gills. Segments III-VI without neurosetae. Segments V and VI
with notopodial capillaries, segment IV usually fused to segment III and
without notosetae, but notosetae often present on segment III. Eleven uncig-
erous thoracic segments. No specialized posterior notosetae. Notopodial cirri
absent. Thoracic uncini with 1 or 2 series of teeth. Between 11 and 18 abdominal
segments with uncigerous pinnules but no rudimentary notopodia.

Type-species: ? Sabella octocirrata Sars 1835. —

sAMYTHOPSIS McIntosh 1885

Prostomium with glandular ridges. Buccal tentacles smooth. Three pairs —
of gills. Segments III-VI without neurosetae. Segments IV—VI with notopodial ~
capillaries. Fourteen uncigerous thoracic segments. Notopodial cirri doubcful. —
About 22 abdominal segments which may have rudimentary notopodia as well —
as uncigerous pinnules.

Type-species: Samythopsis gruber McIntosh 1885.

SAMYTHA Malmgren 1866

Prostomium without glandular ridges. Buccal tentacles numerous and
smooth. Three pairs of gills. Segments III-VI without neurosetae. Segments
IV-VI with notopodial capillaries. Fourteen uncigerous thoracic segments. No
notopodial cirri. Thoracic uncini with 2-3 series of teeth. About 13 abdominal
segments with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Sabellides sexcirrata Sars 1856.

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 117

AMYTHAS Benham 1921

Prostomium without glandular ridges. Buccal tentacles replaced by a
frilly membrane. Three pairs of gills. Segments III-VI without neurosetae.
Segments IV—VI with notopodial capillaries. Fourteen uncigerous thoracic
segments. No notopodial cirri. Thoracic uncini with 2 series of teeth. Fourteen
or more abdominal segments without rudimentary notopodia above the
uncigerous pinnules. !

Type-species: Amythas membranifera Benham 1921.

ARYANDES Kinberg 1867

A questionable and incompletely described genus generally similar to
Samytha but with paleae.
Type-species: Aryandes gracilis Kinberg 1867.

MICROSAMYTHA Augener 1928
Synonym ? Alkmaria Horst 1920.

Note: Both genera are incompletely described and may belong to the family
Terebellidae. More is known of Microsamytha and this name is retained pro tem.

Prostomium without glandular ridges and ‘of the Ampharete type’. Buccal
tentacles smooth. Three pairs of gills. Segments III-VI without neurosetae.
Segments IV, V and VI with notopodial capillaries. Thirteen uncigerous
thoracic segments. Between 13 and 19 abdominal segments.

Type-species Microsamytha rychiana Augener 1928.

MELINNAMPHARETE Annenkova 1937

Prostomium without glandular ridges. Buccal tentacles papillose. Three
pairs of gills. Segments III-VI without neurosetae. Notosetae usually present
on all four segments (III-VI) and often enlarged to form paleae on segment ITI.
A dorsal ridge between segments VI and VII. Twelve uncigerous thoracic
segments. No notopodial cirri. ? number of abdominal segments.

Type-species: Melinnampharete eoa Annenkova 1937.

Neosabellides Hessle 1917

Prostomium without glandular ridges. Buccal tentacles papillose. Three
pairs of gills. Segments III-VI without neurosetae. Notosetae present on
segments V and VI but usually absent from III and IV. No dorsal ridge
between segments VI and VII. Twelve uncigerous thoracic segments. No
notopodial cirri. Thoracic uncini usually with 2 series of teeth. About 19
abdominal segments.

Type-species: Sabellides elongatus Ehlers 1913.

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

PABITS Chamberlin 1919

Prostomium without glandular ridges. Buccal tentacles smooth and borne
on a long probosciform organ. Three pairs of gills. Segments III-VI without
neurosetae. Notosetae present on segments V and VI and usually on IV as well.
Twelve uncigerous thoracic segments. No notopodial cirri. Thoracic uncini
with 2 series of teeth. Number of abdominal segments unknown.

Type-species: Pabits deroderus Chamberlin 1919.

SAMYTHELLA Verrill 1873
Synonyms Eusamytha McIntosh 1685 and ? Eclysippe Eliason 1955.

Prostomium without glandular ridges. Buccal tentacles smooth and not
borne on an elongate tentacular lobe. Three pairs of gills. Segments III-VI
without neurosetae. Notopodial capillaries present on segments IV-VI and
sometimes on III as well. Twelve uncigerous thoracic segments. Notopodial
cirri may be present on the last few thoracic setigers. Thoracic uncini with 1 or
2 series of teeth. Up to 36 abdominal segments.

Type-species: Sampthella elongata Verrill 1873.

GLYPHANOSTOMUM Levinsen 1883
Synonym Amythasides Eliason 1955.

Prostomium without glandular ridges. Buccal tentacles smooth (? always).
Three pairs of gills. Segments III-VI without neurosetae. Segments IV—VI
with notopodial capillaries and segment III sometimes with paleae. Eleven
uncigerous thoracic segments. Notopodial cirri absent. Thoracic uncini with
2 or more series of teeth. Twelve to 25 abdominal segments without rudimentary
notopodia but with uncigerous pinnules.

Type-species Samytha pallescens Theel 1878.

MUGGA Eliason 1955

Prostomium without glandular ridges. Buccal tentacles smooth. Three pairs
of gills. Segments III-VI without neurosetae but all of them may have notosetae.
Nine uncigerous thoracic segments. Notopodium of the last thoracic segment
dorsally situated and bearing modified notosetae. Thoracic uncini with more
than 2 series of teeth. Number of abdominal segments unknown.

Type-species: Mugga wahrbergi Eliason 1955.

AUCHENOPLAX Ehlers 1887

Synonym Melinnoides Benham 1921.

Prostomium without glandular ridges. Buccal tentacles smooth with a
groove along one side. Two pairs of gills. Segments III-VI without neurosetae.
Segments V and VI with notopodial capillaries. Twelve uncigerous thoracic
segments and about 12-15 abdominal ones. First row of thoracic uncini on
segment VII very long. Thoracic uncini with 1-2 series of teeth. Notopodial

cirri absent. Type-species: Auchenoplax crinita Ehlers 1887.

~

4
}
é
t

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 119

SUMMARY

The characters of existing genera are discussed and it is shown that
confusion has arisen because the anterior segments have been telescoped and
there are two different systems of numbering them. It is proposed that the
separation of genera be based primarily on the number of gills and the number

of uncigerous thoracic segments. The main diagnostic characters of 49 type-
| species have been tabulated and it is suggested that 16 genera which are mostly
monotypic be sunk. Keys and definitions of the remaining 33 genera are

provided.

REFERENCES

ANNENKOVA, N. 1928. Ueber die pontokaspichen Polychaeten. 2. Die Gattungen Hypaniola,
Parhypania, Farbicia und Manayunkia. Annu. Mus. zool. Akad. Leningrad 30: 13-20.

ANNENKOVA, N. 1930. Zur Polychaetenfauna von Franz-Joseph-Land. (Melinnexis gen. nov.
arctica sp. n.) Zool. Anz. 95: 269-272.

AnNENKOVA, N. 1937. [The polychaete fauna of the northern part of the Japan Sea.] Jssled.
Mor. USSR 23: 139-216. [In Russian. ]

AuGENER, H. 1918. Polychaeta. In Michaelsen, W., ed. Beitrige zur Kenntnis der Meeresfauna
Westafrikas. 2: 67-625. Hamburg: Friederichsen.

AuGENER, H. 1928. Beitrag zur Polychaetenfauna der Ostsee. @. Morph. Oekol. Tiere 14: 102-104.

BENHAM, W. B. 1921. Polychaeta. Sci. Rep. Aust. antarct. Exped. (c), 6, 3: 1-128.

BERKELEY, E. and C. 1943. Biological and oceanographical conditions in Hudson Bay. 11.
Polychaeta from Hudson Bay. 7. Fish. Res. Bd. Can. 6: 129-132.

Cau.tery, M. 1944. Polychétes sédentaires de l’expédition du Siboga. [I1.] Siboga Exped. mon.
24 2bis: 1-204.

CHAMBERLIN, R. V. 1919. The Annelida Polychaeta. Mem. Mus. comp. Zool. Harv. 48: 1-514.

Day, J. H. 1955. The Polychaeta of South Africa. Part 3: Sedentary species from Cape shores
and estuaries. 7. Linn. Soc. (Zool.) 42: 407-452.

Day, J. H. 1961. The polychaete fauna of South Africa. Part 6: Sedentary species dredged off
Cape coasts with a few new records from the shore. 7. Linn Soc. (Zool.) 44: 463-560.

En ers, E. 1887. Report on the annelids of the dredging expedition of the U.S. coast survey
steamer Blake. Mem. Mus. comp. Zool. Harv. 15: 1-335.

EHLERS, E. 1913. Die Polychaeten-Sammlungen der deutschen Siidpolar-Expedition 1901-1903.
Dtsch. Siidpol.-Exped. 13: 397-598.

Exiason, A. 1955. Neue oder wenig bekannte swedische Ampharetiden (Polychaeta). Goteborg,
VetenskSamh. Handl. (7) 6B (16): 1-17.

FAuvEL, P. 1897. Recherches sur les ampharétiens, annélides polychétes sédentaires. Morpholo-
gie, anatomie, histologie, physiologie. Bull. sci. Fr. Belg. 30: 277-488.

FAUVEL, P. 1927. Polychétes sédentaires. Faune Fr. 16: 1-494.

FAuvEL, P. 1936. Polychétes. Result. Voy. Belgica Zool.: 1-44.

Gosse, P. H. 1855. On new and little known marine animals. Ann. Mag. nat. Hist. (2) 163 31-35.

Grim, O. A. 1877. [The Caspian Sea and its fauna.] Jn Grimm, O. A., ed. [Transactions of the
Aralo-Caspian expedition.| 2: 1-168. S.-Petersburgh. [In Russian. ]

Gruse, E. 1851. Die Familien der Anneliden. Arch. Naturgesch. 16: 249-364.

Gruse, E. 1860. Beschreibung neuer oder wenig bekannter Anneliden. Arch. Naturgesch. 26:
71-118.

Grusz, E. 1878. Einige neue Anneliden aus Japan. Jber. schles-Ges. vaterl. Kult. 55: 104-106.

HeEsste, C. 1917. Zur Kenntnis der terebellomorphen Polychaeten. Zool. Bidr. Uppsala 5: 39-258.

Horst, R. 1920. Polychaete anneliden uit het Alkmaarder Meer. Zool. Meded. 5: 110-111.

Kinser, J. G. H. 1867. Annulata nova. Ofvers. Vetensk Akad. Férh., Stockh. 23: 337-357-

Levinsen, G. M. 1883. Systematik-geographisk oversight over de nordiske Annulata, Gephyrea,
Chaetognathi og Balanoglossi. Vidensk. Medd. dansk naturh. Foren Kbh. 2: 92-348.

McIntosu, W. C. 1885. Report on the Annelida Polychaeta collected by H.M.S. Challenger
during the years 1873-76. Rep. Voy. Challenger 1873-76 (Zool.) 12: 1-554.

I20 ANNALS OF THE SOUTH AFRICAN MUSEUM

McIntTosu, W. C. 1914. Notes from the Gatty Marine Laboratory, St. Andrews No. 36, 4. On
the British Ampharetidae. Ann. Mag. nat. Hist. (8) 13: 77-110.

Ma.wmcren, A. J. 1865. Nordiska Hafs-Annulata. Ofvers. VetenskAkad. Forh. Stockh. 22: 355-410.

Matmcren, A. J. 1866. Annulata Polychaeta Spetsbergiae, Groenlandiae, Islandiae et Scandinaviae
hactenus cognita. Helsingforsiae.

Meyer, E. 1887. Studien tiber den Korperbau der Anneliden. Mitt. zool. Sta. Neapel, 7: 592-741.

MiLne-Epwarps, M. H. 1836-41. Les annélides. In Cuvier, G. Le regne animal distribué d’ aprés son
organisation. [3rd ed.] 9. Paris: Masson.

MULLER, Fr. 1858. Einiges tiber die Anneliden fauna der Insel Santa Catharina an der brasilia-
nischen Kiiste. Arch. 1 Naturgesch. 24: 211-220.

Nixsson, D. 1912. Beitrage zur Kenntnis der Nervensystems der Polychaeten. Zool. Bidr. Uppsala
1: 85-161.

Ostromooumov, A. 1897. [Hydrobiological researches at the mouths of rivers in southern
Russia. Preliminary communication.] Bull. Acad. Sci. St.-Pétersb. 6: 343-362. [In Russian. ]

QuaTREFAGES, A. de. 1865. Histoire naturelle des anneles marins et d’eau douce. 1-2. Annélides et
géphyriens. Paris: Roret.

Sars, M. 1835. Beskrivelser og iagttagelser over nogle maerkelige eller nye i havet ved den Bergenske kyst
levende dyr af Molypernes, Acalephernes, Radiaternes, Annelidernes og Molluskernes classer. Bergen.

Sars, M. 1851. Beretning om en i sommeren 1849 foretagen zoologist reise i Lofoten og Fin-
marken. Nyt Mag. Naturv. 6: 121-211.

Sars, M. 1856. Fauna littoralis Norvegiae . . . 2: 1-24. Nye annelider. Bergen.

THEEL, H. J. 1878. Les annélides polychétes des mers de Nouvell-Zemble. K. svenska Vetensk-
Akad. Handl. 16 (3): 3-75.

Uscuakov, P. V. 1955. [The polychaete worms of the far eastern seas of the U.S.S.R.] Tadl.
anal. Faune U.R.S.S. 56: 1-445. [In Russian. ]

VERRILL, A. E. 1873. Report upon the invertebrate animals of Vineyard Sound and the adjacent
waters, with an account of the physical characters of the region. Rep. U.S. Comm. Fish. 1:
295-778.

Wacner, N. 1885. Die Wirbellosen des Weissen Meeres. Zoologisch Forschungen an der Kiiste —
des Solowetzkischen Meerbusens in den Sommermonaten der Jahre 1877, 1879 und 1882.
1: 1-171. Leipzig.

WESENBERG-LUND, E. 1949. Polychaetes of the Iranian Gulf. Dan. sci. Invest. Iran 4: 247-400.

WESENBERG-LunND, E. 1950. The Polychaeta of West Greenland, with special reference to the
fauna of Nordre Strédmfjord, Kavne- and Bredefjord. Medd. Grd¢nland 151 (2): 1-171.

Wiren, A. 1883. Chaetopdder fran Sibiriska Ishafvet och Berings Haf insamlade under Vega- —
expeditionen 1878-79. Jn Nordenskidld, N. A. E. von. Vegaexpeditionens vetenskapliga
Takttagelser . . . 2: 383-428. Stockholm.

Wiren, A. 1885. Om zirkulations och digestions-organen hos Annelider of familjerna Amphareti-
dae, Terebellidae och Amphictenidae. K. svenska VetenskAkad. Handl. 21: 1-58.

WoLLEBAEK, A. 1912, Nordeuropaeiske Annulata Polychaeta. I. Ammocharidae, Amphicteni-
dae, Ampharetidae, Terebellidae og Serpulidae. Skr. VidenskSelsk., Christ. 1911 (2): 1-144.

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TABLE OF CoNTENTs and SummARy. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 42 in. x 7 in. (7} in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (black type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

Smitu, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmitH, C. D. 1954. South African plonias. Jn Brown, X. Y. Marine faunas. and ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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MUS. COMP, ZOOL
LIBRARY,

JUN 2 4 1964

LIEUWE DIRK BOONSTRA

HARVARD
UNIVERSITY

THE GIRDLES AND LIMBS OF THE
PRISTEROGNATHID THEROCEPHALIA

April 1964 April
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MUS. COMP. ZOOL
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JUN 2 4 1964

THE GIRDLES AND LIMBS OF THE PRISTEROGNAGHIDRD
THEROCEPHALIA UNIVERSITY.

By
LIiEUWE Dirk BOONSTRA
South African Museum, Cape Town

(With 50 text-figures and one Plate)

CONTENTS

PAGE

Introduction : vette RQ
Ainnosaurus paucidens oan. ao sp. nov. ep, Lee
Material .. Py 2 - fe eZ
Historical .. i sid ig Y126
General Morphology 3 MP ae. t26
Pectoral Girdle vd a Sai OG
Humerus ty Ai at) 928
Ulna and Boda a Be 6: HaRZO
Carpus and Manus .. af SRG 0)
Pelvic Girdle .. si be nt i at
Femur ; ve Sion ek
Tibia, Fibula med Fes my, Eanes 627!
Systematic Descriptions of Specimens .. 136
A. Pectoral Girdle .. y eee SO

B. Humerus .. = Bae 7: Be

C. Lower Forelimb ed Pecciaut ae 150

D. Pelvic Girdle me ed ea 515)

E. Femur ; Ser THO

F. Lower Hind Limb rac Bes ee Oo
Discussion .. oy if Ns my) OA
Summary .. : ee is shee’
Acimowledeements he ay sath OS
Keferences .. ai sus Pe cia DOS

INTRODUCTION

In the Tapinocephalus zone of the Karoo the oldest Therocephalia known
are well represented. The Therocephalia form the main component of the
carnivores, the only other carnivores being the very large anteosaurs represented
by five genera and a few small to medium-sized Gorgonopsia represented by
10 genera, of which 30 specimens are known in all.

The Therocephalia are represented in the Tapinocephalus zone by the
families Pristerognathidae, Lycosuchidae, Scylacosauridae and Scaloposauridae.
Of these the pristerognathids are the dominant family and are well represented
in numbers of specimens as well as in the number of distinct species. Hitherto
16 genera with 28 species have been described. Most of the specimens consist of
partly preserved snouts, with good complete skulls few in number. Of the
postcranial skeleton little is known, for only rarely are some of the postcranial

121

Ann. S. Afr. Mus. 48 (5), 1964: 121-165, 50 figs. and 1 Plate.

r22 ANNALS OF THE SOUTH AFRICAN MUSEUM

bones preserved and then mostly associated with cranial parts difficult to
classify.

In this paper I present an account of the girdle and limb material in the
South African Museum mostly collected by myself since 1928.

This collection includes material which necessitates the establishment of
the following new genus and species.

ZINNOSAURUS new genus

Type-species: Kinnosaurus paucidens.

Diagnosis: A medium-sized pristerognathid with the dental formula i.5, c.1,
pce.2; the incisors are fairly weak and well spaced and the postcanines are weak
and lie far apart; the postcanines situated on a thin lappet of the maxilla
clearly demarcated from the general maxillary surface. Scapula with a well-
developed flange-like process on its posterior border immediately above the
glenoid for the scapular head of the triceps. The glenoid widely open and facing
much laterally. Humerus with greatly expanded epicondyles, thin and sheet-
like. Femur with broadly oval caput and strong thickened internal trochanter.

Affinities: Nearest relative Glanosuchus.

Kimnosaurus paucidens new species
(Figs. 1, 19, 28, 46d and 46e)

Holotype. S.A.M. 12185, skull with lower jaw, scapulo-coracoid, humeri, ends of
femora. |
Locality: Meyerspoort, Beaufort West, South Africa.

Horizon: High Tapinocephalus zone, Lower Beaufort Beds, Karoo.

I have great pleasure in naming this new genus after H. Zinn, who has, as
techinical assistant, accompanied me on numerous collecting trips in the
T apinocephalus zone.

The scapulo-coracoid, humeri and ends of femora of the holotype are —
described in the appropriate sections of this paper.

MATERIAL

In the South African Museum catalogue 219 specimens of pristerognathids
from the Tapinocephalus zone are listed. Of these only the following 37 specimens ~
have parts of the girdles and limbs preserved. Taken together one gets a good ©
idea of the family character of these structures. :

The material studied consists of the following specimens listed in the order
of the date of acquisition by the South African Museum collection.

S.A.M. 4335. Pristerognathus sp. Beaufort West. High? Tapinocephalus zone. Coll. —

Whaits 1916? Incomplete coraco-scapulae, distal and proximal ends of a —

humerus and the proximal halves of an ulna and radius. No skull.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 123

S.A.M. 5018. Pristerognathus sp. Abrahamskraal, Prince Albert. Low Tapino-
cephalus zone. Coll. Haughton 1916. Two incomplete ilia, proximal and
distal halves of two femora and of two humeri, proximal ends of tibiae,
radii and ulnae. No skull.

Fic. 1. <innosaurus paucidens gen. et spec. nov. S.A.M. 12185.
Type x4. Skull. a, lateral. 5, dorsal.

S.A.M. 9005. Scymnosaurus major. Holotype. Klein-Koedoeskop, Beaufort West.
Low Tapinocephalus zone. Coll. Boonstra 1929. Interclavicle with proximal
parts of both clavicles, distal ends of both humeri, complete radius and
ulna, associated with a good anterior half of the skull.

S.A.M. 9084. Scymnosaurus ferox. Rietkuil, Beaufort West. Low Tapinocephalus
zone. Coll. Boonstra 1929. A coracoid, proximal and distal ends of a
humerus and proximal ends of an ulna and radius associated with a good

skull.

124. ANNALS OF THE SOUTH AFRICAN MUSEUM

S.A.M. 11458. Pristerognathus sp. Mynhardtskraal, Beaufort West. Low Tapino-
cephalus zone. Coll. Boonstra and Avenant 1939. An interclavicle, proximal
ends of clavicles, parts of coracoids, Pee ends of femur, humerus, radius,
ulna. No cranial material.

S.A.M. 11459. Scymnosaurus sp. Buffelsvlei, Beaufort West. Low Tapinocephalus
zone. Coll. Boonstra and Marais 1939. Part of carpus and tarsus associated
with a poor snout. ,

S.A.M. 11557. Scymnosaurus sp. Die Cypher, Beaufort West. Low Tapinocephalus
zone. Coll. Boonstra 1940. A scapula, humerus, femur, interclavicle,
clavicle, radius, and partial manus. Without skull.

S.A.M. 11558A. Scymnosaurus? Die Cypher, Beaufort West. Low Tapinocephalus
zone. Coll. Boonstra 1940. A pair of well-preserved ischia.

S.A.M. 11695. Scymnosaurus sp. Seleryfontein, Fraserburg. Low? Tapinocephalus
zone, Coll. Boonstra and Jooste 1946. A coraco-scapula, interclavicle,
clavicles and humerus. With good anterior half of skull.

S.A.M. 11794 Cynariognathus sp. Seleryfontein, Fraserburg. Low? Tapinocephalus
zone. Coll. Jooste 1947. Imperfect pelvis, femora, humeri, epipodials and
manus. With a poor skull.

S.A.M. 11888. Thertoides cyniscus. Holotype. Vindragersfontein. Beaufort West.
Low Tapinocephalus zone. Coll. Boonstra and Zinn 1948. Parts of pectoral
girdle, humerus, radius, ulna and part of manus associated with a nearly
complete skull.

S.A.M. 11934. Pristerognathus? Steenboksfontein, Laingsburg. Low? Tapino-
cephalus zone. Coll. Boonstra 1951. Distal end of femur and proximal ends
of tibia and fibula. Without cranial association.

S.A.M. 11936. Pristerognathoides sp. Bosluiskraal, Laingsburg. Low? Tapino-
cephalus zone. Coll. Boonstra 1951. Proximal and distal end of a femur
associated with a snout.

S.A.M. 11942. Ptomalestes avidus. Holotype. Steenboksfontein, Laingsburg. Low ?
Tapinocephalus zone. Coll. Boonstra 1951. Part of pectoral girdle, humeri,
ulnae and radii associated with a good skull.

S.A.M. 11957. Scymnosaurus sp. Abrahamskraal, Prince Albert. Low Tapino-
cephalus zone. Coll. Le Roux 1923. Acetabular parts of ischium and pubis
and proximal end of femur. Without skull.

S.A.M. 12051. Alopecognathus sp. Rietfontein, Laingsburg. Low? Tapinocephalus
zone. Coll. Boonstra and Fourie 1957. Part of scapula, proximal end of
humerus, proximal and distal ends of femur, ends of radii and ulna and
part carpus, tibia and fibula and a complete pes associated with a snout.

S.A.M. 12102. Pristerognathoides sp. Kalkkraal. Prince Albert. Low Tapino-
cephalus zone. Coll. Boonstra and Zinn 1957. Most of the pectoral girdle,
ends of humerus, ulna and radius associated with a complete skull.

S.A.M. 12112. Pristerognathus? Skoppelmaaikraal, Laingsburg. Low Tapino-
cephalus zone. Coll. Botes 1957. Part of ilium, proximal end of humerus and
other fragments without any skull parts.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 125

S.A.M. 12118. Scymnosaurus sp. Palmietfontein of Kruidfontein, Prince Albert.
Low Tapinocephalus zone. Coll. Boonstra and Zinn 1957. Proximal end of
femur, part scapula.

S.A.M. 12185. <innosaurus paucidens gen. et spec. nov. Meyerspoort, Beaufort
West. High Tapinocephalus zone. Coll. Boonstra and Zinn 1959. Part
coraco-scapula, one complete humerus and one proximal half, proximal
and distal end of femur and proximal end of ulna and radius and distal
end of radius associated with a good skull.

S.A.M. 12193. Scymnosaurus sp. Fortuin of Dalajalon, Beaufort West. High
Tapinocephalus zone. Coll. Boonstra and Zinn 1959. Fairly complete
pectoral girdle, humerus, ulna and radius associated with piece of jaw
containing teeth. |

S.A.M. 12204. Pristerognathoides sp. Plaatjiesrivier, Beaufort West. High
Tapinocephalus zone. Coll. Boonstra and Zinn 1959. Fairly complete
pectoral girdle, humerus, ulna and radius associated with the posterior
third of a skull.

S.A.M. 12262. Scymnosaurus sp. Seleryfontein, Fraserburg. Low? Tapinocephalus
zone. Coll. Boonstra and Jooste 1959. Femur, radius, ulna and manus.
S.A.M. Ke218. Alopecognathus sp. Lammerkraal, Prince Albert. High
Tapinocephalus zone. Coll. Boonstra, Zinn and Gow 1959. Weathered

pelvis. :

_ §.A.M. K233A and B. Alopecognathus sp. Palmietfontein, Beaufort West. High
Tapinocephalus zone. Coll. Boonstra, Zinn and Gow 1959. Two incomplete
pectoral girdles.

S.A.M. K223C. Pristerognathoides sp. Palmietfontein, Beaufort West. High
T apinocephalus zone. Coll. Boonstra, Zinn and Gow 1959. A nearly complete
pelvic girdle associated with a skull.

S.A.M. K227. Pristerognathus sp. Louisrus of Dalajalon, Beaufort West. Tapino-
cephalus zone. Coll. Boonstra and Zinn 1959. Two ilia found in association
with a poor snout.

S.A.M. K231. Alopecognathus angusticeps Lammerkraal, Prince Albert. High
Tapinocephalus zone. Coll. Boonstra and Gow 1959. The major part of an
articulated skeleton articulated to a skull.

S.A.M. K234. Pristerognathus sp. Palmietfontein, Beaufort West. High Tapino-
cephalus zone. Coll. Boonstra, Zinn and Gow 1959. Part of pelvis associated
with a snout.

S.A.M. K238A. Pristerognathus? Lammerkraal, Prince Albert. High Tapino-
cephalus zone. Coll. Boonstra and Zinn 1959. A pelvis and pes.

S.A.M. K245A. Pristerognathid? Rietfontein of Vlakfontein, Beaufort West.
High Tapinocephalus zone. Coll. Boonstra, Zinn and Gow 1959. A pelvis
and part of the pes.

S.A.M. K306. Pristerognathus sp. Kranskraal, Beaufort West. Low Tapino-
cephalus zone. Coll. Boonstra, Zinn and Gow 1960. Distal and proximal
ends of two femora.

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

S.A.M. K317. Pristerognathus sp. Bulwater, Beaufort West. Low Tapinocephalus
zone. Coll. Boonstra and Gow 1960. Proximal and distal ends of femur
with parts of skull.

S.A.M. K3309. Alopecognathus sp. Klipbanksfontein, Beaufort West. Tapino-
cephalus zone. Coll. Boonstra and Zinn 1962. Incomplete pectoral girdle
associated with anterior two-thirds of a skull.

S.A.M. K352. Scymnosaurus sp. Skoppelmaaikraal, Laingsburg. Low? Tapino-
cephalus zone. Coll. Boonstra 1962. Femur (isolated).

S.A.M. K353. Scymnosaurus sp. Klein-Koedoeskop, Beaufort West. Low
Tapinocephalus zone. Coll. Boonstra 1929. Proximal end of a large femur.

HIsTORICAL

Hitherto little was known of the postcranial skeleton of the Pristerognathi-
dae.

In 1929 Broom published a figure of the shoulder girdle of Pristerognathus
minor as seen in ventral view. Broom does not state on what specimen this
restoration was based and I have not been able to trace the specimen. In the
sequel I am reproducing figures (fig. 2) of the pristerognathid pectoral girdle
drawn from a model I have carved out of a rectangular block of plaster which
has enabled me to be quite sure that the lateral, anterior and ventral views are
really at right angles to each other. My ventral view differs considerably from
that given by Broom, particularly in the disposition of the scapulo-coracoid.

In 1932 Broom published as a front view of the pelvis of a therocephalian
(possibly Pristerognathus minor) what is in fact a ventral view, but again I have
not been able to trace the specimen on which the drawing was based.

The specimen on which Seeley founded Theriodesmus phylarcus, from the
Endothiodon zone, has by some recent compilators been referred to the Pristero-
gnathidae. Comparison with the pristerognathid material described in this
paper shows that Theriodesmus is not a therocephalian at all, but really a
gorgonopsian.

GENERAL MORPHOLOGY
Pectoral Girdle (fig. 2)

This general account is compiled from facts derived from 18 specimens
representing 6 pristerognathid genera described in the systematic part of this
paper. The material as a whole is not well preserved and in even the best
specimens the constituent bones have been displaced relative to each other and
distorted mostly by dorso-ventral compression. The reconstructions presented
here in semi-diagrammatic form have been drawn from a composite model
carved out in a block of plaster.

As in all primitive reptiles the pectoral girdle is composed of 11 bones—
one unpaired and five paired, but in no case is the cleithrum preserved and its
presence is only indicated by the facet on the scapula to which it was applied.
There is no ossified sternum.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 127

The girdle consists of two sets of bones—replacement and dermal bones.
The replacement bones, forming the so-called scapular girdle, form a scapulo-
coracoid composed of a dorsal plain blade-like scapula, lying practically
vertically but curving slightly round the thorax, and a ventral antero-posteriorly
elongated coracoidal plate, composed of a large precoracoid and a smaller
posterior coracoid, curving gently inwards towards the middle line. The pair of
scapulo-coracoids are held in position relative to each other by their connexion

Fic. 2. Diagrams of the pristerognathid pectoral girdle drawn from a model based on a number of
specimens. a, lateral. b, anterior. c, ventral.

CO—coracoid (posterior). CL—clavicle. CM—cleithrum. g—glenoid. ICL—interclavicle. PCO—
precoracoid (anterior). SC—scapula. scf—foramen supracoracoideus. tr—origin of the scapular head
of the m. triceps (in Zinnosaurus the origin is from a distinct tubercle).

with the clavicular girdle of dermal bones consisting of an unpaired medially
disposed interclavicle and a pair of bracing clavicles. Ventrally the coracoidal
plate rests on the upper face of the interclavicle. Ventrally the spatulate end
of each clavicle curves below around the anterior end of the interclavicle to
fit into an oval hollow on the under face of this bone. Dorsally the stem of the
clavicle sweeps upwards externally of the anterior end of the precoracoid and
the lower part of the scapula and is then applied to the anterior edge of the
scapular blade. Here it meets the cleithrum which is in all probability a splint-
like bone applied to the upper anterior scapular edge.

On the posterior border of the scapulo-coracoid, at the junction of the
scapular and the coracoid, lies the simple, antero-posteriorly shortened, glenoid

3

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

cavity. The glenoid has a dorsal scapular facet which faces ventro-posteriorly
and but slightly externally, and a ventral coracoid facet which faces dorso-
externally and slightly posteriorly. The nature of the glenoid cavity prevents the
humerus from assuming any appreciable downward disposition and also limits
an anterior disposition, but it can be freely directed upwards and backwards.

The area of origin of the scapular head of the triceps muscle from the
postero-lateral surface of the scapula above the glenoid is very indistinct, except
in one specimen, the type of the new genus <imnosaurus, where a prominent
tubercle is developed very similar to that present in the anteosaurian dino-
cephalian genus, Hccasaurus. On the anterior edge of the scapula there is no
indication of an incipient acromion process. The scapula blade is flat with no
indication of any spine.

The precoracoid takes no part in the formation of the glenoid cavity and
just enters into its anterior rim. Immediately anterior to this lies the foramen
supracoracoideus. The anterior extent of the precoracoid is great, forming a
large surface for the origin of the m. supra-coracoideus. Above the precoracoid
lies the thin anterior plate of the scapula from whose outer face the m. scapulo-
humeralis originates.

The coracoid is a smaller but more heavily built bone than the precoracoid,
with a large strong glenoid facet. There is no special process as in the pelycosaurs
for the origin of the coracoidal head of the triceps and this head was probably
absent.

The clavicle of the pristerognathids is very distinctive, differing greatly
from both that of the more primitive pelycosaurs and of the other contemporary
therapsids. It is peculiar in that the ventral spatulate end curving round to the
under face of the interclavicle is sharply bent backwards and extends far
posteriorly along the under face of the interclavicle. The dorsally sweeping
stem is also relatively stronger.

The interclavicle is a long, well-developed bone with the anterior spatulate
end rounded in outline, not much expanded laterally and curving upwards
only very slightly. In these features it differs markedly from the interclavicle
in both pelycosaurs and the other therapsid contemporaries.

The cartilaginous sternum presumably lay above the spatulate posterior
end of the interclavicle and extending to the coracoids.

Humerus (fig. 3)

A dozen fairly complete humeri and numerous proximal and/or distal
ends are available for study, but bad preservation due to both erosion and
post-mortem deformation makes it difficult to get a good picture of the nature
of the pristerognathid humerus. It is particularly difficult to determine the
angle subtended by the proximal and distal ends and thus the rotation on the
shaft.

The pristerognathid humerus varies from a fairly light bone with
moderately expanded ends and a fairly long and slender shaft to a moderately

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 129

heavy bone with greatly expanded ends and a short thickened shaft. The rota-
tion of the ends on the shaft apparently varies from 10° to 25°.

The proximal surface has the processus medialis and processus lateralis
indistinctly demarcated from the caput which is narrowly oval in outline. The
facet of the caput curves a little on to the dorsal surface of the humerus. The
delto-pectoral crest is fairly weak and this subsides into the shaft without
continuing as-an oblique ridge in the direction of the entepicondylar foramen.

cap

dpc

re
a b G

Fic. 3. Diagrams of the pristerognathid humerus. a, dorsal. 6, ventral.
c, proximal.

advl—anterior dorso-ventral line. bf—bicipital fossa. br—origin of the m.
brachialis. cap—caput humeralis. d—insertion of the m. deltoideus. dpc—
delto-pectoral crest. ect—ectepicondyle (radial). ecf—ectepicondylar foramen.
ent—entepicondyle (ulnar). enf—entepicondylar foramen. ld—insertion of the
m. latissimus dorsi. lml—latero-medial line. of—fossa for olecranon (trochlear).
pl—processus lateralis. pm—processus medialis. rce—radial condyle or capitel-
lum. tr—origin of the medial head of the triceps. uc—ulnar condyle.

The bicipital fossa is deep with a strong rounded posterior rim whose posterior
face forms a rectangular area for the origin of the medial head of the triceps.

The proximo-dorsal surface of the humerus is divided by the anterior
dorso-ventral line (ADVL) into two parts. Anteriorly (preaxially) to this line
lies the roughly triangular surface for the insertion of the m. deltoideus. On the
surface posterior (postaxially) of the ADVL there is a weak oblique latero-
median line (LML). Anterior to this line the area of insertion indicates a
_ Strong m. latissimus dorsi. Posterior to the LML a well-developed rectangular
area indicates the origin of a strong medial head of the m. triceps.

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distally the epicondyles are usually moderately developed, except in
Kinnosaurus, where the epicondyles have widely expanded thin flanges of bone.
In this genus the ectepicondyle, which is fused to a well-developed supinator
flange, forms a widely expanded thin sheet of bone. A rounded foramen pierces
the supinator-ectepicondylar flange obliquely. The entepicondyle in this form
forms an extensive distal sheet of bone indicating well-developed flexors.

Distally the condyles are weakly developed. This is particularly so in the
case of the radial condyle whose articulatory face lies practically wholly distally
with hardly any ventral surface and no indication of a ventral bulbous swelling
(capitellum) as is present in most of the contemporary therapsids. On the
dorso-distal surface the trochlear fossa is shallow, which is related to the virtual
absence of an olecranon on the ulna.

The elbow joint in the pristerognathids thus differs markedly from that of
the other therapsids.

The distal position of the distal humeral condyles makes a more upright
dispostion of the limb possible. With the weak development of the olecranon a
deep trochlear fossa is not necessary for the extension of the epipodial.

Ulna and Radius (fig. 4)

The pristerognathid ulna has its proximal end greatly expanded. Proxi-
mally the lateral corner is developed into a short but thickened olecranon with
a rugose surface for the reception of the m. triceps. The sigmoid face for the
reception of the humerus is shallow but the coronoid process is well developed.
Dorsally the sigmoid rim is concave to receive the head of the radius, and the
sigmoid face of the ulna and the proximal face of the radius form a common
articulatory facet for the reception of the humerus. This facet is shallow and, the
humerus having hardly a capitellum, the elbow joint forms a poor hinge joint.

The sigmoid face of the ulna lies on the proximal end of the bone and the
coronoid process on the medial edge is also proximally situated, so that in the
elbow joint the propodial and epipodial meet end-on without the ulna curving
round the distal end of the humerus.

Carpus and Manus (fig. 4)

The proximal row of carpals consists of three elements—a robust radiale, a
lighter elongated ulnare and a small laterally compressed intermedium. A
pebble-like pisiforme lies laterally of the ulna-ulnare articulation.

In the middle row there are two centrals—one lying between the radiale
and the first and second distals, the central one lying wedged in between the
ulnare and radiale and distally articulated with the third distal.

There are four distals—the fourth and fifth are fused and articulate with
the fourth and fifth metacarpals.

The metacarpals are well developed—the first is very short and looks
very much like the first phalanx, the second is nearly twice as long as the first,
the third about 24 times as long, the fourth 3 times as long and the fifth just

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 131

over 3 times as long. The phalangeal formula is 2, 3, 3, 3, 3. The proximal
phalanges are all fairly short and robust and there is no indication of any
further reduction. The ungual phalanges carried talon-like claws and have a
ventral thickening proximally.

The first digit is the shortest, then come the second and fifth of equal length,
then the third and fourth which are again of equal length.

fase) Fic. 4. Diagram of the priste-
rognathid epipodial and manus
in dorsal view.

The purchase of the forefoot on the ground is thus more meso-post-axonic
than pre-axonic.

The digits as a whole are short and the metacarpals and carpus in com-
parison long. The manus can be considered semi-digitigrade.

C,, C,—centrals. [—inter-
medium. P—pisiforme. R—

radius. RE—radiale. U—ulna.
UE —ulnare. 1-5—distals. I-V

metacarpals.

Pelvic Girdle (fig. 5)

Pelves are not well represented and this account is compiled from half a
dozen specimens representing four genera.

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

The pelvis is much lower than the pectoral girdle and the pubo-ischiadic
plate shorter than the interclavicle, but longer than the coracoidal plate.

The three bones of each side are not firmly ankylosed and the junction
between the two sides is weak between the pubis but stronger at the ischiadic
keel.

The acetabulum is large and nearly circular in outline; all three bones
take part, with the ilium contributing about half. The acetabulum faces mainly
outwards; it is shallow, with strong rims for the attachment of the joint capsule

d b

Fic. 5. Diagram of the pristerognathid pelvis. a, lateral. 6, ventral.

a—acetabulum. ap—anterior process of the iliac blade. ife—area of origin of

m. ilio-femoralis. ifi—area of origin of m. ilio-fibularis. it—area of origin of

m. ilio-tibialis. [L—ilium. IS—ischium. isk—ventral keel of the ischia. ist—

ischial tuber. P— pubis. pf— pubic foramen. pp—posterior process of the iliac

blade. pt— pubic tuber. sab—supra-acetabular buttress. san—supra-acetabular
notch.

and ligaments. Dorsally lies a strong supra-acetabular buttress formed by the
iltum, overhanging the acetabulum and forming the main attachment of the
joint capsule. At the dorso-posterior corner of the acetabulum, just posterior to
the buttress, lies the supra-acetabular notch. :

At the level of the iliac buttress the iliac blade is antero-posteriorly
expanded to form a high anterior and a somewhat lower posterior process. In
most specimens the anterior process is much shorter than the posterior process,
but in two specimens the lower corner of the anterior process is prolonged to
form a long but weak process. In the other specimens there is an indication
of this process. More and better-preserved specimens may prove that this
elongation is normal for the family. The outer face of the iliac blade is both
dorso-ventrally as well as antero-posteriorly convex. In antero-dorsal direction
runs a shallow groove and near the dorsal edge in a vertical line with the
posterior end of the buttress there is also a slight hollow. One can thus speak
of an undulating surface. There is no eversion of either anterior or posterior

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 133

edges. The areas of origin of the ilio-fibularis, ilio-femoralis and ilio-tibialis are
thus indefinitely determinable.

The ischium in its acetabular part is massive and forms here, the strongly
thickened postero-ventral segment of the strong acetabular rim. From here it
tapers posteriorly and forms a thickened upper edge but without a distinct
ischial tuber. Extending medially the ventral surface is flattened anteriorly
but just behind the junction with the pubis it carries a strong and prominent
ventral keel.

The pubis in its acetabular part is fairly massive and here forms a fairly
strong acetabular rim. From here it tapers anteriorly and forms a thickened
lateral edge but without a marked pubic tuber. The ventral surface of the
pubis is fairly flat. Just medial to the acetabular rim, near its posterior edge,
lies a well-developed pubic foramen.

The pubo-ischiadic plate is broad and large as in the pelycosaurs, the
antero-posterior dimension of the pubis being relatively larger than in other
therapsids and both anterior and posterior pelvic openings broad and not
V-shaped as in other therapsids; but the posterior opening is incipiently
V-shaped because of the ischial keel.

Femur (fig. 6)

The pristerognathid femur is represented in 20 specimens of 5 separate
genera. Ten femora are fairly complete and there are a couple of dozen poor to
good distal and proximal ends unconnected and usually lacking a shaft.

The femur, always longer than the humerus, is a fairly light bone with
only slightly expanded ends and with a long, fairly slender shaft. There is a
considerable twist on the shaft so that the proximal head and the distal condyles
subtend an angle of 30°-40°. The distal preaxial condyle lies a little further
ventrally than the postaxial distal condyle and a little proximal of its fellow.

The preaxial border is more concave than the postaxial border due to
the femoral head being directed somewhat preaxially.

Proximally the caput femoris is terminal but somewhat preaxially directed ;
it is antero-posteriorly elongated, thick preaxially with a rounded edge;
postaxially it tapers and curves gently towards the external trochanter, into
which it flows. In its thickened preaxial part the caput is convex and rounded
in its postaxial part. The caput femoris is much smaller than the acetabulum.

The external trochanter forms the proximo-postaxial rounded corner of
the femur; it flows with a rounded curve into both the caput and the postaxial
edge.

On the dorsal proximal surface lies a well-developed ridge near the preaxial
border of the femur; it lies parallel to the preaxial border and commencing
from the caput runs for a short distance distally and then fades away. This
ridge serves as the area of insertion of the m. pubo-ischio femoralis internus.
Between this ridge and the external trochanter lies the area of insertion of the
m. ilio-femoralis.

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

On the ventral proximal surface a sharp, prominent longitudinal ridge is
developed. Lying in the middle of the bone, it commences a short distance
distally of the caput and extends distally for a variable distance to fade away
into the general ventral face of the shaft. This ridge is the only part of the
primitive Y system of adductor ridges preserved in the pristerognathids and
represents an internal trochanter.

@n ec
Fic. 6. Diagrams of the pristerognathid femur. a, dorsal. b, ventral.
c, anterior. d, posterior. e, proximal.

ad—insertion of adductor muscle. cap—caput femoris. ec—ecto-

condyle. en—entocondyle. ext—external trochanter (major). ft—

—origin of m. femoro-tibialis. g—origin of the gastrocnemius.

if—insertion of the m. ilio-femoralis. int—internal trochanter.

ifo—intertrochanteric fossa. pifi—ridge on which is inserted the
m. pubo-ischio-femoralis internus.

Between the internal and external trochanters lies a relatively small
intertrochanteric fossa.

The distal condyles lie terminally facing very slightly ventrally and are
well modelled. The intercondylar sulcus is very shallow. Dorsally the inter-
condylar fossa is very shallow, but ventrally the gastrocnemic fossa is quite well
developed.

Tibia, Fibula and Pes (fig. 7)

The posterior epipodial and pes are not as well represented in the material
at my disposal as the corresponding structures of the forelimb.

The posterior epipodial is longer than the anterior one, as we have already
seen is also the case in regard to the propodial.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 135

Both tibia and fibula are lightly built, slender bones, lighter and slenderer
than the radius and ulna. In both epipodials the pairs of bones are approximately
of equal length. Proximally the tibia and fibula form a common terminal
articular facet to meet the femoral condyles end on. Distally the tibia articulates
with a rounded face on the astragalus and the fibula with an elongated facet
on the proximal end of the calcaneum.

tf Fic. 7. Diagram of the
pristerognathid epipodial
and pes in dorsal view.

(c) AS—astragulus. CA—cal-

caneum. C—central. F—
fibula. T—tibia. tf—tarsal

@

AD A 2 foramen. 1-5—distals. I-V
}e : — metatarsals.

0

As in pelycosaurs and therapsids generally the pristerognathid tarsus has a
pair of proximal elements—an astragalus and a calcaneum.

The astragalus is a robust, roughly ovoid bone with a rounded face for
articulation with the tibia; laterally it has an elongated facet articulating with
the calcaneum and distally a curved facet facing the centrale.

The calcaneum is a larger but more lightly built bone. It is a sheet of bone,
thickened proximally to form an elongated facet for the reception of the fibula

4

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

and with a thickened medial edge abutting against the astragalus, but notched
for the passage of the penetrating tarsal vessels, and thickened distally where it
carries a facet to receive the fourth distal carpal. There is no indication of a
tuber-like heel.

One fairly small centrale is present. There are five distal tarsals, but in
one specimen the small fifth distal is fused to the large fourth distal as is usual
in therapsids.

The fourth distal is always large; the third and second distals, when well
developed, have flattened upper faces and are roughly squarish in outline; the
first distal is usually large, articulating with both first and second metatarsals,
but in one specimen it is a small, pebble-like bone.

The first metatarsal is a short, fairly squat bone, the second, also short, has
a more constricted shaft; from the third to the fifth the metatarsals become
progressively longer with well-expanded ends.

The digits are, as in the manus, short, with the phalangeal formula 2, 3, 3,
3, 3. The first phalanx in all the digits is usually fairly long, but in one specimen
that of the first digit and in another that of the fifth digit is greatly shortened,
being disc-like without a constricted waist.

The second phalanx of the second to the fifth digits is short with a con-
stricted waist.

The terminal phalanges are long, curving, claw-like. The first digit is
short and the other four of about equal length. The purchase of the hind foot
thus lies in the postaxial part of the foot.

SYSTEMATIC DESCRIPTIONS OF SPECIMENS

A. PECTORAL GIRDLE
Alopecognathus (figs. 8-10)

The specimen 12051 includes a scapular blade; K339 has a nearly complete
scapula; K223A and K223B each consists of a fairly complete girdle, but both
lack the scapular blades and the upper part of the clavicles and have undergone
distortion and displacement of the constituent elements; K231 includes anearly
complete girdle, but is distorted by dorso-ventral compression. In the accom-
panying figures the distortion has been corrected and the symmetry restored.
In none of these is the cleithrum preserved.

The girdle is a structure of considerable size; high, broad, with its ventral
element—the interclavicle—long. The scapula, as restored, is dorsally directed
or is tilted slightly backwards and curved slightly to follow the contour of the
thorax. The area of origin of the scapular head of the triceps is indistinct—
there is no ridge, mound or tubercle. The coracoidal plate is long, but in two
specimens the anterior edge of the procoracoid is concave, whereas in the other
it forms an even convex curve. The clavicles have a long, posteriorly directed
ventral spatulate end in two specimens, whereas in another this part of the
clavicle is quite short. The interclavicle is large, with a rounded anterior

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138 ANNALS OF THE SOUTH AFRICAN MUSEUM

expansion and a long broad posterior spatula joined by a fairly narrow neck
at the level of the posterior coracoids.

Priesterognathoides (figs. 11 and 12a)

In 12102 the right half of the pectoral girdle is preserved together with an
incomplete interclavicle and parts of the left half of the girdle. The girdle is
partially disarticulated but only slightly distorted in its fall backwards when the
soft tissues decomposed.

In 12204 a disarticulated girdle has a good scapula, coracoid, procoracoid,
interclavicle and the ventral spatulate ends of the clavicles.

The scapula is apparently tilted somewhat backwards and shows a marked
curvature around the thorax. No origin of the triceps can be determined and
there is definitely no tubercle. The coracoid is short, but the procoracoid well
developed. The interclavicle is very similar to that of Alopecognathus.

Pristerognathus (fig. 12b and c)

In 4335 there are a pair of incomplete scapulo-coracoids. The coracoid
is short and the procoracoid well developed. No scar or tubercle for the scapular

o@=e cee © SC S88 S@ee ea &

‘ew eeeerre

Fic. 12. Pectoral girdles. x4. a, Pristerognathoides

Fic. 11. Pristerognathoides sp.S.A.M. sp. S.A.M. 12204 in lateral view. b, Pristerognathus

12102.x4. Pectoral girdle in sp. S.A.M. 4335 in lateral view. c, Pristerognathus sp.
lateral view. S.A.M. 11458 in ventral view.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 139

head of the triceps can be traced. In 11458 there is a good interclavicle which has
a large anterior expansion, but the posterior spatula is only slightly expanded.
The ventral spatulate ends of the clavicles extend well posteriorly. The pro-
coracoid extends anteriorly of the interclavicle.

Scymnosaurus (figs. 13-17)

Five specimens have parts of the pectora! girdle preserved. 8034 and 12193
have a well-preserved coracoid each. Both are massive, and that of 12193 is
probably of S. major. In the type specimen of S. major (g005) there are the
anterior two-thirds of a large interclavicle and the ventral spatulate ends of
both clavicles, which, though massive, have relatively a short posterior extent.
In 11557 there are a disarticulated scapula, interclavicle and clavicle. The
scapula is robust but short; the glenoid facet faces much posteriorly and
laterally; the facet for the cleithrum is clearly shown, but the origin of the
scapula head of the triceps is from the smooth postero-lateral surface just
above the glenoid.

A left clavicle is stoutly built; the dorsal end clasping the anterior border
of the scapula is expanded and strong. The ventral spatulate end is large and
extends far posteriorly along the circular head of the interclavicle. Both the
articulating surface of the clavicle as well as the hollowed face on the inter-

Fic. 13. Bones of pectoral girdles.
x 4. a, Scymnosaurus major. S.A.M.
9005. Type. Ventral view of inter-
clavicle and clavicle. b, Scymnosaurus
ferox. S.A.M. 9084. Right coracoid
in ventral view. c, Scymnosaurus sp.
S.A.M. 12193. Right coracoid in
ventral view.

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 14. Right scapula of Scymnosaurus sp. S.A.M. 11557. X 4. a, posterior. 5, lateral. c, anterior.
d, internal.

Fic. 15. Scymnosaurus sp. S.A.M.
11557. X $. Clavicle and inter-
clavicle. a, ventral. 5, lateral.

141

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA

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142 ANNALS OF THE SOUTH AFRICAN MUSEUM

clavicle bear longitudinal striae, and the backward movement of the clavicle is
limited by a thickened border on the clavicle which abuts against the anterior
edge of the interclavicle. The dorsal stem of the clavicle and its ventral spatulate
end subtend what is nearly a right angle.

The anterior end of the interclavicle is large and nearly circular in outline,
and a strong median ridge separates the two facets for the clavicles. The posterior
end is greatly expanded but short, and is remarkably lightly built.

In 11695 there is a fairly complete shoulder girdle, but with the constituent
bones displaced and distorted by dorso-ventral pressure. The anterior head of
the interclavicle and ventral spatula of the clavicle are both relatively smaller
than in 11557, and the dorsal stem of the clavicle is longer and more slender,
with a bifurcated end. The scapular blade is fairly tall but is relatively lightly
built.

Theriowdes (fig. 18)

In 11888, the holotype specimen, much of the pectoral girdle is preserved,
but here again disarticulation and distortion have made reassembly and
restoration of symmetry, as shown in the figures, difficult. The girdle is very like

Fic. 19. Cinnosaurus pau-
cidens gen. et spec. nov.
S.A.M. 12185. Type.
Fic. 18. Therioides cyniscus. Pectoral girdle. S.A.M. x}. Lateral view of
11888. Type. x }. a, ventral. b, lateral. scapulo-coracoid.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 143

that in Alopecognathus, but smaller and of a lighter build, with a proportionately
large anterior expansion of the interclavicle.

Ainnosaurus paucidens gen. et spec. nov. (fig. 19)

In 12185, the holotype specimen, there is a well-preserved right scapulo-
coracoid without scapular blade, and a left scapular blade. This scapulo-
coracoid differs very markedly from all those hitherto considered. The scapular
blade has its upper anterior part expanded and on this outer face lies a facet for
the upper end of the cleithrum; lower down the cleithrum is applied to the
anterior edge of the scapula.

Immediately above the upper rim of the glenoid there lies a process on
the posterior face of the scapula for the origin of the scapular head of the triceps.
This process forms a strong, prominent flange of bone, laterally compressed
and dorso-ventrally elongated.

The glenoid is a widely open cavity facing largely outwards. Both its dorsal
scapular rim and ventral coracoidal rim are sharp and prominent. Anteriorly
there is no rim on the procoracoidal border of the glenoid so that in its anterior
movement the head of the humerus would ride in a broad groove.

The procoracoid extends far anteriorly.

B. HUMERUS
Alopecognathus (fig. 20)

In 12051 there is a good, apparently undistorted, proximal half of a
humerus. In K231 both humeri are preserved but both are quite obviously
distorted and in the figures I have tried to correct the distortion. The result
is not very convincing and the two humeri have quite distinctive outlines.

In 12051 the central part of the caput quite definitely flows over onto the
dorsal surface and this would obviously affect the nature of the shoulder joint

Fic. 20. Alopecognathus angusticeps. Humerus. x 4. S.A.M. 12051 : a, ventral. 5, dorsal. S.A.M.
K231: c, ventral. d, dorsal.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

as the humerus would tend to be directed more horizontally. In K2g1 the
caput is terminal. In 12051 the delto-pectoral crest is weak with little indication
of the pectoralis insertion, and the processus medialis, caput and processus
lateralis flow into one another.

In K231 the distal condyles are mostly terminal with only a little extension
onto the ventral face. Both supra-trochlear fossa and brachialis fossa are
deeply excavated. The radial condyle is not developed into a rounded
capitellum.

Cynariognathus (fig. 21a)

In 11794 both humeri are imperfectly preserved. The bone is lightly
built and the distal condyles mostly terminal without a bulbous capitellum.

Pristerognathoides (figs. 21b, c and d and 22)

In 5018 both humeri are incompletely preserved and in 12204 isa fairly
good left humerus. In other specimens, e.g. 12102, only poorly preserved
humeral ends are present. In 5018 the humerus is long, with both ends well
expanded and the shaft slender. The caput, mainly terminal, curves slightly onto
the dorsal surface. The distal condyles are a little ventral of terminal, and the
capitellum is moderately swollen. I have not been able to locate an ectepi-
condylar foramen. In 12204 the ulnar condyle is separated from the olecranon
fossa by a well-developed ridge which would appear to limit the extension of the
epipodial. The delto-pectoral crest is weak and the area for the insertion of the
deltoideus small.

ad

b

Fic. 21. Humerus. x}. Cynariognathus sp. S.A.M. 11794. a, ventral.
Pristerognathoides sp. S.A.M. 5018. b. dorsal. c, ventral. d, posterior.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 145

Pristerognathus

A number of humeral ends of Pristerognathus are poorly preserved and
warrant no description except to state that the humerus is very similar to
that of Pristerognathoides—but smaller.

Ptomalestes (fig. 23)

In 11942, the holotype, both humeri are preserved. I am including figures
of both the right and left humerus which show how much they have suffered
from post-mortem distortion. If the distortion is corrected the humerus would
be a fairly robust bone, moderately long with well-expanded proximal and

~

- @ a :

Fic. 22. Humerus. Pristerognathoides sp. S.A.M. 12204. x4. a, dorsal. b,
ventral. c, posterior. d, S.A.M. 12102, ventral.

qd

Fic. 23. Humerus. Ptomalestes avidus. S.:A.M. 11942. Type. x }. a, dorsal. b, ventral. c, posterior.
d, proximal. e, ventral of right humerus.

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

distal ends and a fairly short shaft; both caput and distal condyles are terminal,
with the capitellum weak and little ventrally placed. A ridge separates the
ulnar trochlea from the shallow supra-trochlear fossa.

=<.-<<-"

Fic. 24. Humeri. x 4. Scymnosaurus major. S.A.M. 9005. Type.
a, dorsal. 6, ventral. Scymnosaurus ferox. S.A.M. 9084.
c, posterior. d, ventral.

Scymnosaurus (figs. 24-26)

Of this genus I have two very well preserved distal ends, and one good
complete humerus, which I believe to be undistorted, and this specimen must
be taken to give us the best idea of the pristerognathid humerus as represented
by the largest genus of this family of primitive Therocephalia.

In 9005, the type specimen of Scymnosaurus major, the good distal humeral.
end is, considering the bulk of the animal, only moderately robust. The epi-

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA

147

e d
Fic. 25. Scymnosaurus sp. S.A.M. 11557. x 4. Humerus. a, dorsal. b, ventral. c, posterior. d, anterior.
é, proximal.

—-------

26. Scymnosaurus sp. S.A.M. 11695. x4. Humerus. a,
c, anterior.

dorsal. 5, ventral.

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

condylar expansions are modest, with the confluent supinator flange and
ectepicondyle slightly more prominent than the entepicondylar edge. The
condyles lie chiefly terminal. The radial condyle forms only a weak capitellum,
which only slightly enters the ventral surface. The ulnar condyle forms a
shallow trochlea, which dorsally has a ridge separating it from the very shallow
supra-trochlear fossa hardly functioning as an olecranon fossa. Ventrally there
is hardly a coronoid fossa, but the large oval entepicondylar foramen opens
into a deep hollow bounded postaxially by a strong rounded ridge. The distal
epicondylar edges are strong but not much expanded and their rugose surfaces
give a strong origin for the flexors and extensors. In 9084, which is smaller and
lighter, being a humerus of the less bulky Scymnosaurus ferox, the ectepicondylar
flange is more expanded and curves downwards to form a deep groove. In
11557, which is most probably also of Scymnosaurus ferox, there is a well-preserved
left humerus. Its terminal distal condylar face is less rounded than in 9084 and
has no capitellum to speak of, and in its postaxial part it is concave instead of
convex, indicating the presence of considerable joint cartilege. Its ectocondyle
is without the flange present in 9084. Its supratrochlear fossa is deeper than in
9084 with a strong preaxial border.

11557 is the only Scymnosaurus humerus with a good proximal two-thirds
preserved. Ventrally the bicipital fossa is deep and well demarcated from the
surface of the short shaft. The delto-pectoral crest is fairly strong but the
pectoralis insertion is not localized but diffuse. The postaxial border of the
bicipital fossa is formed by a robust rounded ridge which on its postaxial
face gives a strong face for the origin of the medial humeral head of the triceps.
Dorsally both the anterior dorso-ventral line and latero-medial line are
moderately developed and the areas of insertion of the deltoideus, latissimus
dorsi and both the scapulo-humeralis anterior and posterior are well
developed. |

Proximally the caput is terminal with only a slight extension onto the
dorsal surface; it is a very flat oval and flows evenly into both the processus
medialis and lateralis.

The incomplete humerus of 11695 shows a greater twist on the shaft but is
otherwise very similar to that of 11557.

Therioides (fig. 27)

The one humerus known in the type, 11888, is fairly short with moderately
expanded ends but otherwise shows no special features to distinguish it from
those of the other medium-sized pristerognathids so far described.

Kinnosaurus gen. et spec. nov. (fig. 28)

In the type material (12185) there are a practically complete right
humerus and the proximal half of the left humerus. Both have suffered from
distortion. I am including figures of the right humerus with the distortion
corrected.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 149

The caput is strap-shaped but in its middle part tends to overflow slightly
onto the dorsal face. The processus medialis and processus lateralis flow gently
into the caput. The delto-pectoral crest is long, with its proximal edge thin and
its distal end somewhat thickened for the reception of the pectoralis.

qd

Fic. 27. Theroides cyniscus. S.A.M. 11888. Type. x 4. Humerus.
a, dorsal. 6, ventral. c, posterior.

-

qd

Fic. 28. innosaurus paucidens gen. et spec. nov. S.A.M. 12185. x4. Right
humerus with distortion corrected. a, dorsal. b, ventral. c, posterior. d, anterior.

The distal end is greatly expanded. This expansion is mainly due to the
development in both epicondyles of thin sheets of bone. This is particularly so
in the ectepicondyle, where the confluent supinator process and the epicondyle
form an extensive antero-ventrally sweeping sheet of bone.

The distal condyles are terminal with very little capitellar development,
but the ulnar condyle is quite well developed with a dorsal ridge separating it
from the very shallow supra-trochlear depression.

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

C. THE LOWER FORELIMB AND FOREFOOT
Alopecognathus (fig. 29)

In 12051 proximal and distal ends of both ulnae and radii and one good
carpus are preserved. .

Proximally the head of the radius fits into the concave sigmoid rim of the
ulna so that its facet lies nearly as far proximally as that of the sigmoid facet of
the ulna which is also situated terminally. The olecranon is robust but does not
extend proximally as a process.

Fic. 29. Alopecogna- Fic. 30. Cynariogna-
thus angusticeps. thus’ sp. SAE
S.AsM.r2051f x4: 11794. X+4. Right
Epipodial and epipodial and partial
carpus. manus.

In the carpus the radiale is a stout bone roughly rectangular in outline.
A longitudinal ridge separates a larger preaxial dorsal face from a smaller
postaxial face. Distally a ridge separates the facets for the two centrals.

The intermedium is a laterally flattened small bone wedged in between
radiale and ulna and ulnare.

The ulnare is a long element with expanded ends and a long constricted
waist. Its proximal facet is much smaller than the distal ulnar facet.

Both centralia are well-developed bones, each articulating with a pair of
distals.

There are four distals—the fused fourth and fifth articulating with the
fourth and fifth metacarpals.

Only the proximal ends of the five metacarpals are preserved.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA I5I

Cynariognathus (fig. 30)

In 11794 both anterior epipodials are preserved but the preservation is
not very good. Both the ulna and radius are lightly built and the ulna has no
olecranon process. In the figure the proximal ends of the two bones are shown as
lying next to each other but in life the proximal head of the radius is applied
to the edge of the sigmoid cavity of the ulna so that the two bones have their
proximal facets forming a confluent articulating face. The radial facet meets the
capitellum and in extension the ulna rides on the trochlear facet of the humerus.

Fic. 32. Ptomalestes

avidus. S.A.M.
Fic. 31. Pristerognathoides ? S.A.M. K357. tig42. Type... Xz:
Nat. size. Partial manus. Epipodial.

Only the right manus is in part preserved—the digits are missing.

In the carpus 9 bones are preserved. The proximal row consists of a long
slender ulnare shaped somewhat like a metacarpal, a large laterally compressed
intermedium and a short, broad radiale with distally two well-developed
concave facets for the two centralia.

The two centrals are well-developed bones of irregular shape each with a
good facet for articulation with the radiale.

There are four distalia; the first three articulate with the first three
metacarpals, whereas the fourth articulates with both the fourth and fifth
metacarpals.

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

Of the five metacarpals the last three have only their proximal ends
preserved. They are short, with well-expanded ends. No. 1 is small and light,
and from the second to the fifth they increase rapidly in size so that No. 5 is
quite a strong element.

Pristerognathoides (fig. 31)

In a number of specimens ulnae and radii are preserved, exceptionally
complete but mostly represented by proximal and distal ends. They call for
little comment, except that in all these there is no definite olecranon process,
and that proximally the two facets form a confluent terminal articulating face
for the humeral condyles.

Fic. 34. Scymnosaurus sp. S.A.M. 11557.

x 4. a, radius and manus in dorsal view.

b, lateral view of no. IV_ terminal
phalanx.

Fic. 33. Scymnosaurus major X }.
Epipodial of. S.A.M. 9005.
Type. Carpus of S.A.M. 12193.

Sa oer ae em

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 153

In K357 there is a good carpus with all five metacarpals present. Here the
intermedium is a weak element and articulating with the ulna and ulnare there
is a well-developed pebble-like pisiforme. The ulnare is elongate, the radiale
robust, and the two centrals well developed. There are four distals, with the
fourth articulating with both the fourth and fifth metacarpal.

The first two metacarpals are short, squat bones and the other three much
longer, with the fifth the longest. The proximal phalanx of the fifth digit has
its proximal end greatly expanded.

Ptomalestes (fig. 32)

In the type specimen of Ptomalestes (11942) both anterior epipodials are
preserved. Both the radius and ulna are lightly built and the ulna has no
distinct olecranon process. In the figure the proximal head of the radius is
shown moved out of its articulation with the proximal end of the ulna.

Scymnosaurus (figs. 33-35)

In this genus the epipodial, carpus and manus are represented by some
very good specimens.

The epipodial in the type specimen of Scymnosaurus major (g005) is repre-
sented by a well-preserved left ulna and radius (a mirror image is shown in the
figure). In 9084 there are good proximal ends of the right ulna and radius in
articulation. In 11557 there is a good radius, in 12193 proximal and distal
ends of both ulna and radius and in 12262 a good complete radius and ulna.

In all these specimens the olecranon process of the ulna extends little
proximally of the sigmoid face, and the proximal head of the radius lies anteriorly
(morphologically dorsally) of the proximal end of the ulna and fits securely
against the sigmoid edge forming a confluent articulating face for the feeble
capitellum and the shallow trochlea of the humerus.

The carpus is preserved in part or nearly complete in 12193, 11459, 11557
and in 12262.

In 12193 the radiale is a robust bone; in dorsal view oval in outline with a
convex dorsal and proximal face; concave distally for the first centrale and
with a convex facet for the second centrale; ventrally it has a deep oblique
groove.

The ulnare has its dorsal face with an outline like that of a metacarpal.
Ventrally it has a longitudinal groove and its preaxial edge is convex in contrast
to the dorsal concave preaxial edge.

The two centrals are robust; the first is nearly circular in outline with a
penetrating foramen near its preaxial edge. The proximal central has its
postaxial edge deeply concave, facing the concavity of the ulnare, thus forming
a passage for the vessels penetrating the carpus.

Of the distal carpals only the proximal parts are preserved with the fourth
distal strongly developed.

In 11459 the two centrals, distals together with metacarpals 1, 2 and 3,

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

and the proximal phalanx of the second digit are preserved. The first central
has a groove and a penetrating foramen near its preaxial border. The meta-
carpals increase in length from 1 to 3.

In 11557 there is a nearly complete carpus, but the manus is incomplete
with the preserved bones partly displaced and disarticulated. In the figure

Fic. 36. Thertodes

cyniscus. S.A.M.
11888. Type. X4-
Fic. 35. Scymnosaurus jeies Dorsal view of epi-
S.A.M. 12262. x 4. Epipo- podial and_ partial
dial and manus in dorsal manus.
view.

the constituent bones are shown in natural relation. The radiale is fairly robust,
with a flattish face for the radius; its dorsal surface shows two faces—the
preaxial face nearly rectangular and the postaxial face more squarish, meeting
at an angle of about 130°; the facet for the first central is slightly concave and
for the proximal central convex.

The ulnare is elongated, with a dorsal outline resembling that of a
metacarpal.

—

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 155

The intermedium is a small, laterally compressed bone. The first central
has a depression near its preaxial border and the proximal central is hollowed
out near its proximal wedge-like end.

The first distale is missing, numbers 2 and 3 lie proximally to their respec-
tive metacarpals and the large fourth articulates with both the fourth and fifth
metacarpals.

The first metacarpal is missing; from number 2 to 5 the metacarpals
increase in length with the fifth about 14 times the length of number 2.

The phalangeal formula is 2, 3, 3, 3, 3, 3. The ungual phalanges are
curved, narrow, pointed and claw-like. The others are short, broad and squat,
with well-expanded ends.

In 12262 the forefoot is very well preserved and is complete except that
part of the radiale and the tip of the ungual phalanx of the second digit have
been lost in transit from the field. ‘The general structure is as in all the pristero-
enathids described above, viz. carpal formula 3, 2, 4 plus an ulnar pisiforme, and
phalangeal formula 2, 3, 3, 3, 3. The fourth and fifth metacarpals articulate
with the large fourth distal carpal; the metacarpals increase in length in post-
axial direction and the distal ends of the third and fourth are particularly well
modelled, with a dorsal trochlear fossa and well-rounded distal corners. The
third and fourth digits are the longest and strongest; the first digit is relatively
weak and the fifth strong. The weight is thus carried somewhat more post-
axially than preaxially.

Theriordes (fig. 36)

In the type specimen (11888) much of the right forefoot is preserved. The
carpus is complete; the last four metacarpals are present but only the second
digit is completely preserved.

The structure is typically pristerognathid but the metacarpals are relatively
long and slender.

D. PELVIC GIRDLE
Alopecognathus (fig. 37)

In K218 a fairly large pelvis is preserved. This is, however, badly weathered
and all that-can be determined is that the pelvic plate is large with a distinct
ischial keel and that the outer face of the ilium is convex.

In K231 most of a fairly large pelvis is preserved, but with the edges of
the ilia incomplete. The pelvic plate is long and broad with both ischium and
pubis strong; just more than half is contributed by the ischium; the ilium is
large, with its length nearly as long as the ventral plate (91%). The pelvis is
low and long, with the height 79 per cent of the length.

The ilium is not completely preserved on either side, but the two sides are
complementary so that the figure could be compiled fairly accurately. The
antero-posterior length of the iliac blade is great but its supra-acetabular height
moderate; the height is a little more than half the length (52°). The posterior

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

process of the iliac blade is long but fairly low, whereas the anterior process
is fairiy short but high; although its ventral corner apparently has a long, low
elongation.

The outer face of the iliac blade is in general convex both antero-posteriorly
as well as dorso-ventrally. The m. ilio femoralis (gluteus) thus originated from
a convex area whereas in tetrapods this area’is usually concave. The supra-
acetabular ridge is strong. The ilium forms more than half of the acetabulum.

Fic. 38. Lateral views of pelves.
x 4. a, Cynariognathus sp. S.A.M.

Fic. 37. Pelvis of Alopecognathus angusticeps. 11794. b, Pristerognathoides ? S.A.M.
N-A.M.” Keer.’ x4. a). lateral view. 5018. c, Pristerognathus? S.A.M.
b, ventral view. Koo7,

The pubis is massive in its acetabular part, with a strong dorso-lateral
edge, terminating in a moderate pubic tuber somewhat outwardly directed.
The two pubes form a weak symphysis. A fairly large foramen pierces the bone
near its posterior edge just medial to the strong acetabular rim. The ischium is
longer than the pubis: its acetabular part is strongly developed and its rounded
dorso-lateral edge ends in a moderate tuber. The two ischia form a strong
symphysis which develops a strong medial keel.

The acetabulum is large and faces mainly outwards.

a — “ga;

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 157

Cynariognathus (fig. 38a) |

In 11794 there are preserved, of the right half of the pelvis, the acetabular
proximal parts of the three constituent bones, whereas on the left there are a
fair ischium and the proximal part of the pubis. The iliac blade is not preserved.
Although of lighter build, the pelvis is essentially as in Alopecognathus. The
pubic foramen is large.

Pristerognathoides (figs. 386 and 39)

In 5018 the two ilia are partly preserved and the figure incorporates
features of both sides. The ilium is very similar to that of Alopecognathus, however
its outer face, though also convex, is less so than in Alopecognathus.

Fic. 39. Pelvis of Pristerognathoides sp. S.A.M. K223C. x }. a, lateral. 6, ventral.

In K233C the pelvis lacks only the posterior ends of the ischia. The pelvis
is fairly low but quite long, the height being 75 per cent of the length as recon-
structed. The blade of the ilium is both low and very short, the height being
74 per cent of the length. The outer face of the iliac blade is peculiar. It is not
hollowed out as is usual in all other therapsids, but cannot be described as
convex either. Dorso-anteriorly there is a pronounced bulge separating two
hollows so that the surface can be described as undulating. The anterior process
is short but high, with its anterior edge cut back at the level of the anteriorly
hollowed-out area.

The posterior process is also short but much lower than the anterior
process.

The supra-acetabular ledge is strong.

Both ischia and pubes are robust in their acetabular parts, with strong
circum-acetabular borders. Anteriorly the pubis has a broad, thin anterior
edge with the outer corner showing little of a pubic tuber. The pubic foramen
is large but the symphysis is weak. The ischia form a fairly strong ventral keel.
In its pubic part the pelvis has a flat floor, which in its ischial part becomes
shallowly V-shaped.

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pristerognathus (fig. 38c)

In K227 there is a pair of beautifully preserved ilia. Much smaller than
the ilia in the forms so far described, these ilia are nevertheless very similar.
The undulating outer face is clearly shown; the anterior process is high, with its
anterior edge notched; the posterior process is relatively longer and lower.
The supra-acetabular height of the ilium is about two-thirds of the length of the
iliac blade.

In K238A there are a partial ilium and most of a pair of ischia.

Scymnosaurus

Of this genus only the acetabular parts of an ilium and a pubis which
indicate a moderately robust pelvis with an acetabulum of moderate size
directed outwards are preserved (11957).

In 11558A a pair of good ischia are preserved.

d b

Fic. 40. Pelvis of a pristero-
gnathid. S.A.M. Ke45A.

x4. a, lateral. 6, ventral.

qd

Fic. 41. Femora. x 4. Alopecognathus sp. S.A.M. 12051. a, dorsal. b, ventral. c, anterior.
Cynariognathus sp. S.A.M. 11794. d, dorsal. e, ventral. f, anterior.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 159

An unidentified pristerognathid (fig. 40)

In K245A there is a good pelvis, basically as in the above described forms,
but the ilium is rather peculiar, with a long, low process directed anteriorly.

E. FEMUR
Alopecognathus (fig. 41a, 6 and c)

In 12051 a proximal and two distal femoral ends are preserved. The ends
are moderately expanded, shaft fairly strong, somewhat dorso-ventrally
flattened, and fairly straight.

The well-rounded caput, although terminal, is directed somewhat pre-
axially. The external trochanter is not prominent and is situated well proximally.
The pubo-ischio femoralis internus ridge is well developed, not near the anterior
border. The internal trochanter forms a sharp ridge. The inter-trochanteric
fossa is small and shallow.

In K231 both femora are present and, although badly preserved, are
essentially similar to that figured here.

Cynariognathus (fig. 41d, e and f )

In 11794 both femora are preserved, but are both somewhat weathered and
distorted. The curvatures in the figures on the long axis are thus artificial.

The ridge forming the internal trochanter is long and not so far proximally
situated.

\

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Fic. 42. Femora. x 4. Pristerognathoides sp. S.A.M. 5018. a, dorsal. b, ventral. c, anterior.
S.A.M. 11936. d, dorsal. e, ventral. f, anterior.

160° ANNALS OF THE SOUTH AFRICAN MUSEUM

Pristerognathoides (figs. 42 and 43)

The femur in this genus is represented by the distal and proximal ends of
both femora in 5018 and K306 and one femur in 11936.

In 5018 the internal trochanter forms a long ridge extending on to the
shaft, but it does not extend far proximally, so that the intertrochanteric fossa
is long. The pubo-ischio femoralis internus ridge is low. The distal end is well
expanded. In 11936 the femur is more robust.

In K306 the right femur is a larger, heavier bone than the left and is
certainly not of the same individual and may be of a different species. The distal
ends are only moderately expanded.

f

Fic. 43. Femora. x4. Pristerognathoides sp. S.A.M. Kg306. Right: a, dorsal.
c, ventral. e, anterior. Left: b, dorsal. d, ventral. f, anterior.

:

Pristerognathus

In this genus the femur is represented by poor ends. In K317 the internal
trochanter forms a short, thickened ridge proximal to which lies a long, deep
inter-trochanteric fossa.

Scymnosaurus (figs. 44, 45, 46a, b and c, 47)

In this genus the femur is fairly well represented. In 11557 there is a good,
well-preserved femur, in 11597 a proximal end, in 12118 a proximal end, in
12193 a distal and proximal end, and in 12262 and K352 a poorly preserved
but nearly complete femur.

In the three complete femora there is a greater amount of twist or rotation
of the two ends on the shaft relative to each other than in the femur of all the
other pristerognathids as hitherto described. This twist or rotation is an anti-

ee

oo

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 161

clockwise or postaxial movement of the proximal expansion in relation to the
distal end. The result is that the caput, though terminal, is directed somewhat
dorsally, whereas in the hitherto described forms this direction runs somewhat
anteriorly.

Another difference is that the pubo-ischio femoralis internus ridge in
Scymnosaurus lies on the morphological anterior edge of the bone and not some-

Fic. 44. Left femur of Scymnosaurus sp. S.A.M. 11557. < }. a, dorsal. b, ventral (projection onto
plane of distal condyles). c, ventral (projection onto the plane of the ventral face of the proximal
end). d, anterior. e, proximal.

what away from this edge onto the dorsal surface. The straight, long axis of
11557 is normal whereas the downward curvature of the proximal end in both
12262 and K352 would appear to be due to post-mortem distortion. The internal
and external trochanters are as in the already described Pristerognathids.

K353 consists of an isolated, well-preserved proximal end of a large femur
which may be that of Scymnosaurus major. Although it has an internal trochanter
and pubo-ischio femoralis internus ridge as in the other pristerognathid femora,
both these structures are feebly developed and appear to be out of proportion
to the size of the bone.

Ainnosaurus (fig. 46d and e)

In 12185 a proximal and distal femoral end is preserved. The caput is well
rounded and somewhat anteriorly directed. The internal trochanter forms a

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

robust ridge. The external trochanter is weak and the inter-trochanteric fossa
short but deep.

The pubo-ischio femoralis internus ridge is very weak; it forms the anterior
border of the bone and runs into the anterior edge of the caput.

F, THE LOWER HIND LIMB AND PES
Alopecognathus (figs. 48 and 49a)

In 12051 the left epipodial and pes are completely preserved and in K231
both epipodials and an incomplete pes.

ech d (a

Fic. 45. Femora. x 4. Scymnosaurus sp. S.A.M. 11957. a, ventral. Scymnosaurus sp.
S.A.M. 12118. 5, ventral. Scymnosaurus major. S.A.M. 12262. c, dorsal. d, ventral.
e, anterior.

The tibia is stout proximally with a well-developed facet for the femur.
The fibula is much slenderer and curved to give a wide spatium interosseum.
The proximal facet of the fibula is terminal and not applied to the outer corner
of the femoral condyle but end-on.

The astragalus is a robust, rounded bone with a convex facet for the tibia.

The calcaneum is a large but lightly-built flat bone thickened proximally
at the facet for the fibula. Preaxially it is notched for the passage of the intertarsal
vessels.

The central is quite small.

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 163

There are five distals. The first is large and articulates with both first and
second metatarsals. The fourth is the largest whereas numbers 2, 3 and 5 are
small.

The metatarsals increase in length from number | to 5.

The digital formula is 2, 3, 3, 3, 3, with the fourth the longest, but due
to the short phalanges all the digits are short. The ungual phalanges are curved
and carried sharp claws.

Fic. 46. Femora. x}. Scymnosaurus major ? S.A.M. K352. a, dorsal.

b, ventral. c, anterior. Zinnosaurus paucidens gen. et spec. nov. S.A.M.

12185. d, ventral surface of proximal end. e, anterior face of proximal
end.

a C

Fic. 47. Scymnosaurus major ? S.A.M. K353. x 4. Proximal end of femur. a, dorsal.
b, ventral. c, anterior. d, posterior.

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cynariognathus (fig. 496)

In 11794 a somewhat crushed tibia and fibula are preserved, with features
as shown in the figure. Distal to the tibia lies a bony element of quite intricate

shape, not at all like the astragalus preserved in Alopecognathus and probably
is not this bone at all.

Fic. 48. Epipodial ;

and pes of Alopecog- Fic. 49. Epipodials and pes. x 4.a, Alopecognathus Fic. 50. Pes of a

nathus. S.A.M. — angusticeps. S.A.M. K231. b, Cynariognathus sp. pristerognathid.
12051. X4. S.A.M. 11794. ¢, Scymnosaurus sp. S.A.M. 11459. S.A.M. 245A. x4.

Scymnosaurus (fig. 49¢)

In 11459 part of a pes is preserved. I have identified a small central.
Distals 2 and 3 are small, 4 is much larger, and 5 is still a small, separate
element. Parts of all five metatarsals are present. Number 1 digit has a very
short first phalanx.

DIsCcussION

It would be of interest to compare the girdles and limbs of the pristero-
gnathid Therocephalia with those of the other therapsids of the Tapinocephalus
zone. Unfortunately these structures in the contemporary therapsids are
inadequately known. Only in the Dinocephalia have they been adequately
described, and in the Dromasauria two specimens have these structures

—a—

GIRDLES AND LIMBS OF THE PRISTEROGNATHID THEROCEPHALIA 165

preserved. Of the Gorgonopsia, they have been described in one genus. Of the
Dicynodontia nothing has as yet been published. Of the other therocephalian
families represented in this zone, a shoulder girdle of a lycosuchid has been
figured, of the Akidnognathidae a manus, and of the Scaloposauridae a brief
account has been given of parts of a hind limb.

I have in recent years collected some gorgonopsians and dicynodonts from
the Tapinocephalus zone in which parts of the girdles and limbs are preserved.

As soon as these specimens have been prepared a comparative discussion
will be presented.

SUMMARY

Descriptions are given of the girdles and limbs of the pristerognathid
Therocephalia from the Tapinocephalus zone of the Karoo. The descriptions are
based on 37 specimens in the South African Museum which have parts of the
girdles and limbs preserved. Taken together this material gives a good idea of
the family character of these structures. A new genus and species of pristero-
gnathid therocephalian, Zinnosaurus paucidens, is described.

ACKNOWLEDGEMENTS

The specimens collected in recent years were obtained on Museum
collecting trips, which were in part financed by the C.S.I.R. For these grants
we are grateful.

Some of the later specimens have been prepared for study by Mr. C. Gow,
to whom our thanks are due.

The Trustees of the South African Museum are grateful to the C.S.I.R.
for a grant to publish this paper.

REFERENCES

Broom, R. 1929. On some recent new light on the origin of mammals. Proc. Linn. Soc. N.S.W.
54, 8: 688-694.

Broom, R. 1932. The mammal-like reptiles of South Africa. London: H. F. & G. Witherby.

Hueng, F. vor. 1956. Paldontologie und Phylogenie der niederen Tetrapoden. Jena: Fischer.

SEELEY, H. G. 1888. Researches on the structure, organization and classification of the fossil
Reptilia. III. On parts of the skeleton of a mammal from Triassic rocks of Klipfontein,
Fraserberg, South Africa (Theriodesmus phylarchus, Seeley), illustrating the reptilian inheri-
tance in the mammalian hand. Phil. Trans. R. S. (B) 179: 141-155.

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Ann. S. Afr. Mus., Vol. XLVIII Plate [

A. Photograph of a habitat group with life-sized models modelled by the author and exhibited
in a diorama in the South African Museum. Scymnosaurus feeding on Brachypareia.

STI ree

B. Photograph of life-sized models made by the author and exhibited as a habitat group in the
South African Museum. The Therocephalians (1. to r.) Pristerognathoides, Therioides and Alope-
cognathus feeding on a cadaver of Moschops.

PaaltRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TABLE oF CoNnTENTs and SumMARy. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 4? in. x 7 in. (7$ in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (black type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

Smiru, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmrrH, C. D. 1954. South African plonias. Jn Brown, X. Y. Marine faunas. and ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, Fuly 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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d MUS. COMP. ZOOL
LIBRARY

M.-L. WAPENAAR & F. H. TALBOT MAY 29 1964

HARVARD
UNIVERSITY,

NOTE ON THE RIGIDITY
OF THE PECTORAL FIN OF
MAKAIRA INDICA (CQUVIER)

April 1964 April
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MUS. COMP. ZOOL
LIBRARY

MAY 29 1964

ARVARD
NOTE ON THE RIGIDITY OF THE PECTORAL BIN esis

MAKAIRA INDICA (CUVIER)*

By
Mary-Louise WAPENAAR & FRANK HAMILTON TALBOT
South African Museum
(With 1 plate and 7 figures in the text)

ABSTRACT

It has been known for a long time that the pectoral fin of the black marlin, Makaira indica,
cannot be folded flat against the side of the body as it can in all other istiophorid fishes. The
anatomy of the pectoral girdle of the black marlin is here compared with that of the striped
marlin, M. audax, to determine the mechanism of the rigid joint. The osteology and musculature
of the pectoral girdle of both species is described and discussed, with particular reference to the
articular region. It is concluded that the bony structure of the joint and the strength and
disposition of its fibrous connective tissue sheath together ensure the rigidity of the pectoral fin
in M. indica. It is suggested that the rigid pectoral fin of M. indica is used as a plane of elevation.

I. INTRODUCTION

Ever since Nakamura (1938) and, more particularly, Gregory & Conrad
(1939) showed that the black marlin Makaira indica (Cuvier) has a rigid
pectoral fin which cannot be folded flat against the body, while all other
istiophorids have a folding pectoral fin, the reason for such a difference, and
the structural features that prevent the fin from folding in the black marlin,
have been a source of speculation among ichthyologists.

The pectoral fin of marlins is in the main an inflexible, sickle-shaped
structure with a narrow leading edge formed by the sharp edge of the strong
marginal ray, and a small movable posterior lobe formed by the last ten rays
or actinosts. In all marlins it is set at an angle of about 35° to the horizontal,
so that when extended at right angles to the body it acts as a plane of eleva-
tion by presenting a horizontal or oblique surface to the direction of flow of
the water rather than a vertical surface as in many teleosts. In all istiophorids
other than M. indica the fin can be rotated from the extended position to
lie flat against the side of the body, but in M. indica the fin is permanently
extended and cannot be folded against the body without structural damage,
for which considerable force must be used.

An anatomical study was undertaken in which the pectoral girdle of
M. indica was compared with that of the striped marlin, M. audax (Philippi), to

* This paper formed part of the Oceanography Symposium held during the 61st Annual
Congress of the South African Association for the Advancement of Science at Durban, July 1963.
The association has kindly given permission for publication here.

167

Ann. S. Afr. Mus. 48 (6), 1964, 167-180, pl. II., 7 figs.

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

determine the reason for the rigidity of the pectoral fin in M. indica. The
articular regions of the pectoral girdles of M. albida (Poey) and M. nigricans
(Lacépéde) were also examined.

Nakamura (1938), Gregory & Conrad (1939), LaMonte & Marcy (1941),
LaMonte (1955), J. L. B. Smith (1956), Robins (1957), Robins & de Sylva
(1961), and Talbot & Penrith (1962) and others have referred to the rigidity
of the pectoral fin of M. indica, but only one attempt to investigate this feature
on an anatomical basis has been made (Morrow, 1957).

Morrow states that the rigidity of the fin in M. indica is due to three bony
pads associated with the articular surface of the marginal ray of the fin, giving
it a rigid three-point suspension and thus preventing it from being rotated and
folded back as occurs in the other species. In our opinion this explanation is
untenable, for reasons which are given below.

II. MATERIAL

Makaira indica (Cuvier)

(a) ‘Two loose girdles, fresh, of a 600 lb. fish, length 2,935 mm., taken
aboard the fishing vessel Karimona by longline west of Slangkop, Cape Peninsula,
30/1/1962. S.A.M. Reg. No. 23194.

(b) Anterior half, fresh, of an 800 Ib. fish, length 3,210 mm., taken aboard
the fishing vessel Walvis Pioneer by longline 40 miles west of Cape Point,
28/1/1962. S.A.M. Reg. No. 23193.

(c) Anterior half, fresh, of a 462 lb. fish, length 2,540 mm., taken on rod
and line 25 miles west-north-west of Cape Point, 25/2/1962. S.A.M. Reg. No.
23244.

(d) Prepared girdle of a 1,028 lb. fish taken by longline south-west of
Hout Bay, March 1961. S.A.M. Reg. No. 23054.

Makara audax (Philippi)

' (a) Anterior half, fresh, of a 130 lb. fish, length 2,120 mm., taken aboard
the fishing vessel Overberg by longline west of Cape Point, 2/2/1962. S.A.M.
Reg. No. 23197.

(b) Prepared girdle of small specimen taken west of Hout Bay by longline,
March 1961. S.A.M. Reg. No. 23052.

Makara nigricans (Lacépéde)

(a) Prepared girdle of large specimen taken aboard the fishing vessel
Cape Point by longline 45 miles north-west of Dassen Island, 29/6/1961. S.A.M.
Reg. No. 23104.

Makaira albida (Poey)
(a) Prepared girdle of specimen taken by longline south-west of Hout
Bay, March 1961. S.A.M. Reg. No. 23053.

RIGIDITY OF PECTORAL FIN OF MAKAIRA INDICA (CUVIER) 169

III. OsTEOLOGY OF THE PECTORAL GIRDLE OF JM. audax AND M. indica

The pectoral girdle of marlins (figs. 1, 2) is suspended from the skull by
a three-pronged post-temporal and a long, flat supra-cleithrum, and is typical
in consisting of a complex of three bones. There is an anterior cleithrum which
has medial and lateral flanges or arms, a rod-like ventral process which meets
that of the opposite side in the mid-ventral line, and two dorsal processes, an
anterior, rod-like one and a posterior expanded process. The posterior bone
of the complex is the coracoid, which is roughly triangular. From its antero-
dorsal corner a ridge runs ventro-caudally. Above the ridge is the dorsal
process of the coracoid; the ventral process of the coracoid meets the cleithrum
just above the ventral process of that bone. Dorso-medially between the
cleithrum and the coracoid lies the scapula, a fairly small semicircular bone
perforated by the scapular foramen, through which the nerves supplying the
abductor musculature of the fin pass. The scapula bears the articular surfaces
for the pectoral fin.

No bony pads such as those described by Morrow (1957) were found in
any of the South African specimens of M. indica. If they are not present in all
specimens it seems unlikely that those found by Morrow are of any significance
in the mechanism of the rigid joint.

The pectoral complex is very similar in M. indica and M. audax, with a
major difference in the articulation of the fin (described below in section V),

anterior dorsal process
of cleithrum

posterior dorsal process
of cleithrum

articular surface for
Bb . ; marginal ray

rsal proces:
of coracoid

* cleithrum

scapula

, dorsal process of coracoid

scapular foramen

coracold ridge

medial arm of cleithrum

lateral arm of cleithrum

ventral process
of cleithrum

yentral process of coracoid

Medial view Lateral view :

Fic. 1. Pectoral girdle of Makaira indica.

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

anterior dorsal
process of cleithrum

posterior dorsal process
of cleithrurm

articular surface for

marginal ray *» clelthrum articular surface (i
urface for

marginal ray

scapula ,
coracold ridge
er dorsal process of coracoid
‘Re

dorsal process of coracold scapular foramen

medial arm of cleithrum

lateral arm of cleithrum
coracold ”

- ventral process of
cleithrum

t vanieal prerate of
coracol

Fic. 2. Pectoral girdle of Makaira audax.

and other minor differences. In M. audax the anterior edge of the cleithrum is
concave, while in M. indica this edge is almost straight, with the ventral process
of the cleithrum somewhat backwardly directed. The coracoid ridge is shorter
and heavier in M. indica than in M. audax, and the dorsal process of the coracoid
is pointed in the former and rounded in the latter. The scapular foramen is
relatively larger in M. audax than in M. indica.

~ “eee, » » ie

IV. MuscuLATURE OF THE PECTORAL GIRDLE OF JM. indica AND M. audax
(figs. 3, 4, 5)

The musculature of the pectoral girdle of M. indica and M. audax was
investigated. The body muscles and the muscles between the pectoral girdle ©
and the head and visceral skeleton were found to be identical in M. indica
and M. audax and will not be included here.

The musculature of the pectoral girdle consists as in all fishes of adductor :
and abductor portions (Shann, 1919), which draw the fin towards and away
from the body respectively. Both portions are more or less divided into superficial
and deep parts, although these are often not easily separable. The abductor
musculature arises from the lateral surface of the scapula and the coracoid,
and the lateral surface of the medial flange of the cleithrum, and inserts on
the marginal ray and the bases of the remaining rays of the fin. For most of
their length from their fleshy origin the superficial and deep fibres are not clearly
separable, but towards their insertion they become more distinct and have a
separate tendinous insertion on the bases of the rays, the deep fibres inserting
on the ventral extremities of the bases of the rays while the superficial fibres
insert on a slight ridge a short distance above the bases of the rays. The last
ten rays are movable in relation to the rest of the fin and have a fairly specialized
musculature, some of which inserts on the radials supporting them (described
below, section V). The adductor musculature arises from the medial surface

marginal ray

cleithrum

fateral arm of cleithrum

coracoid

Superficialis

marginal ray

cleithrum

lateral arm of cleithrum

coracoid

Profundus

Fic. 3. Abductor musculature of Makaira indica.

ventral lobe of fin

ventral lobe of fin

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the cleithrum, scapula, and coracoid, and also has a tendinous insertion
on the bases of the rays on the medial side of the fin. As in the case of the abduc-
tor musculature, the adductors of the last ten rays are separate and very well -
developed. An additional adductor, present in many fishes, the coracoradialis

marginal ray

cleithrum

ventral lobe of fin

coraco-radialis muscle

coracold

Superficialis

marginal ray

scapula

cleithrum
* ventral lobe of fin

Profundus

Fic. 4. Adductor musculature of Makaira audax.

RIGIDITY OF PECTORAL FIN OF MAKAIRA INDICA (CUVIER) 173

Marginal ray

cleithrum

ventral lobe of fin

coraco-radialis muscle

coracoid

Fic. 5. Adductor superficialis musculature of Makaira indica.

muscle, arises from the medial surface of the coracoid and inserts on the last
two radials.

In general the abductor and adductor portions are very similar in M.
indica and M. audax (figs. 3, 4, 5). In M. indica the superficial and the deep
abductor of the marginal ray form a bundle clearly separated from the abductors
of the remaining fin rays, while in /. audax the abductors for the marginal ray
are not distinct. In both species the last ten rays form a distinct lobe witha
separate and well-defined musculature; the abductor portions of the muscula-
ture of this lobe are identical in the two species.

There are certain differences in the adductor musculature of the two
species (figs. 4, 5); again in M. indica the adductors of the marginal ray are
clearly defined and in addition have a slightly different origin, arising from
the posterior dorsal process of the cleithrum rather than from the medial arm
of the cleithrum. In general the adductor muscles in M. audax are arranged
so that the fibres are relatively longer than in M. indica. The greatest difference
is in the adductor musculature of the ventral lobe, with which the present

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

discussion is not concerned as it has no bearing on the mechanism of the rigid
joint. This lobe clearly has important hydrodynamic effects on the fin, however,
and one would expect its mode of action to differ in two species in which the
action of the fin as a whole differs strongly.

The adductor and abductor muscles then are responsible for the movements
of the pectoral fin. By contraction of the abductors other than those of the
ventral lobe the antero-ventral tip of the fin-base is pulled forward and down-
ward. In M. audax (pl. II) this movement causes the fin to be pulled away from
the body into the extended position. This actually involves two movements, the
drawing downwards of the antero-ventral tip of the fin base and the drawing
forwards of the fin as a whole so that the direction of the fin tip is at right angles
to the direction of the body, but both movements occur simultaneously in
M. audax. In the extended position in M. audax either movement can be
carried out; by twisting, the plane of the fin to the water can be altered by
about 10°, and the leading edge can move through any arc between the fully
extended position and the folded position.

Fic. 6. Range of movement of fin in Makatra indica.

In M. indica (fig. 6), where the fin is permanently extended, two limited
movements are possible, and are also brought about by the abductor and
adductor muscles. The angle of incidence of the fin plane to the water can be
altered within an arc of about 4°, brought about by the pulling down by the
abductors of the antero-ventral tip of the marginal ray base; the fin is also
capable of an antero-posterior movement in a horizontal plane through an

ee ee

ce at a

RIGIDITY OF PECTORAL FIN OF MAKAITRA INDICA (CUVIER) 175

angle of 12°. It can thus be seen that the range of movement is far more limited
in M. indica than in M. audax.

It would be anatomically unusual if the rigidity of the fin were dependent
on the musculature. This would imply continual sustained muscle activity to
hold the fin rigid during forward movement of the fish; that is, during most
of its life. The total extension of a muscle group is never limited by its own
action alone (Professor L. H. Wells, personal communication). The maximum
extension is normally limited by the structure (ligament, cartilage and bone) of
the joint itself. If they were not limited in this way, the structure of muscle
fibres is such that on relaxation they could expand until damaged. If muscles
held the fin rigid in the black marlin it would also follow that in a freshly
dead specimen the fin should be able to move back against the body, even if
the muscle fibres were damaged in so doing. This is not the case. As would be
expected, therefore, the structure of the joint itself and not the muscles is the
limiting factor to further backward movement. In any joint the arrangement
of the muscles will depend on the amount of movement allowed by the arrange-
ment of the bones and ligaments of the joint. Thus in M. audax, where the
range of possible movement of the fin is more extensive, the muscle fibres are
longer than in M. indica.

The fin musculature of M. indica is very well developed in spite of the small
range of movement of the fin. It is suggested that while the fin of MZ. audax
encounters little water resistance in the initial stages of its extension, water
pressures render considerable muscular effort necessary for moving the fin
when it is extended. It is these latter movements with which the muscles of
M. indica are concerned, so that they must be well developed, as well developed
as in M. audax, in fact. Short fibres are adequate for these short-range
movements, however.

The main point arising from the study of the pectoral musculature is that,
if the abductors of M. audax are contracted, the pectoral fin can be maintained
in the same position as that of MW. indica, but in M. indica the fin is maintained
in this position even when the abductor muscles are relaxed and the adductors
contracted. The muscles do not assist in rendering the joint rigid; as would be
expected, M. indica has developed some method of maintaining the rigid
position of the fin other than by sustained muscular contraction.

V. ARTICULATION OF THE PECTORAL FIN (fig. 7)

The most significant difference in the pectoral complex lies in the arrange-
ment and shape of the articular surfaces for the pectoral fin. In M. audax,
M. nigricans, and M. albida the articular surface for the base of the marginal ray
lies on the dorsal edge of the scapula; its surface is markedly convex and curves
smoothly from the lateral to the medial surface of the bone; in other words it
is saddle-shaped. The base of the marginal ray in these species is concave to
correspond with its articular surface on the girdle. In M. indica the articular

I 76 ANNALS OF THE SOUTH AFRICAN MUSEUM

surface for the marginal ray, although close to the dorsal edge of the scapula,
lies entirely on the lateral surface and is flat, so that a limited amount of
sliding but no rolling movement is possible. The base of the marginal ray in
this species is correspondingly flat. The dorsal edge of the scapula above the
articular surface (which surface is occupied by the inner half of the saddle-
shaped articular surface in M. audax) is here extremely rugose and pitted for
the attachment of connective tissue.

In all four species there is at the posterior end of the articular surface for
the marginal ray a shallow trough, which receives a downward process of the
articular surface of the marginal ray base. This trough lies mainly on the dorsal
edge of the scapula.

Posterior to the articular surface for the marginal ray are the articular
surfaces for the radials (fig. 7). The radials are very similar in all four species.
They are four in number, the first two being short and cubical, and the posterior
two rather long and slender and forming the base of the posterior lobe of the fin.
Their articulation lies mainly on the dorsal edge of the scapula; that of the

anterior dorsal
process of cleithrum

marginal ray

posterior dorsal process
of cleithrum

articular surface

scapula
Y_ articular surface for ;

Zz marginal ray radials

bases of fin rays

dorsal process of coracoid

coracoid ridge
lateral arm of cleithrum

medial arm of cleithrum

Fic. 7A. Articular region of pectoral girdle of Makaira indica.

RIGIDITY OF PECTORAL FIN OF MAKAIRA INDICA (CUVIER) 177

anterior dorsal process posterior dorsal process
of cleithrum of cleithrum

Marginal ray

articular surface

articular surface for
marginal ray

scapula
bases of fin rays

radials

dorsal process of coracoid

lateral arm of cleithrum coracoid ridge

medial arm of cleithrum

Fic. 7B. Articular region of pectoral girdle of Makaira audax.

first two is an area of attachment rather than articulation, most movement
taking place between their distal ends and the bases of the rays. The third and
fourth radials are slightly more movable, and movement is possible between
the second and third radials in all species. The distal ends of the first two radials
are smoothly curved in M. audax, allowing free movement of the rays over them,
but are strongly rugose and pitted in M. indica for the attachment of connective
tissue. The last ten rays are movably articulated on the distal ends of the third
and fourth radials in both species.

In both M. audax and M. indica the pectoral fin is held to its articular
surfaces by a connective tissue sheath, but this sheath shows significant differ-
ences in the two species. In M. audax the ligaments are elastic and loosely
arranged so as to allow maximum movement of the joint, but in M. indica a very
strong sheath of interwoven fibrous tissue is developed which holds the fin
strongly to its articular surfaces; the fibres are short and their area of attach-
ment to the bone is more extensive than in M. audax.

178 ANNALS OF THE SOUTH AFRICAN MUSEUM
VI. Discussion

No features of the osteology of the pectoral girdle other than the articular
region suggested a mechanism for maintaining the rigidity of the fin. The muscu-
lar system, although showing differences connected with the movements
carried out by the fin in the two species compared, is not adapted to hold the fin
away from the body in M. indica.

It is suggested that the only difference between the pectoral girdles in
M. indica and the other species studied which is large enough to be of significance
in the functioning of the joint of the pectoral fin lies in the position and confor-
mation of the articular surfaces of the fin, in particular that for the marginal
ray, and in the development of the connective tissue of the joint.

Owing to the flat, lateral articular surface for the marginal ray base of the
fin of M. indica, the fin cannot roll back so that its base rests on the dorsal edge
_of the scapular, without leaving its articular surface; to prevent it from being
forced off its articular surface by the pressures it encounters in the extended
position, a very strong connective tissue sheath is developed around the joint.
This tough connecting sheath prevents the fin from lying back against the body.

If the fin is forced back against the body in a dead specimen, an operation
requiring considerable force, the connective tissue sheath is torn. The bones,
including the radials, are undamaged, suggesting that there can be no bony
locking or strutting device for maintaining the rigidity of the fin.

After the ligaments have been broken the fin can fold back considerably
farther than before, but not completely as in the other species; it is stopped by
the dorsal expansion of the marginal ray being jammed against the posterior
edge of the posterior dorsal process of the cleithrum, which is slightly thickened
in this species. Unless the ligamentous sheath is torn this position is not reached,
so this is not a mechanism for holding the pectoral fin rigid.

It is suggested that at some time and for some reason in its evolutionary
history M. indica or its ancestors found it necessary to maintain the fin in a
laterally extended position. This was presumably accomplished at first by
muscular contraction sustained over long periods, but in time the tension was
taken by greatly strengthening the connective tissue sheath attaching the fin to
the girdle, and the inner portion of the girdle’s articular surface, which is used
only when the fin lies against the side, was lost.

The pectoral fin of the black marlin appears to act either as a stabilizer,
or as a plane of elevation. The body is very large and deep, and it may be that
some stabilizing factor is necessary during forward movement. The broadbill
swordfish, Xiphias gladius, of similar body shape, also has rigid pectoral fins.
However, if this is the reason for the modification, it is surprising that the blue
marlin, Makaira nigricans, also a large, deep-bodied marlin, does not have rigid
pectoral fins.

The other possibility, that the fin acts as a plane of elevation during forward
movement, deserves consideration.

RIGIDITY OF PECTORAL FIN OF MAKAIRA INDICA (CUVIER) 179

It is interesting to speculate on the possible reasons for the necessity of such
continuous lifting force. In pelagic surface teleosts very small marine species
usually possess a well-developed closed swim bladder, but among many medium
and large sized species, particularly in the Scombridae, it is variable or absent
(Jones & Marshall, 1953). For example, in many species of the genus Thunnus, it
is variably reduced or rudimentary; in Katsuwonus pelamis, Sarda lineolata,
S. chiliensis, the genus Auxis, the genus Euthynnus and most species of the genus
Scomberomorus, it is absent. It is probable that in fast-swimming forms which
change depth rapidly a large swim bladder is a liability because of its necessarily
slow change of volume. The absence or reduction of the swim bladder would
necessitate some upward thrust to counteract the tendency to sink because of
increased density. In such powerfully swimming fishes as the tunas and the
marlins where forward movement may be continuous this presents little
difficulty, and presumably the increased effort has not been too great to offset
the advantage gained in vertical manoeuvrability.

Preliminary examination (dissection of one black marlin and one striped
marlin) showed that the swim bladder structure is very different between the
two species, and that the black marlin seems to have a relatively smaller swim
bladder. This work is being continued.

It is therefore tentatively suggested that the black marlin has a reduced
swim bladder, and that some upward thrust is supplied by the rigid pectoral fins.

Comparison may be made here with the sharks. In this group the pelagic
surface forms are large, and in the absence of a swim bladder upward thrust
is obtained by the broad and rigid pectorals.

ACKNOWLEDGEMENTS

We are indebted to the manager and skippers of the Atlantic Tuna
Corporation and to Mr. F. Slack and Mr. W. Gilmore of the South African
Marlin & Tuna Club for the specimens; to Dr. N. A. H. Millard of the Zoology
Department of the University of Cape Town and Mr. M. J. Penrith of the
South African Museum for advice and help; to Professor L. H. Wells of the
Department of Anatomy, University of Cape Town, for advice on joints; and
to all those who helped with the conveying and handling of the material.

The Trustees of the South African Museum gratefully acknowledge the
grant-in-aid towards the cost of publishing this paper made by the Council
for Scientific and Industrial Research.

REFERENCES

Grecory, W. K., & Conran, G. M. 1939. Body forms of the black marlin and the striped
marlin of New Zealand and Australia. Bull. Amer. Mus. nat. Hist. 76: 443-456.

Jones, F. R. H., & MarsHatt, N. B. 1953. The structure and functions of the teleostean swim-
bladder. Biol. Rev. 28: 16-83.

LaMonte, F. R. 1955. A review and revision of the marlins, genus Makaira. Bull. Amer. Mus.
nat. Hist. 107: 323-365.

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

LaMonte, F. R., & Marcy, D. E. 1941. Swordfish, sailfish, marlin, and spearfish. Ichthyol.
Contr. internat. Game Fish Assoc. 1 (2): 1-24. (Not seen. Information from Morrow, 1957.)

Morrow, J. E. 1957. On the morphology of the pectoral girdle in the genus Makaira. Bull.
Bingham oceanogr. Coll. 16 (2): 88-105.

NakaAmurA, H. 1938. Report of an investigation of the spear-fishes of Formosan waters. U.S.
Fish & Wildlife Serv., Spec. Sci. Rpt. Fisheries, 153, 1955 : 1-46.

Rosins, C. R. 1957. Charles F. Johnson Oceanic Gamefish Investigations, Progress Rpt. 4.

Rosins, C. R., & De Syiva, D. P. 1961. Description and relationships of the longbill spearfish,
Tetrapturus belone, based on western North Atlantic specimens. Bull. Mar. Sci. Gulf and
Caribbean. 10 (4): 383-413.

SHann, E. W. 1919. The comparative myology of the shoulder girdle and pectoral fin of fishes.
Trans. Roy. Soc. Edinb. 52: 531-570.

Suir, J. L. B. 1956. Swordfish, marlin and sailfish in South and East Africa. Ichthyol. Bull. 2,
25-33:

Tatsor, F. H. & PenrirH, M. J. 1962. Tunnies and marlins of South Africa. Nature. 193,
no. 4815: 558-559.

Ann. S. Afr. Mus., Vol. XLVIII Plate II

(6b) Extended position.

Range of movement in Makaira audax.

ee eee
we we —— — ee ee ee

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TABLE oF CoNTENTs and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. x 7 in. (7$ in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (black type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

SmiTH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

Smiru, C. D. 1954. South African plonias. Jn Brown, X. Y. Marine faunas. and ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, Fuly 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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A NEW SPECIES OF FLAT FISH,
MANCOPSETTA MILFORDI, FROM SOUTH AFRICA,
WITH NOTES ON THE GENUS MANCOPSETTA

September 1965 September
Volume 48 Band
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Cape Town Kaapstad

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A NEW SPECIES OF FLATFISH, MANCOPSETTA MILFORDI, FROM

SOUTH AFRICA, WITH NOTES ON THE GENUS MANCOPSETTA
By

M. J. PENRITH*

South African Museum, Cape Town

(With 2 figures in the text and 1 plate)

CONTENTS agi ee

PAGE
Introduction . : F : 181
Mancopsetta milfordisp.n. . Specie ts
Generic grouping : : nan oae ol
Genus Mancopsetta . y oi 166
Distribution . ; ; ; 187
Acknowledgements. : OS tei:
Summary : . : : 188
References : ; ‘ ters

INTRODUCTION

The genus Mancopsetia was proposed by Gill (1881) to replace the pre-
occupied generic name Lepidopsetta used by Gunther (1880) when describing
M. maculata, the first of the bothid fishes found which lacked pectoral fins. This
“species has until now remained the only species in the genus and has been
_ known from a very few specimens found in the Southern Hemisphere. The
related genus Achiropsetta (Norman, 1930) was known from two species, both
‘rare and recorded only from the Southern Hemisphere. Recently the South
African Museum was presented with two specimens of a bothid fish lacking
pectoral fins. These have been assigned to the genus Mancopsetia.

Mancopsetta milfordi sp. n.
(fig. 1, pl. III)

Material :
S.A.M. 24041: trawled by a commercial trawler in 300 fathoms west of Cape

Town. Standard length 393 mm. (Holotype).
S.A.M. 23767: trawled by a commercial trawler in 370 fathoms west of Cape

Town. Standard length 298 mm.

The body is compressed, sinistral and fairly elongate, the maximum depth
being about 22 in body length (excluding the caudal fin). The axial portion
of the body is thickened relative to the rest of the body in the smaller specimen,

* Seconded from the Council for Scientific and Industrial Research Oceanographic Unit,
University of Cape Town. |

181

a ‘
Ann. S. Afr. Mus. 48 (7), 1964, 181-188, 2 figs., 1 pl.

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182 ANNALS OF THE SOUTH AFRICAN MUSEUM

but this seems to become less marked with increase in size as it is not noticeable
in the larger specimen. The caudal peduncle is very short, being only 4 of the
longitudinal diameter of the eye.

There is a single straight lateral line on both sides with about 160 scales
on the ocular side. The scales are ctenoid with normally eight to twelve spinules.
Towards the edges of the fish, especially on or near the fins, the spinules project
at an angle away from the body, but on the main trunk they lie flat. The entire
head, including the jaws and part of the eyes, is covered with fine scales. The
eyes are separated by a low, scaled, bony ridge. All fin rays, including the
caudal, have a row of scales.

The head from the tip of the maxilla to the hindermost edge of the oper-
culum is contained 4 times in standard length, while the greatest diameters
of the eyes (both eyes equal) are 4 the length of the head. The snout is slightly
shorter than the eye diameter. There is a small fleshy growth over the front
of the maxilla. The mouth reaches back to a point almost level with the posterior
edge of the lower eye. Teeth are present on both sides of the jaw but are
slightly better developed on the ocular side. The mouth has a very wide gape
and is protrusible. The membranes on the ocular side are more protrusible
than those on the blind side. There are nine small gill-rakers on the lower arch
of the first gill.

There is no trace of any pectoral fin on either side. The pelvic fin on the
ocular side is larger than that on the blind side. On the eyed side the anterior
base of the pelvic fin is median, while posteriorly the base is twisted to the left,
i.e. on to the ocular surface of the body (fig. 1).

The body proportions and meristic counts of the two specimens are given
in Table I and compared with a specimen of M. maculata from the collection
of the British Museum (Natural History) (Registered No. 1930.5.6: 41).

The colour, preserved in formalin, is brown-grey with the anal and dorsal
fins grey slate/blue. There is no trace of any pattern.

TABLE I

M. milfordi M. milfordi M. maculata
Total length .. wif an << 456 334. - 239
Standard length a Me a 392 298 199
Greatest depth .. a bi ak 143 (364) - .g9'(392) 88 (442)
Head length .. a On ee IOI (258) 76 (255) 53 (266)
Length of snout : : oe 22 (56) 15 (50) 10 (50)
Diameter of eye (hanconn ie ihe 24 (61) 20 (67) 17 (85)
Interorbital width aa ae 54: 8 (17) 5 (15) 3 (15)
Length of Maxillary bs ne 48 (122) 33 (111) 18 (go)
Number of scales in lateral lige es 164 P77 119
‘Dorsal rays... ve ae as 130 133 119
Anal rays fe nd at 109 110 98 ~
Pelvic rays, blind side .. us ste 6 6 5
Pelvic rays, ocular side bite Be G f) ri
Gill-rakers, lower arch ae se 9 9. 12

Body proportions and meristic counts of M. milfordi and M. maculata. Body proportions in
millimetres and in thousandths of standard length in parentheses.

A NEW SPECIES OF FLATFISH, MANCOPSETTA MILFORDI 183

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2p
~ 23>s toys ao:

Ficure 1. Head of M. milfordi showing the twisting of the posterior portion of the left pelvic fin
onto the ocular surface.

Mancopsetia milfordi differs from M. maculata in the number of dorsal and
anal rays and lateral line scales, the twisting of the posterior end of the left
pelvic fin onto the ocular surface and in certain body proportions. It resembles
M. maculata, however, in general body shape, especially head shape, and in the
type of scales.

The species is named for Mr. C. S. Milford, managing director of the
trawling firm, Messrs. Irvin and Johnson (Pty.) Ltd., in recognition of generous
support for marine biological research.

GENERIC GROUPING

According to Norman (1934) there are only two genera of bothids (both
known only from the Southern Hemisphere) which lack both pectoral fins.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

These two genera, Mancopsetta and Achiropsetia are said to differ in the following
characters. .:.’.

(a) Achiropsetta is said to have a flat scaled interorbital space, whereas
that of Mancopsetta is stated to be a naked narrow bony ridge.

(b) Achiropsetta is said to be more elongate and to have a thickened trunk.

(c) In Mancopsetta the pelvic fin on the ocular side is not directed to the
left posteriorly as is said to be the case in Achiropsetta.

(d) In both genera the scales are small and ctenoid, but in Mancopsetta
the 8-14 spinules lie flat, whereas in Achiropsetta there are only one
or two spinules which are directed away from the body.

These differences would seem to be clear cut, but the finding of a second
species of Achiropsetta, A. slavae (Andriashev, 1960) and the present species have
suggested that the differences between the genera are not so marked as was
formerly thought. Neither A. slavae nor M. milfordi fits completely the generic
description of the genus in which it has been placed. The distribution of various
taxonomic characters among the four species of bothids which lack pectoral
fins is shown in Table II below.

TABLE II
Achiropsetta Mancopsetta

tricholepis slavae maculata milfordi
Dorsal rays bi At Me 2 132 114-117 116-119 130-133
Anal rays Le a ” 114 93-96 97-98 109-110
Lateral line scales : ae et ? 115 114-120 164-177
Depth in standard ieceie ar ar 2°4 2 a2 a |
Gill-rakers in lower arch ae ie 9-10 12 12 9
Head in standard length e) a 4°5 ae, 3°8 4
Coloration A Ae Ae plain patterned patterned plain
Fleshy lip on maxilla an ve {<) ) Hpresent present present present
Spinules on scales é say, > wertieal vertical flat flat
Pelvic fin of eyed side poercd ra the left yes no no yes
Maximum known total length (mm.) 105 63 270 456
Smallest known total length (mm.) .. 100 48 134 334

Comparison of the four known species of bothid fishes which lack pectoral fins.

From the table it is clear that the only important character that the two
species of Achiropsetia have in common, and distinct from the two species
placed in the genus Mancopsetia, is the unusual form of the scales with vertically
projecting spinules.

Although A. tricholepis is stated to have a flat scaled interorbital space
(Norman, 1930, 1934), a specimen examined from the collection in the British
Museum (1933.5.8: 1) shows a clear bony ridge. M. maculata on the other hand
was said (Norman, 1934) to have a bony ridge without scales. In the specimen
of M. maculata examined however, scale pockets were clearly present on the
ridge. It seems likely that the scales were rubbed off in the trawl net. A scaled
ridge is therefore present in M. maculata, M. milfordi and A. tricholepis. Nothing
is known concerning this feature in A. slavae.

A NEW SPECIES OF FLATFISH, MANCOPSETTA MILFORDI 185

Another difference, the thickened trunk in Achiropsetta, is most probably
a function of size as it is present in Achiropsetta (of which only small specimens
are known) and the smaller of the two M. milfordi but not in the larger nor in
M. maculata.

From the illustrations in Norman (1934) there is a strong suggestion of
marked differences in the shape of the head in M. maculata and A. tricholepis
but this is caused by illustrating the largest available example of the former
species and the smallest of the latter. Figure 2 gives some idea of the changes of
head shape that may occur in a species with an increase in size.

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Ficure 2. Head shapes of various specimens of the genus Mancopsetta redrawn from various
sources (not to scale).

A. M. maculata of 238 mm. total length (After Norman, 1934).

B. M. maculata of 134 mm. total length (After Gunther, 1880).

C. M. milfordi of 334 mm. total length.

D. M. slavae of 63 mm. total length (After Andriashev, 1960).

E. M. slavae of 56 mm. total length (After Andriashev, 1960).

F. M. tricholepis of 100 mm. total length (After Norman, 1930).

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

From the above it is evident that there is only one important and constant
difference between the two genera Mancopsetta and Achiropsetta, viz. the type
of spinules present on the scales. At the same time, however, there are several
clear points of similarity between A. tricholepis and M. milfordi and again
between A. slavae and M. maculata.

It is extremely probable that the presence of vertically projecting spines
in Achiropsetta is a larval or juvenile character as similar spines occur in many
bothid larvae (Kyle, 1913). Andriashev (1960) holds a similar view and has
stated ‘It is very probable that the peculiar prickliness of the body of Achiropsetta
(that is, the strongly developed, vertically projecting spines) can be considered
as a larval character of a caenogenetic nature’. He also noted the similarity in
fin counts between A. slavae and M. maculata, as well as the fact that these two
species had the most similar distribution. He concluded, however, that the
specimens of A. slavae could not be assigned to the genus Mancopsetta since in the
latter genus the body is not transparent and the scales do not have the vertically
projecting spinules. Andriashev further states that it is not possible to explain
these in terms of development, as the difference in size between the smallest
M. maculata and the largest A. tricholepis is only 29 mm. The greatest similarities
are, however, between M. milfordi and A. tricholepis, where the minimum
difference between the two species is 229 mm., and between A. slavae and
M. maculata, where the difference is 71 mm.

_ While it is not possible at this stage in our knowledge of these fishes to
suggest that A. tricholepis is actually the juvenile stage of M. milfordi or A. slavae
that of M. maculata, it is suggested that for the reasons given above, and bearing
in mind that these four species, in lacking pectoral fins, form a unique group
within the family Bothidae, the species should not be separated into two genera
but should rather be united into one genus, Mancopsetta. The differences
between the species of the former genus Achiropsetta and those of Mancopsetia
being far more likely due to growth rather than major genetic differences.
The generic definition of Mancopsetta will, however, have to be modified from
that given by Norman (1934), not only to include the species formerly placed
in the genus Achiropsetta, but also the new species M. milfordi.

Genus MANCOPSETTA

Lepidopsetta (non Gill, 1864) Gunther, 1880, p. 18.
Mancopsetta Gill, 1881. p. 373.
Achiropsetta Norman, 1930. p. 361.

Body fairly elongate, compressed and sinistral. Eyes large, close together
and with the upper parts of each eyeball densely scaled. Lower eye a little in
advance of the upper. Mouth of moderate size not reaching farther back
than a point level with the posterior edge of the lower eye, maxillary narrow,
not expanded behind. Jaws and teeth about equally developed on both sides.
A small fleshy growth over the front tip of maxilla. Teeth small, conical and
usually uniserial; vomer toothless. Few gill-rakers, those present being short.

A NEW SPECIES OF FLATFISH, MANCOPSETTA MILFORDI 187

Dorsal fin starting far forward on head. All fin rays simple, scaled. Pelvic fin
of eyed side may or may not be posteriorly deflected to the left. No true pectoral
fins but a fin bud may be present in small specimens. Lateral line on both sides
single, straight and well developed.

Type species Lepidopsetia maculata Gunther, 1880.
The genus contains four species:

M. maculata (Gunther) 1880.

M. tricholepis (Norman) 1930.

M. slavae (Andriashev) 1960.

M. milfordi sp. n.

DIsTRIBUTION

Andriashev (1955) has discussed the distribution of these fishes and has
shown that they are known from a few specimens found over a wide area of
the Southern ocean. The following are the only published locality records:

M. maculata. This species is known from three specimens, all caught by
bottom trawling in deep water —

Marion Island: 46° 43’ S. 38° 5’ E. 580 m. (Gunther, 1880).

Falkland Islands: 45° 45’ S. 59° 35’ W. 311-247 m. (Norman, 1937).

South Georgia: 53° 48’ S. 35° 57’ W. 411-401 m. (Norman, 1938).

M. tricholepis. This species is known from only two examples taken by
bottom trawling in shallow water—

Tierra del Fuego: 53° 00’ S. 68° 06’ W. 22-20 m. (Norman, 1930).
Falkland Islands: (No details available) (Norman, 1937).

M. slavae. Four specimens of this species are known and all were caught in
midwater using an Isaacs-Kidd midwater trawl—

Wilkes Land: 64° 51’ S. 114° 17’ E. 725 m. cable (Andriashev, 1960).

(Antarctic) 63° 50’ S. 116° 25’ E. 1,100 m. cable (Andriashev, 1960).
64° 22’ S. 121° 08’ E. 1,100 m. cable (Andriashev, 1960).
64° 10’ S. 127° 46’ E. 1,100 m. cable (Andriashev, 1960).

M. milfordi. Known from only two specimens both taken by bottom trawl
in deep water —

Cape of Good Hope, west of Cape Town. 550 m.

Cape of Good Hope: 33° 40’ S. 17° 30’ E. 675 m.

Some of these records are very widespread, especially M. maculata and
suggest that these fishes are more common than the small number of known
records would suggest. This is undoubtedly due to the very small amount of
trawling that has been undertaken in the Southern ocean. More collecting would
be very useful, as a good set of growth stages would certainly show whether
M. slavae and M. maculata are in fact growth stages of the same species, and
likewise with M. milfordi and M. tricholepis.

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

ACKNOWLEDGEMENTS

I am grateful to Professor J. L. B. Smith of Grahamstown, Dr. A. P.
Andriashev of Leningrad, and Mr. P. R. Sloan of La Jolla for help in obtaining
literature; to the Trustees of the British Museum (Natural History) for the
loan of valuable material for comparative purposes; to Messrs. Irvin and
Johnson (Pty.) Ltd. for presenting these two interesting specimens to the
South African Museum; and to Dr. F. H. Talbot and my wife, Mary-Louise
Penrith, for reading the manuscript.

The Trustees of the South African Museum are grateful to the Council
for Scientific and Industrial Research for the award of a grant to publish this

paper.
SUMMARY

A new species of bothid, Mancopsetta milford: (Pisces: Bothidae), is described
from deep water off the Cape of Good Hope. Four species of bothids are known
which lack pectoral fins. They have been placed in two genera, Mancopsetta
and Achiropsetta. It is shown that the two genera exhibit a marked overlap in
characters and can be divided on only one character, which is probably a
juvenile feature. The genus Mancopsetta is redefined to include all four species.

REFERENCES

ANDRIASHEV, A. P. 1960. Families of fishes new to the Antarctic. 3. Pelagic young of flatfish
(Pisces, Bothidae) off the Antarctic coast. Zool. Zh. 39: 1056-1061.

GiLL, T. 1881. Account of recent progress in Zoology. Rep. Smithson. Instu, 1880: 331-390.

GUNTHER, A. 1880. Report on the shore fishes. Rep. Voy. Challenger 1873-6 1(6): 1-82.

Kye, H. M. 1913. Flatfishes (Heterosomata). Rep. Danish oceanogr. Exp. medit. 1908-10 2 (At).

Norman, J. R. 1930. Oceanic fishes and flatfishes collected in 1925-27. Discovery Rep. 2: 263-370.

NNorMAN, J. R. 1934. A Systematic monograph of the flatfishes Heterosomata I. London, British Museum
(Nat. Hist.).

Norman, J. R. 1937. The coast fishes II. The Patagonian region. Discovery Rep. 16: 3-150.

Norman, J. R. 1938. The coast fishes III. The Antarctic zone. Discovery Rep. 18: 3-104.

Plate II

Ann. S. Afr. Mus., Vol. XLVIII

adA}0[0oH]

Ipsofjiu vyjasqGoounyy

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TABLE OF CONTENTS and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. x 7 in. (74 in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers)
first and last, preceded by a colon (without prefix ‘p’). Thus:

SmitH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmiTH, C. D. 1954. South African Plonias. In Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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FRANK HAMILTON TALBOT AND MARY-LOUISE PENRITEHY
NOV 19 t¥oa

HARVARD

CTENOGOBIUS CLOATUS SMITH, 1666YERSITy
A SYNONYM OF CTENOGOBIUS SALDANHA
(BARNARD, 1927)

September 1965 September
Volume 48 Band
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ANNALS OF THE SOUTH AFRICAN MUSEUM

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Cape Town Kaapstad

P<

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
(Cash with order, post free)

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CTENOGOBIUS CLOATUS SMITH, 1960, A SYNONYM OF CTENOGO-
BIUS SALDANHA (BARNARD, 1927) © :

By

Frank Hamitton Tatsot and Mary-Loutsr Penrrrn~ ~* ~~: ZOOL.

LIBRARY
South African Museum, Cape Town |

Ad \/ H f \ 4 ur
NUV Lg 1900

(With 2 figures in the text and 1 plate)

CONTENTS

PAGE
Introduction ; . 189
Description of ee malta ; 190
Discussion . ; s : : 1QI
Acknowledgements : : - 192
Summary . 4 , ; she gs
References . : 3 : é 193

INTRODUCTION

. New iadbckial in the South African Museum fish dibecons from Port
Elizabeth, False Bay, and Saldanha Bay, has shown clearly that Ctenogobius
‘cloatus Smith, described from Knysna, and Clenogobius saldanha (Barnard),
, described from Saldanha Bay, are synonymous.

. The material examined is as follows:

S:A.M. 23277: 1 specimen, 79 mm., tidal swimming pool, St. James, False Bay;
S.A.M. 21488: 1 specimen, 33 mm., shallow water, Langebaan, Saldanha Bay;
S.A.M. 21489: 1 specimen, 30 mm., shallow water, Langebaan, Saldanha Bay;
S.A.M. 21490: 2 specimens, 34-37 mm., shallow water, Langebaan, Saldanha

Bay;
S.A.M. 22034: I specimen, 40, mm., inter-tidal eal Sea Point, Table aia,
S.A.M. 17355: I specimen, 85 mm., Seldache Bay (type of saldanha Barnard) ;
S.A.M. 23832: 6 specimens, 28-34 mm., dredged in 7 m. water, Saldanha Bay;
S.A.M. 23831: 4 specimens, 51-61 mm., inter-tidal pools, Port Elizabeth;
S.A.M. 24047: 4 specimens, 46 -5~54 mm., tidal swimming pool, Port Elizabeth;
S.A.M. 24048: 3 specimens, 45-5-50 mm., inter-tidal pools, Port Elizabeth;
S.A.M. 23979: 3 specimens, 53-79 mm., pedal pool, Strandfontein, False

Bay;

S.A.M. 24049: 2 specimens, 67 mm., 70 mm., inter-tidal pool, Strandfontein,

False Bay; ,
Rhodes Univ.: 1 specimen, 85 mm., lagoon, Keven (type of cloatus Smith).

189

5
Ann. S. Afri. Mus. 48 (8), 1964, 189-193, 2 figs., 1 pl.

a

1g0 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ctenogobius saldanha (Barnard)
(pl. IV, figs. 1, 2)
Gobius saldanha Barnard, 1927, p. 823.
Bathygobius saldanha (Barnard), Smith, 1949, p. 331.

Monishia saldanha (Barnard), Smith, 1960, p. 304.
Ctenogobius cloatus Smith, 1960, p. 302.

Fin counts: D. VI + I 10-11; A. I 9-10; P. 19-23; C. 15 (branched rays
only). Gill-rakers: 6—7 on lower arch, total 9. Scales 34-38, transverse 11. Depth
4:9-6°8. Head 3-0-3:6 in standard length. Teeth in jaw in several series;
outer row markedly enlarged, inner rows viliform, no canines. Tongue truncate
or feebly bilobed. Pectoral girdle without flaps, but with a low, indented
ridge on the anterior border. Pectoral fin with upper 3-4 rays free, silk-like,
markedly bifurcating (fig. 1). Pelvic fraenum (i.e. membrane connecting
outer rays across base) strong and well developed (fig. 1). Eye 2-9-3-7 in
head. Bony interorbital narrow, less than 5 in eye diameter, eyes adjacent.
Pores and papillae of head shown in fig. 2.

Ground colour whitish, with three very broad, irregular, faintly dusky
cross-bars on body. Sides irregularly spotted with black. Underparts white.
Nape dusky with a lighter transverse bar. Head whitish grey. A few small dark

FIGURE I.

Pectoral fin and pelvic fraenum of Ctenogobius saldanha (Barnard) from (a) Port Elizabeth;
standard length 45.5 mm., (6) Saldanha Bay; standard length 35 mm., and (c) Knysna (type
of cloatus Smith); standard length 85 mm. The fin is more contracted in (c) than in (a) and (6).

CTENOGOBIUS CLOATUS SMITH Ig!

spots on preopercle, and usually four in a straight line along hind margin of
opercle. Pectoral base dusky with several dark spots. Dorsal fins dusky with
rows of black dots forming bands. A large black spot on the membrane between
the fourth and fifth and the fifth and sixth dorsal spines. Pectoral, pelvic, anal
and caudal fins dusky.

Head, nape, preopercle, and opercle naked; pectoral base, prepelvic, and
belly with cycloid scales.

FIGURE 2.

Diagram of pores and papillae on head of Ctenogobius saldanha (Barnard).

Discussion

A re-examination of the type of saldanha shows certain errors in the original
description. The outer row of teeth in each jaw is clearly enlarged (not villiform
throughout) ; there are 11 series of scales between the first anal spine and the
dorsal fin base, not 10 as stated in Barnard’s description; and the bases of the
pectorals are scaled.

Smith’s description of cloatus indicates the following differences from
saldanha: strong development of the pelvic fraenum, absence of free upper
pectoral rays in his specimens, enlargement of the outer row of teeth, number
of transverse scale rows, shape of the tongue, and distribution.

The pelvic fraenum of saldanha shows no differences from that of the type
of cloatus (fig. 1). In both cases the fraenum is well developed.

Smith’s type has ragged fins with many rays broken off, and the fin
membranes are damaged. Nevertheless, this specimen has the upper pectoral

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

rays very similar. to those of saldanha (fig. 1) (both the type of saldanha and fresh
specimens from the type locality). In addition, South African Museum speci-
mens from Port Elizabeth have the upper rays similar to those of the type of
saldanha.

The apparent differences in teeth and in number of transverse scale
rows fall away after re-examination of the type of saldanha.

The tongue in both cloatus and in specimens of saldanha from the type
locality is truncate or feebly bilobed. It is not adnate, and Barnard (1927)
seems to have been correct in his suggestion that the apparently adnate tongue
of the type was due to the fact that the specimen was preserved with the mouth
unusually widely opened. Dehydration may also have played a part in this,
and in the shrinkage of the tongue, which has obscured its shape.

Bohlke & Robins (1960) have shown that the pore system of the head is
important in the classification of gobies. This was found to be identical in all
specimens examined, including Smith’s type specimen of Ctenogobius cloatus
(fig. 2).

As has been found in another intertidal group of fishes, the Clinidae, and
in gobies such as Psammogobius knysnaensis Smith and Coryphopterus nudiceps
(C. & V.), distribution around the Cape from at least as far west as Saldanha
Bay to Algoa Bay or farther occurs commonly.

We can find no difference between east and west coast specimens, and
conclude that-cloatus and saldanha are synonymous.

The well- -developed fraenum, presence of prepelvic scales, naked head
and nape, scale and fin counts, narrow bony interorbital, enlarged outer teeth,
tongue shape, and restricted gill openings, place this species within the genus
Ctenogobius Gill, 1858, as defined by Koumans (1953).

ACKNOWLEDGEMENTS

_ Acknowledgement is made to Professor J. L. B. Smith of the Department
of Ichthyology, Rhodes University, Grahamstown, for the loan of the type
specimen of cloatus, and to Miss R. M. Tietz, of the Port Elizabeth Museum,
and Mr. M. J. Penrith of the South African Museum, who assisted in the
collection of fresh material.

_ The Trustees of the South African Museum are grateful to the Council
for Scientific and Industrial Research for the award of a grant to publish
this paper.

SUMMARY

It is shown that Ctenogobius cloatus Smith is synonymous with Ctenogobius
saldanha (Barnard) (Pisces: Gobiidae). Ctenogobius saldanha is redescribed.

{ha Ce, a

a

CTENOGOBIUS CLOATUS SMITH 193

REFERENCES

BARNARD, K. H., 1927. A monograph of the marine fishes of South Africa Part II. Ann. S. Afr.
Mus. 21: 419-1065.

Bou.kE, J. E., & Rostns, C. R. 1960. A revision of the gobioid fish genus Coryphopterus. Proc.
Acad. nat. Sci. Philad. 112: 103-128.

Koumans, F. P. 1953. The fishes of the Indo-Australian archipelago. X. Gobioidea. Leiden: E. J. Brill.

SmitTH, J. L. B. 1959. Gobioid fishes of the families Gobiidae, Periophthalmidae, Trypauchenidae,
Taenioididae, and Kraemeriidae of the Western Indian Ocean. Ichthyol. Bull. 13: 185-225.

Smitu, J. L. B. 1960. Fishes of the family Gobiidae in South Africa. Ichthyol. Bull. 18: 299-314.

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Ann. S. Afr. Mus., Vol. XLVIII

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DYyUDpIVS sniqosouay’)

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TaBLeE OF ConTENTS and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 4? in. x 7 in. (74 in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers)
first and last, preceded by a colon (without prefix ‘p’). Thus:

SmitH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmitH, C. D. 1954. South African Plonias. In Brown, X. Y. Marine faunas. 2nd ed. 3: 63-065. .
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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ISOPODA AND AMPHIPODA COLLECTED BY ©”
THE GOUGH ISLAND SCIENTIFIC SURVEY

September 1965 September
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ISOPODA AND AMPHIPODA COLLECTED BY THE GOUGH ISLAND
: SCIENTIFIC SURVEY

By
K. H. BaRnarpt
South African Museum, Cape Town

(With 3 figures in the text) LIBRARY
CONTENTS VOV 19 i¥oD
PAGE
Prefatory note. : i + ros HARVARD
Introduction ‘ : : - “1O5 BENTAVG.
Isopoda ’ : : : iets) ai)
Amphipoda . : : f “3 p20
Acknowledgements : : we 2OQ
Summary . : : 3 2” 209
References . : ; : 209

PREFATORY NOTE

The collection described in this paper was made by the Gough Island
Scientific Survey, which worked on Gough Island from 13 November 1955
until 13 May 1956. Attention was largely concentrated on terrestrial biology,
and consequently the marine amphipods and isopods described here were all
obtained from a small number of collecting stations in the littoral and sub-
littoral zones near the mouth of The Glen on the east coast of the island. No
dredging was undertaken. The material was submitted to the author soon
after the return of the Expedition, and this paper was written in 1957. At that
time it was hoped that all the Expedition collections might be described in a
comprehensive report, but the unequal progress of work on different taxonomic
groups, some of which have yet to receive study, has made this impossible.
Rather than delay publication still further it has been decided that all results
should be presented as they stand, and the belated appearance of this paper is

a direct consequence of this decision.
Martin W. Holdgate

INTRODUCTION

The following collection of Isopoda and Amphipoda from the Gough
Island Scientific Survey was entrusted to me by Dr. M. W. Holdgate, to whom
I express my thanks for the opportunity of studying specimens from this remote
and little known locality (Position: 40° 21’ S., 09° 55’ W.). The Collection is
deposited in the British (Natural History), with reference specimens of most
species in the South African Museum.

+ Dr. K. H. Barnard died on 22 September 1964.

195
Ann. S. Afr. Mus., 48 (9), 1965: 195-210, 3 figs.

3

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

The Isopoda were more interesting than the Amphipoda as was to be
expected for Stephensen (1949) had already studied the species of Amphipoda
found at Tristan d’Acunha by the Norwegian Expedition. No species of
amphipod was collected at Gough Island which was not already known from
the sister isle; but not all the 32 (littoral and shallow-water) species recorded
from the latter have been found at Gough Island.

Much less is known about the Isopoda of these islands. Only three littoral or
shallow-water marine species have been recorded from Tristan, two of them
also from Gough Island. One introduced cosmopolitan woodlouse has been
long known from Tristan, and one endemic species since the visit of H.M.S.
Challenger. 'The Discovery II collected one of these marine species from Gough
Island; the other two, also the two terrestrial woodlice, have been obtained by
the present survey party.

In addition six marine species new to the Tristan—Gough Island area were
collected, two of which are described as new species. The rediscovery of the
New Zealand Munna neozelanica is noteworthy.

The isopodan and amphipodan fauna of these islands has relationships, in
general, with South Africa, the Magellanic region, Australasia, and the sub-
antarctic islands. Stephensen (1949) gave some zoogeographical remarks on
the amphipods. An analysis for the isopods may be premature, but for what it
is worth is as follows (excluding the two deep-water species collected by the
Challenger) :

Endemic on Tristan and/or Gough Island: 3 marine species (Paridotea apposita
n. sp., Jaeropsis vibicaria n. sp., and Isocladus tristensis), 1 terrestrial (Stylo-
niscus australis).

Common to Gough Island and New Zealand: 1 marine species (Munna neo-
zelandica).

Common to Gough Island and St. Paul Island: 1 marine species ( Jaeropsis
paulensis).

Common to Tristan and/or Gough Island and South Africa and New Zealand:
2 marine species (Paridotea ungulata, Dynamenella huttont).

Common to Tristan and/or Gough Island and other subantarctic regions: 1
marine species (Jais pubescens, commensal on Sphaeromids).

Cosmopolitan, introduced: 1 terrestrial species (Porcellio scaber).

Mention may be made of one maritime woodlouse whose absence (seem-
ingly) on Tristan and Gough Island is rather remarkable, namely Deito. The
species of this genus are known from South Africa, South America, Australia,
New Zealand and its subantarctic islands, and St. Paul (southern Indian
Ocean) (see Chilton, 1915: 437; and Barnard, 1932a: 220). They are large,
conspicuous, and abundant. It is, therefore, a fair presumption that the genus
does not occur on these islands.

Dr. Sivertsen and Dr. Holthuis are at present studying the isopods
collected on Tristan and neighbouring islands by the Norwegian Expedition.

ISOPODA AND AMPHIPODA COLLECTED BY GOUGH ISLAND SURVEY 197

Their results will give a much clearer picture of the fauna of the whole group
than the analysis here given based on Gough Island alone.

The preponderance of amphipods in the present collection may perhaps
be due, at least in part, to methods of collecting. Kelp is a favourite haunt of
these crustaceans, and numerous specimens can usually be obtained from only
a few handfuls of weed. Isopods, however, often present a more difficult
problem, varying with the type of beach, rock formation, tide range and other
factors, not least Neptune’s co-operation in curbing the turbulence of the waves.

ISOPODA
Family Anthuridae

Paranthura sp.

A single juvenile specimen, 4°5 mm. in length, with only 6 peraeon
segments and 6 pairs of legs. It has styliform mouth-parts, and the 5th joint of
peraeopods 5 and 6 does not underride the 6th. It is therefore probably a
species of Paranthura.

The colour pattern may prove distinctive when more specimens can be
obtained, possibly at Tristan if not at Gough Island. Pale yellowish, a dark
patch on front of head, and a dark transverse band across pleon, faint speckling
on the peraeon segments, eyes black.

Locality: Dell Rocks (A 47. M 54).

Family Sphaeromidae
Isocladus tristensis (Leach)

Exosphaeroma tristense (Leach) Tattersall, 1913: 882, pl., fig. 1 (juv. ¢ and 9).
Tsocladus tristensis (Leach) Barnard, 1914: 384, pl. 33, fig. B (adult 3); 1955: 62 (transverse
profile of body mentioned).

Localities: Beach debris, Glen mouth; shore near Base; Dell Rocks;
stomach of Soldier fish (A 27. M 1; A 47. M 60; B 5. M 83; B 10. M 88; and
B 10. M go).

A 27. M 1 a lot comprising adult and immature males, ovigerous females,
and juveniles. Specimens of the commensal asellid Jats pubescens have been
removed from some of them.

A 27. M g (part) one very juvenile, probably this species.

Previously recorded from Tristan and Gough Island.

A re-examination of I. magellanensis Richardson, 1906, is necessary before
it can be synonymised with tristensis.

Dynamenaella huttoni (Thomson)

Dynamenella huttoni (Thomson) Chilton, 1909: 657. Barnard, 1940: 419 (references).
Dynamenella kraussi Barnard, 1914: 415, pl. 35, fig. B.

These specimens agree with South African specimens in convexity of body,
and absence of a raised rim on frontal margin of head. Although the surface is

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

glabrous (Barnard, 1914), it is not polished except in the mid-dorsal line of
peraeon segments 2-6; on the head, lateral portions of peraeon, and on pleon
and telson it is finely shagreened, and on the dorso-lateral portions of peraeon
finely reticulated. |

Colour (as preserved) corresponding with that of South African specimens
(Barnard, 1914: 417): reddish, with or without a paler stripe (or series of spots)
on each side, and indications of paler patches in middle of back.

The size (male 7 mm., female with embryos 5 mm.) is considerably less
than the average size of South African specimens.

Localities : Dell Rocks and Seal Rocks (A 40. M 25; B 4. M 81; 1504).

Family Idoteidae
Paridotea ungulata (Pallas)
Paridotea ungulata (Pallas) Hale, 1924: 221, fig. 9 e, f. 1929: 320, fig. 325 b (side view of 6th and
7th peraeon segments). Sheppard, 1957: 151, fig. 4 a-e.
(Fig. 1 a)
Tristan d’Acunha and Gough Island. Not taken by the present Survey.

Paridotea apposita n. sp.
(Fig. 1 5)

The parallel-sided body gives this species the appearance of a large form
of fucicola Barnard (1914: 327, pl. 36, fig. E). In other respects closely resemb-
ling ungulata.

Side-plates 5 and 6 postero-lateral angle quadrate, 7 a little produced,
the angle slightly acute.

Ficure 1. Paridotea. Side-plates 4-7 and apex of telson of:
(a) ungulata (Pallas). (b) apposita n. sp. (c) munda Hale.

Pleo-telson scarcely more than -one-third of the total length (ungulata:
two-fifths), apex shallowly emarginate between the rounded postero-lateral
angles, on which a faint keel is traceable. Sternal plates as in ungulata (see
Barnard, 1914: pl. 36, fig. D, st. w.).

ISOPODA AND AMPHIPODA COLLECTED BY GOUGH ISLAND SURVEY 199

Maxilla 1 (of male 30 mm. long) outer plate with 12 spines, with a slender
one between the two groups of 6, 4 of the inner group serrulate, but not the
innermost shortest one, a seta in middle of plate below bases of spines; inner
plate with 4 plumose setae and a simple seta. Maxilliped 7-jointed.

Paraeopods as in ungulata; 3 stout serrulate spines on inner margin of 6th
joint of peraeopod 1, one in each of peraeopods 2-7; 4th—6th joints of peraeo-
pods 1-6, furry on inner margin in the large male, not furry in females or the
smaller specimens.

Uropod peduncle without median keel, a plumose seta at outer (when
appendage is folded in) distal corner; length of ramus slightly less than basal
width (2: 2-3), outer (when folded) margin more oblique (following the outline
of telson) than in ungulata, the truncate distal margin relatively narrower.

Pleopod 2 of male, stylet as in ungulata.

Colour (as preserved): dull pinkish or reddish.

Localities: Beach debris, Glen mouth; stomach of Soldier fish (A 27. M 2)
I immature, 18 mm.; B 10. M 861 ¢ 30 mm., 2 immature Ig and 22 mm.;
B 10. M 89 1 2 31 mm.; B 10. M g0 1 2 27 mm., I immature 18 mm.
(head missing).

The 31 mm. female shows the brood lamellae beginning to develop as tiny
lobes, 0-5 mm. long, on segments 2—5; the 27 mm. female shows only slight pits
in the positions where the lobes will appear at the next moult. Neither female
shows any sign of the anterior peraeon segments broadening.

Thanks to the kindness of Dr. H. M. Hale, then Director of the South
Australian Museum, I was able to examine a specimen of P. munda Hale 1924,
a species which merits comparison with ungulata and the Gough Island species.
The accompanying figures (fig. 1) of the posterior side-plates and the telsonic
apex will illustrate the differences in the three species. Hale’s figures of the side-
plates (1924 and 1929) do not seem to show the correct shapes, in comparison
with the specimen I have seen. This specimen also has a narrower notch at the
telsonic apex. (fig. 1 c).

The Cape species fucicola Barnard (not figured here) has a narrower
telsonic notch than munda, side-plates 5 and 6 do not reach as far back as the
hind margin of their segments, and the narrowly rounded apex of side-plate 7
extends only slightly beyond its segment.

Family Jaeridae
Iais pubescens (Dana)

Tais pubescens (Dana) Chilton, 1909: 649. Barnard, 1914: 435, pl. 37, fig. C (typ. err. fig. of plp.
3 has been inverted and labelled ‘mxp’. See corrigendum 1916: 106). Nordenstam, 1933:
178, figs. 41 a—c (references). Menzies J. L., Barnard, 1951: 138, pls. 42, 43.

Nordenstam (1933) stated that my figure (Barnard, 1914) of the mandible
differed from that given by Stebbing (1900: pl. 38), apparently without
noticing that the former was the male, notwithstanding that he himself

4

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

examined a male. He was also unaware of the corrigendum relating to the
‘mxp’
Paraeopod 1 triunguiculate (or as worded by Menzies & J. L. Barnard

(1951), ‘biunguiculate, the inferior claw bifid’) as in the other peraeopods.

Localities: Stream, at 150 ft. altitude, free-living; Beach debris, Glen
mouth, on Isocladus; stomach of Soldier fish, on Isocladus (A 11. G. 64; A 27.
M 1; B 10. M 88).

The occurrence free-living in a stream at 150 ft. altitude is interesting.
Chilton (1909: 650) found them free-living at the mouths of fresh water
streams in the Sounds on the west coast of New Zealand, often out of range of
ordinary high tides. But at 150 ft. altitude the Gough Island individuals would
be probably far above even an occasional splash of salt water.

Previously recorded as a commensal on the same Sphaeromid from
Tristan.

Note on South African specimens. Menzies & J. L. Barnard (1951: 148) have
relegated the specimens recorded from the Cape and Durban to ‘species
inquirendae’, and state that the former are ‘possibly not Jas’. This latter
statement may perhaps have been due to the unfortunate typ. err. regarding
the 3rd pleopod mentioned above.

I have re-examined Cape material, and find that the 1st peraeopod is
triunguiculate as in the other peraeopods, and therefore these specimens can
be assigned to pubescens as diagnosed by the collaborating authors. The enlarge-
ment of the mandibles in fully adult males, of which I have found two additional
specimens, is also confirmed. Males with pleopods developed, and thus
presumably functional, are much less common than females, and fully adult
males with enlarged mandibles are rare.

A single specimen from Durban was in poor condition, and consequently
indecisive. Possibly specimens from Natal and from Inhambane, Portuguese
East Africa (University of Cape Town Ecological Survey) and from Ceylon
may prove referable to singaporensis Menzies & J. L. Barnard.

Genus JAEROPSIS Koehler

Only the austral species are here considered. According to the position
and shape of the uropods two groups of species can be separated (fig. 2). In
marionis, patagoniensis, intermedius and palliseri they are inserted only a short
distance apart, project straight backwards, and their inner margins are straight
(or nearly so) and fit closely together. In paulensis and curvicornis (of Stebbing)
they are inserted well apart, fold obliquely inwards, and even when fully
folded their apices do not touch one another. The shape of the telsonic apex is
correlated: triangular and more or less acute in the first, rounded in the second
group. Other taxonomic characters seem to be variously distributed. Chilton’s
descriptions (1892a and 1912) of neozeylanicus are inadequate (see also Hurley,

1957: 19).

+ ~atte-

a atiied

a

ISOPODA AND AMPHIPODA COLLECTED BY GOUGH ISLAND SURVEY 201

Lateral margin 2nd anten- Uropod Maximilliped
head telson na joints of apex of and joint
First group : peduncle peduncle _ of palp
-marionis serrate serrate linear hooked ste eh 1
patagoniensis smooth 1 tooth enlarged hooked linear
intermedius serrate serrate enlarged hooked lobed
(anteriorly)

vibicaria smooth smooth enlarged rounded lobed
palliseri smooth smooth enlarged rounded ?
Second group
curvicornis (Ceylon, smooth serrate enlarged hooked lobed

S. Africa)
paulensis (St. Paul) smooth smooth enlarged rounded ?lobed*
paulensis (Gough Is.) smooth smooth enlarged rounded lobed

* Vanh6ffen’s fig. 59 f/ shows a linear joint but the palp has been displaced, either in
dissection or mounting, and consequently no reliance can be placed on the shape, as drawn,
being the true shape.

Jaeropsis paulensis Vanh.
(Fig. 2 5)
Jaeropsis paulensis Vanhoffen, 1914: 531, fig. 59 al.

Anterior half of head grey, forming a transverse bar, posterior half and
upper lip creamy-white, peraeon irrorated with grey, paler than the bar on head,
pleo-telson faintly dusted with grey, appendages creamy-white, eyes black.

Males 4—4°5 mm., ovigenous females 3—4°5 mm.

Appears to agree very closely with paulensis from St. Paul Island (southern
Indian Ocean). The discrepancy in the palp of the maxilliped is probably
explainable as suggested in the footnote to the table above.

The apex of the uropod is rather broadly rounded and has no hook, but
the inner margin appears to be minutely serrulate.

Localities: Dell Rocks; West Point Reef (A 40. M 33; A 47. M 60; B 1.
M 69; 1541).

a: b Cc

Ficure 2. Jaeropsis. (a) vibicaria n. sp., pleo-telson, with uropod further enlarged.
(b) paulensis Vanh. apex of telson, with uropod further enlarged. (c) ‘curvicornis’
(South Africa), apex of telson, with uropod further enlarged.

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

Jaeropsis vibicaria n. sp.
(Fig. 2 a)

Although only a single specimen is present, a new species seems unavoid-
able; its main taxonomic characters are given in the above table. Nordenstam
(1933) had both sexes of patagoniensis and intermedius, consequently the smooth
telsonic margin and lack of apical hook on the uropod cannot be attributed to
sexual differences.

Antenna @, left missing, right mutilated; 4th and 5th peduncular joints
enlarged, with scarious outer edge which appears to be crenulate. Maxilliped,
lobe of and joint of palp extending only one-third length of 3rd joint; 6 coupling
hooks.

Peraeopods and pleopods as for genus.

Uropods inserted close together, directed backwards, their inner margins
straight and approximate, apex rounded, with 2-3 setae (?plumose); both
rami minute, especially the outer ramus, each with a tuft of setae.

A dark grey triangular patch covering head except hind corners, con-
tinued as a mid-dorsal somewhat moniliform stripe on peraeon, expanding
into a subtriangular patch on pleo-telson, but not covering the anterior corners
or the posterior third; rest of body, and appendages creamy-white, eyes black.
5°5 mm.

Locality: Dell Rocks (A 47. M 60). One male.

The colour pattern resembles that figured for patagoniensis Richardson

(1909: 421, fig.).

Jaeropsis curvicornis Stebb. ( ?non Nicolet)
(Fig. 1 ¢)
Jaeropsis curvicornis (?non Nicolet) Stebbing, 1905: 51, pl. xi, fig. C. Barnard, 1914: 224, pl. 20,
fig GC:

Since my description of one male from the Cape, I have seen more
material: St. James (False Bay) and Sea Point (Cape Town) (coll. K.H.B.),
Lambert’s Bay and Saldanha Bay (both on west coast) and Mossel Bay (coll.
University of Cape ‘Town Ecological Survey).

The lateral margin of head is smooth, that of pleo-telson serrate, both in
juveniles and adults.

The uropod has an apical hooked tooth,* and the inner distal margin
serrulate; each ramus with a tuft of setae. A better figure than my 1914 one is
given here.

The colour resembles that of paulensis, and the dark crossbar on head
offers a ready means of picking out this asellid from other species.

Presumably Stebbing was able to compare his Ceylonese specimen with
Nicolet’s description and figure; but it would be much more satisfactory to
have a full description of specimens from the original Chilean locality. Until

* Stebbing mistook this for the inner ramus; as also did Miss Richardson. (1909) in her
description of patagoniensis.

ISOPODA AND AMPHIPODA COLLECTED BY GOUGH ISLAND SURVEY 203

that is available I feel some hesitation in identifying Cape poe with
Nicolet’s curvicornis, but none in identifying them with the ‘curvicornis’ of

Stebbing.

Family Munnidae

Munna neozelanica Chilton

Munna neozelanica Chilton, 1892a: 267. 1892b: 2, pl. 1. figs. 1-8a, pl: 2, figs. 85-15, Monod, 1931:
14, figs. 8c, 9 c-e.

Haliacris neozelanica (Chilton) Chilton, 1909: 650, fig. 14 a (prp. 1 3). 1912a: 132. Stephensen,
1927: 357:

Largest male 4 mm., smallest, with hammer-shaped 1st peraeopod 2:5 mm.;
ovig. females 2-5-2-75 mm.

Peraeopod 1 of male, ‘hammer’ more strongly calcified than the other
joints. The other peraeopods setose in male, especially 4th—6th joints of
peraeopods 6 and 7.

Chilton and Monod figured the 1st peraeopod of a juvenile male (Monod’s
specimen 1-5 mm. in length), closely resembling that of the female figured by
Chilton. Chilton figured three stages in the development of the ‘hammer’
(figs. 8b, 8c, 8d). giving the magnification of the figures but without stating the
respective lengths of the specimens from which the appendages were removed.
At what size, therefore, the change in shape occurs cannot be stated; in the
present smallest male, 2-5 mm. in length, the peraeopod has already assumed
its final shape, but of course is smaller than in the 4 mm. male. Chilton’s
largest specimen was 3 mm. in length.

No other Munnid has been described with the 1st peraeopod of the male
shaped as in the New Zealand species, and the present specimens agree with
the descriptions and figures of Chilton and Monod. Hitherto recorded only
from New Zealand (south island) and the subantarctic Auckland and Campbell
Islands.

Localities: Dell Rocks and shore nearer Base (A 40. M 27; A 47. M 33;
B5. M 84).

Family Trichoniscidae
Genus sTyLoniscus Dana
Vandel, 1952: 14.
Styloniscus australis (Dollfus)

(Fig. 3)

Trichoniscus (?) australis Dollfus, 1890: 6.
Styloniscus (?) australis (Dollfus), Vandel, 1952: 17.

Sparsely and minutely granulate, chiefly on posterior margins of head and
‘peraeon segments. Eye composed of 3 ocelli in a triangle, contiguous or almost
so. )

Antenna 2, 5th joint with a basal and a subapical conical spine sometimes

204 , ANNALS OF THE SOUTH AFRICAN MUSEUM

a Ob d C e

FiGurRE 3. Styloniscus australis (Dollfus). (a) distal joints of peraeopod 1. (6) distal joints
of peraeopods 6 and 7. (¢) penial process. (d) pleopod 1 g. (e) pleopod 2 ¢.

I or 2 smaller intermediate ones, and an apical spine on outer margin; flagellum
3-jointed, with apical tuft of setae, slightly shorter (excluding apical tuft) than
5th joint.

Maxilla 1, outer plate with 9 spines, the inner ones simple, and 2 accessory
setae scarcely extending beyond the longest spines and extremely finely setulose,
outer margin entire, setose. Maxilliped, basal plate with convex outer margin,
apically lobed, palp (outer plate) unjointed, inner plate composed of 1 joint with
a small terminal joint, the latter much smaller than in magellanicus, cf. verrucosus
Budde-Lund (1906: pl. 4, figs. 10, 11) and Vandel (1952, fig. 9).

Peraeopod 1, 4th joint with one long and one short spine on outer margin,
4 spines on inner margin, the 3rd longest; 5th joint with 5 spines on inner
margin, the 4th longest and apically trifid; 6th joint with 2 small spines on
outer margin, one on inner margin. No sexual difference.

Peraeopods 6 and 7, 5th joint with 5 or 6 spines on inner margin, the
subapical one longest and apically trifid; 6th joint with 4 spines on inner
margin, a series of about 15 close-set spines on outer margin. No sexual
difference. ‘

Penial process with short, conical, wrinkled process in the apical emargi-
nation. Pleopod 1 of male, outer corner of basal plate serrulate. Pleopod 2 of
male, see fig. 3¢. Pleopod 5 of male, exopod grooved.

3°5-4 mm. X 1:3-2 mm. Mottled grey.

Localities: Gough Island: various, from Beach debris, Glen mouth, to
summit of Expedition Peak, alt. 2950 ft.; and Penguin Island (A 1. G 5 Ss
G 13; A2. G 23; A 8. G 56 [A 9 on label]; A 12. G 194; S [stc? =A] G 88;
A 14. G 80; A 27.G 149; A29.G 171; A 29. G 216; A 30. G 184; A 30. G 191;
A 32. G 200; A 35. G 211; A 37. G 226; A 39. G 246; Agi. G 252; A 43. G;
268; B 3. G 305; B 3. G 315; B 8. G 338; B 13. G 373).

ISOPODA AND AMPHIPODA COLLECTED BY GOUGH ISLAND SURVEY 205

Tristan d’Acunha: Jenny’s Watrin.

Vandel (1952: 99 and chart 76) said in regard to the geographical distribu-
tion of Styloniscus: South Africa (southern coastal area), Madagascar, Mauritius,
Patagonia, Australia, Tasmania, New Zealand and southern islands, Crozet
Is., and ‘perhaps Tristan d’Acunha and Juan Fernandez’. Chilton’s ‘Marion
Island’ (obiter dictum 1915: 454), is surely a laps. cal. for Crozets, which is
_ correctly listed in his Subantarctic Islands New Zealand (1909: 799).

The rediscovery of this species, first taken by H.M.S. Challenger on Tristan
d’Acunha, is therefore welcome. :

Family Porcellionidae
Porcellio scaber Latr.

Porcellio tristensis White, 1847: 99.
Porcellio scaber Latr., Barnard, 1932a: 252, fig. 21 a-—c.

Localities: Gough Island: various, from the Glen up to Michael’s Ridge,
m@meeroooit. (A1.G5;A2.G13;A2.G23;A4.G30;A8.G53;A13. G 73;
mise = — Al 13.G 68; A 14. G 80; A 16. G 84; A 16.G 93; Aa1.Gi117;A
eit 110; A 24. G 127; A27.G 140; A 28. G 164; A 35. G a11; A 37. G 229;
A 38. G 236; B 3. G 305; B11. G 350; B 13. G 365; B 13. G 373; B14. G 380;
B 15/16. G 397). :

Tristan d’Acunha: Jenny’s Watrin.

Collected on Tristan by Capt. Dugald Carmichael (1816) and described
as a new species by White (1847); later recorded from Tristan by Barnard
(1932a).

The Gough Island material comprises uniform slaty-grey and variously
mottled specimens; some, e.g. from the Open Phylica forest by Lower Waters-
meet (A 8. G 53) are very sparsely spotted with grey on a yellowish ground
colour.

AMPHIPODA

Schellenberg, 1931; Barnard, 1932b:; Stephensen, 1949; Macnae, 1953.

Family Lysianassidae
Parawaldeckia kiddert (S. 1. Smith)

Parawaldeckia kidderi (S. 1. Smith) Stephensen, 1927: 300, fig. 2. Schellenberg, 1931: 6 and 22
(contrasted with Socarnoides kergueleni). Stephensen 1949: 5.
Stephensen (1927) gave figures of uropod 3 in the two sexes: in the male
it carries a brush of long setae.

Locality: Dell Rocks (B 2. M 75).
Recorded from Tristan and neighbouring islands (Nightingale and
‘Inaccessible).

206 _ ANNALS OF THE SOUTH AFRICAN MUSEUM

Acontiostoma marionis Stebb.

Acontiostoma marionis Stebbing, 1888: 709, pl. 30. Chilton, 1912b: 462. Barnard, 1932b: 32.
Acontiostoma magellanicum Stebbing, 1888: 714, pl. 31. (= juv.)

Gough Island, 100 fathoms (Chilton: Scotia).

Family Stenothoidae
Stenothoe sivertsent Steph.

Stenothoe sivertsent Stephensen, 1949: 9, figs. 2, 3. Macnae, 1953: 1026.

Locality: from Macrocystis kelp (M 25. Misc.).
Previously recorded from Tristan and neighbouring islands.

Family Pontogeneiidae
Pontogeneia tristanensis Brnrd.

Pontogeneia tristanensis Barnard, 1932b: 199, figs. 118 m, 120. Stephensen, 1949: 15.

Localities : Midshipman Rock, Dell Rocks, outer fringe kelp, kelp between
Reef Point and Dell Rocks (A 42. M 34; A 44. M 34; A 44; B 1. M 65; BM.
M 77; M. 25. Misc.).

Previously recorded from Tristan and neighbouring islands.

Paramoera tristanensis Brnrd.

Paramoera tristanensis Barnard, 1932b: 209, figs. 118 k, 127. Stephensen, 1949: 16. Macnae, 1953:
1026.

Localities: Dell Rocks and Half-way Cave (A 40. M 26; A 47. M 42; B 4.
M 80; B 5. M 82).
Previously recorded from Gough Island, Tristan and neighbouring
islands.
Family Gammaridae
Melita tristanensis n. sp.

Melita gayi (non Nicolet) Stephensen, 1949: 22. Macnae, 1953: 1027.

Agreeing with Stephensen’s Tristan material in having no medio-dorsal
tooth on pleon segment 4, and thus disagreeing with the South African orgasmos
Barnard, 1940. Stephensen said pleon segment 5 had ‘one pair of small subdorsal
teeth’; the Gough Island specimens resemble orgasmos in having a pair of
denticles with a seta between them on each side of the median line. I think one
may assume that the two denticles in the Tristan specimens were actually
double. .

Side-plate 1 strongly expanded forwards as in orgasmos, but postero-
inferior angle of pleon segment 3 quadrate with a small point (as in palmata).

In view of these conflicting characters it is difficult to decide what specific
name to attach to the Tristan-Gough Island specimens. Schellenberg (1931:
203) separated gay: Nicolet (incl. inaequistylis auct.) from the northern palmata
mainly because of lack of material for comparison; he made no mention of such

ISOPODA AND AMPHIPODA COLLECTED BY GOUGH ISLAND SURVEY 207

critical characters as the dorsal dentation of the pleon, or the shape of the hind
corner of pleon segment 3.

In 1940 (p. 454) I revised my opinion as to the identity of ‘inaequistylis’
and its possible synonyms, and distinguished two species orgasmos Barnard
and zeylanica Stebbing in the South African material previously recorded as
inaequistylis. 1 consider it desirable to recognize distinct species in regions which
have been separated for a considerable period, until material from each
region has been thoroughly examined.

Therefore, because no detailed description has yet been given of the South
American gayi, I attach a new name to the Tristan-Gough Island specimens.

A further question is: which characters are critical for defining the species ?
Typically orgasmos has a dorsal tooth on pleon segment 4, but sometimes
(Barnard, 1940: 455) this is absent!

Colour (as preserved): purplish-brown, eyes black.

Locality: Dell Rocks (A 40. M 26; A 47. M 60).

Previously recorded from Tristan and neighbouring islands.

Family Talitridae
Orchestia platensis Kroyer

Orchestia platensis Kroyer, Stephensen, 1944: 57, figs. 15, 16. 1949: 25. Macnae 1953: 1027.
Orchestia tristensis (Leach MS.) White, 1847: 86.

Locality: Beach, opposite Hut (A 13. G ga).
Previously recorded from Tristan and Inaccessible Island.

Orchestia scutigerula (Dana)
Orchestia scutigerula (Dana) Stephensen, 1949: 26, figs. 11, 12; Macnac, 1953: 1027.
Localities: Penguin and Midshipman Islands (A 38. G 232; A 39. G 242;
390. G 247; A 42. G 261).

Previously recorded from Nightingale and Inaccessible Islands.

_ Hyale hirtipalma (Dana)
Hyale hirtipalma (Dana), Stephensen, 1949: 30, fig. 13.

Only one Male (in Bz. M 75) has the strongly setose flagellum of antenna
2 (cf. Barnard, 1916: 234); per contra the hand of case: 2 lacks the
characteristic brush of setae.

Localities: Beach debris, Glen mouth; Dell Rocks, Penguin Island, Half-
way Cave, West Point Reef (A 27. M 11; A 28. M 18; A 28.M 19;B2.M 75;
B 4. M 79; B 5. M 82; 1541; 1543).

Previously recorded from Tristan and neighbouring lone:

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hyale grandicornis (Kroyer)

Hyale grandicornis (Kroyer), Chilton, 1912b: 508. Stephensen, 1949: 33. figs. 14, 15. Macnae,
1953: 1028.

Localities : Glen mouth and Dell Rocks (A 28. M 19; B1. M 72; B2. M 75).
Previously recorded from the shore of Gough Island by Chilton (1912),
and by Stephensen (1949) from Tristan and Nightingale Island.

Allorchestes tristanensis Macnae
Allorchestes tristanensis Macnae, 1953: 220.

One immature female in the Discovery collection was assigned to this genus,
and later collecting has confirmed its occurrence on Tristan. Stephensen (1949)
regarded specimens as a variety of Hyale grandicornis, but Macnae (1953) has
described them as a species of Allorchestes, commenting on the slight difference
between the two genera.

Family Aoridae
Aora typica Kroyer
Aora typica Kroyer, Stephensen, 1949: 41, fig. 18. Macnae, 1953: 1032.

Locality: Dell Rocks (A 40. M 26; B 1. M 69).
Previously recorded from Tristan and neighbouring islands.

Family Photidae
Eurystheus remipes Brnrd.

Eurystheus ?afer Stebbing, Chilton, 1g12b: 510, pl. 2, figs. 30-34.
Eurystheus remipes Barnard, 1932b: 229, fig. 143.

Gough Island, trawled in 100 fathoms (Chilton: Scotia).
Falkland Islands, 105-115 metres (Barnard: Discovery).

Family Ampithoidae
Ampithoe brevipes (Dana)
Ampithoe brevipes (Dana), Barnard, 1932b: 239, fig. 150. Stephensen, 1949: 44.

Locality: Dell Rocks, from kelp (A 44. M 34; B 1. M 69).
Previously recorded from Tristan and neighbouring islands.

Family Jassidae
Ischyrocerus longimanus (Haswell)

Ischyrocerus anguipes var. longimanus (Haswell), Barnard, 1932b: 243.
Ischyrocerus longimanus (Haswell), Stephensen, 1949: 45 (references). Macnae, 1953: 1032.

Locality: from kelp (M 25 Misc.; M 27).
Previously recorded from Tristan and neighbouring islands.

Stephensen (1949) considered that the true anguipes is confined to the
northern hemisphere.

4
%
4

ISOPODA AND AMPHIPODA COLLECTED BY GOUGH ISLAND SURVEY 209

Family Caprellidae |
Caprella acutifrons Latr.

Caprella acutifrons Latr., Barnard, 1932b: 300. Macnae, 1953: 1032.
Caprella acutifrons var. natalensis Mayer, Stephensen, 1949: 53.

Localities: from kelp (A 44. M 35; A 44. M 36; A 44 outer fringe kelp;
M 24. Misc.; M 26; M 27).

Previously recorded from Tristan and neighbouring islands.

ACKNOWLEDGEMENT

The Trustees of the South African Museum acknowledge gratefully a grant
from the South African Council for Scientific and Industrial Research for the
publication of this paper.

SUMMARY

An account is given of the Isopoda and Amphipoda of Gough Island
(40° 21’ S., 09° 55’ W.) based on the collections of the Gough Island Scientific
Survey, 1955-1956. Eleven species of Isopoda and sixteen species of Amphipoda
are recorded, including six marine species of Isopoda new to the Tristan—
Gough area. Two new species of Isopoda, Paridotea apposita and Faeropsis
vibicaria, and one new species of Amphipoda, Melita tristanensis, are described.

REFERENCES

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BARNARD, K. H. 1916. Contributions to the crustacean fauna of South Africa. 5. Amphipoda.
Ann. S. Afr. Mus. 15: 105-302.

BarNARD, K. H. 1932a. Contributions to the crustacean fauna of South Africa. Terrestrial
Isopoda. Ann. S. Afr. Mus. 30: 179-388.

BARNARD, K. H. 1932b. Amphipoda. Discovery Rep. 5: 1-326.

BARNARD, K. H. 1940. Contributions to the crustacean fauna of South Africa. 12. Further
additions to the Tanaidacea, Isopoda, and Amphipoda, together with keys for the identifi-
cation of the hitherto recorded marine and freshwater species. Ann. S. Afr. Mus. 32: 381-543.

BARNARD, K. H. 1955. Additions to the fauna-list of South African Crustacea and Pycnogonida.
Ann. S. Afr. Mus. 43: 1-107.

BuppE-Lunp, G. 1906. Die Landisopoden der Deutschen Siidpolar-Expedition, 1901-1903.
9: 69-92.

Cuitton, C. 1892a. Notes on some New Zealand Amphipoda and Isopoda. Trans. N. &. Inst.
24 (1891): 258-269.

Cuitton, C. 1892b. A new species of Munna from New Zealand. Ann. Mag. Nat. Hist. (6) g: 1-2.

Cuitton, C. 1909. The Crustacea of the subantarctic islands of New Zealand. In PHILOso-
PHICAL INSTITUTE OF CANTERBURY. The subantarctic islands of New Zealand, ed. by C. Chilton,
&c. 2: 601-671. Wellington, N.Z.: Government Printer.

Cuitton, C. 1912a. Miscellaneous notes on some New Zealand Crustacea. Trans. N.£. Inst.
44 (1911): 128-135.

Cuitton, C. 1912b. The Amphipoda of the Scottish National Antarctic Expedition. Trans. roy.
Soc. Edinb. 48: 455-520.

CuitTon, C. 1915. Deto, asubantarctic genus of terrestrial Isopoda. 7. Linn. Soc. ( Zool.) 32:435-456.

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Do.trus, A. 1890. Isopodes terrestres du ‘Challenger’. Bull. Soc. Etud. sci. Paris 12: 1-7.

Hate, H. M. 1924. Notes on Australian Crustacea. No. 2. Trans. roy. Soc. S. Austr. 48: 2-6.

Hate, H. M. 1929. The crustaceans of South Australia. 2: 201-380. Adelaide: Government Printer.

Hour ey, D. E. 1957. Some Amphipoda, Isopoda, and Tanaidacea from Cook Strait. Zool. Publ.
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Macnag, W. 1953. On a small collection of amphipods from Tristan da Cunha. Proc. zool. Soc.
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Menzigs, R. J. and BARNARD, J. L. 1951. The isopodan genus Jais. Bull. S. Calif. 50: 136-151.

Monon, T. 1931. Tanaidacés et isopodes sub-antarctiques de la collection Kohl-Larsen du
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~~.

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TABLE oF CONTENTS and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. x 7 in. (74 in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers)
first and last, preceded by a colon (without prefix ‘p’). Thus:

SmitH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

Smit, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

‘ i

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. ~~ ~ead

MARY-LOUISE PENRITH

MUS. COMP ZOO!
LIBRARY

NOV 19 i965
HARVARD
STUDIES ON THE SOUTH AFRICAN CLINIDAE I.
DESCRIPTION OF A NEW SPECIES OF
PAVOCLINUS, AND REDESCRIPTION OF
GYNUTOCLINUS ROTUNDIFRONS (BARNARD)

September 1965 September
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STUDIES ON THE SOUTH AFRICAN CLINIDAE. I. DESCRIPTION
OF A NEW SPECIES OF PAVOCLINUS, AND REDESCRIPTION OF
GYNUTOCLINUS ROTUNDIFRONS (BARNARD)
By
US. COMP. ZOOL

MUS. Ct

Mary-Louise PENRITH LIBRARY
South African Museum, Cape Town Si ee a Wa
NQV £3 ty0e

(With 4 figures in the text and 1 plate)

HARVARD
CONTENTS UNIVERSITY
PAGE
Introduction. : . : ‘ : ae Qin
Description of Pavoclinus litorafontis n. sp. ; Pee ye
Discussion : : ‘ : : ; 2. 2s
Redescription of Gynutoclinus rotundifrons (Brnrd) . 215
Discussion , ; ; ; ; : 2. 216
Acknowledgements. F : : ; ey 2l7
Summary. : { ! : ‘ : Mee 6s,
References : ‘ F ; ; ‘ PAROS My,
INTRODUCTION

During the course of a systematic and biological study of the South African
fishes of the family Clinidae, extensive collecting has been carried out in the
intertidal region of the south-western Cape coast.

In May 1963 two specimens of a clinid which could not be identified with
any of the species known at present were collected with hand-nets from the
dense beds of the green alga Caulerpa filiformis at Strandfontein, in False Bay.
Eleven more specimens were subsequently collected from the same area, and
recently a single specimen was collected at Onrus River mouth, near Hermanus.

The specimens were placed in the genus Pavoclinus Smith, 1945, on account
of their lack of a supra-orbital tentacle, possession of a raised crest consisting of
the first three dorsal spines, which is not separated from the rest of the fin by a
notch in the membrane between the third and fourth dorsal spines, and their
toothed vomer.

While collecting was being carried out in kelp-filled pools at the bottom
of the intertidal zone at Lambert’s Bay in January 1964, a male specimen of
Gynutoclinus rotundifrons (Barnard, 1937) was caught. This species was pre-
viously known only from a single female specimen collected at Oudekraal in
1934 by the University of Cape Town. In March 1964 a juvenile specimen was
collected from kelp in a gully at Lambert’s Bay. A redescription of the species
is desirable, since the holotype was not fresh by the time it was received by Dr.
Barnard and the colouring had therefore completely disappeared, the male
was unknown, and there are a few errors in the original description.

211
S,
Ann. S. Afr. Mus. 48 (10), 1964: 211-217, 4 figs., 1 pl.

a

oT? ANNALS OF THE SOUTH AFRICAN MUSEUM

Pavoclinus litorafontis n. sp.
(Pl. V, figs. 1, 2)

(litorafontis from Litus, -oris: beach; fons, -ntis: fountain)

Material: (a) From Caulerpa beds in the intertidal zone, Strandfontein: S.A.M.
23876, 2 specimens, male and female, 91 mm., 110 mm.; S.A.M. 23877, 2
males, 102 mm., 175 mm.; S.A.M. 23952, 4 specimens, 1 male, 3 juveniles,
46-132 mm.; S.A.M. 23962, two juveniles, 41 mm., 43 mm.; S.A.M. 23972,
one male, 136-5 mm.; S.A.M. 24052, one male, 164-5 mm. (holotype);
(b) from an intertidal pool at Onrus River mouth, in Bifurcariopsis brassicaeformis ;
S.A.M. 24081, one female, 64. mm.

nS the

Description: D. XXIX-XXXIII 6-8; A. II 20-23; P. 12; V. I 3; C. 19.

Depth 4:5-5:75; Head 3:75-5 in standard length; eye 3°5-4:25 in head.
Upper jaw (from angle of jaw to snout tip) 33°5-41°5% of head length (2-4-3

(a) Ventral view. (6) Lateral view.

FicureE 1. Intromittent organ of Pavoclinus litorafontis.

in head). Caudal peduncle length 58-5—75% of head length; caudal peduncle
depth 22°5-33-5% of head length.

First three spines of dorsal fin elevated to form a crest, decreasing in
relative height with increase in size of fish. Not even a shallow notch in mem-
brane between third and fourth spines. Upper pectoral rays short, pectoral fin
roughly elongate-ovate. Inner pelvic ray stout, equal to others. Caudal peduncle
long. Caudal fin subtruncate.

Body somewhat elongate, compressed, covered with small imbricating
scales. Snout bluntly conical. Eye rounded, protuberant, equal to snout. No
supra-orbital tentacle. Nasal cirrus a small flap. Lips moderately thick.

Lateral line of minute double and alternating single pores to the post-
pectoral curve in the line, thereafter of minute, non-alternating single pores.
Intromittent organ of male with a large, conical tip and a very short basal
portion; a single pair of fleshy crescentic lips ensheathing the lower part of the

tip (fig. 1).

STUDIES ON THE SOUTH AFRICAN CLINIDAE 213

Colour: Mainly green, with darker green, yellow, and iridescent silver longi-
tudinal stripes. Usually a longitudinal silver or yellow stripe across the cheek
and operculum. Fins green, dorsal and anal sometimes with translucent
patches, always translucent patches in the caudal fin. Underparts green. The
colouring is often much like that of the specimen figured in Smith (1949), No.
1001 (middle), as one of the colour variations of Pavoclinus heterodon (C. & V.),
but the specimen is shown with a short caudal peduncle and a reduced inner

(a) (5) (c)

FicureE 2. Left pelvic fin of (a) Pavoclinus litorafontis, (6) Pavoclinus heterodon, (c) Pavoclinus
pavo.

pelvic ray. It is common for species of this genus to have silvery or pearly
markings, often arranged longitudinally.

Table 1. Fin counts and body proportions of the four species of Pavoclinus.

litorafontis heterodon pavo profundus
Dorsal spines .. a: os Fs 29-33 30-35 30-35 30
D. rays .. ¥ vi a5 Shy 6-8 4-6 2-4 4
mE TAYS: . me ai wh as 20-23 21-24 20-23 21
feadinS.L. .. Be ns we 375-5 3°5-4°75 3°5-4°75 3°9
Depth .. o mene AFG 3°25-5 3°75-5 4°75
Upper jaw (% ue head), : 33°5-41°5 30-36 22—36°5 36
Caudal peduncle length (% of bead)s: 58°5-75 26°5-38°5 40-465 36
Ist d. spine (% of std. L. oa fi ws Q-16°5 6°5-11 5°5-10°5 5-7
Eyein head... sha 2. 2°75-4.5 2°5-3°75 3-4 2°75

Discussion

Only one specimen of Pavoclinus profundus was available for comparison,
since this species is known only from the single specimen, taken in relatively
deep water at Knysna (Smith, 1960). However, the specimen examined differs
from P. litorafontis in a number of ways: P. profundus has a rather even dorsal
fin, the first three spines not forming a crest; the inner pelvic ray is reduced;
the caudal peduncle is short; there are fewer dorsal soft rays than in P. litora-
fontis.

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

For a comparison of the fin counts and body proportions of the four
species of Pavoclinus see table 1.

Pavoclinus litorafontis differs from P. heterodon and P. pavo in that (i) the
inner pelvic ray is not reduced, (ii) the caudal peduncle is very elongate,
(iii). the dorsal soft rays are more numerous.

Pavoclinus heterodon and P. pavo both have the inner pelvic ray reduced to — :

half or less than half the length and thickness of the other two rays (fig. 2),
while in P. litorafontis all the specimens had the inner pelvic ray as stout as the
other two and equal to them in length.

From table 1 it can be seen that the caudal peduncle of P. litorafontis is
relatively considerably longer than that of P. heterodon or P. pavo. The caudal
peduncle of P. heterodon is not elongate, the average caudal peduncle length for
most of the South African Clinidae being about 25-35% of the head length.
P. pavo has the caudal peduncle somewhat elongate, but in none of the specimens

in the South African Museum collection did the caudal peduncle length

approach that of P. litorafontis.

P. pavo has 2—4 dorsal soft rays, P. heterodon 4-6, and P. litorafontis 6-8 (see
table 1), thus both P. heterodon and P. litorafontis may have 6 dorsal soft rays.
However, by far the majority of the specimens of P. heterodon examined had 5
dorsal soft rays, while only one of the specimens of P. litorafontis had 6, so that
there is little overlap between the two species in this respect.

P. litorafontis most closely resembles P. heterodon in that both are relatively
large species with a bluntly conical snout, while P. favo is a small species with
an acutely pointed snout. P. litorafontis is more elongate and compressed than
P. heterodon, and has a higher dorsal crest. This is particularly noticeable in the
juvenile specimens of 40-60 mm., which in P. litorafontis have a high crest (12—
16-5°% of standard length), while the crest is lower in juvenile P. heterodon
(7°5-10% of standard length). The differences in caudal peduncle length and
in the development of the inner pelvic ray between these two species are
particularly striking, and the specimens can almost invariably be separated by
the soft dorsal ray count as well. Male and female specimens of P. heterodon of a
similar size to the specimens of P. litorafontis were collected from the Caulerpa

beds at Strandfontein at the same time and were clearly different from them —

in all the respects mentioned above, so that P. lztorafoniis cannot be regarded
as a local or sexual variation or a geographical subspecies of P. heterodon.

It is rather unusual to find an undiscovered species of intertidal fish in a
well-known area. However, no intensive collecting of Clinidae has been done
in this area for many years, and probably none at Strandfontein, and as this
species is well camouflaged, evidently fairly rare, and possibly restricted with
regard to distribution, it is perhaps not surprising that it has only now come to
light. Strandfontein is an extremely interesting area zoologically; during the

present study another clinid, Petraites brevicristatus, previously known from only ~

a few specimens, was found to occur in quite large numbers at Strandfontein,
as well as the brotulid Bidenichthys capensis, also previously considered rare.

STUDIES ON THE SOUTH AFRICAN CLINIDAE 215

Gynutoclinus rotundifrons (Barnard, 1937)
(Figs. 3, 4)

Clinus rotundifrons Barnard, 1937: 63.
Gynutoclinus rotundifrons Smith, 1945: 358.

Material: S.A.M. 18587, one female, 83.5 mm., from kelp in an intertidal pool
at Oudekraal, W. coast of Cape Peninsula (holotype) ; S.A.M. 24009, one male,
43°5 mm., from kelp, intertidal pool, Lambert’s Bay; S.A.M. 24082, one
juvenile male, 26 mm., from kelp, intertidal gully, Lambert’s Bay.

Description: D. XXX-XXXII 8; A. II 22-23; P. 12-14; V.1 3; C. 13. Depth
3°8, 3°95 in larger specimens, 5-2 in juvenile. Head 3.7-4:15 in standard
length; eye 35-4 in head. Upper jaw 42°5-54:5% of head length. Caudal
peduncle length 28-6-36-4°/, of head length ; caudal peduncle depth 28-6—32-2%
of head length.

Figure 3. Gynutoclinus rotundifrons (Barnard), g, 43°5 mm. (Drawn as in life
from specimen contorted on preservation).

First four dorsal spines slightly elevated, particularly in the two smaller
specimens, to form a low, rounded crest, the second and third spines being the
highest. Crest not separated from the rest of the fin by a notch in the membrane.
Profile of dorsal fin undulating. Pectoral fin rounded. Inner pelvic ray reduced,
not more than half of other two rays. Caudal peduncle short, about as broad as
long. Caudal fin subtruncate.

Body compressed, not elongate, more so in the juvenile specimen than in
the larger ones; covered with minute cycloid scales, not imbricating. Head
spherical, inflated, broad, with mucus pores opening on conspicuous papillae.
Eye rounded, somewhat protuberant. A minute, simple papilla over the eye.
Anterior nostril tubular, nasal cirrus large, deeply bilobed. Posterior nostril
conspicuous, surrounded by short, skinny flaps. Mouth rather large. Vomer
toothed. Lips very thin.

Lateral line of single, non-alternating pores throughout, obsolete posteri-
orly in the holotype, but distinct on the caudal peduncle in the smaller speci-
mens. Intromittent organ of male with a long basal portion and a club-shaped

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

tip, ensheathed at the tip by a pair of thin, crescentic lateral lips and a minute
pair of more or less confluent dorsal lips (fig. 4).

Colour: (a) Male, 43-5 mm. Ground colour pale brown with about seven
darker brown irregular cross-bars, edged with iridescent blue in fine broken
lines. A very dark brown narrow vertical line behind and above the pectoral ©
axil, and another at the end of the caudal peduncle. Cross-bars of body con-
tinued on to the dorsal fin, with translucent patches between them; translucent
areas with fine black dots. Dorsal fin uniformly dark brown posteriorly; a

(a) Ventral view. (6) Lateral view.

FicureE 4. Intromittent organ of Gynutoclinus rotundifrons.

single small translucent patch at the base near beginning of dorsal soft rays.
Caudal fin translucent with very faint brown cross-bars, darkening at the
margins. Pectoral fin translucent with four very fine dark brown cross-bars, the
proximal one curved. Anal fin mainly dark brown, with two translucent
patches near the base. Pelvic fins light brown with dark brown cross-bars.
Head mainly light brown below. A dark brown stripe from the eye forwards
in front of cheek to angle of jaw. Head above and opercular region deep pink.
Snout pink. with a darker pink bar between the eyes and another above the ©
upper lip, a fine darker pink line down the middle. Eye silvery with golden- ©
brown radii. Chin and lips light brown mottled heavily with darker brown. —
Branchiostegal membranes and jugular region silvery grey with fine black
speckling. Belly silvery with a golden-brown sheen. Intromittent organ of male
greyish, with fine black speckling.

(b) Juvenile, 26 mm. Whole body and head yellow, without cross-bars or —
markings, underparts lighter yellow. Dorsal fin with alternating pink and
translucent patches. Anal fin yellow. Caudal and pectoral fins translucent. —
Pelvic fins yellow with brown cross-bars.

Discussion

Neither Barnard (1937) nor Smith (1945) noticed the supra-orbital
papilla. This is not surprising, as it is very small, and in the holotype had

STUDIES ON THE SOUTH AFRICAN CLINIDAE 217

shrunk owing to desiccation in the preservative; in life it is erect and quite
distinct even to the naked eye. None of the other South African Clinidae
described have a simple supra-orbital papilla; either there is a multifid supra-
orbital tentacle, or a supra-orbital outgrowth is completely lacking.

Gynutoclinus rotundifrons is apparently a weed-dwelling species, as all three
specimens so far found were taken from amongst fronds of kelp. It is the only
species which has not been recorded east of Cape Point, but if it is as rare as it
appears to be, it is possibly more widely distributed.

ACKNOWLEDGEMENTS

Acknowledgement must be made to Dr. F. H. Talbot of the South African
Museum for advice and help; to Professor J. L. B. Smith of the Department of
Ichthyology, Rhodes University, for the loan of the unique specimen of
Pavoclinus profundus; to my husband Mr. M. J. Penrith of the South African
Museum, who collected most of the specimens studied, and to Mr. C. Berrisford
for the Onrus River mouth specimen.

The Trustees of the South African Museum acknowledge the award of a
grant by the Council for Scientific and Industrial Research for the publication
of this paper.

SUMMARY

A new species of clinid, Pavoclinus litorafontis (Pisces: Clinidae), is described.
Gynutoclinus rotundifrons (Barnard), previously known only from a unique
specimen, is redescribed.

REFERENCES

Barnarp, K. H. 1937. Further notes on South African Marine Fishes. Ann. S. Afr. Mus. 32 (2):
41-67.

Smitru, J. L. B. 1945. The fishes of the family Clinidae in South Africa. Ann. Mag. nat. Hist.
(11) 12: 535-546.

Smiru, J. L. B. 1949. The sea fishes of Southern Africa. Central News Agency Ltd., South Africa.

Smitn, J. L. B. 1960. A new species of South African clinid fish. Ann. Mag. nat. Hist. (13) 3:

689-691.

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Plate V

Ann. S. Afr. Mus. Vol. XLVIII

‘(yyueg “ff ‘Ww :ydes80j,0yd) gs] qISsIA Jou ore seare poquoursidun oy} 23~7d sty) UI Ysnoyye “uy
9Y} JO YISUZT 9y1 NOYSno1y) snonuNuod si sXhe1 puv sourds [esiOp 9Y} Us9MJoq 9UBIqUIOU 9Y J,

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INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TaBLeE oF ConTENTs and SumMARyY. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

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REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers)
first and last, preceded by a colon (without prefix ‘p’). Thus:

Smitu, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmitH, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

hy ay Lape to umn

MUS. C SOMP. ZOOL
LIBRARY

NOV 19 i900

F. W. GESS

CONTRIBUTION TO THE KNOWLEDGE OF ‘THE
SOUTH AFRICAN SPECIES OF THE GENUS
CERAMIUS LATREILLE (HYMENOPTERA:

MASARIDAE)

September 1965 September
Volume 48 ~~ Band

Part. “41 Deel

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

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
(Cash with order, post free)

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word uitgegee in dele op ongereelde tye na beskikbaarheid

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OUT OF PRINT/UIT DRUK

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Printed in South Africa by In Suid-Afrika gedruk deur
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Court Road, Wynberg, Cape | Courtweg, Wynberg, Kaap

CONTRIBUTION TO THE KNOWLEDGE OF THE SOUTH AFRICAN
SPECIES OF THE GENUS CERAMIUS LATREILLE (HYMENOPTERA:

MASARIDAE)
By MUS OMP ZOOL,
F. W. GEss
; Nid j (GOO
South African Museum, Cape Town
OARVARL
( With 1 map) UNIVERS!
CONTENTS
PAGE
Introduction . . 219
Descriptions of oa oat Be pahiion ecieds: . 219
Discussion of the Distribution of the Genus Ceramius . 228
Summary i ; : ; ’ ‘ : . 230
Acknowledgements . : : é : : Hy Rgr
References A ; : F : ; ‘ HST
INTRODUCTION

While arranging the Masaridae in the collection of the South African
Museum according to the recent revision of the family by Richards (1962), it
was found that this material, which was unfortunately not seen by Richards,
supplemented his account in several i oca especially in the genus Ceramius
Latreille.

In the present paper, dealing with the above genus, the hitherto unknown
@ of clypeatus Richards and ¢ of metanotalis Richards are described; the ¢
assigned by Richards to peringueyi Brauns is removed from that species and
together with a matching 9 is described as a new species, richardsi, and lastly,
rex de Saussure is resurrected. The locality records of all the specimens in the
collection are given in the hope that these may aid in presenting a more
complete picture of the distribution of the various South African species.
Finally the distribution of the genus as a whole is discussed in the light of the
biology of these wasps.

The sequence of species followed is nearly the same as that adopted by
Richards, and this paper closely follows the presentation set by the latter in his
revision. South African Museum is here abbreviated to S.A.M.

DESCRIPTIONS OF SPECIES AND DISTRIBUTION RECORDS

Ceramius cerceriformis de Saussure

Ceramius cerceriformis de Saussure, 1853: xxi, §; Richards, 1962: 97, 3; 9.
Ceramius (Ceramioides) cerceriformis de Saussure, 1854: pl. 4, nee 1, 0; 1855: 72, 3.
Cerceris vespiformis de Saussure, 1855: 79, @.

219
Ann. S. Afr. Mus. 48 (11), 1965: 219-231, I map.

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

No specimens were found in the South African Museum collection that
could be assigned to this species.

Ceramius schulthessi Brauns

Ceramius schulthessi Brauns, 1902: 182, 2; Brauns, 1913: 196, pl. 2, fig. 6, g, 2; Richards, 1962: gg.

Specimens examined: Cape Province: Willowmore, no date, 399 (one
marked as co-type), I. xii. 1904, 9, xi. 1917, 2 99 (Dr. Brauns) ; Augusfontein,
Calvinia, ix. 1947, 3 99 (S.A.M. Staff); Oudtshoorn, Zebra, x. 1951, 3 99
(S.A.M. Staff); Touws River—Hondewater (18 miles E. of Touws River), xii.
1962, 2 (S.A.M. Staff); Bloutoring Station (30 miles E. of Touws River), xii.
1962, 4 9° (S.A.M. Staff).

Namaqualand: Between Kamieskroon and Springbok, x. 1939, 4 go
4 99 (S.A.M. Staff); Wallekraal, x. 1950, 2 (S.A.M. Staff).

Ceramius peringueyi Brauns

Ceramius peringueyi Brauns, 1913: 194, 2; Richards, 1962: 100 [9 only].
[non] Ceramius peringueyi Brauns, Richards, 1962: 100, ¢ [ = richardsi sp. n.]}.

Specimens examined: Cape Province: Stellenbosch, x. 1888, holotype 2
(L. Peringuey) (Transvaal Museum) ; Het Kruis, x. 1947, 3 99 (S.A.M. Staff) ;
Paleisheuwel, xi. 1948, 9 (S.A.M. Staff).

Ceramius clypeatus Richards
Ceramius clypeatus Richards, 1962: 99, ¢.

Q. Black; raised disk of clypeus to a variable degree, inner orbits at
deepest portions of ocular sinus, spot of variable size on tempora, usually a
small spot on prepectus, variably sized spot at apex of scutellum, rarely a
minute spot on propodeal spine, occasionally a short basal streak along outer
side of fore tibiae and rarely indicated basal spots on outer side of mid and
hind tibiae, spots at sides of gastral tergites 1-4 and usually also on 5, sometimes
produced inwards on 2-5 either forming narrow continuous or interrupted
bands, creamy-white to yellowish.

Underside of antennal joints 4-12 though sometimes fewer, legs except
coxae, trochanters and base of femora, orange. Wings light fuscous, veins brown.
Length 14-16 mm., length of fore wing 10°5—11°5 mm., hamuli 18-24.

Altogether very similar indeed to the male, the peculiarly modified
clypeus, for example, except in its greater width in the female, being virtually
identical in the two sexes. The chief secondary sexual structural differences are
the following: eyes somewhat smaller, further apart, interocular distance at
level of sockets twice length of scape (without radicle); antennal scape less
curved and less robust, segment 3 shorter, only half as long as scape (without
radicle), 4-10 progressively wider, 10 about 14 times as wide as long, 11-12 a
little narrower; fore trochanter simple.

CONTRIBUTION TO KNOWLEDGE OF CERAMIUS LATREILLE 221

From the specimens examined it appears that in both sexes the longer spur
of the hind tibia is not originally simple but may become so by the loss or
wearing away of the fine spines situated near the tip of the spur. While the
trifid condition is common, there is a specimen with a 4-spined spur and others
with all graduations to the simple condition.

Specimens examined: Cape Province: Clanwilliam, Nardouw, ix. 1941,
II gd, 2 99 (S.A.M. Staff); Het Kruis, x. 1947, 7 3d (S.A.M. Staff) ; Citrusdal
Dist., xi. 1948, 5 92 (S.A.M. Staff); 4 miles S. of Clanwilliam, ix. 1961, 2
(S.A.M. Staff).

Ceramius richardsi sp. n.
Ceramius peringucyt (non Brauns) Richards, 1962: 100 [partim, $ only].

The ¢ of this new species, which has been adequately described by
Richards, was unfortunately assigned by him to peringueyi Brauns to which
species it most certainly does not belong. This misidentification was made
obvious by the discovery in the collection of the South African Museum of a
hitherto undescribed 9 which closely matches Richards’ ¢ in all important
characters such as the unusual form of the clypeus. That this latter character
is of value in associating the sexes is demonstrated by the discovery, also in the
South African Museum collection, of the hitherto undescribed @ of clypeatus
Richards, a closely related species, the ¢ of which was used by Richards as a
comparison in describing what he thought was the ¢ of peringuey: Brauns. In
clypeatus Richards the form of the clypeus is common to both sexes. I have
pleasure in naming the ¢ under consideration and the here described matching
Q after the author, Professor Richards. The true 3 of peringueyi Brauns is thus
still unknown.

2. Black; spot on mandibles near base, large discal spot on clypeus, inner
orbits to top of ocular sinus, streak at top of tempora, two widely separated
streaks on hind margin of pronotum, dot at apex of scutellum, moderately
large spot on prepectus, small anterior spots at base of fore tibiae and end of mid
femora, narrow lateral spots on gastral tergites 1-5 (tergite 6 hidden) produced
inwards and forming narrow incomplete bands on tergites 2-3 and centrally
widened ones on 4-5, creamy-white.

Antennal segments 4-12 beneath, portions of mandibles, anterior vertical
portion of clypeus beneath disk, whole of legs except coxae of all legs and
_ trochanters and bases of femora on mid and hind legs, reddish.

Wings fuscous, venation dark brown.

Length 15 mm., length of fore wing 10 mm., hamuli 21.

Head, thorax and gaster with long, rather dense, whitish hairs. Mandibles
strongly striate distally, ending in two large blunt teeth with a smaller more
dorsal one. Clypeus elongate, strongly raised, anteriorly falling at right angles
towards ventral margin; length of vertical part 3 length of disk; from just
below point of inflection two small curved teeth project upwards; ventral

1

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

margin produced, somewhat lamellate, truncate and slightly emarginate; disk
narrower at apex than at base; anterior-lateral margins raised and slightly
lamellate. Antennal sockets separated by 34 times their diameter; interocular
distance at level of sockets twice the length of scape (without radicle) ; total
length of scape 34 times greatest width (at apex); segment 2 very short, 3
slightly shorter than 4-+5+6, 4-12 all of about same length, 4-10 becoming
gradually and progressively wider, 11 slightly narrower than 10, 12 narrower
still, rounded at apex. Disk of clypeus and small, roughly triangular area above
it and between antennal sockets almost smooth; frons punctured; posterior
ocelli a little in front of hind margin of eyes; distance between eye and posterior
ocellus: distance between posterior ocelli = 10 : 7; occiput behind eye some-
what wider than interocellar distance; occipital keel present.

Thorax with fairly coarse, separated punctures, the interstices shining,
about as wide or wider than the punctures. Mesoscutum shining; prescutal
furrows deep over their entire length, especially so behind. Raised disk of
scutellum with rounded edges and without a central keel.

Metanotum with a central prominence, lateral depressions fairly deep but
open. Tegula smooth and shining, only the base punctured. Propodeum with
fairly long blunt spines; posterior surface almost flat; spiracle with anterior
margin strongly produced backwards. Fore tibial spur regularly curved, tip
somewhat recurved. Mid and hind tibiae with two spurs, longer spur of hind
tibia simple. Inner keel of hind coxa present on proximal half only. Claws
simple. Gaster shining; tergite 1 constricted posteriorly, more finely punctured
than thorax; tergites 2-6 becoming progressively finer punctured, 2 somewhat
contracted at base.

Specimens examined: Cape Province: Clanwilliam, ix. 1928 (Dr. Brauns),
Holotype g (Transvaal Museum, Pretoria); Paleisheuwel, xi. 1948 (S.A.M.
Staff), Allotype 9 (S.A.M.).

The female has ten mites present on the axillae and lateral depressions
of the metanotum.

Ceramius nigripennis de Saussure

Ceramius (Paraceramius) nigripennis de Saussure, 1854: pl. 3, fig. 4, 9; 1855: 60.
Ceramius nigripennis de Saussure, Richards, 1962: 100.

Ceramius hesset Turner, 1935: 296, 3, 2.

[non] Ceramuis nigripennis of other authors.

Specimens examined: Namaqualand: Kamieskroon, ix. 1930, ¢ holotype,
allotype of C. hessei Turner, 24 99 (S.A.M. Staff) ; between Kamieskroon and
Springbok, x. 1939, ¢, 2 (S.A.M. Staff) ; Bowesdorp, xi. 1931, 3, 1x. 1941, 2 gd
(S.A.M. Staff); Outiep, Garies, ix. 1953, 5 (J. du Toit).

Ceramius toriger von Schulthess
Ceramius toriger von Schulthess, 1935: 383, 2; Richards, 1962: 101.

Specimens examined: Cape Province: Augusfontein, Calvinia, ix. 1947,
19 99 (S.A.M. Staff); Tankwa Karoo, Waterval, xi. 1952, 10 99 (S.A.M. Staff) ;
5 miles N. of Nieuwoudtville, ix. 1961, 2 99 (S.A.M. Staff).

oe ee

;

CONTRIBUTION TO KNOWLEDGE OF CERAMIUS LATREILLE 223

A single female from Namaqualand: Knersvlakte, x. 1950 (S.A.M. Staff),
differs from the description of this species and from the above listed specimens
in that the light coloured markings are more extensive and are yellow, not ivory.
Structurally there are no differences. The distribution of the yellow markings
is given below. |

Black; large spots on basal half of mandibles, clypeus (except for oblique
black streaks arising from bottom of antennal sockets, infuscation between ends
of these streaks and lateral margins, and ferruginous lateral and apical margins),
a large pentagonal spot enclosing a black triangle on frons above clypeus, inner
orbits to centre of ocular sinus, spots behind eyes, uninterrupted pronotal band
extending onto sides, large spots on mesopleura, sides of mesonotum next to
tegulae, a small median spot in posterior region of mesoscutum, posterior
quarter of scutellum and centre of metathorax, streaks on axillae, whole of
propodeum behind level of spiracles, broad apical bands widened laterally on
tergites 1-5, tergite 6 except for depressed area, whole of sternite 2, wide apical
bands on sternites 3 and 4, portions of femora and tibiae, yellow.

In addition, the antennae are much lighter in colour, the scape being
largely yellow.

Ceramius braunsi Turner
Ceramius braunsi Turner, 1935: 294, 5, 9; Richards, 1962: ror.

Specimens examined: Cape Province: Olifants River, between Citrusdal
and Clanwilliam, x—xi. 1931, ¢ holotype, 2 allotype, 3 gd, 32 9? (S.A.M.
Staff); Pakhuis Pass, Clanwilliam, ix. 1942, 9 (S.A.M. Staff); 4 miles S. of
Clanwilliam, ix. 1961, 2 gg, 2 99 (S.A.M. Staff).

Ceramius jacoti Richards

Ceramius jacoti Richards, 1962: 101, d, @.
Ceramius nigripennis (non de Saussure) Brauns, 1913: 201, pl. 2, fig. 3, 3.

Specimens examined: Cape Province: Hex River, i. 1884, 2; Oudtshoorn,
Zebra, x. 1951, 17 92 (S.A.M. Staff); Ouberg Pass, S.E. of Touws River, xi.
1962, 9 (S.A.M. Staff); Verkeerde Vlei, Touws River—Hottentots Kloof, xii.
1962, 11 99 (S.A.M. Staff); Constable, xii. 1962, 9 (S.A.M. Staff); Touws
River—Ouberg Pass, xii. 1962, 9 (S.A.M. Staff); 8 miles N.E. of Touws River,
xii. 1962, 2 (S.A.M. Staff) ; Bloutoring Station, 30 miles E. of Touws River, xii.
1962, 98 99 (S.A.M. Staff).

Ceramius beyert Brauns
Ceramius beyeri Brauns, 1903: 69, do, 9; Richards, 1962: 102, figs. 105-9.

Specimens examined: Cape Province: Willowmore, no date, 9 cotype,
2 99 (Dr. Brauns) ; Somerset East, 25-30. xi. 1930, 3, (R.E. Turner) ; Nieuveld

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

Escarpment, Rietvlei, 1. 1949, 9 (S.A.M. Staff); Tankwa Karroo, Renoster
River, xi. 1952, 116 99 (S.A.M. Staff); Constable, xii. 1962, 3 99 (S.A.M.
Staff); Matroosberg Station, xii. 1962, 9 (S.A.M. Staff); Touws River—
Hondewater (18 miles E. of Touws River), xii. 1962, 2 99 (S.A.M. Staff);
Bloutoring Station, 30 miles E. of Touws River, xii. 1962, 2 99 (S.A.M. Staff).

Ceramius damarinus Turner
Ceramius damarinus Turner, 1935: 293, 3, 9; Richards, 1962: 102.

Specimens examined: S.W. Africa: Ongandjera, ili. 1923, type g, 2 gd
cotypes, 3, type 9 (S.A.M. Staff); Kamanyab, iii. 1925, 2 gg (S.A.M. Staff).

Ceramius lichtensteini (Klug)

Gnatho lichtensteinii Klug, 1810: 36, 38, pl. 1, fig. 3, e and f.

Ceramius lichtensteinii (Klug), Klug, 1824: 225; de Saussure, 1855: 73, 2; Brauns, 1913: 193;
Bequaert, 1928: 145; Richards, 1962: 102.

Ceramius macrocephalus de Saussure, 1854: pl. 3, fig. 2, 2; Brauns, 1903: 65, 68, g, Q.

Ceramius rufomaculatus Cameron, 1906: 325, 9.

[non] ?Ceramius rex de Saussure, Richards, 1962: 102.

Specimens examined: Cape Province: Willowmore, 15. xl. 1899, dg, 10. 1.
1900, 2 (Dr. H. Brauns); Dunbrody, 1go00, g, 2 (Rev. O’Neil), 1901, 9 (J. A.
O’Neil) ; Uitenhage, Dunbrody, no date, 9 (Rev. O’ Neil); Pearston, 1905, 2 99
(Dr. Broom) ; Aberdeen, xi. 1935, 2 (S.A.M. Staff) ; Tankwa Karroo, Waterval,
X1. 1952, 21 9g, 9 99 (S.A.M. Staff); Bloutoring Station, 30 miles E. of Touws
River, xii. 1962, 2 99 (S.A.M. Staff).

South West Africa: Damaraland, 1890, 2 (R. Lightfoot). It is very doubt-
ful whether this last record is correct.

Ceramius caffer de Saussure

Ceramuus caffer de Saussure, 1855: 76, 2; Richards, 1962: 104, figs. 110-113, 115, d; Q.
Ceramius consobrinus de Saussure, 1855: 77, 2; Brauns, 1913: 198, 3, 2.

Specimens examined: Cape Province: Stellenbosch, x. 1888, 3 gg, 2 29,
no date, 9 (L. Peringuey), 1908, 26 99 (C. P. Lounsbury).
All the specimens carry mites in the acarinarium.

Ceramius metanotalis Richards
Ceramius metanotalis Richards, 1962: 106, fig. 114, 9.

¢. Black; whole disk of mandibles, labrum, clypeus except very narrow
margins, roughly rectangular area above and between antennal sockets
(separated from clypeus by a narrow black line at suture and produced slightly
upwards at inner margins of antennal sockets and with a small black tubercle
at centre), narrow orbits up to centre of eye emargination, small occipital spots
behind upper portion of eyes, underside of antennal scape, underside of 2nd

———— $4 4 oH ee bo 6 ee Se eee ee Ae

CONTRIBUTION TO KNOWLEDGE OF CERAMIUS LATREILLE 225

segment and basal two thirds of 3rd segment, pronotal band (narrowly inter-
rupted at centre and produced on to humerus and to tegula), a minute spot on
postero-lateral corner of mesoscutum and inner corner of axilla, sometimes a
small spot at tip of scutellum, sometimes a very narrow streak on lower part of
axilla, a spot at angles of propodeum, a single dorsal spot on mesopleuron, legs
(except dorsal portions of coxae and trochanters, hind surface of middle and
hind femora and tips of tarsal segments 3, 4 and 5 of above legs), inner margin
of tegula, distal part of humeral plate, large spot on each side of first gastral
tergite (not or only narrowly joined at hind margin), bands on tergites 2-5
(strongly widened at sides and plano-convex medially at hind margins),
_ posterior portion of tergite 6 and small spot at sides of tergite 7, almost all of
sternites 2—5 and sides of sternites 6 and sometimes 7, pale yellow.

Antennal flagellum except some black dorsal suffusion on all segments
bar the last, ferruginous. Wings faintly brownish, veins brown. Length 17, 18, 19
mm., length of fore wing 12, 13, 13 mm., hamuli (20, 21), 22 (20, 22).

The chief secondary sexual structural differences are the following: Sides
of clypeus more converging ventrally; margin narrower and very slightly
concave. Eyes larger, a lot closer together; interocular distance at level of
sockets = 1-5 times length of scape (without radicle) (2-2 in 2). Antennal scape
strongly widened, curved as in 9; segment 3 flattened in side view, narrow
except at apex, a little shorter than scape (without radicle) and slightly longer
than 4+5+6; 4-11 becoming progressively wider; 12 forming a powerful,
long, flattened and fairly wide hook, curved at base and at apex; inner surface
of hook with a low, off-central, longitudinal carina on distal half; 8-10 with a
shining, slightly raised transverse swelling beneath; entire underside of 11
swollen and shining. Fore trochanter with a very large, crescentic lobe, curving
outwards, outer edge transparent, somewhat sinuate. Segment 1 of mid tarsus
longer, curved; 2-5 strikingly laterally compressed, wide in side view; 3 and 4
almost oval in outline. Gaster with tergite 7 elongate, apically with a wide,
shallow, angular emargination; sternite 3 with disk transversely swollen,
raised on each side into a mound ending in a blunt tubercle, without raised
preapical lateral keels; sternite 4 unmodified in structure; disks of both sternites
3 and 4 covered with dense white pubescence; sternites 7 and 8 very similar to
those of caffer de Saussure; prominence on 7 more pronounced.

Specimens examined: Cape Province: Bulhoek, Klaver—Clanwilliam,
X. 1950, 3 gd, 27 29 (S.A.M. Staff).

Six of the 27 females have mites in the acarinium. The three males are free
of mites, however.

Ceramius rex de Saussure

Ceramius rex de Saussure, 1855: 75, 9; Turner, 1935: 290.
Ceramius lichtensteinii (non Klug), Richards, 1962: 102.

A single female specimen from Namaqualand: Klipvlei, Garies, XI. 1931
(S.A.M. Staff), believed to be this species, bears the label ‘Ceramius rex Sauss. 9,

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

det. Turner’. Concerning this specimen, Turner (1935: 290-1) correctly stated
that it corresponds to the description of rex de Saussure, but measures only 19
mm., not 24. His further statement that it is allied to lichtensteinii (Klug) is
incorrect for the specimen is entirely different from the latter species, being
allied to caffer de Saussure and metanotalis Richards, though distinct from both
of these. Richards (1962: 102) treated rex de Saussure as a doubtful synonym of
lichtensteinit (Klug) but did not see the specimen now under consideration
which appears to be the true rex de Saussure. The following is a description of
this specimen.

Q. Black; clypeus, a large, broad pentagonal spot between the antennae
and above the clypeus (from which it is separated by a narrow black line at
suture), a narrow streak on inner margins of eyes below, streak in ocular sinus,
spot on mandibles near base, underside of scape, spots behind eyes (joined
along occipital margin), pronotal band produced onto humerus and to tegula
(leaving a triangular black area on side), spot at postero-lateral corner of
mesoscutum, inner corner and streak on lower part of axilla, posterior part of
scutellar disk and flap of posttegula, central area of metanotum and portion
below acarinarium, whole of propodeum (except for black region anterior to
spiracles on sides, black lateral streaks on posterior surface near junction with
metanotum and two black marks just above oriface), a large spot on meso-
pleuron, spots on anterior surface of coxae 2 and 3, portions of trochanters,
greater part of femora and tibiae of all legs, inner margin of tegula, wide
posterior bands widening on sides on gastral tergites 1-5, almost whole disk of
6, two small streaks on posterior area of sternite 1, whole of sternites 2-5 except
extreme base of 2, pale yellow.

Anterior margin of clypeus, distal half of mandibles, antennal flagellum
(except some black dorsal suffusion), some suffusion on tibiae and entire tarsi,
suffusion on gastral sternite 6, ferruginous. Wings faintly brownish, veins brown.

Length 19 mm., length of fore wing 13:8 mm., hamuli 23.

Head, thorax and base of first gastral segment with long whitish hairs,
densest on head, dense on pleura, sides and angles of propodeum;; rest of gaster
with very fine tomentum-like pubescence.

Clypeus one third longer than wide at ventral margin, moderately coarsely
punctured; ventral margin truncate, sharply angled, with a fairly wide, smooth
border. Antennal sockets separated by 4:1 times their diameter; interocular
distance at level of sockets twice the length of scape (without radicle) ; scape
(without radicle) almost 3 times as long as greatest width; segment 2 very short,
broader than long; 3 half as long as scape (without radicle), twice as long as
greatest width, of same length as 4-++-5-+6; 4-10 progressively wider; 10 about
twice as wide as long; 11 and 12 a little narrower. Frons dull, finely punctured;
POL : OOL = 1 : 1-7; posterior ocelli about half ocellar diameter in front of
hind margin of eyes; occipital keel absent. Pronotum with spiracular lobe well
defined by a furrow; lateral furrow partially obscured by legs, apparently
rather weak; anterior margin apparently not markedly keel-like; dorsal surface

CONTRIBUTION TO KNOWLEDGE OF CERAMIUS LATREILLE 227

finely and closely punctured. Mesoscutum dull, finely, confluently punctured;
prescutal furrows well marked over their entire length, especially behind;
parapsidal furrows fairly weak; median notal suture marked anteriorly by a
smooth, shining, unpunctured line, posteriorly marked by a furrow. Suture
between axilla and scutellum with a pit ventrally. Raised disk of scutellum
convex, in profile smoothly arcuate to posterior margin, with a well marked,
raised, median keel and moderate lateral keels posteriorly, dull, punctured like
mesoscutum; lateral declivities more shining, feebly punctured. Metanotum
with central prominence smooth; acarinaria present laterally; entrance to
acarinarium large, larger than that of caffer de Saussure, about 4 times longer
than wide, width nearly constant throughout, only very slightly wider laterally;
part of metanotum in front of slit of about same width as latter. Mesopleuron
dull, with very close microscopic punctures and sparse coarse ones. Meta-
pleuron dull, with similar but less pronounced puncturation and with some
microscopic transverse striae on dorsal half; as in metanotalis Richards with a
marked bean-shaped depression dorsally in the posterior boundary. Propodeum
with angles rounded; posterior surface shallowly concave; spiracle long and
narrow with anterior margin produced backwards. Fore tibial spur regularly
curved, slightly recurved at apex; mid tibia with two spurs; larger spur of hind
tibia bifid. Claws with a small tooth. Gaster dull, with exceedingly minute and
close punctures, smaller than those of metanotalis Richards, and with very fine
tomentum; larger punctures completely absent from all tergites; tergite 1 very
transverse, nearly 34 times wider than long, a little constricted posteriorly,
with a hyaline border; tergite 2 constricted anteriorly, maximum width 14
times greater than maximum width of tergite 1; tergite 3 of equal width
anteriorly as tergite 2 posteriorly; tergites 3-6 becoming progressively narrower ;
sternites nearly flat, very closely and finely punctured.
Four mites are visible in the acarinaria.

Ceramius bicolor (Thunberg)

Philanthus bicolor Thunberg, 1815: 131, 289 [].

Ceramius karooensis Brauns, 1902: 282, 9, 373, 3.

Ceramius bicolor (Thunberg), Schulz, 1912: 68-69, 99; Bequaert, 1929: 79; Richards, 1962:
115, figs. 131, 132.

Specimens examined: Cape Province: Aberdeen, xi. 1935, 3 dd; 8 92
(S.A.M. Staff); Murraysburg Dist., xi. 1935, 6 dd, 2 92 (S.A.M. Staff);
Augusfontein (Calvinia), ix. 1947, g, 2 99 (S.A.M. Staff) ; Oudtshoorn, Zebra,
x. 1951, ¢ (S.A.M. Staff); Moordenaars Karoo, Lammerfontein, x. 1952, 3 2?
(S.A.M. Staff); Willowmore—Vondeling, x. 1952, 2 (S.A.M. Staff); Rooinek
Pass, x. 1952, ¢ (S.A.M. Staff); Tankwa Karoo, Waterval, xi. 1952, 25 dd;
85 99 (S.A.M. Staff) ; Touws River-Hondewater (18 miles E. of Touws River),
xii. 1962, 5 dd, 5 99 (S.A.M. Staff); Bloutoring Station (30 miles E. of Touws
River), xii. 1962, 9 99 (S.A.M. Staff).

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ceramius linearis Klug

Ceramius linearis Klug, 1824: 227, 3; Richards, 1962: 115, figs. 128-130, 113 (b).
Ceramius (Paraceramius) linearis de Saussure, 1855: 71, 3.
Ceramius fumipennis Brauns, 1902: 275, 3, 2; Bradley, 1922: 397 (correction of Brauns).

_ Specimens examined: Cape Province: Algoa Bay, 25. xii. 1898, 3, 2
(Dr. H. Brauns); Dunbrody, 1g00, 9 (Rev. O’Neil); Aberdeen, xi. 1935, 9
(S.A.M. Staff).

Ceramius capicola Brauns
Ceramius capicola Brauns, 1902: 278, 3, 9; Bradley, 1922: 397; Richards, 1962: 117.

Specimens examined: Cape Province: Willowmore, 1. x. 1899, 2 (Dr. H.

Brauns); Somerset East, 25-30. xi. 1930, 2 99, g (R. E. Turner); Aberdeen, —

X1. 1935, 736d; 3 9P (S.A.M. Staff); Murraysburg Dist., xi. 1935, 5 dd, 11 29
(S.A.M. Staff); Teekloof, Fraserburg Dist., xi. 1935, 5 dd, 2 (S.A.M. Staff);
Oukloof, Fraserburg Road, xi. 1936, 2 99 (S.A.M. Staff); Richmond Dist., xi.
1939, ¢ (S.A.M. Staff); Oudtshoorn, Zebra, x. 1951, 5 dd, 9 (S.A.M. Staff).

Ceramius socius Turner
Ceramius socius Turner, 1935: 297, 5d, 2; Richards, 1962: 117, figs. 113 (a), 134-136.

Specimens examined: Cape Province: Montagu, x—xi. 1919, 2 (Collector’s
name not recorded) ; Worcester, ix. 1921, 2 dg, 2 99 (R. E. Turner) ; Verkeerde
Vlei (Touws River—Hottentots Kloof), xii. 1962, 3, 13 92 (S.A.M. Staff);
Constable, xii. 1962, 3 99 (S.A.M. Staff); Matroosberg Station, xii. 1962, J,
® (S.A.M. Staff); 8 miles N.E. of Touws River, xii. 1962, 9 (S.A.M. Staff).

DISCUSSION OF THE DISTRIBUTION OF THE GENUS Ceramius

The genus Ceramius occurs in two widely separated geographical regions
in the Old World, one being the extreme south-west of the Ethiopian Region
and the other that portion of the Palaearctic bordering on the Mediterranean
Sea. Thus, in the Ethiopan Region, the genus is in the main restricted to the
Cape Province where it is found in Little Namaqualand, the South Western
Cape, the Little Karroo and the southern parts of the Great Karroo. It does not
extend further east than the Great Fish River. Outside the Cape Province, one
species (damarinus Turner) is endemic to South West Africa (Kaokoveld and
Ovamboland), and one Eastern Cape species (capicola Brauns) has been recorded
from two localities (Kroonstad and Thaba Nchu) in the Orange Free State.
This distribution has been plotted by means of a 4° square grid system on an
outline plotting map (Map 1). In the Palaearctic, the genus occurs in Algeria,
Morocco, Gibraltar, Portugal, Spain, the: south of France, Greece, Turkey,
Russian Armenia and probably Israel. |

Climatically the above areas are characterised by a predominantly
winter rainfall while the vegetation is generally low and semi-desert in nature.

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Mar |. Distribution of the genus Ceram

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

In South Africa, Ceramius favours those parts of the Karroid and False Karroid
areas (as defined by Acocks, 1953) which lie within the winter rainfall region,
though two of the nineteen species, damarinus Turner and capicola Brauns, have
been recorded from summer rainfall regions.

A study of the records shows that in the southern hemisphere no species
flies earlier than September or later than March, while in the northern hemi-
sphere no species flies earlier than March or later than August. In other words,
flight is restricted to the dry spring and summer months succeeding the winter
rainy season.

Having ascertained where and when Ceramius occurs it seems of interest to
examine the reasons. In this connection the biology of the genus has to be taken
into account. As stated above, the adults fly during the dry months of the year
succeeding the winter rainy season. During this time mating takes place and
burrows lined with mud pellets are built in the ground and surmounted by
mud chimneys. In these the eggs are deposited and, in the South African
species at least, it is recorded by Brauns (1910, cited by Richards, 1962: 29)
that the female continues to provision the young with pollen and nectar until
these larvae are ready to pupate, when she seals the opening to the nest with a
plug of mud. There is only one generation per year.

Two requirements for the successful run of the life-history are immediately
apparent: there must be a copious supply of pollen and nectar at the time the
young are being reared and there must be an extended dry period, not only to
allow the adult to collect this pollen and nectar but also on account of the fact
that the burrows in the ground remain unplugged during the larval stage.
Despite some possible protection afforded by the chimneys, the larvae would
have little chance of survival in case of heavy rain.

The combination of winter rainfall followed by an abundance of suitable
flowers rich in pollen and nectar during an extended dry period is obviously
the clue to the distribution of the genus Ceramius. The only areas fulfilling these
requirements are those in which Ceramius occurs. In this connection it is likely
that the flowers visited by Ceramius will prove to be low-growing Compositae and
mesembryanthemums (Aizoaceae) which, following the winter rains, are such
a striking feature of the semi-desert areas inhabited by Ceramius in South Africa.

SUMMARY

The account of the South African species of Ceramius included in the

revision of the Masaridae by Richards (1962) is supplemented by the study of

the material in the South African Museum collection.

One new species, richardsi, and the hitherto unknown @ of clypeatus Richards
and ¢ of metanotalis Richards are described as is also a specimen thought to be
rex de Saussure. The locality records of all the specimens in the collection are
given and the distribution of the genus in South Africa is mapped. Lastly the
distribution of the genus as a whole is discussed with reference to the biology of
these wasps.

ba 6 ow

CONTRIBUTION TO KNOWLEDGE OF CERAMIUS LATREILLE 231

ACKNOWLEDGEMENTS

I wish to thank Dr. G. van Son of the Transvaal Museum, Pretoria for the
loan of type material of peringuey: Brauns, clypeatus Richards and metanotalis
Richards. The Trustees of the South African Museum are grateful to the
Council for Scientific and Industrial Research for a grant to publish this paper.

REFERENCES

Acocks, J. P. H. 1953. Veld types of South Africa. Mem. bot. Surv. S. Afr. 28: i-iv, 1-192.

BEQuaAERT, J. 1928. A study of certain types of Diplopterous wasps in the collection of the
British Museum. Ann. Mag. nat. Hist. (10) 2: 138-176.

BEQuAERT, J. 1929. A new Pscudomasaris from California, with some considerations on the
Masarid wasps (Hymenoptera). Psyche 36: 61-88.

BraDLey, J. C. 1922. The taxonomy of the Masarid wasps, including a monograph on the North
American species. Univ. California Publ., Tech. Bull. 1: 369-464.

Brauns, H. 1902-3. Beitrage zur Kenntnis siidafrikanischer Masariden (Hym.). <. Hymen.,
Dipt. 2: 181-185, 275-282, 373-374 (1902); 65-73 (1903).

Brauns, H. 1910. Biologisches iiber siidafrikanische Hymenopteren. Z. wiss. InsektBiol. 6:
384-387, 445-447.

Brauns, H. 1913. Dritter Beitrag zur Kenntnis der Masariden (Hym.) von Siidafrika. Ent. Mitt.
2: 193-209.

Cameron, P. 1906. Descriptions of some new species of Hymenoptera from Pearston, Cape
Colony. Trans. S. African phil. Soc. 16: 323-333.

Kuuc, F. 1810. Einige neue Piezatengattungen. Mag. Ges. naturf. Freunde, Berlin 4: 31-45.

Kuuc, F. 1824. Entomologische Monographien. Berlin.

RicHarps, O. W. 1962. A revisional study of the Masarid wasps (Hymenoptera, Vespoidea). London.
British Museum (Natural History).

SaAussurE, H. de 1853. Note sur la tribu des Masariens, et principalement sur le Masaris vespi-
formis. Ann. Soc. ent. France (3) 1: Bull.: xvii—xxi.

Saussure, H. de 1854-5. Etudes sur Ja famille des Vespides, III. [Masaridae on pp. 1-96
( = 1855) and on plates 1-5 ( = 1854).]

ScHULTHEss, A von 1935. Some more South African Masaridae (Vespoidea). Ann. Mag. nat.
Hist. (10) 16: 383-390.

Scuutz, W. H. 1912. Aelteste und alte Hymenopteren Skandinavischer Autoren. Berl. ent. Z.
57: 52-102.

THUNBERG, C. P. 1815. Philanthi, generis insecti Hymenopteri, monographia. Nova Acta Soc.
Sci. Uppsala 7: 126-139.

Turner, R. E. 1935. Notes on the Masarid wasps of the genus Ceramius. Ann. Mag. nat. Hist. (10)

15: 290-299.

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SmitH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
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in arabic numerals, with wavy underlining; pagination of paper; place of publication; name

of publisher. Thus:

SmirH, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: Page
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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L. D. BOONSTRA MUS. COMP. ro

OL

LIBRARY
THE RUSSIAN DINOCEPHALIANa&avasy
DEUTEROSAURUS UNIVERSITY

September 1965 September
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THE RUSSIAN DINOCEPHALIAN DEUTEROSAURUS
By

L. D. Boonstra
South African Museum, Cape Town

CONTENTS 7
PAGE oe
Introduction . : » | (283 |
Deuterosaurus . . aud tog UY Lg
Mnemeiosaurus . ; ic ge BOS
Summary : , 286 MARV A
Acknowledgements . 3.) 286 UNTY
References : : #286
INTRODUCTION

In recent compilative publications in English, German, Frenchand Russian
the compilers Romer, von Huene, Piveteau and Orlov, in following the latest
views of our Russian colleagues Efremov and Orlov, have added the weight of
their authority to perpetuate some rather serious mistakes.

I have, during a four weeks’ stay in Moscow, had the opportunity of study-
_ nig the dinocephalian material housed in the Palaeontological Museum of the
Academy of Sciences of the U.S.S.R. and feel that a preliminary note pointing
out these errors would be welcomed by those who have not had the opportunity
of seeing the material for themselves.

DEUTEROSAURUS

Orlov in 1954 gave a full account of all the material he considered warran-
ted inclusion under the appellation Deuterosaurus.

This material consists of parts of two skulls, teeth and many postcranial
bones of which there is little evidence of having been found in association. In
fact, except in the case of some postcranial bones, there is definite evidence that
they were not found in association. These separate finds have on various
grounds been considered to belong to one and the same genus.

This lumping together started on its false course seriously when Seeley in
1894 described specimen No. 2 and identified it as a second specimen referable
to Eichwald’s type (No. 1) of Deuterosaurus and produced a composite drawing
of a skull in which the type lower jaw, teeth and partial occiput were fitted to
the distorted partial skull of No. 2 and associated with this some vertebrae, a
femur, a radius, part of a humerus and scapula and parts of the pelvis.
Eichwald’s type specimen (No. 1) consists of a lower jaw from which the nature
of the incisors, canines and postcanines as well as that of the upper incisors and
canines can be determined and fitting on to this are the posterior part of the
lower jaw, suspensorium, parts of the occiput, braincase and subtemporal and
suborbital arches. |

233
Ann. S. Afr. Mus. 48 (12), 1965: 233-236.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

This specimen (No. 1) constitutes the type of Eichwald’s Deuterosaurus
biarmicus.

The Diagnostic Characters of the Genus DEUTEROSAURUS:

1+?
Gan 9
2. Incisors and canines of the upper and lower jaws intermesh.
Lower canine passes outward of upper jaw margin.
. Incisors fairly long, with slight lingual step, unequally developed in the ©
different teeth and separated from the talon. :
5. Postcanines with bulbous spatulate crown; series short, probably not more
than 8.
Quadratojugal no longer a bone of the outer lateral surface.
Quadrate posteriorly situated and lower jaw long.
No boss on the angular.
Slight indication of upward sweep of the premaxillaries.
Snout probably higher than broad.
Infra-temporal bar fairly weak with deep temporal fossa.

Taxonomic Position of DEUTEROSAURUS:

1. Dental formula: 12 te
4

= oS

I
I

eae

It is evident that the type is poor and important diagnostic features are

not preserved.
Of the determinable characters none are typically tapinocephalian, but
rather represent a mixture of anteosaurian and titanosuchian features.
The anteosaurian features are:
the short series of bulbously spatulate postcanine teeth;
the fairly long incisors, but the lingual step is more pronounced than
is usual in the better known anteosaurians:
the medial shift of the quadratojugal, which is no longer a lateral
surface bone;
the slight upward sweep of the alveolar border of the premaxillaries;
the snout probably higher than broad;
the weak infra-temporal bar and deep temporal fossa.
The titanosuchian features are:
the intermeshing of the upper and lower canines with the lower
canine passing outside the upper jaw;
the absence of the angular boss and the general little pachyostosis.
I would thus place Deuterosaurus in a separate family —Deuterosauridae—
in the Infra-order Anteosauria.

Postcranial Bones referred to DEUTEROSAURUS:

The femur is represented by a number of specimens (Nos. 59, 13, 1/1326,
294/20, 72). All indicate a slender curved femur quite distinct from the femur
of both Tapinocephalia and Titanosuchia, but ee similar to the few
known femora of the South African Anteosauria.

THE RUSSIAN DINOCEPHALIAN DEUTEROSAURUS 235

If the type jaws of Deuierosaurus are, as I am convinced, anteosaurian and the
femora also anteosaurian it may very well be that the above specimens are
correctly referred to Deuterosaurus, but they could equally well be referred to
one or other of the other known Russian anteosaurians as e.g. Titanophoneus,
Doliosauriscus, Syodon, Admetophoneus.

The distal end of a humerus (No. 33) and the tibia (No. 86) also appear
to be anteosaurian.

MNEMEIOSAURUS

Specimen No. 2, which Seeley (1894) mistakenly referred to Eichwald’s
Deuterosaurus, consists of an incomplete and distorted skull from which few
diagnostic features can be determined. :

Diagnostic Characters of the Genus MNEMEIOSAURUS :
? I I+?

1. Dental formula: ts c> pe

2. Crown of postcanine tooth spatulate.
3. Intertemporal width small, with high and sharp parietal crest flanked by
the postorbital.

4. Anterior to the temporal fossa proper there is a sloping surface, below the

level of the dorsal surface proper, which is formed by the postfrontal,

frontal and postorbital (cf. Phthinosuchus).

Pachyostosis moderate.

Skull and particularly the snout, high and apparently short.

Orbit large.

Postorbital bar slender, but forming a wide flange of bone Sspieelne the

posterior bony face of the orbit (cf. Phthinosuchus).

g. Vomers narrow, choanae short.

10. Pineal canal of moderate length.

11. Lacrimal with antero-dorsally directed process as in Ulemosaurus.

12. Vomers narrow and vaulted as in primitive gorgonopsians (Phthinosuchus),
but posteriorly spatulate.
Mnemeiosaurus, on the scanty evidence, is difficult to place taxonomically,

but is probably a tapinocephalian as suggested by Nopcsa. The presence of a

canine indicates a primitive form but otherwise it appears quite specialised.
Provisionally it may be placed with the moschopids, where a form like

Avenantia also has a narrow intertemporal region.

COM DE

Teeth referred to DEUTEROSAURUS:

The teeth numbered No. 3, 4 and 5 and figured by Efremov (figs. 19, 20,
21) were not found in association with either specimens No. 1 or No. 2.

They are quite distinct from the incompletely preserved incisors of the
type of Deuterosaurus (No. 1) and no comparison is possible with the one incisor
root preserved in Mnemeiosaurus (No. 2).

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

The teeth in Deuterosaurus (No. 1) are slightly modified anteosaurian teeth. —

Tooth No. 4 approaches most nearly to the incisor teeth of the Tapino-
cephalidae.

Teeth No. 3 and No. 4 may very well be of a form very close to Ulemosaurus.

The incisor teeth hitherto considered as deuterosaurian are thus really
typical of the Tapinocephalia and the real deuterosaurian incisors are really
anteosaurian incisors.

SUMMARY

~The Russian dinocephalian Deuterosaurus, is discussed after study of the
material in Moscow and it is shown that some material has been erroneously
included in this genus. It is considered that the type species, Deuterosaurus
biarmicus Eichwald should be placed in the family Deuterosauridae, in the
infra-order Anteosauria. Mnemeiosaurus, which was referred to Deuterosaurus,
appears to be a tapinocephalian which might be placed provisionally with the
moschopids. The true affinities of postcranial bones and teeth that had been
referred to Deuterosaurus are considered.

ACKNOWLEDGEMENTS

_. The author’s visit to Moscow was in part made possible by a special grant
from the Department of Education, Arts and Science of the Government of the
Republic of South Africa. In Moscow my work was greatly facilitated by the
kind helpfulness of my colleagues Orlov, Tchudinov, Shishkin and Tatarinov.
To them my sincerest thanks.

_. The Trustees of the South African Museum are grateful to the Council
for Scientific and Industrial Research for a grant to publish this paper.

REFERENCES

BoonstrA, L. D. 1954. The cranial structure of the titanosuchian Anteosaurus. Ann. S. Afr. Mus.
/ 42: 108-148.

BoonstrA, L. D. 1957. The moschopid skulls in the South African Museum. Ann. S. Afr. Mus.
44: 15-38.

Erremovy, I. A. 1954. [Fauna of terrestrial vertebrates in the Permian cuprous sandstones of the
western part of the Ural Mountains regions.] Trud. Paleont. Inst., Akad Nauk SSSR 54: 1-416
(In Russian). .

EICHWALD, E. von. 1848: Ueber die Saurier des kupferfiihrenden Zechsteins Russlands. Bull.
Soc. Nat. Moscou 21 (2): 136-204.

Nopcsa, F. 1928. Palaeontological notes on reptiles. Geol. hung. (Palaeont.) 1 (1): 1-84.

SEELEY, H. G. 1894. Researches on the structure, organization and classification of the fossil
Reptilia. VIII. Further evidences of the skeleton in Deuterosaurus and Rhopalodon from the
Permian rocks of Russia. Phil. Trans. (B) 185: 663-717.

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London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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LIEUWE DIRK BOONSTRA FEB 7 1986

HARVARD
UNIVERSIFY

THE GIRDLES AND LIMBS OF THE
GORGONOPSIA OF THE TAPINOCEPHALUS
ZONE

November 1965 November
Volume 48 Band
Part. 13° . Decl

ANNALS OF THE SOUTH AFRICAN MUSEUM

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THE GIRDLES AND LIMBS OF THE GORGONOPSIA OF
THE TAPINOCEPHALUS ZONE

By mus. COMP. ZCOL
LIC RARY,
LiEUWE Dirk BooNsTRA FEB ZY 19¢ ;
Mo a bel
South African Museum, Cape Town rm
: . HARVARD
(With 11 figures in the text) UNIVERSITY,
CONTENTS

PAGE
Introduction . SE OE |
Material. : ‘ - 2e77
Pectoral Girdle : . = 238
Pelvic Girdle . : 236
The Humerus . ‘ ire
Phe Femur. eanzAe
The Anterior Bpipodial ) Ag:
The Manus . 2) 244
Posterior Epipodial . . 246
Mine Pes: 4 ‘ - 246
Discussion : : 5249
Summary : sae V24D)
teed eesneats ‘ 12) 249

INTRODUCTION

From the Tapinocephalus zone twelve gorgonopsian species have been
described, each based on a single specimen, the number of genera being eleven.
Of these eleven Watson has put eight into five separate families with the
remaining three uncertain as to family. Only six species are based on adequate
Skulls. Hitherto the postcranial skeleton in only one specimen has been described.

In the South African Museum we have eighteen specimens and of these
parts of the girdles and limbs are present in only six specimens.

MATERIAL

Specimens with parts of girdles and limbs preserved are:
8.A.M. 8950. Hipposaurus boonstrat. Holotype.
Skull and most of the skeleton much damaged in oignasiiion by a labourer
under direction of S.H. Haughton.
Klein-Koedoeskop, Beaufort West, Low Tapinocephalus zone. Collected
Boonstra 1928.
S.A.M. 9012. Gorgonopsian.
Isolated proximal end of a femur.
Klein-Koedoeskop, Beaufort West.
Low Tapinocephalus zone. Collected Boonstra 1929.

237
Ann. S. Afr. Mus. 48 (13), 1965, 237-249, 11 figs.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

S.A.M. 9081. Hipposaurus major. Holotype.
Skull, incomplete pectoral and pelvic girdles, distal end of front epipodial
and incomplete manus.
Klein-Koedoeskop, Beaufort West.
Low Tapinocephalus zone. Collected Boonstra 1929.
S.A.M. 9084A. ?Hipposaurus major.
An isolated humerus lacking the proximal head.
Rietkuil, Beaufort West. Low Tapinocephalus zone.
Collected Boonstra 1929.
S.A.M. 12010 Galesuchid?
Bloemhof of Voélfontein, Prince Albert.
Low Tapinocephalus zone. Collected Boonstra and Zinn 1956.
S.A.M. 12118A. Galesuchid.
Part of pelvis associated with a snout.
Palmietfontein of Kruidfontein, Prince Albert.
Low Tapinocephalus zone. Collected Boonstra and Zinn 1957.

PECTORAL GIRDLE

(Figs. 1 and 2)

In the two species of Hipposaurus the two specimens have the pectoral
girdle adequately preserved, but in neither is the cleithrum and in only one is
the ossified sternum preserved in part.

The coraco-scapula is well developed with the scapular blade lying at
right angles to the vertebral axis but curving slightly around the thorax. The
coracoidal plate is large and long and rests on the interclavicle. There is no
supraglenoidal buttress or foramen. The scapular facet of the glenoid faces
downwards and backwards and slightly outwards.

The coracoidal facet faces upwards and slightly outwards. The precoracoid
forms the anterior corner of the glenoid. Immediately anterior to the glenoid,
but in a slightly higher level, the precoracoid is pierced by a fairly large foramen
supracoracoideum.

Above the glenoid on the posterior edge of the scapula is an indistinct
scar for the origin of the scapular head of the triceps muscle. The coracoid is
without..a process or even a scar for the origin of a coracoidal head of the
triceps.

The dermal clavicular girdle is well developed, except the cleithrum which
was apparently a slender splint-like bone judging by the facet on the anterior
edge of the scapular blade.

The interclavicle is well developed with a broad spatulate anterior expan-
sion curving slightly upwards, a constricted waist and a fairly long and broad
tongue-like posterior part.

The anterior spatulate end is underlain in its lateral part by the broad
ventral end of the clavicula, which ends well away from the middle line and

GORGONOPSIA OF THE TAPINOCEPHALUS ZONE 239

Fig. 1. Hipposaurus boonstrai. Holotype. S.A.M.

8950.
Pectoral girdle x 4. a. Lateral. b. Ventral

Fig. 2. Hipposaurus major. Holotype. S.A.M. 9081.
Pectoral girdle x 4. a. Lateral. >. Ventral.

which has no posteriorly directed process to underlie the interclavicle in
posterior direction as is the case in the pristerognathid clavicle.

From its broad, ventral expansion the clavicle sweeps upwards with a
rather slender process lying externally and extending slightly anterior to the
curved anterior edge of the scapulo-coracoid. Its upper extremity is applied to
the anterior scapular edge, where it also meets the ventral end of the cleithrum.

The sternum is ossified as a flattish disc-like element lying above the
posterior end of the interclavicle. No facets for the ribs can be seen.

PELvic GIRDLE
(Figs. 3 and 4)

The two specimens of the two species of Hipposaurus have the pelvic
girdle preserved in part. In both the iliac blade is poorly preserved, but the

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

eee mem em
ead Ries ‘e
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.
.
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Wess “i J

Fig. 4. Galesuchid. S.A.M.
12118 A.

Pelvic girdle x 4. a. Lateral

b. Ventral.

Fig. 3. Hipposaurus major. Holotype. S.A.M. 9081.
Pelvic girdle x 4. a. Lateral. b. Ventral

. two ventral elements are adequately represented. Associated with a snout
identified as a galesuchid species there is the left half of a pelvis with a damaged
iliac blade, a good pubis and an incomplete ischium.

From this inadequate material it would appear that the oldest gorgonop-
sians of the Tapinocephalus zone had a low, long pelvic girdle, flattish in its
pubic part and V-shaped in its ischiadic part.

The ilium was apparently low. Its blade has little or no anterior process,
but with a well developed long but low posterior process. The supra-acetabular
edge is sharp with only a weak supra-acetabular buttress, but with a distinct
supra-acetabular notch.

The ilium forms about half of the large outwardly facing acetabulum.

The pubes form a flat pelvic floor, long but not very broad. The pubic
tubera are weakly developed, but the lateral edge in Hipposaurus is strong and
girder-like. Each pubis is pierced by a fairly large foramen lying some distance
anterior to the posterior pubic edge, facing ventrally it is obscured in lateral
view by the girder-like thickened outer pubic edge. Ventrally the two pubes do

M
s
a

GORGONOPSIA OF THE TAPINOCEPHALUS ZONE 241

not form a keel, where they meet in the median line. In the median line a
diamond-shaped lacuna separates the pubes from the ischia.

The ischia meet in the middle line as a strong symphysis forming a well
developed ventral keel. Together they are broader than the pubes and also
longer. The postero-lateral corners, although thickened, are hardly tuber-like.
The ischium forms a strong posterior rim to the acetabulum.

THE HuMERuS

(Figs. 5 and 6)
In the holotype of Hipposaurus boonstrai there is a good complete right,
humerus and the distal end of the left humerus. In addition I have the distal.

three-quarters of a larger humerus found unassociated with any other part of
the skeleton, which I tentatively identify as that of ?AHipposaurus major.

a b E

Fig. 5. Hipposaurus boonstrai. Holotype. S.A.M. 8950.

Humerus x 4}. a. Ventral in orthoprojection on the distal expansion. b. Dorsal in
orthoprojection on the distal expansion. c. Posterior in orthoprojection on the distal
expansion.

Ulna and radius x 4. d. Anterior. ec. Median. f. Lateral or outer view of the ulna.

The hipposaurid humerus is relatively a long slender bone with a long
slender shaft and both the proximal as well as the distal ends moderately
expanded. There is a considerable ‘twist’ on the shaft so that the ends subtend
an angle of about 70°.

In the figures the views called dorsal, ventral and posterior are ortho-
projections on to the plane in which the distal condyles lie.

The caput is terminal, narrowly oval, flowing into the processus medialis
and lateralis, both indistinctly demarcated. The delto-pectoral crest is of
moderate size with no distinct pectoralis tuber. The bicipital fossa is deep and

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

long. The surface for the origin of the medial humeral head of the triceps is well
developed. Midway on the shaft there is on the ventral surface a strong ridge
lying somewhat posteriorly, flanked by a groove on its postaxial side. This
would appear to serve for the insertion of the m. coraco-brachialis longus.
The distal end is remarkable in that no epicondylar foramina are present.
In ventral view a longitudinal groove can be seen near the postaxial edge,
which is sharp and bent backwards, and this entepicondylar groove housed the
median vessels usually passing through the entepicondylar foramen. A similar
edge and groove, but of much weaker development on the radial side carried
the radial vessels which pass through the ectepicondylar foramen when this is
present. Distally the entepicondyle is better developed than the ectepicondyle.

Oe

Fig. 6. ?Hipposaurus major. S.A.M. 9084A.
Humerus x 4. a. Ventral. 6. Dorsal. c. Posterior.

Both condyles are well modelled. The radial condyle forms a prominent
oval capitellum which faces mostly ventrally. The ulnar condyle lying further
distally curves around the end of the bone on to the dorsal surface, where
proximally lies a deep, well excavated trochlear fossa. Ventrally the brachialis
fossa is very shallow, giving the hipposaurid humerus a quite different appear-
ance from that of the other therapsids where there is a well developed entepi-
condylar foramen.

THE FEMUR
(Fig. 7)

In the type specimen of Hipposaurus boonstrai there are a good right femur
and the proximal and distal ends of the left femur. The only additional material

GORGONOPSIA OF THE TAPINOCEPHALUS ZONE 243

at my disposal is a proximal end found as an isolated fragment and thus only
subordinately identifiable.

The hipposaurid femur is a long lightly built bone with very moderately
expanded proximal and distal ends. It is sigmoidally curved with the ends
subtending an angle of about 30°. The caput femoris is terminal, but through
the preaxial curvature on the shaft, it is somewhat preaxially directed. The
external trochanter is not very clearly demarcated.

In ventral view the proximal end of the femur is most peculiar.

Halfway along the bone the ventral surface of the shaft has a low curvature
in section. In proximal direction in the midline a low ridge becomes pro-
gressively developed, with the surface both pre- and postaxially becoming
only slightly hollowed out in section. This ridge then ends abruptly with a
sharp oblique edge. This ridge representing a remnant of the primitive ‘Y’

q b ‘¥ d @ fi

Fig. 7. Femora x 4. Hipposaurus boonstrai. Holotype. S.A.M. 8950.
a. Dorsal. 6. Ventral. c. Anterior.

Gorgonopsian. S.A.M. go12A.
d. Dorsal. e. Ventral. f. Posterior.

system, represents the internal trochanter. Proximal to the internal trochanter
the preaxial edge is strongly rounded and postaxially a hollow represents the
intertrochanteric fossa. Distally of the internal trochanter there is no indication
of a separate fourth trochanteric ridge for the m. coccygeo-femoralis.

On the dorsal proximal surface there is a weak ridge near the preaxial
edge for the insertion of the m. pubo-ischio-femoralis internus and externally
of this ridge a shallow depression for the insertion of the m. ilio-femoralis. The
distal condyles for the reception of the tibia are terminal with the fibula
articulating with the outer face of the ectocondyle.

The other gorgonopsian femur available differs considerably from that of
the above hipposaurid.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

The external trochanter is clearly demarcated and on the dorsal preaxial
edge there is a distinct pubo-ischio-femoralis internus ridge flanked by a
groove.

On the ventral surface there is a distinct low mound distally of the inter-
trochanteric fossa constituting a separate internal trochanter. Distally of this
lies a long low ridge to which the m. coccygeo-femoralis was attached and this
constitutes the fourth trochanter.

Tue ANTERIOR EPIPODIAL
(Figs. 5d, e, and f and 8)

In Hipposaurus boonstrai there is a good left and parts of the right epipodial,
whereas in H. major only the distal ends of the radius and ulna are preserved.

Both radius and ulna are long slender bones. The radius has a well
modelled proximal facet, cup-like to fit closely on to the well modelled
capitellum of the humerus. On the postaxial corner of the radius there is a well
developed flange for the insertion of the biceps. The proximal postaxial edge
of the radius fits against the rim of the sigmoid fossa of the ulna.

The ulna is much longer than the radius, with its sigmoid fossa and
olecranon process lying proximally of the proximal radial facet.

The sigmoid fossa is formed by the preaxial surface of the olecranon and
the proximal surface of the coronoid process. It fits accurately round the
trochlea of the humerus to form an efficient hinge-joint.

Proximally of the sigmoid fossa the olecranon carries a very well developed
process, broad but dorso-ventrally compressed. In extension this process
passes into the deep olecranon fossa on the dorsal surface of the humerus.

In the type specimen of Hipposaurus major the distal end of the radius is
seen to have a large oval facet for its articulation with the radiale. A similar
facet on the ulna articulates with both intermedium and ulnare.

THE MANus
(Fig. 8)

In 1935 I gave a figure and description of the badly damaged carpus of
Hipposaurus boonstrai. Now I have the carpus of Hipposaurus major prepared by
myself with the aid of adequate equipment and from this specimen one gets a
better picture of the structure of the hipposaurid carpus.

In the proximal row there are three bones, with an additional large disc-
like pisiforme postaxially. The radiale is a stout bone with a large oval proximal
facet for the radius and distally a smaller facet for the first centrale. On its
curved postaxial surface the intermedium and second centrale are articulated.

The intermedium has a well developed dorsal surface as the bone is not
compressed from side to side. Proximally a good oval facet faces the inner part
of the distal ulnar facet and a similar facet distally meets the second centrale
distally. |

The ulnare is an elongated fairly flattened bone with expanded ends and a

GORGONOPSIA OF THE TAPINOCEPHALUS ZONE 245

Fig. 9. Hipposaurus

boonstrai. Holotype.

S.A.M. 8950 x #4.
Hind epipodial.

¥ @ ‘ ‘ Fig. 8. Epipodial and manus in
ve iy dorsal view x 4. Hipposaurus
she oT major. Holotype. S.A.M. 9081.

-

slightly constricted waist. The proximal facet is a long oval and distally a
similar facet meets the enlarged fused fourth and fifth distal carpals. The first
centrale is a curiously shaped bone, distally it has two facets for the first two
distal carpals, a proximal facet for the radiale and postaxially a curved face is
applied to the concave face of the second centrale. The second centrale is
larger, but also curiously shaped with two distal facets, one for the third distal
and the other for that part of the fused element constituting the fourth distal.
Proximally it has good contact with the intermedium and preaxially two
faces fit against the radiale and first centrale respectively.

There are four distal carpals. The first large, the second and third smaller
and the fourth a large element representing a fusion of the fourth and fifth.

All five metacarpals are completely and well preserved.

The first metacarpal is the shortest with an expanded proximal end
matching the large first distal; but the distal end is unexpanded.

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

The second metacarpal is longer with both ends expanded and a long
constricted shaft.

The third metacarpal is even longer with a well expanded and well
modelled distal end.

The fourth metacarpal is the longest bone of the metapodium, with both
ends expanded and the distal well modelled.

The fifth metacarpal is of distinctive shape; its proximal end has its facet
for the fused fourth and fifth distal directed preaxially and its distal end is not
expanded.

Of the digits only the proximal ends of the first four phalanges are
preserved.

POSTERIOR EPIPODIAL
(Fig. 9)

The right epipodial is well and completely preserved and the left incom-
pletely in the type specimen of Hipposaurus boonstrai. It is composed of two long
bones with the tibia fairly robust and the fibula of lighter build. The tibia is a
straight bone, but the fibula is much curved so that there is a good spatium
interosseum. In the fibula the two ends are expanded with the proximal end
the stronger. In the tibia the distal end is only moderately expanded, whereas
the proximal end is quite massive. This meets the femoral condyles end on,
whereas the proximal end of the fibula is applied in a sliding joint to the post-
axial epicondyle.

On its dorsal face the tibia has proximally a strongly developed cnemial
crest, which extends far proximally fitting into the intercondylar sulcus of the
femur and gives a good surface for the insertion of the tendon of the femoro-
tibialis and associated muscles.

Distally the tibia is applied to the rounded knob-like facet on the astragulus
and the fibula to a similar facet on the calcaneum.

Tue PEs
(Fig. 10)

In the holotype of Hipposaurus boonstrai there are a good right pes and parts
of the left foot. Since my original description in 1934 I have been enabled by
better equipment to expose the tarsal elements more fully. This applies par-
ticularly to the astragulus, which is now also visible from the plantar surface.

The calcaneum is most peculiar, but its structure can be easily derived
from the typical disc-like element found in most therapsids. In its distal part
the calcaneum is typically therapsid and its distinctive shape is due to the
additional development of a strong well modelled tuber proximally. Distally
the calcaneum has a slightly domed dorsal surface and a broad distal facet for
the large conjoined fourth and fifth distal tarsals. Further proximally the
dorsal surface carries a strong rounded facet for the fibula. Proximally of this

GORGONOPSIA OF THE TAPINOCEPHALUS ZONE 247

facet a strong hook-like curved tuber is developed, with postaxially a strong
rounded ridge running from the fibular facet to the extremity of the tuber.
Between this ridge and the fibular facet there is a saddle-shaped excavation.

Preaxially the calcaneum is applied to the astragulus so that the fibular
and tibial facets lie in the same plane as also the distal facets of the two bones.
The cruro-tarsal articulation thus lies in one plane. In plantar view it is seen
that the calcaneum in its preaxial part is overlain by the astragulus.

The astragulus has on its proximal dorsal surface a strong rounded knob-
like facet for the tibia. Distal to this knob there is a deep transverse groove,
with distally a shallowly concave upper surface passing distally into the distal
facet for the centrale. In my original description I thought that this groove

Fig. 10. Hipposaurus boonstrat. Holotype. S.A.M. 8950 x #4.
a. Dorsal view of pes as reconstructed. 6. Plantar view of
astragalus and calcaneum.

indicated a line of junction between two formerly separate elements, but the
exposure of the plantar surface does not support this view. Preaxially it can
now be seen, particularly in plantar view, that the astragalus extends further
than shown in my original figure. In plantar view this preaxial extension is
seen to form a strong process with a downwardly directed thick rounded edge.

The single centrale is a peculiar wedge-shaped bone lying transversely
between the astragalus and the first two distal tarsals.

The greatly enlarged fused fourth and fifth distal tarsal has a very similar
wedge shape and lies between the calcaneum and the third distal and articu-
lating with the fourth and fifth metatarsals.

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

The first distal tarsal is of most unusual shape. It is a large elongated bone
looking much more like a shortened metatarsal than a distal tarsal.

The second distal is a small squarish bone articulating with the centrale
and the second metatarsal. The third distal is larger and lies between the
fused fourth and fifth tarsal and the third metatarsal.

The metatarsals are all long bones with expanded ends and slender shafts,
except the first which is quite short.

The phalanges are only completely preserved in the fourth and fifth
digits. The phalangeal formula can thus be given as 2, 3, 4, 4, 3.

The second phalanx of the third digit is much reduced and in a near
descendant of Hipposaurus one can expect a formula of 2, 3, 3, 4, 3 which would
later become 2, 3, 3, 3, 3.

In another specimen, S.A.M. 12010, which is in all probability a galesuchid,
and prepared for study since the above was written, there are associated with
fragments of skull and vertebrae much weathered parts of the limbs, including
the proximal part of a tarsus, an ulna and the middle part of a manus. The
ulna has its sigmoid face not situated medially, nor does it have the well
developed proximal process to the olecranon as in Hipposaurus. The ulna thus
agrees more with the ulnae as known in later gorgonopsians.

The proximal tarsals merit description (fig. 11).

qd b

Fig. 11. Hipposaurid. S.A.M. 12010 X 1.

The proximal tarsal bones of the left hind foot. a. Dorsal. 6. Ventral.
As—astragalus
Ca —calcaneum

fe — facet for the centrale

ff — facet for the fibula

f4 — facet for the fourth distal
t . — tuber calcis

tf — facet for the tibia.

The calcaneum is a large element which still retains some of the characters
of the primitive flattened disc-shaped structure, but in its proximal part shows

GORGONOPSIA OF THE TAPINOCEPHALUS ZONE 249

highly advanced characters, viz. the facet for the fibula no longer lies proxi-
mally but is shifted on to the dorsal surface and protruding backwards it shows
a quite well developed tuber separated from the fibular facet by a deep groove.

In ventral view it is clearly seen that the astragalus overlies the calcaneum
medially. Here the calcaneum has a well developed process extending under
the astragalus and this process can be considered a sustentaculum tali. The
gorgonopsians from so low down as the Tapinocephalus zone are thus the first
therapsids to show this typical mammalian structure.

The astragalus is a much smaller bone than the calcaneum and has a
most remarkable facet for the tibia raised well above the base of the bone as a
high eminence.

The distal facets of both the calcaneum and astragalus for the fourth
distal and centrale, respectively, are very well modelled.

DIscussION

A comparative account will be given in a subsequent paper, when I have
completed my study of the other therapsids of the Tapinocephalus zone.

SUMMARY

Descriptions are given of the girdles and limbs of the Gorgonopsia of the
Tapinocephalus zone in South Africa. Of the eighteen specimens in the South
African Museum, six have parts of the girdles and limbs present, and this
account is based on these specimens.

ACKNOWLEDGEMENTS

The Trustees of the South African Museum are grateful to the Council
for Scientific and Industrial Research for a grant to publish this paper.

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first and last, preceded by a colon (without prefix ‘p’). Thus:

SmirH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmitH, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).
Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:

Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.

When another species has been called by the same name:

[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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MUS. COMP. ZCOL

LIEUWE DIRK BOONSTRA LIERARY.
FEB @ (009

HARVARD

THE SKULL OF UNIVERSITY

STRUTHIOCEPHALUS KITCHINGI

November 1965 November
Volume 48 Band
Fart. ) 14." Deel

ANNALS OF THE SOUTH AFRICAN MUSEUM

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S : Aes 1 gpa tne | MUS. COMP. ZCOL

y. LIERARY
BEB 1905
HARVARD
THE SKULL OF STRUTHIOCEPHALUS KITCHING] “VERS

By
LIEUWE DIRK BOONSTRA

South African Museum, Cape Town
(with 11 figures in the text)

CONTENTS

PAGE
Povroduction (204. 5) "50's 3! Q5T
Description eat aitice y oahti te “rane
PUANEVINSIONE LT G4 hi nit sh)! ol) ie tas, 'BB5
Summary . Terediwans 94.205
a eas ay ay L205
PReIPPenCeg! 2.4/4 ie pa. «&, 205

INTRODUCTION

Since Brink’s description in 1958 of the type skull (B.P.I. 284) found on
De Bad, Beaufort West by Kitching, the author with H. Zinn and H. Boonstra
had the good fortune to excavate a second skull (S.A.M. K272) on Perdefontein,
Beaufort West in 1960.

This skull was found in situ lying upside down with parts of the lower
jaws exposed and the rest intact in a mudstone matrix. Unfortunately the
exposed lower jaws have suffered from weathering which has destroyed the
posterior parts of both rami. The palatal surface of the skull was firmly encased
in a jacket of reinforced plaster and the massive skull could be lifted in one
piece. On preparation the skull proved to be very well preserved and only
very slightly distorted.

On study it became apparent that this specimen shows a number of
structural features much better than the type specimen does. Certain additions
to Brink’s description can thus be made and in the sequel it will also become
clear that I differ in interpretation on a number of points, even if allowance
is made for considerable individual variation.

A correction has also to be made in regard to the type species due to
Brink having misread Broom. The species whaitsi has as its holotype the skull
(S.A.M. 2678) from Vivier, Beaufort West and the second specimen mentioned
by Broom consists of a skull and much of the skeleton (S.A.M. 3012) from
Abrahamskraal, Prince Albert. Broom rightly thought that the second specimen
belonged to the same species as the holotype skull. dines are thus two skulls
and one skeleton.

251

Ann. S. Afr. Mus. 48 (14), 1965, 251-265, 11 figs.

252 ANNALS OF THE SOUTH AFRICAN MUSEUM

DESCRIPTION
Build and carriage of the skull

The fronto-nasal boss in my specimen is very like that of the type, but I
do not think that it represents a horn-core. In the collection of the South
African Museum there are over two dozen skulls of Struthiocephalus and Kerato-
cephalus. ‘These all have a fronto-nasal boss developed to a varying degree and
in all of them the sculpturing of the boss is very similar to that of the skull in
general. If the boss were a horn-core one would have expected its surface to
differ from that of the general skull sculpturing. That the boss could have
been used as a battering ram is most probable, but then without a special
horn covering.

There is no doubt about the hang-dog carriage of the skull in Struthio-
cephalus. As a matter of fact this applies to all the tapinocephalians in general
—also to those without a naso-frontal boss. The struthiocephalines, with their
long snouts and anteriorly directed upper front teeth, undoubtedly fed on
softer vegetable matter than the moschopines. Their limbs have also been
shown to be more adapted to marshy conditions and Brink’s suggestion that
the struthiocephalines may even have fed duck-like under water is quite
probable. The surface moulding of the bone around the nostrils—especially
of the septomaxilla—suggests the presence of musculature for the closing off
of the nostril.

The structure of the skull

Although the present skull is very well preserved and prepared a number
of sutures cannot be traced with absolute certainty. This is due to a number
of factors such as the rugose nature of the outer surfaces, closure of sutures,
fusion of elements and small displacements. When comparing the figures
given here with those of the type skull, and also with those of other species of
the genus the effects of the pachyostosis should be born in mind. The pachy-
ostosis in the tapinocephalians is to some extent individual with differences
often seen between the relations and extent of the two bones of a pair in the
same skull. Age is also a factor. Differences that have been given as specific
are often due to differences in the tempo of the pachyostosis in adjoining bones.
Thus a strong pachyostotic development in, for instance, the postfrontal,
causes an overlap or overgrowth over the adjoining bones and affects the
relative size and shape of the outer surfaces of these bones. Where, however,
authors show a radical difference in the relations of bones errors of observation
and/or interpretation must be considered probable.

Lateral and dorsal surfaces. (Figs. 1 and 2)

In comparing my figures with those of Brink it is manifest that the type
skull has been subjected to dorso-ventral compression. The effects are especially «
obvious in the nature and disposition of the lateral pterygoid flange and the

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 253

quadrate. It should, moreover, be borne in mind that my figures are ortho-
projections and not perspective drawings. This would in part account for the
fact that in my figures the tabular is visible in dorsal view and in lateral view
_ forms much of the posttemporal bar. But in my specimen the tabular is really
~ much more developed and in its forward growth overlaps much of the lateral

Fic. 1. Struthiocephalus kitchingi. S.A.M. K272 x 1/6. Lateral view. Orthoprojection
on to the sagittal plane.
AN—angular. AR —articular. D—dentale. eam—external auditory meatus.
F—frontal. [P—inter- or postparietal (dermo-supraoccipital). J—jugal. L—lacri-
mal. M—maxilla. N—nasal. P—parietal. PAL—palatine. PM—premaxilla.
PO —postorbital. POF—postfrontal. PRF—prefrontal. PT—pterygoid. Q—
quadrate. QJ—quadratojugal. SA—surangular. SM—septomaxilla. smf—septo-
maxillary foramen. SQ—squamosal. T—tabular. TR—transversum (ecto-

pterygoid).

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

tongue of the parietal which is wedged in between the tabular and the upsweep-
ing process of the squamosal. In my specimen the postorbital is more developed
and this at the expense of the postfrontal. This particularly affects the
appearance of the postorbital bar as seen in dorsal view.

Fic. 2. Struthiocephalus kitchingt. S.A.M. K272 x 1/6. Dorsal view.
Orthoprojection on to the alveolar plane.

Brink figures the squamosal as entering the ventral orbital border, whereas
in my specimen the jugal extends posteriorly, ventral to the postorbital and
forms nearly all of the ventral rim of the orbit with a small contribution by
the postorbital. This is the normal relation, not only in the tapinocephalids

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 255

but also in all the other Dinocephalia (anteosaurids, titanosuchids, styraco-
cephalids and the Russian brithopids). Brink’s figure indicates an error in
observation.

In the present specimen the outer surface of both septomaxillaries is
well preserved and this shows a distinctive moulding of the posterior border
of the nostril to form a rounded swollen rim set off by the presence of a lateral
groove. I suggest that this moulded structure indicates attachments for a
valvular closure of the nostril when the animal is feeding with the snout

submerged.

Occiput. (Fig. 3)

The occiput is low and broad with the squamosals bulging laterally to
form prominent ‘cheeks’. A strong rounded thickened upper and lateral border
formed by the interparietal and the tabulars prominently demarcates a deep
bipartite area of origin for the nuchal muscles. The interparietal and the
tabulars have large posterior faces, whereas the supraoccipital is very low. The
large face of the paroccipital is directed much more ventrally than posteriorly.
The ridge on the squamosal bounding the external auditory meatus medially
is very prominent. The posttemporal fossa is all but closed by the downgrowth

Fic. 3. Struthiocephalus kitchingt. S.A.M. K272 x 1/6. Occipital
view. Orthoprojection at right angles to the sagittal and alveolar
planes.

BO — basioccipital. EO—exoccipital. fq—quadrate foramen.
jf—jugular foramen. mp—mastoid process of the paroccipital.
pin—pineal (parietal) foramen. POC—paroccipital. ptf—post-
temporal fenestra. ptpocf—pterygo-paroccipital fenestra.
SO—supraoccipital. ST—stapes. stf—stapedial foramen.
Other lettering as for Fig. 1.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the tabular. The stapes is seen to lie diagonally with the distal end lying
low down in the recess on the quadrate. Little is seen of the posterior face of
the quadrate in this view because of its horizontal disposition. The occipital
condyle is directed much ventrally with the exoccipitals forming most of the
posterior face and the basioccipital facing mainly ventrally.

Ventral surface. (Fig. 4)

Comparing my figure of the ventral surface to that given by Brink a
number of differences are apparent, apart from the fact that mine is a projection
and Brink’s a perspective drawing. The differences are in main due to the fact

that the present specimen being uneroded is in a much better state of preserva- —

tion. The regions mainly affected are the transverse processes of the pterygoids,
quadrate and stapes, the occipital condyle and the relations of the supra-
occipital, interparietal, tabular and squamosal.

In the present specimen the lateral flanges of the pterygoids form well
demarcated deep transverse ridges extending far ventrally in their lateral
parts, lying far below the level of the quadrate rami.

In its lateral part the quadrate ramus forms a deep vertical flange of

bone which meets the quadrate along a large synchrondrotic face. More ©

medially the pterygoid is deeply vaulted and sends a process posteriorly which
abuts against the paroccipital at a level higher than the quadrate process of
the paroccipital.

The quadrates are well preserved and carry cotyli shaped as shown in the
figure. The stapedial recess is well preserved on both sides.

Both stapes are well preserved and are seen to lie diagonally with the
distal ends lying well anterior to the plane of the fenestrae ovales. The occipital
condyle is a prominent large knob roughly circular in outline with its articular
face directed only slightly posterior off the ventral. The exoccipitals form
more of the articular face than is shown in Brink’s figure.

The paroccipital has a large ventral face. Medio-anteriorly it has a
process meeting the prodtic above the level of the fenestra ovalis. Latero-
anteriorly there is a long quadrate process applies to the quadrate and ter-
minating posterior to the stapedial recess. At a higher level the quadrate

TF

|
t
:
:

process of the paroccipital has a thin flange of bone whose anterior edge meets ©

a process of the quadrate ramus of the pterygoid. Posterolaterally the par-
occipital develops an everted thickened edge underlying the squamosal to
form a mastoid process. The supraoccipital is wide but low and the posttemporal

fossa all but obliterated by the overgrowth of the large paroccipital. The ~
tabular has a large ventral face which, however, does not extend anteriorly —

between the paroccipital and squamosal as shown by Brink in his figure of the ©

type.
The squamosal carries a prominent ridge, roughly comma-shaped in

?

outline, and this forms the internal limit of the external auditory meatus j

developed as an antero-posteriorly directed groove.

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI

257

<RHBVBYV®

Fic. 4. Struthiocephalus kitchingi. S.A.M. K272 x 1/6. Ventral

view. Orthoprojection on to the alveolar plane.

fm—foramen magnum. fo—fenestra ovalis. ic—internal
carotid foramina.

iptv—interpterygoid vacuity. PBS—para-
basisphenoid. PROT —prodtic.

qp—quadrate process of the
paroccipital. rt—replacing teeth. V—vomer. Other lettering

as in previous figures.

The position of the stapes. (Fig. 5)

In order to describe the relations of the stapes to the bones with which it

makes contact I have sketched the various bones disarticulated but lying in
their relative positions.

258

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 5. Struthiocephalus kitchingi. S.A.M. K272 x 1/3. The stapes and
contiguous bones as disarticulated. Ventral view. Orthoprojection on
to the alveolar plane.
con—condyles of the quadrate. cpt—contact of pterygoid (quadrate
ramus) with the quadrate, paroccipital and para-basisphenoid. cpot—
contact of paroccipital with the prodtic. cq—contact of quadrate with
the pterygoid and the stapes. csq—contact of the squamosal with
the quadrate and the quadratojugal. np—notochordal pit in the
basioccipital.
Other lettering as in previous figures.
1— wedge of the pterygoid applied to the medial edge of the quadrate
‘(4)-

2—area on the posterior process of the quadrate ramus of the pterygoid
overlapping the distal end of the stapes.

3 —surface of the quadrate overlapping the stapes dorsal to the stapedial
recess proper.

4—area on the medial edge of the quadrate making contact with the
posterior process of the quadrate ramus of the pterygoid.

5—surface of the quadrate underlapped by the paroccipital.

6—area on the dorsal process of the stapes in contact with the
paroccipital (7).

7—area on the paroccipital in contact with the dorsal process of the
stapes (6).

8—area on the paroccipital underlapped by the stapes.

g—area on the paroccipital underlapped by the proximal end of the
stapes.

10—area of the protic underlapped by the stapes.

11—hollow in the distal end of the stapes which receives a process of
the quadrate.

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 259

The footplate of the stapes lies fitted into the fenestra ovalis on both sides
so that its proximal end cannot be seen.

Antero-dorsally the proximal end of the stapes is applied to a face on
the pro6étic and further posteriorly underlies a face on the paroccipital. Both
these contacts are not very intimate—definitely not synostotic—but rather
syndesmotic or synchondrotic. Ventrally the footplate is in contact with the
edges of the fenestra ovalis formed by the paroccipital, basioccipital and the
para-basisphenoid.

The distal end of the stapes makes contact with the quadrate and
paroccipital.

Postero-laterally the distal end of the stapes has a fairly short truncated
process—the tympanic process—whose tip is connected directly or by the
intercalation of a cartilaginous extra-stapes to the tympanum.

Antero-medially of this tip the distal end of the stapes fits into an elongated
groove-like stapedial recess in the quadrate, lying in a plane nearly parallel
to the quadratic cotyli. The fit is not tight and movement in this diagonal plane
is possible even with the presence of cartilage or connective tissue.

The anterior part of the distal surface of the stapes has an oval hollowed-out
articulatory face which makes contact with a process of the quadrate lying
anterior to the elongated stapedial recess and posterior to the surface of contact
with the end of the quadrate process of the pterygoid.

On the postero-distal surface of the stapes, posterior and medial to the
tympanic process, and at a higher level than the main shaft of the stapes lies
the dorsal process of the stapes, which makes contact with the quadrate process
of the paroccipital. This contact was apparently synchrondrotic.

Stapes. (Fig. 6)

In ventral view the stapes presents a main portion consisting of a proximal
end forming the footplate, a greatly expanded distal end and a shaft with a
waist-like constriction. From the posterior edge of this main portion the rest
of the bone lies at a higher level. About halfway along the shaft a large oval
stapedial foramen pierces this flange. Postero-distally of the foramen there is a
process which meets the paroccipital—this is the dorsal process.

The distal end bears a process directed postero-distally—this is the
tympanic process. Anterior to the tip of the tympanic process the distal end
has an elongated face which fits into the stapedial recess in the quadrate.

Anterior to this the distal end presents a concave oval articulating face,
which articulates with a convex process on the quadrate lying immediately
posterior to the contact face on the quadrate, which receives the quadrate
process of the pterygoid.

In dorsal view the stapes has the surface of its main portion and the
surface of the dorsal process lying in the same plane. On the upper face of the
dorsal process there rises a conical protuberance with its tip directed medially.
This I have labelled the medial dorsal process. The function of this process

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

Vee

hee
er :

Fic. 6. Right stapes of Struthiocephalus kitchingt. S.A.M. K272 x 1/3.
A—anterior. B—posterior. C—dorsal. D—ventral. FPOC—facet on
the dorsal process which is applied to the quadrate process of the
paroccipital. FQ—facet fitting into the elongated stapedial recess in
the quadrate. FQP—concave facet applied to a convex process on the
quadrate lying medial to the stapedial process proper. MDP—medial
dorsal process. PDP—proximal dorsal process. STF—stapedial
foramen. ‘TY—tympanic process.

would appear to be to receive a tendon probably attached to the inner edge
of the quadrate.

In anterior and posterior view this median dorsal process is seen to be
both strong and prominent.

Quadrate. (Fig. 7)

The quadrate is a large bone—robust in its cotylar region and in the
parts making contact with the squamosal, quadratojugal and the quadrate
ramus of the pterygoid. But its inner portion overlying the paroccipital consists
of a thin sheet of bone with a free edge.

Its stout pterygoid process bears a large roughly oval area for the reception
of the quadrate ramus of the pterygoid, which at a higher level has an additional
process meeting the quadrate in a groove on the lower part of the inner edge
of the quadrate. The articulatory area on the pterygoid process is a hollowed
out area with, in its middle part, a longitudinal ridge. This is matched on the
quadrate ramus of the pterygoid by an articulating face bearing a median
groove flanked by two longitudinal ridges.

The nature of this joint is such that it admits of movement between the
quadrate and the pterygoid in a parasagittal plane. The roughness of both
articulating faces indicates the presence of synchondrotic cartilage.

Just above the quadrate-pterygoid articulation there is a transverse
everted flange of bone. The upper edge of this flange carries a transverse
groove which thus lies between two lips. This groove is the stapedial recess.
The outer end of this everted flange develops a thin free standing process to
which the tympanum was probably attached.

.

261

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI

‘(radoaid ssooas Tetpodejs) sodvys ay} Jo uoNdoda1 94} JOF ssa091 poresuoja 9y3— LS ‘sodvys ay Jo pus [eysIp
9Y} UO IVF DAVUOD JY} YIM UONL[NOYIe OF ssao01d xaAU0I—g JS *ss00r perpodeys oy} Jo diy 1o110}sod oy} Sururs0j 9Sp2]—"T.LS
‘jeSnfoyerpenb oy} Jo pus [esJUSA oY} SUTATODOI ss9001— (OY ‘uouresoj o3erpenb ay} jo Jopioq JoUUT 9y} SuIUIoy YoI0U—NO
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AjpeorSojoydiouz) so1s0yue—(] ‘TeIpsui Jo tguUT—) ‘(ore Ajyeor8o0joydaour) tes1op—g *(101103sod Ayyeorsojoydsouz) ,e1yUsA —V
E/1 x hoy W'V'S “WUuyILy snpoyfasorynaygy JO ajyerpenb iysry °4 ‘Ory

20

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

In between the quadrate-pterygoid joint and the outer lip the everted
flange has a rounded process to which is articulated the hollow face on the
distal end of the stapes.

On its posterior face the quadrate is greatly thickened where it is applied to
the squamosal and quadratojugal. Medial to this thickened area lies a wide
groove with a smooth surface. This housed part of the body of the m. capiti-
mandibularis.

A moschopid and jonkertid quadrate. (Figs. 8 and 9)

I am including comparable figures of a quadrate of a moschopid and a
jonkeriid. Essentially these are of the same type as that of Struthiocephalus,
except that in the jonkeriid quadrate a very definite recess is developed on the
posterior face well above the stapedial recess. I can offer no suggestion as to
what was received in this recess.

Fic. 8. Left quadrate of Moschops sp. S.A.M. 11701 X 1/3.

The Tympanum

A depressor mandibuli muscle originating from the prominent ridge on
the squamosal just medial of the groove housing the external auditory meatus
and inserted on the process of the articular would allow ample room for a
tympanum with a diameter of about 30 mm.

The tympanum having one point of attachment on the free tip of the
lip of the ledge bounding the stapedial recess could lie in a parasagittal plane
and receive the tympanic process of the stapes meeting it at right angles.

The external auditory meatus extending from the tympanum along the
groove in the posterior face of the squamosal could have its opening just above
the level of the upper limit of the squamosal ridge.

The nature of the joint of the stapes with the quadrate as described above
would allow of movement of sufficient amplitude for the conduction of sound
waves.

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 263

Fic. 9. Right quadrate of Jonkeria haughtoni. S.A.M. 4343 X 1/3.

Dentition. (Fig. 10)

Brink maintains that in the type specimen the marginal teeth are disposed
in a double row both functional at the same time. This view is manifestly
incorrect. The fact is that the replacing teeth arise lingually of those in use
and each new tooth is thus a younger member of the same tooth family of

264. ANNALS OF THE SOUTH AFRICAN MUSEUM

Cc D

Fic. 10. First left incisor of Struthiocephalus kitchingi.
S.A.M. K272 x $.

A-— posterior. B—anterior. C—labial view.

D—lingual view. H—heel. R—root. T—talon.

the tooth it replaces. The position is thus as in the Titanosuchia. The structure
of the teeth is of the talon-and-heel type thoughout the series, with decrease in
size in posterior direction.

Fic. 11. Skull of Moschosaurus longiceps. Type.
S.A.M. 3015 x 1/6.
A—dorsal. B—lateral.

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 26 5

Discussion

In a recent paper I have suggested that all the described species
of Struthiocephalus could well be considered conspecific and I arranged them in
a growth series.

I would now go further and suggest that Moschosaurus could very well be
the youngest form of such a series. (Fig. 11)

SUMMARY

A detailed description of the skull of Struthiocephalus kitchingi is given,
based on a second skull from Beaufort West. This specimen shows a number of
features much better than the type specimen and leads to some differences in
interpretation.

ACKNOWLEDGEMENTS

The skull here described was obtained on a collecting trip by the South
African Museum which was in part financed by a grant from C.S.I.R. to
whom we tender our thanks.

The Trustees of the South African Museum are grateful to the Council
for Scientific and Industrial Research for a grant to publish this paper.

REFERENCES

Boonstra, L. D. 1963. Diversity within the South African Dinocephalia. S. Afr. 7. Sci. 59:
196-207.
Brink, A. S. 1959. Struthiocephalus kitchingi sp. nov. Palaeont. Afr. 5: 39-56.

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of publisher. Thus:

SmirH, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plenia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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LIERARY:

Scoala ry
FEB ¢ lwo

HARVARD
UNIVERSITY
A NEW PALAEONISCID FISH FROM THE

WITTEBERG SERIES (LOWER CARBONIFEROUS)
OF SOUTH AFRICA

November 1965 November
Volume 48 Band
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MUS. COMP, ZOOL:

LIERARY,
FEB @ 1065

A NEW PALAEONISCID FISH FROM THE WITTEBERG SERTERVARD
(LOWER CARBONIFEROUS) UNIVERSITY
OF SOUTH AFRICA
By
R. A. JuBB

Albany Museum, Grahamstown.

(With 1 plate and 2 figures in the text)

CONTENTS

PAGE
Introduction .. Lith. LeGy
Genus Mentzichthys nov. . . 270
Mentzichthys walshi sp. nov. . 270
Discussion. . He ran ea 2
Acknowledgments IP W272
Summary. . a Mer ore) 2g
References soit ig se Ls 7S

INTRODUCTION

In May 1963 Mr. G. Walsh, an official of the Sundays River Irrigation
Board, discovered a deposit of fossil fishes in the mountains just south of Lake
Mentz and close to the entrance of Karroo Poort. Mr. J. J. G. Blignaut, a
retired geologist, visited the site immediately and confirmed the importance
of its discovery. Since the beds are well exposed, from the upper portion of
the Witteberg quartzites through the complete sequence of Upper Witteberg
Shales to the Dwyka Tillite, it was possible for Mr. J. A. Marais of the Geologi-
cal Survey Department to determine the geological horizon on which the
fossil fish occur. The fossiliferous bed, which is nearly vertical and from three
to six inches thick, occurs 740 feet above the topmost white quartzite of the
Witteberg and in the upper shales which were formerly known as the Lower
Dwyka Shales. These shales are regarded as of lacustrine origin.

Lithologically, according to Haughton (1963), the Series, as now defined,
consists of fine-grained thinly bedded whitish quartzites intercalated with
dark green or blue (buff when weathered) micaceous shales and flagstones.
Locally lenses of grey sandy limestone or calcareous quartzite, sometimes
carrying plant fragments, are present, as well as bands of coarse grit. The
uppermost member (formerly Lower Dwyka Shales) consists mainly of greenish
and bluish shales and flagstones with some hard sandstone bands.

The most characteristic fossil in the quartzites of the series is the spiral

267
Ann. S. Afr. Mus. 48 (15), 1965: 267-272, 1 pl., 2 figs.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

impression known as Spirophyton which is characteristic of the Middle Devonian
of America. Other fossils previously discovered, although scanty, include
Bothrodendron, Cyclostigma (characteristic of the Carboniferous of Queensland
and New South Wales), Didymophyllum, Halonia, Knorria, Lepidodendron, Lepido-
strobus, Singularia, Selaginites and Stigmaria. Fragments of the body segments of
a eurypterid named Hastimima have also been discovered, as well as some fish
scales. A recent discovery by Mr. J. A. Marais is a species of Psilophyton. In
age Du Toit (1954) considers these non-marine beds, about 4,000 feet in
_ thickness, to embrace the whole time interval from about the Middle Devonian
to the Lower Carboniferous, and it is to this latter period that the Lake Mentz
fish fossil bed appears to belong.

An outstanding feature of the exposed section of the fossil bed is the
concentration of fish impressions it contains. This concentration spoils what
could have been, in many cases, perfect impressions. As stated above, the
maximum thickness containing fishes varies from three to six inches but the
extent of the bed is unknown as it dips and disappears. Under normal circum-
stances sediments of this thickness would represent an appreciable interval
of time, but, from the great number of fishes overlapping one another, and
the disposition of some, it is evident that a mass mortality involving a large
concentration of Palaeozoic fishes occurred. Such a mass mortality, followed
by perfect preservation, could only have taken place if the primeval pool of
lake they were inhabiting was completely frozen over in the first place. Total
freezing would have resulted first in the concentration of fish in the bottom
layers, then asphyxiation followed by preservation in the bottom muds under

<<

ae

freezing conditions before being finally entombed by some major earth or _

glacial movement. The open mouths of many of the fishes suggest asphyxiation
due either to deoxygenation of the water by the fishes themselves, or to their
being forced down into a deoxygenated hypolimnion.

Although the Witteberg Series is associated with lacustrine conditions,
certainly non-marine, it would be misleading to regard the Lake Mentz fossils
as being those of entirely freshwater species. Of the Palaeozoic and Mesozoic
fishes described from southern Africa many belong to widely distributed
genera and families which are associated with both marine and non-marine
formations. Rayner (1958) quoting Gunter points out that, as a general rule,
marine fishes are more tolerant of fresh waters than vice versa. There is ample
evidence of this today where marine species such as Carcharinus leucas, Pristis
microdon, Megalops cyprinoides and Glossogobius giuris are all found up to 300 miles
by river from the sea in many of the rivers of the east coast of Africa.

The Lake Mentz fossil fishes represent two distinct species and possibly a
third. Of the two more complete species one is a platysomid-like fish with very
large dorsal ridge scales, and the other a slender stream-lined palaeoniscid
which is described below as representing a new genus and species. The generic
name refers to the type locality and the specific name is in honour of the
discoverer of the fossil bed.

269

NEW PALAEONISCID FISH FROM WITTEBERG SERIES

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2'70 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Palaeoniscidae
Genus MENTZICHTHYS nov.

Definition: Body fusiform and elongate; principal rays of pectoral fins not
articulated till towards their terminations; dorsal and anal fins situated pos-
teriorly, moderate in size, triangular in shape, the former originating slightly
ahead of the latter; caudal fin heterocercal, inequilobate and deeply cleft,
the axial lobe being well developed; distinct fulcra on all fins; head with
prominent rostrum and relatively large anteriorly placed orbit with large
postorbital; suspensorium very oblique, gape wide, teeth consisting of short
well-spaced cutting teeth and fine conical teeth on the maxillary, and numerous
fine conical teeth on dentary; opercular large, larger than subopercular,
branchiostegal rays fairly numerous, head bones sculptured with striae and
tubercles; scales rhombic, of moderate size, with large peg-and-socket articula-
tion, entire posteriorly, ridge-scales running from the occiput to the origin of
the dorsal and along the leading edge of the caudal fin.

Mentzichthys walshi sp. nov.
(Figures 1, 2. Plate VI A, B)

Specific diagnosis: Slender fusiform fishes whose maximum total length appears
to be about 18 cm., the length of the head being contained about four and a
half times, and the greatest depth six and a half times in the total length of
the body.

The Skull

The shape and arrangement of the bones of the skull are shown in Figure
2, which has been prepared, allowing for distortion, from three impressions of
heads which were cleaned and then polished with a nylon brush before photo-
graphing. Plasticine casts were also used for reconstructing the skull, broken
lines being used in the final drawing where sutures could not be determined
with confidence.

The ornament of the frontals, supratemporals, parietals and post-temporals
consists of striae running more or less antero-posteriorly. The postrostral is
ornamented mostly with tubercles which extend down to the nasals (see
Plate VI A). The branchiostegal rays are not ornamented but the large and
prominent postorbital bone is markedly rugose. Numerous transverse striae
ornament the maxillary and particularly the dentary. The suspensorium is
very oblique. The opercular and subopercular are also ornamented, the
former being much larger than the latter. The teeth showing on the dentary
are minute and numerous, similar teeth appear on the maxillary but there
are well spaced short cutting teeth as well which are confined to the anterior
and middle section.

NEW PALAEONISCID FISH FROM THE WITTEBERG SERIES 271

he Paired Fins

_ Preservation is not good enough to detect ornamentation of the supra-
eithrum or cleithrum. The pectoral fin, situated low down ventrally, is
dium in size with a base length about one third that of the longest lepido-
| tri hia. The lepidotrichia, which number about 19 are unarticulated and
|unbranched until their distal ends. The anterior edge of the pectoral fin is
ered with small fulcra. The pelvic fin, whose origin is about midway
een the origin of the pectoral and the origin of the anal, is relatively
aall with about 13-15 lepidotrichia which are articulated and distally
branched. The anterior edge is covered with small fulcra.

* 2 Unpaired Fins

g The dorsal fin, which is situated nearer to the caudal fin than to the head,
is one and a quarter times as high as the length of the base. It is triangular in
shape with from 28-35 lepidotrichia which are articulated and branched
ey. The anal fin, whose origin is posterior to that of the dorsal, has a base
onger than its height. It is triangular in shape with 35-45 lepidotrichia which
a articulated and branched distally. Small fulcra are present along the
anterior edge. One of the outstanding features of this new species is the large
jal lobe of the caudal fin. This fin is distinctly heterocercal, deeply cleft

id inequilobate, the leading edge being covered with ridge scales.

_ Squamation

Not many scales are available for examination as, in most of the samples,
ust the underneath pattern of the squamation appears in the impression.
he scales are medium in size and rhombic in shape with pronounced peg-
d-socket articulation. The few scales available, mostly immediately posterior
e head, show transverse furrows with pits, the posterior edges being rough
entire. There are from 16-18 rows of scales from the dorsal to the ventral
flace at the point of maximum girth, and 52-58 scales along a lateral series.
oderately enlarged ridge-scales, numbering about 21, extend from the
ciput to the origin of the dorsal fin. Ridge-scales of a similar size anteriorly
tend along the caudal peduncle and up the leading edge of the caudal fin.

t

faterzal

_ The holotype is an impression on a slab No. A.M./64/4562 in the Albany
Museum, Grahamstown. A photograph of this is shown in Plate B. This
, from two to three inches thick, has numerous impressions on both faces.
her samples of importance in constructing the illustrations in Figures 1 and
re A.M. 64/4563, A.M. 64/4564 and A.M. 64/4565, in the Albany Museum.
her material, S.A.M. Kr169 and S.A.M. Ki170 is preserved in the
South African Museum, Cape Town.

Pag (e'4 ANNALS OF THE SOUTH AFRICAN MUSEUM

DiIscussION

Reasonably entire specimens of M. walshi have the following extrem
measurements:
Total length: 90 mm. Std. length: 70 mm.
Total length: 145 mm. Std. length: 110 mm.
From these measurements it has been possible to estimate the maximur
size of incomplete fossils to be about 180 mm. The type, Figure 1 and Pla
B, has a total length of 145 mm.
M. walshi belongs to the group pf palaconseids having a very obliqu
mandibular suspensorium (Woodward, 1891). The remarkable developmer
of the axial caudal lobe, the smaller orbits and the large postorbital bone:
distinguish this new genus from Rhadinichthys Traquair, 1877 as re-defined
by Moy-Thomas and Dyne (1938). These features, as well as the larger oper=
cular, distinguish Mentzichthys from the genus Cycloptychius Young, 1865, as
re-defined by the same two authors. Of the known palaeoniscids it woul
appear that Mentzichthys is most closely related to those of the Lower Carbonife- 3
rous assigned to the genus Cycloptychius. ‘ |

ACKNOWLEDGMENTS

I wish to thank Professor Orvar Nybelins of Stockholm, Sweden, for
reading this paper in its draft form and for providing valuable comments
Dr. H. V. Eales of the Department of Geology, Rhodes University, assisted by
his team of senior students, provided the material from which the holotype of i
M. walshi was selected. This project, carried out in collaboration with Dr.
T. H. Barry, was part of a research programme sponsored by the C.S.1.R., :
who also granted funds for the publication of this work. 4

SUMMARY

A new palaeoniscid fish Mentzichthys walshi n. gen. et sp. is described from —
the Witteberg Series (Lower Carboniferous) of South Africa, just south of
Lake Mentz in the eastern Cape. It is suggested that a mass mortality of thesé
fishes took place in lacustrine conditions under ice.

REFERENCES

Du Torr, A. L. 1954. The geology of South Africa. 3rd ed. Edinburgh & London: Oliver & Boyd
Haucuton, S. H. 1963. Stratigraphic history of Africa south of the Sahara. Edinburgh & Londo nt
Oliver & Boyd.
Moy-Tuomas, J. A. & Dynes, M. B. 1938. The actinopterygian fishes from the Lower
Carboniferous of Glencartholm, Eskdale, Dumfriesshire. Trans. roy. Soc. Edinb. 59: 437-480
Rayner, D. H. 1958. The geological environment of fossil fishes. Jn WeEsTOLL, T. S., ed. Studie
on fossil vertebrates: 129-156. London: Athlone Press.
Woopwarp, A. S. 1891. Catalogue of the fossil fishes in the British Museum (Natural History).
London: British Museum.

Ann. S. Afr. Mus., Vol. XLVIII. Plate VI

A. Portion of a skull of Mentzichthys walshi, partly obliquely crushed, showing pronounced
postorbital bone and ornamentation.

B. Holotype of Mentzichthys walshi.

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REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

SmitH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

Smiru, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, Fuly 1958, are to be observed (particularly articles 22 and 51).
Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:

Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species: _
Plonia natalensis (non West), Jones, 1956: 18.

When another species has been called by the same name:

[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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HARVARD
HIPPARION FROM LANGEBAANWEG, UNIVERSITY

CAPE PROVINCE AND A REVISION OF THE
GENUS IN AFRICA

November 1965 November
Volume 48 Band
Fact) 26 Deel

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

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
(Cash with order, post free)

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
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(Kontant met bestelling, posvry)

OUT OF PRINT/UIT DRUK

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6(1, t.—p.1.), 7(1; 3), 8, 9(1-2), 10(1-3),
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Price of this part / Prys van hierdie deel
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE AND A
REVISION OF THE GENUS IN AFRICA!

By

EpouARD L. BONE

B pakiah : Ss. MP. ZOOL:
Department of Vertebrate Palaeontology, University of Louvain” MUS, COMP. ZOOL

LIERARY,
and FEB ¢. 1865
3
RONALD SINGER HARVARD
Department of Anatomy, University of Chicago, U.S.A. UNIVERSITY:
(With 24 tables, 16 text figures and 13 plates)
CONTENTS

PAGE

INTRODUCTION ; : : , : : : ; é : , : 2° 276
General ; : F } : ; : : : é F - 276
The Penerocanves deposits ; : Sa eer : : : 5 : By 2g
Localization of the sites . : : ‘ : é ; , : ; Wenner)
History of the discoveries : : ; c ; : : : : a7
Geology d : : : ‘ ; : : : - | 270
Faunal assemblage for Langebaanweg, : : : : , as : : . 280
Methods and nomenclature . : 280
GEOGRAPHICAL DISTRIBUTION OF AFRICAN SITES WHERE HIPPARIONIDS HAVE BEEN RECORDED 281
Morocco . 4 : : : : ; ‘ : : i : . - 205
Algeria ; : ; : : : : , . : : : : . |) BOT
Tunisia ; ; : d : ; : : ; ; d , 5 . 289
Fayum ‘ é : ; ‘ : : ; ; ‘ : : ; . 282
Ethiopia. ‘ , : : : : ; : ; : : 5 5) 202
Central Africa. ; : : ; : : ; : s : ; eo?
East Africa : ; : , : : : : ; , : : 3 283
Uganda d : ; : ‘ : 4 , : ; ; ; =) 208
Kenya ; : 5 ‘ ‘ ; , : : : : : e208)
Tanganyika . : : , ‘ : ‘ : ; : : ‘ te 208
South Africa ; ‘ : : : : é : ; : F 2 . 283
Cape Province : : : : : ; A ‘ : : ; - 283
Orange Free State : ‘ ‘ ‘ : b : : : . 208
Transvaal. : : : : 203
SUMMARY OF PUBLISHED GEOLOGICAL EVIDENCE FROM AFRICAN HIPPARIONID SITES . 2 £209
Miocene. ; : : , : ; ; : ; ‘ 5 : WL P2G9
Pliocene : ; ; ; ; 3 ‘ : : ; ; : s so 200
Pleistocene . : : ‘ : : : ; : : : : . 286
North Barca’ : ; : A : : ; : : g : . 286

1 The cost of publication of this paper has been met, in part, by grants from the ‘Fondation
Universitaire’ of Belgium, and the Wenner-Gren Foundation for Anthropological Research,
Inc., New York.

2 Present address, Facultés Universitaires, Namur, Belgium.

3 Honorary Curator of Physical Anthropology, South African Museum, Cape Town.

273

Ann. S. Afr. Mus. 48 (16), 1965: 273-397, 24 tab., 16 figs., 13 pls.

274

East Africa
South Africa .

ANNALS OF THE SOUTH AFRICAN MUSEUM

LISTS OF FAUNAL ASSEMBLAGES AT THE AFRICAN HIPPARIONID SITES

Individual hipparionid
Oued el Akrech

sites

Oued el Hammam and Niareeeu

Ain Hanech
Garet Ichkeul
Tozeur ;
Wadi Natrun
Omo

Koro Toro
Kaiso
Kanam.
Eyasi

Laetolil (South Serengeti

Olduvai

Vaal River sannep ee

Cornelia (Uitzoek)
Kromdraai

Bolt’s Farm
Makapansgat

Elandsfontein (Hopefield)

Synopsis (Table 1)

COMMENTARY ON THE PUBLISHED FAUNAL ASSOCIATIONS AT HIPPARIONID SITES IN AFRICA

Miocene

Pliocene

Pleistocene .
North Giles

East and Central Africa

South Africa .

Conclusions based upon the faunal andl foflnee endenee in oe iicetaee

CHRONOLOGICAL RANGE
Upper limit
Lower limit

OF Hipparion

GENERAL DESCRIPTION OF © Eppa TEETH

Upper dentition
Lower dentition

Summary of the sevaciensnes of Cennanen teeth

Upper dentition
Lower dentition
REVIEW OF ADDITIONAL
Description
Ectostylid
Ptychostylid .
Protostylid
Hypostylid
Entostylid
Frequency
Ectostylid
Ptychostylid .
Protostylid
Hypostylid

ENAMEL ELEMENTS (ee OF THE LOWER TEETH

Evolutionary history of ie Soyhids

Evolutionary significan

Ecological considerations

ce of the cones and poids

Taxonomy based on stylids

REVIEW OF PUBLISHED

AFRICAN Hipeaion CRANIAL MATERIAL

~

-~» wet © ees tae

!

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 275

Miocene. : : : ; : : : : W335
Oued el eerie aaa Marceau 5 : ‘ . : : 5 : «| 335
Camp Berteaux . : . : : : : : : : : - 335
Other sites. .. : HY AO
Discussion on the ica pheracter ae aenarin eiennin Reordbenr 1959 - 336

Enamel plications . : ; : : ‘ : : : gor
Protocone : : 5 ; , : : : : : i BD oe
The double knot. : : : : : é : ; : Be eK)
Stylids . ‘ ; é : 4 : : : : : é 330
Hypsodonty . : ; ‘ F ‘ : : - : ‘ OBS

Eocene . : : : ‘ : : : . 5 : é ac 10)
Mascara d : : d : : é d ‘ : ‘ /. 339
St. Arnaud Cemetery : ; : 2 : : : : : . - 339
Ain el Hadj Baba . : : P ‘ : ‘ , ‘ ‘ : - 339
Utique (northern Tunisia) : ; j é . ; : : E | 340
St. Donat oe) : ; : ‘ , : : 5 é ; 340

Pleistocene ‘ : ’ : : ‘ : : f g ‘ - 340
North Afiea - ‘ : ; . , : : : : : 3 e340

Oran. ; ‘ ‘ 2 : , ‘ : : : : ig4o
Ain Jourdel . P : : A d , ‘ : : : 2a 4A
Beni Foudda : ‘ ; ; : : : : : : Le Ba
Wadi Natrun ‘ } : , ‘ ; : , 5 : y egar
Central Africa ; ; : : ‘ ; ; : ; : ; -. 342
more Toro . ‘ , : : : : : . : : jb PGE
East Africa . : ; : t : ; , , ; d ; ago
Oma. ... : ; ; : : : - 342
Olduvai Garkading unpublished eta) : : é : ‘ ‘ - 343
South Serengeti : : 3 ’ : : : ; ; Syhgey!
Lake Eyasi_. : ; : j : : . : f : «346
Lake Albert . : 5 , : : ' : ; : ‘ - 346
South Africa . ; : ; ; ; : : : . ; ‘ BE VAS
Namaqualand : ; : s ; ; : ; : ; - 346
Christiana : ; , i A ; ‘ BAF
Transvaal Pisticlipitheenie cave Mireapind : : , ) ‘ 5 we gag
Cornelia (Uitzoek) . , ; : d ’ : : : : 34
Note on Aipparion incisors é : : LORS

A SUMMARY OF PUBLISHED POSTCRANIAL REMAINS OF AFRICAN jee 2 P joe Bayi

Unpublished data. : : : : : : ; ; - | 352

THE ECOLOGICAL ADAPTATIONS OF Peatusion : : : : : ce

Ecological and functional significance of owe ise ; ; : ; ~ 354
Overall dimensions of the skeleton . : : f : : i - 354
Relative dimensions of cheek teeth and incisors . . . ; ‘ : aya
The development of the preorbital fossa. y . : : : : - 354.
The isolated protocone . : é ; : : : ; s : - 354
Enamel plications . : ; : ' i , ; : ; : i (355
Hypsodonty . : . : ; ; ; ; § : : : - 359
The double knot. : : : ‘ 4 : : : 3 4 . 360
Total tooth structure : : , d i : : ) ; : . 360
Whe extremities < : - 361

The Upper Miocene and Bigistocenc hae Beenie by Afeean fappanio nade : 361

Effects of bionomical conditions on the evolution and the if aaes of Abscat
hipparionids ‘ : : 4 Be

UNPUBLISHED MATERIAL FROM THE VAAL RIVER DEPOSITS, SOUTH AFRICA. : . 365

Sydney-on-Vaal and Pniel_ . : ; : : : -» 1365

MATERIAL FROM LANGEBAANWEG, CAPE ee SOUTH AFRICA. : A 5 « 307

Description of Hipparion teeth ; : . : ; p : : : ee GOW.
Milk dentition : ‘ " ; 2 ; : , ; : gas (o7/

Lower permanent peaution ; ‘ : : i : , ‘ : - 369

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

General description ; ; ; ; : ‘ : : ; , - 370
Upper permanent dentition . : ; ; : ; ; ; «. 372
General dimensions (mm) of P22 ‘ : ; : : ; 3 . 472
Description of selected individual teeth ; ; ; : : s) 9949
Appearance of the tooth sectioned just above the cuddle ; ‘ P s+ 3945
Typical features of the upper teeth . ; , , ‘ ; : - 376
Postcranial equid remains : ; j ; , : : i ; . 378
ORIGIN AND DIFFERENTIATION OF AFRICAN HIPPARIONIDS . ; d . 5 . 382
TAXONOMIC STATUS OF THE AFRICAN HIPPARIONIDS . ’ : ; , y x 36%
Discussion . : ' F , y , , ; : ; : 7). Soe
ACKNOWLEDGEMENTS. : : , ; ; ; ; ; : A - » 392
SUMMARY . : : : ; , : : 7 ; ; ; : - 393
REFERENCES . ; ; : ; P ; ‘ ; ; ‘ : : -. 393
INTRODUCTION
GENERAL

The discovery of the genus Hipparion de Christol 1832 in Africa was first
mentioned as late as 1878 (Pomel) and then again in 1884 (Thomas), but it
was not properly described before 1897 (Pomel). These early papers dealt with
specimens found in the Maghreb, North Africa. Additional finds were then
made in the Oran and Constantine areas, and, somewhat later, specimens
were discovered in Tunisia, in the Nile Valley at Wadi Natrun, in Ethiopia,
in the Kaiso Beds of Uganda, and at Cornelia (Uitzoek) in South Africa. The
scanty stratigraphical data and faunal associations of these discoveries led to
an overmultiplication of genera and species, based mostly on isolated single
teeth.

Subsequently more complete discoveries and more detailed descriptions
were published. In 1937 Hopwood presented some conclusions of the Oldoway
Expedition in a monograph on the fossil Equidae. Dietrich (1942) described
an important hipparionid! series from South Serengeti, to which Arambourg
(1947) added important collections from Omo (Ethiopia). The more extensive
material gave rise to a more unified knowledge of the African Hipparion. It
enabled Gromova (1952), in her revision of the genus Hipparion, to point out
the characteristic features of the African forms, as well as to sketch their
general evolutionary and migratory patterns within Africa.

The lack of stratigraphical correlations and insufficient knowledge of
conditions of exposure and of faunal associations at the various sites where
hipparionids have been discovered, only permitted attempts at relative dating
of the various sites and their fossil horizons. Furthermore, these sites belong
exclusively to Upper Pliocene and Pleistocene epochs. Therefore, it was
generally accepted that Hipparion had appeared in America in Pliocene times

1 The term ‘hipparionid’ refers in a general sense to all forms of African tridactyl equidae.
Previously Dietrich (1942, p. 94) erected the sub-family ‘Hippariinae’ for the same purpose
but without taxonomic definition. Furthermore he also used (1942, p. 94) ‘Hipparion de Christol,
1.w.S.’ in a non-generic sense.

As the present authors have come to the conclusion that all African forms belong to a
single genus Hipparion, the general term ‘hipparionid’ corresponds to the taxonomic one.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 2g) |

and had migrated to Eurasia where the differentiation of the ‘Pontian fauna’
had taken place, and that only subsequently, as late as Pliocene times, Hipparion
had reached Africa as a kind of refuge area. This view was still supported
less than ten years ago when material was discovered in lacustrine limestone
at Mascara (Arambourg, 1956).

Recently Hipparion material was discovered in Algeria in undoubted
Upper Miocene beds. This led Arambourg (1959) to propose the relationships
between the several African hipparionids, and to reduce drastically the former
several genera and species to a more sober and unified taxonomy of three
different groups, namely H. africanum, H. sitifense and Stylohipparion sp. ‘These
formed a chronological succession from the Upper Miocene through the
Upper Pliocene to the ‘Villafranchian’ or early Pleistocene times. This classifi-
cation was more satisfactory and had definite merits and advantages which will
be commented on later in this paper.

THE LANGEBAANWEG DEPOSITS

Recent discoveries (Singer, 1961) of a fairly extensive series of Hipparion
teeth at the Langebaanweg fossil sites, from which Stegolophodon sp. had pre-
viously been reported (Singer & Hooijer, 1958), necessitated a thorough study
and comparative review of the African hipparionids. The unique and uniform
nature of the Langebaanweg material adds materially to our knowledge of
the African hipparionids and also provides new evidence for revising the
relationships between the several forms already recognized in the literature.

Thus the purpose of this paper is twofold:

(a) to record a description and diagnosis of the Hzpparion remains from

Langebaanweg, and
(b) to review and revise the taxonomy of the African hipparionids.

Localization of the sites

The Langebaanweg fossil sites are situated, like the ‘Elandsfontein’ (Hope-
field) site, in the vast Sandveld of the Western Cape, and are located about
75 miles (120 km.) northwest of Cape Town (fig. 1) and 12 miles (19 km.)
from the late Middle Pleistocene—early Upper Pleistocene site at ‘Elands-
fontein’. The former are 150-200 feet (46-62 metres) above sea-level, while
the latter are situated at 300 feet (Singer, 1961).

The fossils are mainly located in and around two quarries being commer-
cially exploited for phosphates by the African Metals Corporation (AMCOR).
‘The ‘Varswater’ site is situated on the farm ‘Langberg Suid’, and the ‘Baard’s’
site is on the farm ‘Remainder of Langberg’, the two being subdivisions of
‘Langberg’ (fig. 2).

Estory-of the discoveries

The first hipparionid teeth were recovered from Baard’s Quarry in 1958.
In 1961, Mr. Robin Warren, an employee of AMCOR, recovered two excel-

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

Great pers ie
LANGEBAANWEG L

SALDANHAg*° 6 NOPE IECe
BAY 4

er

A Sout bee

Elandsfontein

© CAPE TOWN

English miles

Fic. 1. Map indicating the position of Langebaanweg relative to Cape Town.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 279

_lently preserved milk molars at Varswater. Thereafter the remainder of the

specimens were found at various sites in the two quarries.

Geology

The geology of the more than 4 square miles of fossiliferous deposits is
not easy to interpret. There is little with which to compare it, as virtually
nothing is known of the Quaternary geology of the Cape Province.

The surface siliceous sands, possibly aeolian, forms a discontinuous cover
of 4 to 5 feet, which may extend to more than 15 feet in a few places. Below
this layer, at both quarries, there occurs a Dorcasia-type caliche, discontinuous
and stratigraphically above the phosphates. There is also ferricrete, younger
than the phosphates and intimately associated with them. |

The Dorcasia-type surface limestone also occurs on the solid calcrete
dunes at ‘Elandsfontein’, at Saldanha Bay, at False Bay (Singer and Fuller,

1962) and elsewhere. The distribution of the caliche is patchy, and is

considerably stripped by erosion.

At Varswater the phosphate, a variety of collophane, occurs as the cement-
ing material of the consolidated sand, probably as the result of diagenetic
replacement of interstitial limes. There are three main layers which vary
laterally in thickness within a range of 3 feet to 6 feet.

At Baards, the phosphate occurs as nodular to bouldery phoscrete (up to
5 feet thick). The phoscrete is a hard, compact mass of phosphatized sand
consisting of sand grains which have been cemented and partially corroded
by an amorphous calcium phosphate cement. It overlies the phosphatized
nodular sands, sandy clays, and clayey sand. The phoscrete and nodular
phosphate sands, represent replacement of older shelly sands by phosphate
solutions, the phosphate of which is probably derived from guano. A significant
feature is that at Varswater the phosphate has remained unaltered as lime
phosphate (Singer & Warren, in preparation).

The discovery of shark teeth and other marine forms (vide infra) at the
same level as and with the land mammals indicates earlier invasion of the
area by the sea. Studies on borehole cores and the presence of shelly sands and
gravels suggest the formation of sand bars, especially at Varswater, while the
clay and clayey sands at Baard’s indicates a possible lagoonal stasis. Further
studies are being carried out in an attempt to clarify the geological genesis and
morphology of the region.

The major portion of the phosphatic deposits is located at both sites at
up to 100 feet above sea-level. The archaic fossils are related to or found just
above these deposits. The ‘rolled’ nature of much of the fossil material suggests
some transport prior to final deposition. The upper phoscrete and calcrete
layers, as well as the clayey deposits, contain the faunal forms represented
also at ‘Elandsfontein’ (Ceratotherium, Equus, Sivatherium, Homowoceras and various
antelopes), typical of the late Middle Pleistocene—early Upper Pleistocene
(“Hopefield Period’).

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

On a comparative basis with other sites, it seems likely that the stegolo-
phodont—stegodont—archidiskodont complex and the hipparionids derive
from a period extending from the Lower Pleistocene to the early Middle
Pleistocene. The stegolophodont and stegodont specimens together indicate a
very early phase of the Lower Pleistocene. Elsewhere (e.g. at Olduvai)
archidiskodont material overlaps this phase and extends up to the Middle
Pleistocene. It will be demonstrated below that the hipparionids from Lange-
baanweg belong to the same species as that from the lower layers of South
Serengeti, so that their presen e is suggestive of the earliest phases of the
Pleistocene.

FAUNAL ASSEMBLAGE FROM LANGEBAANWEG

The following identifications have been made thus far:

MOLLUSCA ARTIODACTYLA
Trigonephrus sp. Hippopotamus sp.
| Stvatherium(*) olduvatense
CARNIVORA cf. Oreotragus sp.
cf. Arctocephalus cf. Damaliscus sp.
Canis sp. cf. Redunca sp.
Hyaena brunnea Homotoceras sp.
| REPTILIA
PROBOSCIDEA Cf lieeiaca
Stegolophodon sp. Buide
Stegodon sp.
Archidiskodon sp. cf. Struthio
Palaeoloxodon sp. SELACHII(?)
Isurus cf. glaucus
PERISSODACTYLA Lamna nasus
cf. Ceratotherium Carcharias sp.
Eipparion (Hipparion) albertense baardi C’. ferox
subsp. nov. Rhinoptera cf. dubia
Equus helmer Glopias vulpes

(1) According to Arambourg’s recent re-description (1960) of Pomel’s type specimen from
St. Charles (Oran), Libytherium maurusium has precedence (see also Singer and Boné, 1960).
(?) Identified by Dr. B. Schaeffer, American Museum of Natural History, New York.

METHODS AND NOMENCLATURE

For the description of the occlusal surfaces of the teeth, the nomenclatures
used by Arambourg (1947, 1959), Cooke (1950), and Sondaar (1961) have
been modified (see pp. 325-8) and synthesized (figs. 10, IT).

The measurements of the teeth have been taken according to the method
described by Sondaar (1961).

SALDANHA

BAY

ATLANTIC

~ To

LANGE BAAN

Vredenburg

t=)
Varswater

quarries
%

LAGOON

SY

4
ss
3s

NN
XQ quarry
C)

»

y

7 LANGEBAANWEG

=/andsfontein

SS Principal roads

Sx. Railway line

_yo— Contours (Elevations in feet; vertical
interval 50°)

8 Principal fossil sites

Sand dune area on the farm
Elandsfontein

SCALE
0 1 2

Miles 1
4000 O 4000 2000 3000 4000
Yards becacecens te ed

Fic. 2. Contour map showing the location of the quarries at Langebaanweg.

oy f

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 281

The indices devised by the authors are described at appropriate places
in the text.
The following abbreviations refer to the collections and sites of the source
material:
Archaeol. Surv. = Archaeological Survey of South Africa, now in the Univer-
sity of the Witwatersrand, Johannesburg, South Africa.

B.M.N.H. = British Museum (Natural History)

C = Cornelia

L = Langebaanweg

M.M.K. = McGregor Memorial Museum, Kimberley
Nas. Mus. = Nasionale Museum, Bloemfontein

Old. = Olduvai

S.A.M. = South African Museum, Cape Town

w-C: = University of California, Berkeley.

GEOGRAPHICAL DISTRIBUTION OF AFRICAN SITES WHERE HIPPARIONIDS
HAVE BEEN RECORDED (figs. 3-7)

Site Specimen Reference

Morocco
Camp Berteaux (Gara Ziad) (a) Molar teeth
Femur fragment with proximal

epiphyses Bourcart, 1937
(6b) Left M? Ennouchi & Jeannette,
1954
Oued el Akrech (near Argoub
el Hafid) Ennouchi, 1951
Fouarat (south of Kenitra Choubert, Ennouchi &
(Rharb)) Marcais, 1948
ALGERIA
Tafna (Guiard, Prov. Oran) Dalloni, 1915
Oued el Hammam Adult skull no. 141 Arambourg, 1959

Adult skull no. 125

Adult skull with dentition no. 116

Juvenile palate with dentition
no. 122

Adult mandible no. 143

Adult half mandible no. 89

2 juvenile mandibles with milk
teeth nos. 95-96

Several teeth series, juvenile and
adult

Isolated teeth

Humerus, distal fragment no. 159

Radio-ulna no. 123, fragments
no. 13, no. 22 and no. 27

Femur: fragment without epiphyses
and distal fragment

Tibia: distal fragment

Pelvic fragment

282

Site

Oued el Hammam (cont.)

Marceau
Mascara

St. Donat
St. Arnaud (Cemetery)

Ain el Hadj Baba

Ain el Bey
Mansourah
Setif

Beni Foudda (Ain Boucherit)

Ain Hanech
Oran (Puits Kharouby)
Ain Jourdel

TUNISIA

Garet Ichkeul
Tozeur Beds

Utique
Djebel M’dilla

FAYUM

Wadi Natrun, Gart el Moluk

Hill

ETHIOPIA

Omo Valley

CENTRAL AFRICA

Tchad, Koro Toro region

Goz-Kerki
Koula
Bochianga
Quadi Derdeney
Koula Ri Katir

(It is not clear at which of

these five sites

Stylo-

hipparion has been found)

ANNALS OF THE SOUTH AFRICAN MUSEUM

Specimen

Astralagus, 2

Calcaneum, 1

Numerous complete metapodials
(5 Mc III, 5 Mt III), many (6)
of them with lateral digits

P4 and Ma, P,

Skull fragments of one individual:
palate with nearly complete left
dental series, and half right den-
tal series; symphysis with incisors

“Teeth of two jaws’

(a) upper molars
(b) lower molars
Metapodials with developed lateral
digits
Upper teeth: right P4*-M?
left M?

’ Lower teeth

Ma, left P; or Py, Ma
M;, P

Left upper P (P* ?)

(a) 1 incisor, 5 M,, 1 M@ (brought
back by Brumpt from the du
Bourg de Bozas Mission, 1903)
(6) right P? no. 396

left M? no. 395

left M@ fragment no. 370
right M, or M, no. 3 & no. 4
right humerus fragment

Reference

Arambourg, 1959

Arambourg, 1956
Joleaud, 1936
Arambourg, 1956
Pomel, 1897
Arambourg, 1956

Thomas, 1884

Thomas, 1884
Thomas, 1884
Thomas, 1884
Pomel, 1897
Arambourg, 1949
Pomel, 1897
Thomas, 1884

Arambourg, 1949
Roman & _ Solignac,

1931
Solignac, 1927

Solignac, 1927

Andrews, 1902

Haug, 1911; Joleaud,
1933

Arambourg, 1947

Coppens, 1960

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE

Site
EAST AFRICA

Uganda: East shore of Lake
Albert in Kaiso layers

Kenya: Kanam lower beds.
Kanjera
Tanganyika: North-east shore
of Lake Eyasi (W. and N.
of Mumba Hill)

South Serengeti (various loca-
lities)
Olduvai
SOUTH AFRICA
Cape Province:
(i) Namaqualand (40 mi.
east of Springbok)

Sydney-on-Vaal

Pniel, near Barkly

West

(c) Vaal River at Chris-
tiana

Orange Free State:

Cornelia (Uitzoek)

(ii) (a)
(0)

Transvaal:
Kromdraai Cave
Bolt’s Farm
? Makapansgat

Specimen

I incomplete upper molar
BM Mr12615

Teeth

Loose teeth, fragmented mandibles

Left P,-M, and M,

Right P,, M,, M, and part of
M;
(above specimens are S.A.M.

9982)
Left M? (MMK 431)

Left P+ or M, (MMK 5225)
Left M? or M? (Arch. Surv. 113)

(a) right M? (or M?) (Nas. Mus.
C558)
left M® (Nas. Mus. C555)
left M, + M, (Nas. Mus.
C556)

(6) symphysial fragment and in-
cisors

(c) right M, (Nas. Mus. C797)
left P, (Nas. Mus. C796)

(d) left P,-M, (Nas. Mus. C795)

2 lower teeth (discovered by Broom)

NOTE: M? indicates upper molar; M, lower molar, etc.

283

Reference

Hopwood, 1926

Reck & Kohl-Larsen,
1936

Dietrich, 1942
Hopwood, 1929, 1937

Haughton, 1932

Cooke, 1950

Van Hoepen, 1930

Van Hoepen, 1930
Van Hoepen, 1932

Van Hoepen, 1932
Cooke, 1950

Cooke, 1963
Cooke, 1963

SUMMARY OF PUBLISHED GEOLOGICAL EVIDENCE FROM

AFRICAN HIPPARIONID

SITES

The present state of our knowledge of the geological evidence from sites
in Africa where hipparionids have been discovered indicates that they first
appeared in Upper Miocene deposits. A summary of the data is considered
useful for the assessment of the status of the Langebaanweg material.

MIOCENE

The continental layers of vertebrate fossils at OUED EL HAMMAM
(between Mascara and Mercier Lacombe, upstream from Bou Hanifia,

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 3. Map of Africa indicating major areas of discovery of hipparionids. Details of areas
A-C are shown in figs. 4-6, while area D is enlarged in fig. 1.

Algeria—figs. 3, 4) occur between two marine horizons, namely, (1) a Burdi-
galian one, characterized by its mollusc and echinid fauna and (2) an Upper
Miocene Lithotamnium limestone. On this stratigraphical basis, the Hipparion
horizon has been dated as Tortonian, i.e. Upper Vindobonian (Late Miocene)
immediately antedating the Tortono-Sahelian transgression which occurs at
the end of the Miocene marine period in this area (Arambourg, 1959).

The MARCEAU deposit, and especially its lignite and lacustrine clay
and limestone which yield the Hipparion specimens, are referred to a period
anterior to the Pliocene transgression, i.e. they belong to the Upper Miocene
and may be equated with Oued el Hammam (Flandrin, 1942).

ee ees en ee ee

a ee Tg ee ee ee ae es

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285

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE

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286 ANNALS OF THE SOUTH AFRICAN MUSEUM

There is good evidence that CAMP BERTEAUX (near Taourirt, Eastern
Morocco) belongs to this period. The Hipparion layers, from which Mastodon
cf. angustidens has been recovered (Bourcart, 1937), form the base of a lacustrine
series of ‘argiles smectiques’ which are lying on marine horizons with Tortonian
fauna and are covered with Pontian deposits. Choubert and Ennouchi (1946)
have dated the Hipparion layers as Upper Tortonian, being a sub-continental
phase at the end of the Miocene period. Thus, they are somewhat younger
than those from Oued el Hammam (Arambourg, 1959).

The brack water deposits of TAFNA (near Guiard, Province of Oran)
show a ‘Sarmatian facies’ of the Upper Vindobonian. On faunal and litho-
logical grounds, they were first referred to the ‘classical Pontian’ (Dalloni,
1914), but recently they were re-studied and correlated with the Oued el
Hammam horizon (Perrodon and Tempere, 1953). However, the geological
evidence is not clear.

PLIOCENE

Pliocene palaeontological sites are rare in the Maghreb. Some fossiliferous
lacustrine formations in the SETIF-CONSTANTINE area have been referred
to this period (Middle Pliocene). Included are also the limestone deposits of
the ST. ARNAUD Cemetery and of the MASCARA region, AIN EL BEY,
AIN EL HADJ BABA, ST. DONAT, and possibly UTIQUE in Tunisia.

Most of these exposures correspond to the fluviolacustrine filling of a
vast depression between the Tellian and Sahara—Atlas. According to Arambourg
(1956) the Aipparion sitifense layers of St. Arnaud Cemetery, Ain el Hadj Baba
and Mascara are lithologically different (sands and lacustrine limestones)
from the Stylohipparion and Equus numidicus deposits of Ain Boucherit and Ain
Jourdel (gray marls and red conglomerate). While these represent the base

of the ‘Villafranchian’ (similar to Lac Ichkeul and Fouarat), they are referred —

to the Middle Pliocene. No satisfactory geological evidence is available for
TOZEUR (Tunisia): it was referred by Boule (1910) and Solignac (1931) to
the ‘Pontian’ stage because of the presence of Merycopotamus and Hipparion.
Now that the occurrence of these forms is known in Upper Miocene deposits
in India and North Africa, there is a special need for obtaining better
stratigraphical evidence for dating this site.

At WADI NATRUN (Gart el Moluk Hill) in the Nile Valley, fluvio-
marine gypsiferous clays (containing, among other rare vertebrates, Hipparion
albertense) are interbedded with limestones and shales, and deposited in an
arm of the Mediterranean in the Nile Valley. On a stratigraphical basis they
are dated as Middle Pliocene.

PLEISTOCENE
North Africa

The Hipparion fossiliferous horizons in the Maghreb correspond strati-
graphically to the marine regression following the major Pliocene extension.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 287

The most significant exposures are St. Arnaud and Oran in Algeria, and Lac
Ichkeul, near Bizerta, in Tunisia (figs. 3, 4). ‘The sequence has been extensively
described and discussed (Howell, 1959). The general feature is that of a
300-foot series of marls and lacustrine clays and silts, with interbedded
fossiliferous sandstones, gravels and calcareous conglomerates.

At LAC ICHKEUL the base is formed by deep-water Plaisancian and
lagoonal Astian marine deposits capped with fresh-water beds from the early
Pleistocene, with intercalation of a one-foot thick conglomerate.

At BEL HACEL eroded dune sandstones overlying transgressive Astian
marls and sands are bearing the ‘Villafranchian’ fauna: they are further overlain
by the 100-metre high Sicilian beach.

In the ST. ARNAUD area, two distinct fossiliferous horizons are visible.
The lower one (Ain Boucherit, i.e. Beni Foudda) is a coarse calcareous conglo-
merate; the upper one (Ain Hanech) is a cracked, rather sandy or gravelly
clay, intercalated in the thick mar] filling the old marshy or lacustrine depres-
sion. Faunal and stratigraphical considerations make the St. Arnaud upper
deposit somewhat younger than Lac Ichkeul.

At ORAN (ST. CHARLES), where Libyhipparion (?) libycum was first
recovered and named by Pomel (1897), the consolidated dune sandstones and
clays are concordantly overlying the calabrian sandstones, of which they
represent the regression facies. The calabrian itself is in direct contact with the
Flabellipecten flabelliformis sands and sandstones of the marine Pliocene; and at
the base, the sequence reaches the Tortono-Sahelian marks and red stone of
the Upper Miocene (Arambourg, 1960).

Ligniferous clays of the PUITS KHAROUBY near Oran from which
H. massoesylium Pomel was recovered, has usually been placed—on the basis
of its mollusc fauna—in the Upper Pliocene, but there is little geological

_ evidence for it being older than Lower Pleistocene.

East Africa

Most of the fossiliferous horizons of East Africa (figs. 3, 5) are linked with
tuffs, ashes or lapilli projected and deposited by volcanic eruptions. This
volcanic activity spans over a long period of time, probably from Lower
Miocene up to Upper Pleistocene and even to historical times. Therefore it is
difficult, on purely geological grounds, to date the different fossil deposits.
Usually it is only by convergence of tectonic, stratigraphical, lithological and
chemical methods on the one hand, and by faunal studies on the other that it
is possible to propose a satisfactory date.

At KANAM, on the slope of the dissected volcanic centre of Homa

- Mountain, the lacustrine brown calcareous fossiliferous clays, with their

interbedded fine tuffs from intermittent volcanic explosions, are commonly
referred to the Lower Pleistocene.

The KAISO series is complex. It has been repeatedly studied (Wayland,
1926; Fuchs, 1934), and recently Lepersonne (1949) subdivided it into three

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

A dl Po victoria @
Lake Kivu ; A é

sath, B
A] Pree
x

has been described, namely, 1. Lake Albert; 2. Kanjera; 3. Mumba Hills, Lake Eyasi; 4. Olduvai

|
:
:
|
: 4 % “Zs AS. SS SS
M7 ” 250
Fic. 5. Map of Central-East Africa (B in fig. 3) indicating the localities from which Hipparion :
Gorge; 5. South Serengeti, Laetolil beds.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 289

main stages of sedimentation, the middle one being highly fossiliferous. It is
a clayey deposit, alternating with thin beds of fine sands and sandstones, and
discontinuous ironstone horizons and limonite lenses (Kaiso bone beds),
containing silicified wood and a high proportion of vertebrates. It overlies a
silty lower deposit with some minor gravel horizons, resting on a basal ironstone
layer (laterite) capped by unstratified sands above the down-faulted peneplain
surface. This lower part of the Kaiso series, including the fossil horizon, is
usually placed in the Lower Pleistocene (Howell, 1959; Bishop, 1963).

The OMO beds are lithographically comparable to the Kaiso deposits.
The ‘Villafranchian’ fauna, and the Hipparion material in particular, occur in
sandstone horizons between lacustrine volcanic clayey tuffs. In the evolution of
the Rudolph depression in its Nilotic or open phase, Arambourg (1943) has
distinguished two major extensions of the lake during the Pleistocene: (1) the
vertebrate beds occur in a sand and sandstone horizon of the lower Omo
Valley, representing a first phase of lacustrine deposition prior to the general
tectonic uplift of the end of the Lower Pleistocene. (2) Posterior to the tectonic
dislocation, the second phase corresponds to the series of horizontal terraces
on the slope of the Rudolph Basin—they are not fossiliferous. This distinction,
confirmed by the study of the fauna, permits the fossil beds to be dated as
Lower Pleistocene, probably younger than Kaiso and Kanam.

It is more difficult to specify the geological location of the SERENGETI
Hipparion material. The Laetolil beds in the Vogel River area show a sequence
of upfaulted subaerially deposited yellow and gray tuffs (Kent, 1941). Unfor-
tunately the fauna collected by Kohl-Larsen ‘in den Schluchten und Wasser-
rissen’ of the South Serengeti was probably recovered from various horizons
which were altered and mixed-up by subsequent erosion (Dietrich, 1942). The
gray tuffs, about 20 m. thick, which appear to contain the ‘older fauna’ (Kohl-
Larsen, 1939), are dated as Lower Pleistocene, more or less contemporary
with Omo and Olduvai I. Efforts have been made to identify this stratigraphic-
ally older fauna by such means as the type of mineralization and colour.
Conclusions based on this method can only be accepted with reserve because
of the fact that possible weathering may have altered the original appearance
of the fossils. Nevertheless, Dietrich (1942) believed that all his Hypsohipparion
material derived from the lower gray tufts.

The vast sequence of stratified beds exposed in the OLDUVAI GORGE
(northern Tanganyika) has been extensively discussed in recent years, and
lately by Pickering (1960) and Leakey (1963). Independently of the chemical
dating of the beds, which provides evidence of a great duration and antiquity
of Bed I, dating has also been attempted on studies of the fauna- and on a
climatological basis. Little has been added from the strictly geological point
of view. It is important to note that the major climatic, geological and appar-
ently faunal ‘break’ occurs near the base of Bed II and not, as was previously
thought, between Beds I and II. Most workers agree that Olduvai I belongs
to the Lower Pleistocene, being more or less contemporary with (and probably

ce '

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

prior to) Omo and Laetolil, while Beds II, III, IV are considered to belong to
the Middle Pleistocene.

The KANJERA exposures, on the low cliffs of the Homa Mountain close
to Lake Victoria, consist of a tripartite series of basal greenish ash and tuffs,
succeeded by clays with limestone and by upper transgressive beds of brown-
greenish clay. Most of the fossils are obtained from and in close association
with the lower tuff and ash layer, but it is not possible to be certain whether
or not all the material came from this source (Kent, 1942). However, there
is sufficient evidence that the Hipparion may be considered to be more ancient
than the human fossil remains which were recovered from the middle beds.
The Kanjera faunal beds are referred to the Middle Pleistocene, roughly
contemporary with Olduvai IV.

The LAKE EYASI beds are usually referred to an early Upper Pleistocene
period, more on a palaeontological than on a geological basis.

According to Reck and Kohl-Larsen (1936), it is probable that the
Hipparion teeth recovered west and north of Mumba Hill, in the north-east
section of the Eyasi basin (see p. 283), do not belong to the original beds. Indeed
they are very much rolled, which is not the case for the presumably contem- ©
poraneous material, and constitute a secondary deposition. If this interpretation :
is correct, these teeth are probably more ancient than the typical Eyasi fauna,
and should be dated to a period prior to the Upper Pleistocene.

South Africa

The only relatively satisfactory stratigraphical information concerning
HMipparion in South Africa is obtained at the YOUNGER GRAVELS OF THE

CHRISTIANA

z= Sy dney- .
on-Vaal

Pniel
Estates

Sca/e in miles

ZKIMBERLEY

Fic. 6. Map of northern Cape Province, South Africa (C in fig. 3) indicating localities from M
which Hipparion has been described, namely, Sydney-on-Vaal, Pniel and Christiana. —

elie! 5 ss let En NS

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291

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE

terre terranes ~ — as

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

VAAL RIVER (Christiana; Pniel; Sydney-on-Vaal: fig. 6). In this horizon,
which actually presents the first fossil occurrence in the Vaal area (as fossils
of the oldest gravels have probably been destroyed), three phases have been
recognized from their elevation and their stone artefact associations. However,
most of the fossils seem to have derived from Phase II and III, although it is
not impossible that the fossils found below Barkly West may have been contained
in Phase I of the Younger Gravels (Cooke, 1963). On the basis of a climatologi-
cal interpretation (Séhnge, Visser and Lowe, 1937; amended by Cooke, 1947,
and Lowe, 1952), the Younger Gravels were correlated with the Middle
Pleistocene Olduvai Beds II and IV.

The CORNELIA beds, an open site on the fous ‘Uitzoek’ in the
Orange Free State, consists of exposures of clays, gravels and sand, which
represent an old ‘pan’ filling. Oakley (1954) emphasizes the Pleistocene nature
of the site by referring to a ‘lower Fauresmith industry’ from the beds. There is
no reliable date based purely on stratigraphy for this site.

At present it is also impossible to make any tentative geological dating
for the Notohipparion material from NAMAQUALAND. It was recovered from

a granite level in a well, 60 feet down in the surface limestone, 40 miles east of |

Springbok. It is known that the infilling of the valleys with sand has been
occurring in Namaqualand from Upper Cretaceous times, with probable
breaks in the sedimentation, but there is no information on the actual sequence
of gravels and limestone in the well (Haughton, 1932).

The Hipparion occurrence in cave fillings at KROMDRAAT, (?) MAKA- —
PANSGAT and BOLT’S FARM has been dated by faunal and climatological
methods (Ewer, 1957; Brain, 1958), and nothing further can be added from
a purely stratigraphical point of view.

LISTS OF FAUNAL ASSEMBLAGES AT THE AFRICAN HIPPARIONID SITES

It is considered useful to include lists of all the diagnosed, published fauna
from the various sites where Hipparion has been recovered. Omission of particular
sites is due to lack of information, and Hopefield is included because of the
richness of fauna and for comparison with contemporaneous sites. The faunal
assemblage of Langebaanweg is listed on page 280. The sites are listed according
to the order presented on pages 281-3. Wherever possible, generic and specific —
names are retained as they appear in the publications quoted.

INDIVIDUAL HIPPARIONID SITES
Oued el Akrech (Morocco) (After Arambourg and Choubert, 1957)

PROBOSCIDEA PERISSODACTYLA
+Anancus osiris +Stylohipparion sp.
+Elephas africanavus

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 2903

Oued el Hammam (Oran) and Marceau (Algiers) (After Arambourg, 1963)

PRIMATES

+ Macaca flandrin
RODENTIA

+ Hystrix sp.
CARNIVORA

+ Hyaena algeriensis
TUBULIDENTATA

+ Orycteropus mauritanicus
PROBOSCIDEA

+ Turicius sp.

+ Mastodon sp.

PERISSODACTYLA

{Dicerorhinus primaevus

+Hipparion africanum

ARTIODACTYLA

+Palaeotragus germaini
+Samotherium sp.
{Damalavus boroccor
tGazella praegaudryt
+ Tragocerus sp.
+Cephalophus sp.

AVES

{Struthio sp.

Ain Hanech (Algeria) (After Arambourg, 1947; 1949)

CARNIVORA
Hyaena sp.
Canis sp.
PROBOSCIDEA
+ Anancus osiris
tElephas cf. planifrons
+E. aff. meridionalis or recki
PERISSODACTYLA
tAltelodus aff. sumus
tStylohipparion libycum
Equus sp.

(4) According to Leakey, 1958= Mesochoerus.

ARTIODACTYLA

Hippopotamus amphibius
+ Omochoerus(*) sp.
Giraffa sp.
tLibythertum maurusium
Bos sp.
{Bubalus sp.
Oryx sp.
Alcelaphus sp.
+tGazella sitifensis

Garet Ichkeul (Tunisia) (After Arambourg, 1949)

PROBOSCIDEA

tAnancus osiris

tElephas cf. planifrons

PERISSODACTYLA
tAtelodus aff. simus
{Stylohipparion libycum

Equus

ARTIODACTYLA

Hippopotamus sp.

+Libytherium maurusium

Bos sp.

Oryx sp.

Alcelaphus sp.
tGazella aff. sitifensis

Redunca sp.

Tozeur (Tunisia) (After Roman & Solignac, 1934)

PROBOSCIDEA
{Mastodon cf. longirostris

PERISSODACTYLA

+Hipparion sp.

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

ARTIODACTYLA + Tragocerus amaltheus
+ Merycopotamus aff. dissimilis +Hemitragus perimensis
+Capreolus matheron Antilopinae

Wadi Natrun (After Studer, 1898; Andrews, 1902; Arambourg, 1947 and 1963)
MAMMALIA ARTIODACTYLA
+ Hippopotamus protamphibius
+ Stvachoerus giganteus
Camelidae gen. sp. indet.

PRIMATES
+Libypithecus markgraft

CARNIVORA tLibythertum maurusium
+Machairodontidae gen. sp. indet. + Hippotragus (?) cordieri
Phocidae gen. sp. indet. REPTILIA
Lutra sp. :
Crocodilus |
HN: Oliaaih + Euthecodon nitriae i
? Leporidae + Sternothaerus dewitzinanus 4
PROBOSCIDEA + Trionyx pliocaenicus :
+ Mastodon sp. PISCES ‘
PERISSODACTYLA Protopterus $
+Hipparion sp. Synodontis :
Omo (After Arambourg, 1947)
PRIMATES tNotochoerus capensis (*)
+ Dinopithecus brumpt +Phacochoerus africanus fossilis(*)
CARNIVORA Giraffa camelopardalis
+Homotherium (?) ethiopicum t Giraffa gracilis q
PROBOSCIDEA {Sivatherium olduvaiense(*)
| Deinotherium bozasi i Manors Dries sigh.
+ Elephas (Archidiskodon) recki Hobus, (hobus) appears
+ BlPhie | deceoden) ch een
+ Redunca ancystrocera
PERISSODACTYLA Alcelaphus sp. :
P
tAtelodus cf. germano-africanus + Strepsiceros imberbis ;
+ Stylohtpparion albertense Aepyceros melampus ;
Equus cf. zebra Antidorcas sp. q
ARTIODACTYLA Oryx cf. gazella }
+Hippopotamus (Tetraprotodon) prot- Tragelaphus nakuae vy
amphibius Taurotragus cf. procanna |
+t Omochoerus heseloni(*) Gazella praethomsoni
+ Metridiochoerus andrewsi(*) Syncerus aff. brachyceros 4
(1)= Mesochoerus heseloni (Leakey, 1958). (?) = Pronotochoerus jacksoni (Leakey, 1958).

(3) = Notochoerus euilus (Leakey, 1958).

(*) According to Leakey, 1958= Tapinochoerus meadowsi, but does not belong to the Omo
Beds.

(5) See footnote to Langebaanweg fauna (p. 280).

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 295

Koro Toro (Tchad) (After Abadie, Barbeau and Coppens, 1949; Coppens,

1960)
MAMMALIA PERISSODACTYLA
PRIMATES + Stylohipparion sp.

; Ceratotherium si
}‘Australopithecus’ sp. lie

ARTIODACTYLA

+ Hippopotamus cf. protamphibius
+ Sivatherium(*) sp.

CARNIVORA

Hyaena cf. striata

ee CIDE! Giraffa camelopardalis
tArchidiskodon africanavus(*) Alcelaphus sp.
tA. recki(?)
tAnancus sp. REPTILIA
+Stegodon sp. Crocodilus niloticus
1) According to Cooke, 1960= Elephas africanavus.
g
(?) According to Cooke, 1960= Palaeoloxodon recki.
(*) See footnote to Langebaanweg fauna (p. 280).
Kaiso!
CARNIVORA + Metaschizotherium hennigi
+? Homotherium ethiopicum Ceratotherium simum
PROBOSCIDEA
ARTIODACTYLA

+ Stegodon kaisensis

tArchidiskodon exoptatus tHippopotamus imaguncula

+H. protamphibius

PERISSODACTYLA + Notochoerus euilus
+ Stylohipparion albertense + Mesochoerus limnetes
Kanam!
CARNIVORA PERISSODACTYLA
Crocuta crocuta + Stylohipparion albertense

+ Equus oldowayensis
Ceratotherium simum
Diceros bicornis

PROBOSCIDEA ARTIODACTYLA
tAnancus kenyensis + Hippopotamus imaguncula
+Stegodon kaisensis + Metridiochoerus pygmaeus
tArchidiskodon subplanifrons +. Nyanzachoerus kanamensis
TA. exoptatus Giraffa camelopardalis
+Deinotherium bozasi +Libytherium olduvatense

1 Compiled from Hopwood, Leakey & McInnes in Leakey, 1951; Dietrich, 1950; McInnes,
1953; Cooke, 1963; Leakey, 1958.

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

Eyasi!
PRIMATES
Homo sp.
RODENTIA
+Pedetes surdaster
Thryonomys swinderianus
CARNIVORA
Crocuta crocuta
Caracal caracal
Panthera leo
Panthera pardus
TUBULIDENTATA
+ Orycteropus aethiopicus
PERISSODACTYLA
Equus burchelli
+ Stylohipparion sp.

Diceros bicornis

Laetolil (South Serengeti)?
PRIMATES
+ Simopithecus oswaldi

RODENTIA

+Pedetes surdaster
+Hystrix galeata

CARNIVORA
Canis mesomelas
C. mesomelas latirostris
+Canis africanus
Crocuta crocuta
Caracal caracal
? Panthera pardus

TUBULIDENTATA
+ Orycteropus aethiopicus

PROBOSCIDEA

tAnancus kenyensis
tArchidiskodon subplanifrons
tA. exoptatus

tA. recki

+ Deinotherium bozasi

1 See footnote p. 295.

ARTIODACTYLA
Hippopotamus amphibius

Potamochoerus kowropotamus

Giraffa camelopardalis
Strepsiceros strepsiceros
Taurotragus oryx
Syncerus caffer
+Homotoceras nilsoni
Kobus ellipsirymnus
+ Adenota kob
}Redunca redunca
Pelea sp.
t Oryx beisa
Aepyceros melampus
+tGazella granti

PERISSODACTYLA

+ Stylohipparion albertense
Equus burchelli

+ Metaschizothertium hennigi
Ceratothertum simum

+ Serengeticeros efficax

ARTIODACTYLA
Hippopotamus amphibius
+. Notochoerus euilus
+ Okapia stillea
Giraffa camelopardalis
+Libytherium olduvaiense
+ Tragelaphus buxtoni
Taurotragus oryx
Syncerus caffer
? Kobus ellipsirymnus
+Redunca redunca
+ Hippotragus equinus
+Damaliscus angusticornis
Aepyceros melampus

} a, epaceglic ay neh Sa et

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 297

Olduvai (After Leakey, 1958, and Cooke, 1963)
Numbers I, II, III and IV refer to the various beds

PRIMATES
+ Simopithecus oswaldi (1, II, III, IV)
+S. jonathani (II, ? IV)
+Paranthropus boiser (1, IT)
+Homo sp. (1, 11)

CARNIVORA

Canis mesomelas (II, IV)

C. mesomelas latirostris (II, IV)
+Canis africanus (II, IV)

Aonyx capensis (11)

Crocuta crocuta (1)

Panthera leo (II, I11)

P. pardus (? I)

PROBOSCIDEA
tAnancus kenyensis (1, 11)
+ Archidiskodon exoptatus (1)
ta. reo (1, Tf, III, TV)
+ Deinotherium bozasi (I, II)

PERISSODACTYLA

+Stylohipparion albertense (I, II, III,
IV)

Equus burchelli (11, 111, TV)

TE. aff. grevy: (I, II, III, IV)

TE. oldowayensis (1, II, III, IV)

+Metaschizotherium hennigi (1, I1)

Ceratotherium simum (I, II, II, IV)

tSerengeticeros efficax (II)

Diceros bicornis (1, II, III, IV)

ARTIODACTYLA
tHippopotamus gorgops (1, II, III,
IV)
+ Notochoerus hopwood: (III, IV)
+N. compactus (11)

+ Mesochoerus olduvaiensis (1, II, III,
IV) |
+Potamochoerus majus (1, II, III, IV)
Phacochoerus africanus (II, III, IV)
+P. altidens altidens (11, III, IV)
+P. altidens robustus (I, IT)
+ Tapinochoerus minutus (IV)
+T. meadows: (I, II, III, IV)
tAfrochoerus nicoli (II, III, IV)
+ Metridiochoerus andrews: (1)
Orthostyonyx brachiops (11)
+ Okapia stiller (? 1)
Guiraffa camelopardalis (II, 1V)
+G. gracilis (11)
tLibytherium olduvaiense (1, II, III,
IV)
Strepsiceros strepsiceros (? I, II, IV)
tStrepsiceros imberbis (? I, II, IV)
+ Tragelaphus buxtont (11)
Taurotragus oryx (1, II, IV)
+ Homoioceras nilssoni (IV)
+Bularchus arok (11, III, IV)
+Adenota kob (I, II, 111)
+Hippotragus equinus (III, IV)
+H. nro (II, IV)
t Oryx beisa (1)
+ Damaliscus angusticornis (II, IV)
+D. teste (I, II, IV)
+ Alcelaphus kattwinkelt (II, III, IV)
+Beatragus huntert (1, II, IV)
+Gorgon taurinus semiticus (I, II, III,
IV)
{Gazella gazella praecursor (1, II, IV)
+G. granti (I, II, TV)
+Phenacotragus recki (1V)
| Pultiphagonides africanus (1, IT)
+Pelorovis oldowayensis (II, IV)

Vaal River Younger Gravels (After Cooke and Wells, 1946; Cooke, 1949, 1963;

Wells, 1964)
CARNIVORA
cf. Crocuta crocuta

3

PROBOSCIDEA

+Gomphotherium sp.

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

+*Archidiskodon’ subplanifrons +Stylochoerus compactus

+‘Archidiskodon’ broomi +Phacochoerus aethiopicus

+*Archidiskodon’ transvaalensis Phacochoerus africanus

+‘Palaeoloxodon archidiskodontoides’ Hippopotamus cf. amphibius

Ca Terrecktian) + Stvatherium (?) cingulatum(?*)
t‘Loxodonta’ cf. atlantica + Stvatherium olduvatense haughtoni(*)
cf. Loxodonta africana cf. Alcelaphus caama

PERISSODACTYLA +‘Alcelaphus robustus’

tcf. Megalotragus eucornutus

Connochaetes cf. gnou

cf. Connochaetes sp.

cf. Damaliscus sp.

cf. Sylvicapra grimmia

cf. Aepyceros melampus

cf. Antidorcas marsupialis
tGazella wellsi

+Stylohipparion steytleri

+ Equus helmet

tEquus plicatus

+‘Equus sandwith’ (=E. plicatus ?)
Equus cf. burchelli
Equus cf. quagga
cf. Diceros bicornis

ARTIODACTYLA cf. Hippotragus sp.
+ Mesochoerus paiceae cf. Strepsiceros strepsiceros
+ Notochoerus capensis Taurotragus cf. oryx
+ Tapinochoerus modestus Syncerus cf. caffer
+* Tapinochoerus’ meadowst +*‘Homotoceras’ cf. baini

(4) See footnote to Langebaanweg fauna (p. 280).

Cornelia (Uitzoek) (After Cooke, 1963)

PERISSODACTYLA T aurotragus Oryx
+Stylohipparion steytlert +‘Homoioceras’ baini
Equus burchelli + Kobus venterae
+E. plicatus Damaliscus cf. albifrons
+ Eurygnathohippus cornelranus +Damaliscus sp.
ARTIODACTYLA Alcelaphus caama
+? A. helmet
ao P | anus ene ink : +tConnochaetes laticornutus
as ae ids | Megalotragus eucornutus
Phacochoerus africanus
i Gnhaioiens +tGazella wells
Bi ee +G. helmoedi

Giraffa camelopardalis
tLibytherium olduvaiense
Strepsiceros strepsiceros

+Gazella sp.

+ Antidorcas marsupialis

Kromdraai (After Cooke, 1963)

INSECTIVORA Crocidura cf. bicolor
+Proamblysomus antiquus Suncus cf. etruscus
+ Elephantulus langi +? Myosorex robinsont

tO i i

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 299

PRIMATES
+ Parapapio jonest
+Papio robinsont
+Paranthropus robustus

LAGOMORPHA
Lepus capensis

RODENTIA

+ Mystromys antiquus
Tatera cf. brantsi
?Desmodillus auricularis
Grammomys cf. dolichurus

+Rhabdomys cf. pumilio
Mastomys cf. natalensis
Malacothrix cf. typica
Steatomys cf. pratensis

+Palaeotomys gracilis

tCryptomys roberisi

CARNIVORA

Canis mesomelas pappos
{Canis atrox
+Canis terblanchet
Vulpes pulcher

Bolt’s Farm (After Cooke, 1963)

INSECTIVORA
+Proamblysomus antiquus
tAtelerix major

- tElephantulus langi
{Elephantulus antiquus
E. cf. brachyrhynchus

Suncus cf. etruscus

{| Myosorex robinson

CHIROPTERA

Rhinolophus cf. capensis
tcf. Myotis sp.

PRIMATES

+ ?Parapapio broom
1? P. whiter
tCercopithecoides williamsi

RODENTIA
Pedetes cf. caffer

+ Herpestes mesotes
tCrossarchus transvaalensis
tCrocuta spelaea

tC. ultra

+Hyaena bellax

+Felix crassidens

+ Therailurus piveteaur
+Panthera aff. leo

t? P. whiter

TP. shaw

+ Megantereon eurynodon

HYDRACOIDEA

+Procavia antiqua
+P. transvaalensis

PERISSODACTYLA

+Stylohipparion steytlera
? Equus burchelli

TE. plicatus

TE. helmet

ARTIODACTYLA

+Potamochoerops antiquus

+Mystromys hausleitneri
Tatera cf. brantst
+Dasymys bolti
+Rhabdomys cf. pumilio
+ Thallomys debruyna
Leggada cf. minutoides
L. cf. major
Malacothrix cf. typica
+Palaeotomys gracilis
Hystrix africae-australis
tCryptomys robertsi

CARNIVORA

Canis mesomelas

C’. mesomelas pappos

Aonyx cf. capensis

Suricata suricatta
+Crossarchus transvaalensis
+Hyaena bellax
+Leptailurus spelaeus

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

+ Therailurus barlowi

+Panthera aff. leo

+ Machairodus transvaalensis
PROBOSCIDEA

+Loxodonta atlantica

HYDRACOIDEA

Procavia capensis

PERISSODACTYLA
+Stylohipparion steytlert
Equus burchellii
TE. plicatus
ARTIODACTYLA

+ Tapinochoerus meadowsi

Makapansgat (After Cooke, 1963)

INSECTIVORA

tChrysotricha hamilton

| Elephantulus lange
Suncus cf. etruscus

| Myosorex robinsont

PRIMATES

+Simopithecus darti
+Parapapio jonesi

+P. broomi

+P. whiter

+Papio robinsoni

+ Australopithecus africanus
tCercopithecoides williamsi

LAGOMORPHA
Pronolagus randensis

RODENTIA

+ Mystromys hausleitnert

tM. dart
? Tatera cf. brantst
Grammomys cf. dolichurus
Pelomys cf. fallax

+tRhabdomys cf. pumilio
Aethomys cf. namaquensis
Mastomys cf. natalensis
Leggada cf. minutoides

+ Potamochoeroides shawt

+P. antiquus
Tragelaphus scriptus
Taurotragus oryx
Syncerus caffer
Damaliscus cf. pyrgatus
? D. cf. lunatus

+ Alcelaphus robustus

tA. helmer
Connochaetes taurinus

+ Makapania broom

+? Raphicerus campestris

{Gazella wellsi

+ ? Phenacotragus vanhoepent

{ ?Antidorcas marsupialis

Dendromus cf. mesomelas
+ ?Malacothrix makapani

Steatomys cf. pratensis
+Palaeotomys gracilis
+Hystrix major

Hi. africae-australis
+ Xenohystrix crassidens
+Gypsorhychus makapant
tCryptomys robertst

CARNIVORA

? Canis mesomelas pappos
tCynictis penicillata brachyodon
tCrocuta cf. brevirostris
+ Hyaena makapani
+ Therailurus barlowi
+ Megantereon sp. nov.

HYRACOIDEA

+Procavia antiqua
+P. transvaalensis
+Procavia sp.

PERISSODACTYLA

+? Stylohipparion steytler

+Equus helmet

+ Metaschizotherium (?) transvaalensis
Ceratothertum stmum
Diceros bicornis

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 301

ARTIODACTYLA C. caerulus

Hippopotamus amphibius +Redunca darts

+ Notochoerus euilus Redunca arundinum

+Potamochoeroides hypsodon R. fulvorufula

+P. shawt Oryx gazella
Giraffa camelopardalis tAlcelaphus robustus

+Libytherium cf. olduvaiense +A. helmet
Strepsiceros strepsiceros Connochaetes taurinus
Tragelaphus angasi + Oreotragus major
Taurotragus oryx +Makapania broomi
Syncerus caffer Aepyceros melampus

+S. cf. makapani +tGazella gracilior

tCephalophus pricer +Phenacotragus vanhoepent

‘ELANDSFONTEIN, HOPEFIELD (SOUTH AFRICA)
(Modified from Singer, 1957)

PHOLIDOTA +Panthera leo spelaea
Manis sp. + Megantereon gracile
PRIMATES PROBOSCIDEA
{Simopithecus oswaldi hopefieldensis tLoxodonta (Palaeoloxodon) cf. anti-
+Homo sapiens rhodesiensis (‘Saldanha quus recki
Man’) +? Archidiskodon sp.
LAGOMORPHA PERISSODACTYLA
Lepus sp. (cf. capensis) +Equus (Hippotigris) plicatus
+E. helmer
oot bil +E. cf. sandwithi
Bathyergus Sp. (cf. suillus) Ceratotherium simum
Georychus sp. (cf. capensis) Diceros bicornis
HAystrix sp. (cf. africae-australis) ee
Otomys sp. (cf. saundersiae)
Parotomys sp. (cf. brantsz) (eas ee
tM. paiceae
CARNIVORA + Tapinochoerus meadowst
Canis mesomelas Hippopotamus amphibius
Canis adustus + Sivatherium olduvaiense(*)
{Lycaon pictus magnus +Giraffa cf. gracilis
Mellivora capensis +Homotoceras sp.
Herpestes sp. (cf. ichneumon) Taurotragus oryx
Herpestes eogale Redunca arundinum
FHyaena brunnea Raphicerus campestris
tCrocuta spelaea Antidorcas marsupialis
Lynx caracal + Antidorcas sp.

Leptailurus serval - Tragelaphus(?) sp.

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

cf. Hippotragus niger Connochaetes sp.
tcf. H. leucophaeus +Lunatoceras sp.
+ Hippotragus sp. _ tPelea sp.
+Damaliscus cf. dorcas tcf. Gazella wellsi
+Damaliscus sp. tGazella sp. nov.

(1) See footnote to Langebaanweg fauna (p. 280).
(7) In the other faunal lists Tragelaphus has not been substituted for the earlier labelling
Strepsiceros.

Refer now to table 1 on pages 303 to 313.

COMMENTARY ON THE PUBLISHED FAUNAL ASSOCIATIONS AT
HIPPARIONID SITES IN AFRICA
MIOCENE

Oued el Hammam and Marceau

Arambourg (1959) has clearly demonstrated that these two sites contain
fundamentally the same fauna: both include Hipparion africanum, Samotherium
sp. and Hyaena algeriensis, and cannot but be contemporaneous, as is confirmed
by their stratigraphy.

It seems clear that these assemblages correspond to a very special and,
so far, little-known stage of development of the fauna of Africa, posterior to
the Burdigalian and anterior to the ‘classical Pontian’. The fauna from these
sites are very different from the well-represented assemblage of the Lower
Miocene in East Africa (Losodok, west of Lake Rudolph; Rusinga and Moboko
Islands in the Kavirondo Gulf of Lake Victoria) and South West Africa. The
typical Burdigalian assemblages contain, inter alia, Mastodon cf. longirostris,
Deinotherium hobleyi, Aceratherium, Teleoceras, and Anthracothertidae, Propalaeo-
choerus, Bunolistriodon, Dorcatherium, Creodonta, none of which are found at
Oued el Hammam, nor are the anthropomorpha which constitute the typical
African ‘touch’ of this Burdigalian fauna. The only similarity between the sites
and a Burdigalian fauna is the presence at Oued el Hammam of an orycterope,
namely, Orycteropus mauritanicus which is comparable with Myorycteropus McInnes
of East Africa. The separation between Oued el Hammam and the Lower
Miocene fauna of East and South West Africa is emphasized by the presence
(at Oued el Hammam) of Giraffidae (Samotherium and Palaeotragus), the
development of Bovidae (Damalavus, Gazella, Tragocerus, Cephalophus) and the
appearance of Hyaena and especially of the equids with Hipparion africanum.

On the other hand, the assemblages from the two North African localities
are at first sight characteristic of the so-called ‘classical Pontian’ of Eurasia
(Eppelsheim, Pikermi, Mont Luberon). However, on detailed study and
comparison of the several groups involved, Arambourg has shown that Eurasian
‘Pontain’ assemblages and the material from Oued el Hammam may belong
to the same ensemble, but they are not identical: specific and generic differences
exist and there is not even a single species in common (table 2).

393

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314 ANNALS OF THE SOUTH AFRICAN MUSEUM

These differences indicate that the fauna of Oued el Hammam is more
primitive than that of the Eurasian ‘Pontian’, thereby suggesting an ancient
biogeographical autonomy of Africa. The presence of a fauna more primitive
than the ‘classical Pontian’ does not necessarily mean that it antedates the

Pontian. However, as is indicated in the section on the geological aspects of

the sites (vide supra p. 283), the Oued el Hammam deposit lies under lacustrine
or marine deposits. These contain clearly recognized Upper Miocene fauna of
the Sarmatian or of the Tortono-Sahelian age, which were formed by the
marine transgression at the end of the Miocene. As indicated by Arambourg,
there is no possible correlation between Oued el Hammam and the ‘classical
Pontian’ which is formed posteriorly to the Sarmatian.

PLIOCENE

The Hipparion-bearing sites in North Africa and in the Nile Valley,
which are usually considered to be Pliocene, cannot be satisfactorily dated on
the basis of the fauna alone. The faunal assemblages at Tozeur, Ain el Bey
and Mascara are rather poor. Their antiquity is suggested by the presence of
Merycopotamus, Helladotherium and Rhinoceros pachygnathus. The Pliocene nature
of these sites can only be determined from their geology.

At Wadi Natrun, none of the rare fossils recovered is characteristic of a
particular period. Determination of stratigraphical relationships is necessary
for the assessment of the Pliocene date of the fossiliferous beds.

PLEISTOCENE

The simple comparison of lists of fauna from numerous sites often results
in misleading or incorrect conclusions for the following reasons:

1. The assemblages may represent a sampling of fauna from different
biotopes, and furthermore, the sampling (i.e. collecting) may have been made
by means of different methods and for differing requirements. For example,
in the case of small rodents, they may be an incidental part of a general collec-
tion recovered during a thorough investigation, or they may have been sought
for exclusively from, say, breccia by a specialist. However, they may have
been overlooked in the field when the investigator was merely collecting the
larger bones. Thus they would be absent from the collected assemblage, but
this would not reflect the fact that they may have been present in situ. Thus
someone studying or re-studying a particular collection, and being unaware
of the conditions of recovery of the material, may accurately record frequencies
but these may yet be misleading.

2. Different palaeontologists may use criteria and terminology at variance
with others, often because of a familiarity with material from particular areas
or countries. In addition the fragmentary nature of certain specimens may be
responsible for differing interpretations.

3. Just as. different biotopes may express slight differences in similar
forms, so climatic differences may influence similar forms even in adjacent

hee Beg geist AF able 2

Swear cS

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 315

regions. These differences may be interpreted on a specific rather than on a
subspecific level, thus influencing statistical analyses.

Consequently, a practical method of comparing sites within a geographical
region is by studying the results obtained by either a single palaeontologist
or by a particular institute. This requirement is partially met for the ‘Villa-
franchian’ sites of North Africa, for the australopithecine breccias in South
Africa, for Olduvai Gorge, and possibly for Laetolil and Rawi in East Africa.
It is to be remembered that the success of the investigation within each of these
three geographical units varies.

The North African ‘Villafranchian’ fauna is characterized by the associa-
tion of the latest mastodonts with primitive elephants, of the Sivatheriinae
with Giraffa, and of the hipparionids with the modern Equidae.

However, there is sufficient evidence available to suggest that such associa-
tions, found in several places in Africa, are not necessarily contemporaneous.
Unfortunately, the term ‘Villafranchian fauna’ is loosely used with a different
interpretation in different areas, but it may only be applied, sensu stricto, in
a chronological sense to the Mediterranean basin.

Arambourg (1947) proposed a relationship between the fossil-bearing
sites of Africa according to the presence or absence of archaic elements, such
as Archidiskodon planifrons, Anancus, Stegodon and Chalicotheridae. During the
Lower Pleistocene some of these forms seem to have disappeared, being replaced
by A. recki, while other primitive groups with tertiary affinities like Stylohipparion,
Deinotherium and Libytherium persisted.

By means of the Proboscidea sequence, a subdivision of the ‘Villafranchian’
can be formulated:

A. Lower ‘Villafranchian’

Characterized by the PRESENCE of Anancus, Stegodon, Stegolophodon (?),
and the appearance (in a stratified deposit) of Archidiskodon (or Elephas sub-
planifrons, planifrons, africanavus, and exoptatus): Kanam, Kaiso, Garet Ichkeul.

B. Middle ‘Villafranchian’

Characterized by the extinction of Stegodon, the persistence of Anancus,
Mastodon, Archidiskodon (or Elephas) planifrons or subplanifrons and exoptatus and
the first appearance of Palaeoloxodon recki (or E. meridionalis): Koro ‘Toro,
Laetolil, Ain Hanech, Olduvai I.

C. Upper ‘Villafranchian’

Characterized by the presence of Anancus, Stegodon, Mastodon, and the
continued presence of A. exoptatus, E. africanavus and even Deinotherium: Omo.
The post-‘Villafranchian’ horizon commences when Anancus, Stegodon, and
Mastodon as well as primitive forms of Archidiskodon (exoptatus) and Elephas
(planifrons, africanavus) become totally extinct, coincidental with the appearance
of Loxodonta africana and atlantica as well as the further development of P. reckz.

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

North Africa

In a comparison of Garet Ichkeul with Ain Hanech, the fauna of which
are substantially the same, Arambourg (1949) concluded that Garet Ichkeul
was the more ancient (Lower ‘Villafranchian’) because of the more abundant
E. planifrons and Stylohipparion at the former site, while the latter had remains of —
P. recki.

There is good evidence that there is close identity between Ain Hanech
(and Bel Hacel) on the one hand, and Garet Ichkeul, Ain Boucherit (i.e. Beni
Foudda of Pomel) and Oued el Akrech (Arambourg and Choubert, 1957),
on the other. On the basis of the fauna alone, it does not seem possible to
compare these in any great detail with the other North African Pleistocene
Hipparion-bearing sites.

Total number of — Extinct Extant Extinct Extant
identified species species species Total Extinct
% %o
SOUTH AFRICA
Panne ec. si ais 34 25 9 73°5 36-0
Sterkfontein va ey a7 26 II 70°4 42°3
Makapansgat .. i 70 43 27 61°4 62°7
Swartkrans - He 39 32 7 82 21°9g
Kromdraai se ee 41 29 12 70"7 4AI*4
Bolt’s Farm ce Le 56 35 21 62-6 60-0
Vaal River ae, ae 35 20 15 57 751
Cornelia .. ee sil 23 17 6 74 35°3
Hopefield a AN 45 23 22 51°2 95°6
Florisbad ate ai 32 15 iy 46-9 113°4
Vlakkraal. . oe By 18 9 9 50 100
Cave of Hearths 5 45 10 35 22°2 350°0
Wonderwerk _... ye 25 8 17 32 213°0
RHODESIA
Chelmer .. me aM II 6 5 — —
Broken Hill KE it 31 yi 24 22°6 34.2
Mumbwa ue ais 18 3 15 16°6 500
EAST AFRICA
Kaiso ve ah is 10 8 2 80 25
Kanam .. is ae 16 12 4 75°0 oa
Omo is ae ah 21 15 6 71*4 40
Laetolil .. Aye aye 33 20 13 60-6 65-0
Olduvai I Be wis 34. 28 6 82°2 21°4
Olduvai II ue ae 49 38 II Vir nae § 29°0
Olduvai III a de 23 18 5 78-4 27°8
Olduvai IV a ve 39 30 9 76-9 30°0
Rawi a ie Sie 7 4 3 — —
Kanjera a Bi 17 14 3 82°3 Q21°4
Olorgesailie Ay he 9 Gi 2 — =
Eyasi oe by M. 24 9 15 37°5 166-0
‘Gamblian’ a ate 12 7) 5 58°4 7I°4

TABLE 3. The relationships between identified extinct and extant fauna from African Pleistocene
sites.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 317

East and Central Africa

The OLDUVAI GORGE has been extensively studied. From the four
Beds, 55 species have been identified (table 1), Bed II and IV containing
respectively 49 and 39 of them (table 3). However, recent discoveries and
more detailed analyses will considerably alter these figures. Dietrich (1942),
Arambourg (1947), Hopwood (1951) stressed the general Middle Pleistocene
character of the Olduvai faunal assemblage. It has been pointed out that no
available faunal evidence suggests that the lower part of Olduvai is of Lower
Pleistocene age. This view will probably be considerably modified when
descriptions and analyses of recent discoveries are published shortly. In this
paper, only the available published data are assessed. Leakey (1963) emphasizes
the point that “Bed I covers a very long period of time with a gradual change
from a very wet climate at the beginning to savanna conditions and then
subdesert ones’. It is no longer possible to speak simply of ‘the fauna of Bed
I’. He also points out that the fauna from Bed I, although ‘Villafranchian’
and older than Omo, does not conform with the Lower ‘Villafanchian’ fauna
as represented in East Africa at Kanam East and West. Furthermore, it is no
longer certain whether Bed I contains Hippopotamus gorgops, Taurotragus oryx,
Tragelaphus strepsiceros, and even Palaeoloxodon recki which is super-abundant in

Bed II and Bed IV.

Olduvai I i a. DMA Cave of Hearths .. she) 1 E92
Swartkrans * am aE S Broken Hill. . ne ce 2220
Olduvai III ae nee) 25°o Wonderwerk re i) (B20
Olduvai II .. a Ria. 2ato Eyasi ae 5 uh da ge
Olduvai IV eh i 30°0 Florisbad_ .. He Be 242)
Cornelia... rk ml aS Hopefield .. ge YE vi ES
Taung ag es we 2 36" 0 Vaal River a alent ap@
Omo Bi we ee AO) Lattolit ".. ns 3 O06
Kromdraai ie wet) Ard Makapansgat 2 eh Omer
Sterkfontein a” wale Caen Bolt’s Farm. . oe teh, 2 ™6
Bolt’s Farm. . Se Hat KOOSO Sterkfontein on Yin OT
Makapansgat i et MO 2 7 Kromdraai Ae ee oy]
Paetohil | .. “4 ia: G50 Omo Bee ua a ig TAN
Vaal River Bes ne 75:1 Taung aS ae ee 7/939 Ie
Hopefield .. ae aie ens 0 Cornelia... wi eM Mae 7 ao)
Florisbad .. A is RLS A Olduvai IV ee ae 76°9
Eyasi 8 ate .. 166-0 Olduvai II ae ms Tigied:
Wonderwerk ie ae) 2130 Olduvai III a ms 78° 4
Broken Hill. . ne aie AcBie-'O Swartkrans oe bel NOHO
Cave of Hearths .. ea SOO Olduvai I S re oz

TABLE 4. List of sites in the order

of increasing proportions of extant

species. Ratios (extant/extinct) are

determined only for those sites where

more than 20 species have been
identified.

TABLE 5. List of sites in the order of

increasing proportions of extinct

species. Ratios (extinct/total) are

determined only for those sites

where more than 20 species have
been identified.

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 319

OMO has classically been equated with Olduvai I and II but recently a
proposal has been made that it is more similar to the base of Olduvai II (tables
4, 5). The extinct fauna common to Omo and Beds I and II are 30-3% and
20:4% of the total respectively (table 6). There is no substantial difference
between the faunal assemblages except for the presence of Anancus and hominids
at Olduvai. The absence of <injanthropus and ‘Homo habilis’ at Omo, does not
per se imply that Omo is younger than Bed I, because the Olduvai Lake shores,
permitting the possible development of living sites, favoured a concentration
of hominids.

The SERENGETI (Laetolil) fauna is closely related to both Olduvai and
Omo. The extinct species at Serengeti in common with Olduvai I are 23:1%
with Olduvai II are 23-4%; and with Omo are 20:6% (table 6). The essential
difference seems to be the presence at Laetolil of Anancus and Archidiskodon
subplanifrons, both of which are absent at Omo, and also the presence of Meta-
schizotherium hennigi (which is also present at Kaiso). Leakey (1958) believes
that the differences between the Laetolil and Bed I faunas do not imply a
temporal separation but rather illustrate a difference in ecological conditions
existing contemporaneously. He indicates that it has not been proved that
Laetolil is older than Olduvai I. Furthermore, it should be pointed out that
Laetolil probably contains a mixed assemblage of different faunal stages, as
was already suggested by Dietrich.

KAISO and KANAM are badly represented from a faunal viewpoint:
10 (8 of which are extinct) and 16 (12 of which are extinct) species respectively
have been identified. In spite of the fact that information about their recovery
is unsatisfactory, there is no doubt that a real archaic character is attached
to these sites, from which Chalicotherium, Stegodon, Anancus, Stegolophodon and
Hippopotamus imaguncula have been recovered. These forms are considered by
most authorities to represent the most ancient East African Pleistocene fauna
known at present.

KORO TORO consists of five ‘apparently equally old’ deposits which
contain about 30 identified species of mammals. Abadie, Barbeau and Coppens
(1959) and Coppens (1960), studying the fauna of the lowest level, conclude
that it belongs to the Lower ‘Villafranchian’ because of the contemporaneity of
Mastodon, Stegodon and Elephas. However, the rarity of both stegodont and
mastodont (more typical of the base of the Lower Villafranchian and the
Pliocene) and of P. recki (more typical of the Kamasian), and at the same time
the abundance of E. africanavus suggest that we are dealing with an intermediate
stage between the very base (e.g. as at Kaiso where africanavus is present without
reckt) and the top (e.g. as at Omo where recki and africanavus are abundant
but stegodont and mastodont are absent).

Kent (1942) and Arambourg (1943, 1947) emphasized the great similarity
between the Olduvai and the Omo faunal assemblages, and indicated that
they considered the Omo fauna to be slightly older. Arambourg suggested
that the various East African deposits may be considered to correspond to

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

different phases of the transition between the Upper Pliocene and the Lower
Pleistocene: the Serengeti, Kaiso and Kanam tuffs being the most ancient
and being the equivalent of the ‘Villafranchian’ horizons in North Africa
and Europe, while Omo and Olduvai I should be considered as being younger
and corresponding to the first true Pleistocene deposits. This view has been
partially modified because of the recent discoveries at Olduvai. Leakey (1963)
describes a major climatic, faunal and geological ‘break’ near the base of
Bed II, the lowest part of which (overlying the ‘marker bed’ at the top of Bed
I) he considers to be of Upper ‘Villafranchian’ age and comparing very
closely with that of Omo. According to Howell (1959), ‘the faunas (from the
Villafranchian sites of Central and East Africa) differ somewhat in composi-
tion, that from Laetolil beds being probably the youngest, overlapping basal
Olduvai and that from Kaiso being perhaps the oldest. The Omo fauna overlaps
both Laetolil and Kaiso and that from Kanam is probably broadly equivalent.’

South Africa

The fauna of the TRANSVAAL cave breccias (Taung, Sterkfontein,
Makapansgat, Kromdraai and Swartkrans) has been extensively studied.
The fauna of these sites show a very high degree of similarity: 11-°8—32-6%
of the total number of extinct species are common to the different sites (table 6).
A similar range (15°:4—35°6%) has been calculated for the BOLT’S FARM
faunal assemblage found in the vicinity of Sterkfontein. The six sites show an
extinct/total ratio of species of 61-74% (table 4) and an extant/extinct ratio
lower than 63% (table 5), which indicates a considerable antiquity. The
conditions of accumulation at and the geology of these sites are also similar.
On the basis of the fauna (Ewer, 1957), as well as of mineralogical and climato-
logical studies of the breccia (Brain, 1958; Robinson, 1961), a relative age
sequence has been derived, namely, Sterkfontem—Taung, Makapansgat,
Swartkrans and Kromdraai, extending from the Lower to the early Middle
Pleistocene.

HOPEFIELD and FLORISBAD faunas seem to have rather similar
frequencies: not only are 15:1% of the total of extinct species common to
both sites, but their close faunal relationship is expressed by a similar ratio of
extinct/total species (51% and 47% respectively). However, the extant/extinct
ratio (viz. 96% and 113% respectively) suggests a greater antiquity for Hope-
field. Furthermore, there are considerable differences in the types of hominids
and artefacts recovered from these two sites. Although Hipparion has not been
recovered from these two sites, they are included as important Middle-Upper
Pleistocene sites.

From purely a consideration of the fauna, the YOUNGER VAAL RIVER
GRAVELS and CORNELIA could fit satisfactorily in the chronological
sequence between the Transvaal cave breccias and Hopefield—Florisbad with

the following intermediate ratios (tables 4, 5): 57-74% of extinct species;

35-75% of extant/extinct species; and 27-6% of all the extinct species being

-
¢
#

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 321

common to both sites (table 6). According to Cooke and Wells (1946) ‘while
it is possible that some of the living species may post-date the Younger Gravels
phase of deposition, it appears most probable that the bulk of their material
forms a fairly coherent whole, representative of the fauna of the later part of
the first wet phase’. The fauna from the Younger Vaal River Gravels and
Cornelia deposits could be broadly considered as a Middle Pleistocene fauna,
especially in the light of the recent reinvestigation by Cooke and Wells who
indicated that the Cornelia fauna is comparable with the Younger Vaal
River Gravel material (unpublished; quoted by Cooke, 1963, page 96; see
also Wells, 1964).

Although the Lower Pleistocene sites of East and of South Africa have
ratios of extinct species higher than 60% and an extant/extinct ratio lower
than 70%, direct comparisons between East and South Africa have severe
limitations. From an analysis of the fauna (table 1), based on the lists drawn
up by Cooke (1963), it appears that not a single species as recorded is common
to both East and South African Lower Pleistocene sites. We believe that this
lack of relationship is really less marked than indicated because different
names have been given to the same species in East and in South Africa. This
is mainly because the taxonomy in the two regions has been developed by
independent investigations, e.g. the species of Simopithecus (Freedman, 1957;
Leakey and Whitworth, 1958; Singer, 1962). To some extent this shows the
poor state of our knowledge of the African fauna as a whole. Furthermore,
Pickering (1960) warned that a critical approach will always be required
when comparing a plains fauna, such as that found at Olduvai, with an assem-
blage obtained from a cave deposit which is usually the case in the Lower and
early Middle Pleistocene of South Africa.

There is some affinity between the Younger Vaal River Gravels, Cornelia
and Hopefield assemblages on the one hand, and the East African early Middle

Pleistocene collections on the other, ranging from 2-9-11:8% of the total

number of extinct species for any two particular sites among these groups

(table 6).

CONCLUSIONS BASED UPON THE FAUNAL AND GEOLOGICAL EVIDENCE IN THE
LITERATURE

The large amount of data discussed in previous chapters may be utilized
for the construction of a tentative illustration of the relationships between the
sites at which Hipparion has been discovered in Africa (fig. 8). By and large this
supports the views expressed by Cooke (1963) and Bishop (1963).

CHRONOLOGICAL RANGE OF Hipparion

Apart from the actual dating of the geological deposits from which Hipparion
has been recovered in Africa, there is the general question of its first appearance
and its ultimate disappearance on the continent.

It seems that Africa was not the area of origin of hipparionids, as the

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

| NORTH AFRICA | AFRICA EGYPT] ETHIOPIA | CENTRAL Waters AFRICA SOUTH AFRICA
AFRICA & RHODESIA

arene Viakkraal
Wonderwerk
Eyasi Cove of Hearths
Chelmer Florisbad
Vaal

Olorgesailie B. Hill Hopefield
Kanjera
Ternifine
Kromdraai

Pei eA Swartkrans
Bel Hacel Sterkfontein ext.
Makapansgat
Koro Toro “e Sterkfontein
i it} Garet Tee
Fouvarat} O.e. Akrech |Ichkeul : 1?
Lergeeratep

Utique
Mascara
St. Donat
Ain el Bey |Tozeur
A.e.Hadj Baba Wadi
Camp Natrun
Berteaux
bist eS

O.e.Hammam
Marceau
Fic. 8. Tentative correlation between Hipparion sites in Africa. A few other sites have been
incorporated for reference. B. Hill = Broken Hill; O.e. = Oued el; A.e. = Ain el; Gamblian =
Gamble’s Cave; Bolt’s = Bolt’s Farm.

Cornelia

PLEISTOCENE

—---0o-%0r-

PLIOCENE

evidence points to probable migrations from Eurasia. Nevertheless, it is some-
times claimed that Africa contains the evidence of the early, if not the earliest,
existence of Hipparion in the Old World.

At the other end of the chronological range, the hipparionids survived
much longer in Africa than in any other area known at present. With very
few exceptions, most of the sites where African Hipparion have been located
are of Pleistocene origin, i.e. a period when they no longer existed in Eurasia
and in America.

Consequently it is considered necessary to compare the geological range
of occurrence inside with that outside Africa.

UPPER LIMIT

According to available information, the upper limit of Hipparion outside
Africa seems to correspond roughly to the Upper Pliocene. It is generally
accepted that the Villafranchian is defined by the appearance of some modern
genera, among which is Equus. However, there is no reason to deny, a priori,
a coexistence of Hipparion with modern Equus which developed on a parallel
line from Pliohippus. Such co-existence has actually been demonstrated for
Africa. Outside Africa, Equus not only heralds the Pleistocene, but it he
replaces the more primitive Hipparion in its habitats.

j

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 323

AMERICA| CHINA INDIA MIDDLE- SE: CENTRAL & AFRICA
EAST EUROPE W. EUROPE

Azov Sea Val d’Arno, Seneze
(Khopry)

Kanjera

Cornelia

a Vaal ie)
w |
e) d
ie) Koro Toro U
e v
” a
= Ain Hanech Omo i
A

a Laetolil

Ain
Fouarat,Garet Ichkeul, Boucherit
Kanam, Kaiso

Blanco Pinjor
w
Z Patrot ;
rs} Roussillon Wadi Natrun
© | Hemphill Godallo, Berechti A.e. Hadj Baba
= Malouchteni Mascara

Maragha
Polgardi, Vellés
Mont Lubéron

Dhok Patan Pikermi

Samos

Upper-
Clarendon

Taraclia

Valles - Pénédés
Rhéne- Valley
Teruel

Sebastopol
Odessa

Pao-Te (Red Clays)

Clarendon

Oued el Harnmam
Marceau

Eppelsheim

UPPER-MIOCENE

Fic. 9. A diagrammatic representation of the probable chronological range of Hipparion in
different continents.

The youngest formation bearing Hipparion remains in America is the
Blanco formation (Upper Pliocene), where its presence is even questioned by
some palaeontologists. Even if accepted, remains are very scarce.

In China, the latest occurrence is in the Nihowan deposits of the Sang
Kan Ho Valley, east of Pekin (fig. 9); in India, in the Tatrot and the Pinjor
zones of the Siwaliks, while the records are still very insufficient for Mongolia.
In the U.S.S.R. (including its Asian portion), the most recent occurrence is to
be found in the Azov Sea shore deposit at Khopry; in South and Central
Europe at Berechti and Malouchteni in Rumania, and Gédollé in Hungary.
In western Europe, Hipparion has not been found in the Villafranchian deposits
of the Val d’Arno and Senéze; and it is rather uncommon in the Roussillon,
although it is still found in Perrier where it is rather exceptionally associated
with Equus.

Without going into details which are irrelevant in the framework of this
paper, it may be stated that outside Africa, Hipparion does not seem to have
extended into the Pleistocene. In many instances, it was already becoming
rather scarce during the Middle Pliocene and definitely more rare in the
Upper Pliocene.

LOWER LIMIT

The lower limit of Hipparion, or the time of its first appearance, presents a
more difficult problem, as it is not at all easy to correlate the chronological

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

interpretations of all the localities. In addition, a different faunal basis (marine
or continental) is applied to various areas, and a direct comparison of the
fossil associations is not always possible. Furthermore, some geological terms,
like ‘Pontian’, have been used in a different context and with a different
meaning by various authors, e.g. as a facies, or a stratigraphical or a faunal
horizon. Consequently it is especially difficult to appreciate the meaning of a
particular statement, factual though it may be, without danger of misinterpreta-
tion. Therefore, the soundest approach may be, first, to locate the earliest
occurrence of Hipparion, within a region, and then, to correlate as far as possible
the interregional data.

——— EEE

In America, Hipparion is not found in the Barstov formation, appearing —
first in the Clarendon formation (fig. 9). It is also found right at the base of —

the Mint Canyon formation, and in the Hemphill formation. The upper
Clarendon and the Hemphill formations are unanimously considered as ‘Lower
Pliocene’ and correlated with the European ‘Pontian’. But for more than thirty
years, the Mint Canyon has been a major topic of discussion and argumenta-
tion. While most scholars follow Stirton’s opinion (1939) and locate the conti-
nental basal horizon of Mint Canyon in the Pliocene, Maxson (1930), among
others, basing his opinion on the debatable malacological fauna of the overlying
marine Cierbo beds, considers it to be middle Upper Miocene age.

In China, Hipparion appear in the red clays of Chan-si, Chen-si and
Kansou. These Pao-Te formations have not been properly subdivided. At
different times, investigators have pushed them back to the Upper Miocene
(Teilhard and Young, 1931), or restricted them to the Lower Pliocene (Teilhard
and Leroy, 1942).

In India, the first Hipparion are recovered in the Chinji zone of the Siwaliks,
which is referred either to the Middle (Pilgrim, 1938) or to the Upper (Lewis,
1937) Miocene, or even to the Lower Pliocene (Colbert, 1935).

In the U.S.S.R., the earliest occurrences are those of Moldavia and Odessa
and the Sebastopol fauna (Borissiak, 1914). Russian geologists refer the former
deposit to the Middle Sarmatian, and the latter two to the Upper Sarmatian,
i.e. the Upper Miocene. Similar information was recently obtained from the
Upper Sarmatian in the Istanbul vicinity (Chaput and Gillet, 1938; Yalcinlar,
1952).

In central and southern Europe, because of the exceptional associations of
H. primigenium with the typical Miocene Anchitherium, Eppelsheim might be
considered as one of the first known areas of occurrence of the genus. But in the
Rhone Valley (Denizot, 1939), at Vallés-Pénédés in Catalogne (Villalta and
Crusafont-Pairo, 1946, 1947, 1948) and in the Teruel Basin (Sondaar, 1961),
abundant remains of Hipparion have been recovered in deposits which are
dated as Tortonian. Maragha, Pikermi, Samos, Polgardi, Baltavar, Vélés
and Mont Luberon, referred to as typical ‘Pontian’ sites, are probably somewhat
younger, and should be placed in the Lower Pliocene.

In spite of many unsolved problems in correlating these sites, it seems

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 325

difficult to deny that there is in North America, in Asia and in Europe constant
and repeated indications of the appearance of Hipparion in the Upper Miocene

(fig. 9).

GENERAL DESCRIPTION OF Hipparion TEETH
UPPER DENTITION

The pattern of horse teeth has been described many times and detailed
structures have been extensively discussed in previous publications. The
distinctive characters of horse teeth that have been described are based on
the original description proposed by Osborn (1907). In 1918, he successfully
applied this description in his study of the North American Equidae. Subsequent

UPPER TOOTH

4 ‘

/ \
(P- prf}

Fic. 10. Features of the occlusal surface of an upper molar tooth of hipparionids. Abbreviations:

B. prf—bouclé prefossette; Eclo—ectoloph; Hy—hypocone; Hyg—hypoglyph or hypoconal

groove; Hys—hypostyle; Me—metacone; Mel—metaconule; Melo—metaloph; Mes—meso-

style; Mets—metastyle; Pa—paracone; Pas—parastyle; p. c—pli caballin; p. hys—pli hypo-

style; p. plo—pli protoloph; p. prf—pli prefossette; p. prl—pli protoconule; p. ptf—pli post-

possette; Pr—protocone; Prf—prefossette; Prg—preprotoconal groove; Prl—protoconule;
Prlo—protoloph; Ptf—postfossette.

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

alterations and additions have been proposed by, among others, Stirton (1941),
Arambourg (1947, 1959), Gromova (1952), Hopwood (1937) and Cooke

(1950). It is not proposed to make any further modifications here, but it is
necessary to comment on the dental nomenclature, without any phyletic —

implications, so as to outline the basis adopted in this monograph.

Typically three crests are recognized. A mesiodistal ectoloph, joining
paracone and metacone on the buccal surface, an anterior protoloph and a
posterior metaloph. The latter two are more or less transverse in the primitive
condition but in advanced Equids they are half-moon-shaped. Being lopho-
dontic specializations of the protoconule and the metaconule, they build the
protoselene and the metaselene, and they meet the ectoloph at parastyle and
mesostyle, respectively (fig. 10).

Lingually deflected from the main selenic lophs, but more or less attached
to them there is a protocone and a hypocone. Their rather deep bordering
grooves, filled with cement, tend to isolate them from the crests. These grooves
are the pre- and post-protoconal grooves (valleys, sinuses) which lie anteriorly
and posteriorly to the protocone, respectively, and the hypoconal groove
(sinus) or hypoglyph which is related to the hypocone (the posterior being
usually ney well marked, and the ony one noticeable). Sometimes the protocone
shows a ‘spur’: the “protoconal spur’.

When the pre- and post-protoconal grooves are maximally deepened,
they become confluent in a medivallum or internal depression. Thereby they
produce complete isolation of the protocone. This is one of the major and
characteristic features of the upper molars of Hipparion.

Elevations of the cingulum have also been described: parastyle, mesostyle
and metastyle along the ectoloph; hypostyle on the posterior surface which
ultimately develops a cusp, seemingly independent of the cingulum.

The protoloph and the metaloph crests enclose, more or less completely,
the pre- and postfossettes. The more constant and deeper plications in the
enamel wall of the fossettes (‘marks’) and also isolated ‘horns’ have been
assigned special names: anteriorly, the pli protoloph and the pli postfossette,

:

respectively; posteriorly in the anterior mark, the pli prefossette and the pli

protoconule which isolates a ‘boucle préfossette’ or ‘prefossette loop’ (Stirton,
1955), while posteriorly in the posterior mark is the pli hypostyle (fig. 10).

Furthermore, a pli caballin appears constantly in the post-protoconal
groove or in the internal depression. It is a lingual extension from the outer
border of the selene, and eventually, after wear, it displays a two- or threefold
division.

LOWER DENTITION

A distinction is commonly drawn between a mesial trigonid, with buccal
protoconid and lingual metaconid, and a distal talonid, with hypoconid,
entoconid and hypoconulid, separated on the buccal aspect at the level of a
fairly constant external depression (fig. 11).

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 327

LOWER TOOTH

‘Mefd’
(Aid)?
Fic. 11. Features of the occlusal surface of a lower molar tooth of hipparionids. Abbreviations:
Ecsd—ectostylid; End—entoconid; Ensd—entostylid; Enfd (Pid)—entoflexid (Post. int.
depression); Exd—external depression; Hyd—hypoconid; Hyld—hypoconulid; Med—meta-

conid; Mefd (Aid) —metaflexid (Anter. int. depression); Mesd—mesostylid; Pasd—parastylid ;
Prd—protoconid; Prsd—protostylid; Ptysd—ptychostylid.

The stylids on the buccal side are protostylid, an inconstant ectostylid
and a ptychostylid, while on the lingual aspect there are the metastylid and
the entostylid.

On unworn teeth, a crest, the paralophid, can be distinguished extending
from the protoconid to the protostylid in the anterolingual corner.

On the lingual aspect there are two prominent internal depressions or
invaginations, the metaflexid anteriorly and the entoflexid posteriorly.

The metastylid and the metaconid are two rounded formations joined by
a narrow isthmus, giving the appearance of a bow tie and forming the so-called

‘double knot’.

SUMMARY OF THE CHARACTERISTICS OF HIPPARION TEETH

Based on Gromova’s detailed description (1952, pp. 70-6), the following
characteristic features are noted.

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

Upper dentition

1. Less hypsodont. But note that the more recent (African and American)
Hipparion are hypsodont.

2. Pli protocone is present.

3. Enamel plications are usually more developed.

4. There is a rather high percentage of open marks.

Lower dentition

. Relatively less hypsodont.
2. Double knot:
(a) ‘caballus’ type in African Hipparion.
(b) ‘stenonis’ type in American Hipparion, and also in some Chinese.
(c) ‘Hipparion’ type in Europe and Asia.

3. External depression: this is rather profound in the Miocene forms.

4. Posterior internal depression: this is elongated, curved anteriorly and
lingually. Hence its oblique and ‘broken’ appearance.

5. Anterior internal depression: not only does this have an antero-external
angulation (like in Equus), but it also has a postero-external angulation,
with long sharp ‘horns’ directed towards the buccal aspect.

At the posterior extremity there is a deep invaginated plication.

6. Talonid is bifid on Mg.

7. Anterior depression often shows plications at both extremities.
8. There is a tendency to build stylids (proto-, ecto-, hypostylids) in milk
and permanent molars.
REVIEW OF ADDITIONAL ENAMEL ELEMENTS (STYLIDS)
OF THE LOWER TEETH
DESCRIPTION

Ectostylid=Ectostylid of Gromova (1952), Arambourg (1959), Hopwood
(1937), Sondaar (1961).
=Protostylid of Osborn (1907), Stirton (1941).
=Buitestyltjie of Van Hoepen (1930).
#Ectostylid of Osborn (1918), Sefve (1912, 1927).

This stylid arises from the basal cingulum. It is an accessory external
column or pillar that is always independent and located on the buccal side,
close to the mesio-vestibular border of the hypoconid and in the external
groove between hypoconid and protoconid, i.e. between talonid and trigonid.
In early wear it is seldom apparent on the occlusal surface because it rarely
reaches more than a short distance above the crown-root junction and it is
usually embedded in very thick cement. It can usually be observed better
on the buccal aspect where the cement is thinner.

Ptychostylid =Ptychostylid of Arambourg (1947).
=Pli caballinid of Stirton (1941).
=Ectostylid of Osborn (1918).

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 329

This is not an independent pillar, but a fold on the mesial border of the
hypoconid, in the interlobar angle on the buccal side. It may be present when
the ectostylid is present, but there does not seem to be any relationship between
these two formations.

Protostylid =Parastylid of Gromova (1952), Van Hoepen (1932), Stirton (1941).
=Protoconid fold of Cooke (1950).
#Protostylid of Osborn (1907), Stirton (1941).

This is a broad laminated pillar, compressed mesiodistally, found close
to the protocone where it is located on its antero-buccal side. With increasing
wear it fuses fairly rapidly with the protocone. This stylid also arises from the
cingulum and it may develop into an independent element on the antero-labial
angle of a tooth.

Hypostylid

This is a postero-external plication, appearing as a raised element on the
talonid. Sometimes it may develop into an independent pillar facing the
talonid.

Entostylid

This is a rare and inconstant enamel formation in the entoflexid. It may
be completely isolated in the cement (pl. 8, A) or connected to the mesio-lingual
corner of the entoconid.

FREQUENCY

Independent of the degree of attrition, which may tend to hide the presence
or the development of these formations, the stylids are very differently developed
in various genera, species and even in individuals. The stylids may only be
observed on some of the teeth of a particular jaw, and occasionally they are
found to be present only on one tooth in a complete dentition. Observations
have been made by various authors since the time when these stylids were
first described by Gaudry (1862) and Weithofer (1888) on the milk and
permanent teeth of Hipparion mediterraneum from Pikermi.

According to Gromova (1952), these additional elements are usually
poorly developed in the non-hipparionid equids. ‘The stylids are best developed
in the Hipparion group in which they are always present on the milk teeth,
but to a varying degree on the permanent teeth. In this respect the African
Hipparion are the most progressive.

Ectostylid

Statistical analysis of Hipparion elegans provides a frequency of 12%
ectostylids on the premolars and 1% on the molars, and in H. moldavicum
frequencies of 7:5°% and 6:7% respectively. Sondaar (1961) presents the
following data on the presence of ectostylids:

5

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

H, periafricanum: o

H. concudense aguirrei: o

H. gromovae: 1 in 200 specimens

H. concudense: 5 in 84 specimens

H, koenigswaldi: 20 in 150 specimens
FH. primigenium: 23 in 80 specimens

Therefore it can be concluded that the ectostylid is not quite a rarity
in Eurasiatic Hipparion permanent teeth, especially on the premolars. Neverthe-
less, the ectostylids appear to be much more frequent in the later forms of
Hipparion, i.e. in Africa, where it has often been stated to be ‘a constant feature’
(Arambourg, 1956, 1959). Comments on this will be given below. Furthermore,
it can be stated without dubiety that the ectostylids have been commonly
observed in most of the specimens from Pleistocene deposits: not only are
these stylids very frequent, but they are occasionally strongly developed when
they reach along the whole length of the crowns of the teeth.

Ptychostylid

This is the external hypolophid fold which is quite common among all
hipparionids, reaching a maximum development in Neohipparion eurystyle Cope
found in America, in which as many as four may occur. These stylids do not
arise from the cingulum and cannot be considered as true stylid cusps. There
does not seem to be any direct association between these stylids and ectostylids.

Protostylid

According to Sondaar (1961), this stylid is almost a diagnostic feature of
the genus Hipparion even though it varies quite considerably in its extent. It is
often a fold attached to the protoconid, but it may develop into an isolated
pillar. Here again, the African Hipparion seem to present the most progressive
features: it is found in them as early as the Upper Miocene (i.e. H. africanum)
and as late as Notohipparian namaquense, in which it is a very tall and isolated
pillar.

A high frequency is also found in some Eurasiatic forms, namely, in H.
elegans 755% is noted in P,—P, and 80% in M,—M,; in H. moldavicum 88% is
noted in P,—P, and 96% in M,—-M,

Sondaar (1961) states that it is a common, although not constant, feature
in all the Spanish species of Hipparion, except H. truyolsi. Sometimes a double
protostylid has been observed, and this has been especialy noted in H.
koenigswaldi, in which there is a high frequency of ectostylids (vzde supra).

In connection with the American material, Stirton (1942), who used
‘parastylids’ for ‘protostylids’, points out that ‘the statement that the parastylid
does not appear in the American Hipparion is not supported by the evidence.
Though the isolation is not as persistent nor as complete in the New World
forms as in some Eurasiatic Hipparion (H. platyodus Sefve), it does appear in

=
—
>t

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 331

early stages of wear in some teeth. This is true not only in Neohipparion, but in

Nannippus and Pliohippus, though extremely rare in the latter.’

Hypostylid
This is very frequently observed in Equus stenonis and in zebra but it is

very rarely seen in Hipparion. However, it is sometimes found in the African
forms, the most typical example of which is a M, from Omo (Joleaud, 1933).

EVOLUTIONARY HISTORY OF THE STYLIDS

The development of these stylids seems to be influenced by a common
factor: a higher frequency of ectostylids in a particular group is correlated
with a more precocious appearance of the protostylid (i.e. a higher protostylid)
in the same population, and vice versa.

In a particular group it is noted that the protostylid seems to be more
frequently, more strongly and more permanently developed than the ectostylid.

There seems to be an evolutionary pattern in the tendency to build stylids,
from Merychippus (possibly from Parahippus which already shows a weak proto-
stylid) through Miocene times up to Hipparion. Arambourg (1959) notices
that this ‘tendency to develop cingular formations’ is already present in the
Upper Miocene H. africanum from Oued el Hammam. The tendency seems to
have developed further during Pliocene and early Pleistocene times, being
most obvious in the African Hipparion, in which the most recent forms show the
strongest development.

Protostylids and ectostylids appear on the milk teeth of Merychippus in
which they show slight development. They are constantly present and show
a fair degree of development on the milk teeth of all Hipparion, and they appear
on the permanent dentition of the African Pleistocene Hipparion.

Along another evolutionary line of the Equidae, the Pliohippus—Equus
sequence, it has been shown that protostylids and ectostylids remain weakly
developed, the hypostylid being temporarily more developed, as, for example,
in Equus stenonis. With the appearance of E. caballus, all the stylids have
practically disappeared.

EVOLUTIONARY SIGNIFICANCE OF THE CONES AND STYLIDS

In a phyletic perspective, it has been established that the additional
stylids have originated from enamel buds on the basal cingulum of brachyodont

_ teeth. They developed first on the milk dentition and only later in the evolu-

tionary sequence, and in particular groups, did they become permanent
features of the adult dentition, appearing initially on M, and M, and eventually
on the premolars. For example, it is known that the parastylid was present
in Parahippus, but only on the milk teeth as an ill-developed feature. Later it
became a constant and developed characteristic, even of permanent teeth, in
Merychippus and later equids of the same phylum. The ectostylid is present in
Merychippus, but only on the milk teeth, and then only occasionally. It becomes

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

an element of the permanent dentition in Hipparion, where it appears to be
constant in some of the latest representatives of the group, i.e. in the Pleistocene
forms of Africa.

The tendency to raise enamel buds and isolate them is thus an early
ontogenetic trend, which in evolution has progressively influenced the later
(adult) stages of individual development.

Later in this paper it is suggested that this trend was necessitated by the
special architecture of the lower dentition with its characteristic development
of highly individualized conids.

The same trend, both ontogenetic and phylogenetic, seems to be recogniz-
able in the formation and the isolation of the major cones in the upper dentition.
It is really remarkable that the separation of the protocone and the hypocone
from the main lophs is a gradual process, both ontogenetic and phylogenetic.
The isolation of the protocone is rather ancient, and in the permanent teeth
of Hipparion it has extended right down to the base of the crown. Contrariwise,
the isolation of the hypocone is hardly noticeable on the permanent teeth,
but in very early stages of wear it has been observed (vide infra p. 373). However
in milk teeth the isolation of the hypocone reaches a much more characteristic
degree in similar hipparionid groups, e.g. in the South Serengeti Hypsohipparion
and in the Langebaanweg specimens (see p. 368).

The difference between the expression of cones and stylids in milk and
permanent dentitions on the one hand, and the more explicit individualization
of these features among later representatives in some phyletic lines, on the
other hand, seem to be linked with the increasing hypsodonty of the Equidae
teeth in Upper Cenozoic times.

ECOLOGICAL CONSIDERATIONS

Gromova (1952) outlined the ecological significance of the development
of the styles (stylids). She promoted the idea that they played a roleinstrengthen-
ing the tooth so as to meet the heavier requirements of coarser food in a dry
country where the grass was becoming very tough. The presence of additional
pillars and folds probably increases the trituration power of the teeth and
their resistance to pressure forces. Thus their appearance on the milk teeth
would prevent or decrease rapid wear.

Furthermore, Gromova developed a very suggestive correlation between
elongation of the protocone, development of the stylids, thickening of the
enamel, reduction of the external groove and higher hypsodonty. These
functional features are different methods of improving the efficiency of a tooth
with increased power of trituration. Consequently it was suggested that the
styles were developed as an adaptation for improving the grinding of coarser
food in drier climates.

Sondaar (1961), in turn, described a similar correlation among the
Spanish Hipparion which have the highest frequency of stylids. In H.
koenigswaldi the maximal development of plications in the upper molars increased

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 333

the enamel surface with a double hypoconal groove, while the lower teeth
display wavy internal depressions—entoflexid and metaflexid. However, H.
elegans, with the same functional features, only displayed a very high frequency
of protostylids. Most of the African Hipparion developed the stylids and a
maximal hypsodonty, but not the extreme plications. There seems to be a
balanced compensatory effect between stylids, plications and other elements
so that, dependent on the specific region, one or other of these elements would
develop to a varying degree. In America, Hipparion adapted to a hard and
tough grass by an increased hypsodonty and a thickening of the enamel. ‘There
can be no doubt that numerous plications and deep invaginations of the
enamel pattern, as well as elevations of folds and stylids, provide the teeth with
a more efficient trituration surface, and are very suitable for herbivorous
animals in an area of increasing aridity (Stirton, 1941).

TAXONOMY BASED ON STYLIDS

The observation of the presence of stylids, both proto- and ectostylids,
provided a new basis for generically differentiating the various African
hipparionids. Haughton (1932) decided to create the genus Notohipparion
because of the appearance of these stylids. Van Hoepen (1932) followed the
same line of reasoning and erected the genus Stylohipparion, a distinction
supported by Joleaud’s publications of S. lzbycum from Omo (1933).

To some extent Dietrich (1942) followed the same tendency and, on the
basis of the presence or absence of ectostylids, he divided the Serengeti LOWER
cheek teeth into two groups. The group without stylids he termed Hypso-
hipparion and he referred those with stylids to Stylohipparion. However, Dietrich
did not attach a true taxonomic value to this separation because he found it
impossible to distinguish between the upPER teeth of the two groups. He
indicated that the frequency and development of styles (-ids) can be influenced
by environment or selective pressure. Consequently he suggested that Stylo-
hipparion might not represent a true genus, but only a variety formed under a
strong developmental pressure (‘Entwicklungswucht’).

However, outside Africa, the variation in the development of stylids did
not lead to such taxonomic differentiation. Arambourg (1947) questioned the
validity of a generic difference (Stylohipparion) based upon the presence of a
feature which did not seem to separate clearly the Eurasiatic Pliocene forms.
Nevertheless, he opined that the very high frequency and the strong develop-
ment of the ectostylid in African forms supported the validity of the generic
differentiation.

Gromova also discussed this matter. She did not consider a true generic
differentiation of African hipparionids valid. Ectostylids are not an entirely
new characteristic in them exclusively but only represent a further development
(either in frequency or structure) of a feature not at all exceptional (albeit
irregular and even rare) in Eurasiatic Hipparion. Gromova proposed a sub-
generic rank Hipparion (Stylohipparion) for the African group on the basis of

334 - ANNALS OF THE SOUTH AFRICAN MUSEUM

obvious peculiarities of this geographical unit. However, she writes of these
ectostylids: ‘the importance as a diagnostic feature and generic character has
been somewhat exaggerated’.

This criticism can be expanded (see pp. 387-92) and even the sub-generic
status of Stylohipparion can be questioned. In spite of the ‘air de famille’ (Aram-
bourg, 1959) of all African hipparionids, the major reason for placing the
Pleistocene African hipparionids in a special genus or sub-genus is precisely
the presence and constancy of a very high frequency (which the present authors
doubt) of the ectostylids. Although this is generally accepted, it is a highly
debatable basis of separation. As mentioned above, Dietrich (1942) described
a very large collection of lower teeth without ectostylids and placed them in
the ‘genus’ Hypsohipparion. However, Arambourg (1947) suggested that these
teeth had ‘erroneously’ been attributed to a hipparionid and that, in fact, they
belong to F. zebra. This suggestion tends to remove Hypsohipparion from the
hipparionid scene so that the ‘frequency’ of the ectostylids among African
hipparionids still remains artificially high. Arambourg’s suggestion and his
deletion of this group seem to have been largely accepted. Gromova’s mono-
graph does not include the Serengeti material and does not refer to this paper
of Dietrich (1942). Arambourg (1956) has maintained his viewpoint and
speaks of ‘the typically African genus Stylohipparion characterized by the
presence of a broad ectostylid’. Naturally this does not positively exclude
Hypsohipparion, but in 1959, in a revision of the ‘few’ fossil African hipparionids,
Arambourg explicitly limits the Pleistocene material to Stylohipparion,
‘characterized by a permanent ectostylid’.

However, it can now be stated that the Serengeti Hypsohipparion material
can no longer be excluded because of the assumption that it belongs to zebra.
More than 90% of the small series of Langebaanweg equid upper teeth
unquestionably belong to Hipparion so that it cannot be asserted that all the
lower teeth belong to zebra merely because they do not possess an ectostylid.
Not only is this statistically highly unlikely but also the morphological features,
for example, the division of the talonid in Mg; (on Lg938, the only Mg available
at Langebaanweg) would definitely exclude this group from belonging to zebra.
Consequently, as the Langebaanweg material, in which the ectostylid ‘is
constantly absent, belongs to Hipparion, there are no longer sufficient grounds to
exclude a priori the Serengeti Hypsohipparion teeth from the hipparionid group
only on the basis that they do not possess an ectostylid.

Furthermore, it can now also be stated that the concept that African
Hipparion possess a very high frequency or permanent presence of ectostylid is
no longer applicable. Consequently the validity of the generic status of Stylo-
hipparion becomes highly questionable. Later in this paper this will be discussed
more fully in the light of the evidence presented. At present it suffices to state
that in Africa the Pleistocene Hipparion may be found with and without
ectostylids.

:
¢
é
4
’

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 335

REVIEW OF PUBLISHED AFRICAN Hipparion CRANIAL MATERIAL

MIOCENE

Oued el Hammam and Marceau

The presence of Hipparion in the Miocene of the Maghreb has been known
since 1932 when Suess discovered a mammalian fossiliferous deposit upstream
from Bou Hanifia (Oran) during the construction of the Oued el Hammam
dam. On several occasions brief comments have been made about this material
by, among others, Arambourg (1947, 1951, 1954). A rather rich collection
of cranial and postcranial material, with fairly complete milk and permanent
dentitions, as well as isolated teeth (table 7), have now been recovered from
two Upper Miocene sites in Algeria (figs. 3 & 4): Oued el Hammam (Oran)
and Marceau (Algiers). A detailed description was given by Arambourg
(1959): he discussed their relationships to and differences from Eurasiatic
forms of similar age, and other African forms of the Plio-Pleistocene period.
He suggested that this material should constitute a new specific form, namely
Hipparion africanum for which he proposed the following diagnosis:

Hipparion with a large skull, medium-sized limbs and heavily built extremi-
ties. Face and snout are elongated; nasal aparture is long and broad;
orbits are situated far back; pre-orbital fossae are long, simple and distant
from the orbit. Dentition of medium size: P?-M? = 141—154 mm.
Upper molars with strongly plicated enamel; compressed elliptical or
lenticular protocone. Cingular formations developed on lower milk
teeth but a laminated protostylid sometimes persists on permanent molars.
The limbs are rather short, with robust metapodials that have
well-developed lateral digits.

Arambourg emphasizes the ‘African character’ (vide infra) of the dentition
of this new species: enamel plications and narrow elongated protocone, as
well as a tendency to develop cingular formations. Because of these features
which distinguish it from all Eurasiatic forms, and because of its stratigraphical
location in the Upper Miocene, he suggested that this species should belong
to an independent African stock, isolated on the continent of Africa since the
end of the Miocene.

Camp Berteaux

Bourcart (1937) mentioned the discovery of Hipparion at Camp Berteaux
(Taourirt, Eastern Morocco). This material and additional specimens, described
by Ennouchi and Jeannette (1954), are now considered to belong to Hipparion
africanum.

Other sites

It is not possible to comment on the Aipparion material that was merely
mentioned by Dalloni (1915) and Solignac (1927), which was derived from

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

UPPER Specimen Pe Ps Ps M? M? M?#
A-P length get be 141 33°0 26-0 25°0 23°0 23,°0 20°0
Mar.* 27°0O 24°0
Transv. breadth .. Bua 122 22 *'O) 24°0 23°0 16-0
141 22°5 24°5 25e 23°0 20°5 18°5
Mar.* 26-0 25°7
Height 122 48°0 58-0 60-0 47°0
Protocone length et 141 7°8 g°2 10°2 7°32 7°5 7°6
Mar.* 8-4 8
Protocone breadth a 141 5:0 4°0 a5 3°79 3°5 2°5
Mar.* 5:0 4°0
LOWER Specimen Py Pa P, MM, M, Mo eae ee ee,
A-P Length ai! 143 30°0 .29"6. 2510. 25-2) 2h ome
89 27°0. 29°O. 25°0 29°@ 22700 Beam
3 29°O 24°0 29°5 20°97 21" s7ea
Mar.*)" ares
6 30°5 24°2 26-0
105 31°6 29°0 30°74
Transv. breadth .. Mar.* 18:0
6 14*O 1474 13°5
105 12°O 12:0 9g

*— Marceau: no number given.
TaBLE 7. Measurements (mm.) of the teeth of Hipparion africanum (from Arambourg, 1959).

the Upper Miocene layers of the lower Tafna Valley (near Guiard, Oran),
and the Djebel M’Dilla ‘Pontian’ horizons (of Tunisia) respectively.

Discussion on the ‘African character’ of Hipparion africanum Arambourg, 1959

The relationships of H. africanum, of the Upper Miocene of the Maghreb,
to other African hipparionids have been discussed by Arambourg. He dis-
tinguishes three species, stratigraphically separated: H. africanum of the Upper
Miocene, H. sitifense of the Pliocene of the Constantine and Oran area, and
Stylohipparion libycum of the ‘Villafranchian’ of North, East and South Africa.
Despite the great temporal span, Arambourg believes that there is a ‘family
air’ or unity common to these three forms, which distinguishes them from
the Eurasiatic material, and which stresses the common origin of the African
hipparionids, isolated since the end of the Miocene.

From the description given of the skeletal material of H. africanum, it
seems obvious that there is a marked difference between it and the Eurasiatic
forms. Until the present, only teeth have been available for comparison, so that
the lack of adequate comparative postcranial material has made it impossible

gh om

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 337

to determine to what extent H. africanum shares common skeletal features
with the African hipparionids of the Pliocene and Pleistocene. Dr. Leakey
indicates (personal communication) that considerable postcranial material
has now been recovered from Olduvai. These are to be described by Dr. R. H.
Stirton of Berkeley, California, who will elucidate the skeletal relationships.*

On the basis of the teeth alone, it is difficult to acknowledge a close
resemblance between the three African species. As there is a long span of time
separating H. africanum from Stylohipparion libycum, one expects to find some
differences, but it is first necessary to assess and appreciate the importance of
the resemblances stressed by Arambourg before it is possible to evaluate the
phyletic implications.

Of the ‘family air’ features linking H. africanum and the later African
species, Arambourg considers two as most typical: (i) In the upper cheek
teeth, the enamel plications and the shape of the protocone of the molars;
(ii) In the lower molars, there is a tendency to develop additional cingular
stylids. Two other classical features which will be discussed are the hypsodonty
of the cheek teeth and the shape of the double knot in the lower molars.

Enamel Plications : It is commonly believed that the enamel plications have
little phylogenetic significance. The plications are very variable, being much
influenced by functional and ecological conditions. In any event, because of
the vast temporal and spatial differences, and especially climatic and biotopical
environments, this feature cannot be used as a basis of comparison. This is
just an a priori consideration.

However, Arambourg writes (p. 93) that H. africanum is characterized by
the complex pattern of enamel plications, which are more numerous than on
most of the European forms. Although there is quite considerable variation in
the complexity of the enamel at a particular period and in a particular area
(compare H. crassum and concudense; Greek and Samos Pliocene forms), it
cannot be denied that the Eurasiatic forms normally possess a rather plicated
enamel mark-wall pattern. Furthermore it is certainly far more developed
than that of the African Pliocene and Pleistocene species, which are actually
characterized by their small number of plications.

The multiple pli caballin seen in many teeth does not seem to be typically
African either: it is observed in many other groups on both sides of the Mediter-
ranean Sea, and also elsewhere—compare H. sitifense (Mascara), H. gracile
(Pikermi), H. koenigswaldi, H. concudense (South Aragon), Hypsohipparion albertense
(Serengeti), and even in America in Neohipparion eurystyle Cope, in which as
many as four may occur on the anterolabial border of the hypoloph.

Protocone: The elongated and narrow protocone is undoubtedly a typical
and constant feature of all the African hipparionids so far known from the
Pliocene and Pleistocene deposits. It also seems to be a distinct, strongly
expressed evolutionary trend in the whole group (see pp. 354-5; table 14;

* See Olduvia Gorge 1951-1961, 1 by L. S. B. Leakey (1965) which was published while
this paper was in press.

338 ANNALS OF THE SOUTH AFRICAN MUSEUM

figs. 12, 13). A satisfactory comparison of individual teeth at different stages
of wear is difficult, and published sketches of this feature may be deceptive.
Nevertheless, it is obvious that a comparison of the indices of the length and
shape of those protocones available for study definitely confirms this ‘African
feature’ of H. africanum. For all teeth, it shows a constant greater ‘African’
index than any of the European species considered (except for the breadth/
length index of P?, which in any event, is a rather atypical tooth). It is important
to note that such a progressive feature, characteristic of the African phyletic
line, can be traced back as far as the Upper Miocene.

The double knot: Gromova (1952), in a recent review of all the available
evidence, indicates that the original type of knot is the ‘stenonis’ one: it presents
a very different metaconid and metastylid, the latter being strongly angulated,
and separated from the metaconid by a narrow deep valley. This original
type, found in the Merychippus ancestor, has been retained mostly in the
American Hipparion. However, there seems to be a constant modification
amongst Old World Hipparion: towards a ‘Hipparion’ type of knot in Eurasiatic
forms, characterized by a symmetrical and rounded metastylid and metaconid
separated by a wide and shallow valley; and towards a more ‘caballus’ type
in Africa, where the metastylid is sub-triangular. The description and drawings
of H. africanum indicate clearly that it has a ‘Hipparion’ type of knot, more
typical of the Eurasiatic form and bearing no resemblance to the African
forms. As H. africanum is the oldest species of Hipparion known in Africa, it
needs to be decided whether it has acquired this ‘progressive’ feature very
rapidly or whether the ‘Hipparion’ type of knot is not as progressive as was
previously considered, having already been present in some Merychippine
ancestor, and developed in parallel with the ‘stenonis’ type.

Stylids: The tendency to develop stylids is quite marked in many, but not
all, African forms. It is said to be expressed in H. africanum by a small laminated
or rounded, rather isolated protostylid on the permanent P;-M, of specimen
no. 2, and by a small ectostylid on the vestibular aspect of most of the milk
teeth. The tendency is found to be constant among all Hipparion. The stated
degree of development of these structures does not seem to be particularly
‘African’: it is certainly surpassed by some Eurasiatic species, for example,
H. koenigswaldi from Teruel. Of course, it should not be overlooked that the
feature seems to have been expressed rather weakly in Merychippus times. The
minor development that it shows in the Upper Miocene of Algeria may hardly
be considered ‘African’ when compared with the Stylohipparion of the ‘Villa-
franchian’. However, the feature suggests a rather strong and precise
evolutionary trend at work.

Hypsodonty: H. africanum is stated to be poorly hypsodont. The indices are
by no means comparable to the ‘African’ specimens of the Pliocene and
Pleistocene times, and they are of the same magnitude as the other Hipparion
groups of the Mediterranean basin. H. africanum is even less hypsodont than
H. sitifense which is stated to have low teeth.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 339

PLIOCENE
Mascara

A palate with an almost complete dentition and a symphysial region with
incisors, probably belonging to the same skull, have recently been described
by Arambourg (1956).

This Pliocene form is characterized by very small dimensions of the teeth
(table 8), which also show few and simple plications and little hypsodonty. The
protocone, completely independent of the protoloph right to the base of the
crown, is relatively compressed.

The specimens have been referred to H. sitifense Pomel with which they are
stated to share ‘almost identical morphological features and size’.

pe Ps p4 Mt?! M? M3 P2—Ms
1. MASCARA
A-P length dee 3 $1.30 20°5 20 1995) 19 18°8 129
Transv. breadth .. oe tars i Se We 28) 20 19°5 16 18-6
Crown height 7 ve Peewee OD 20"% ~ TO"F 55 13
Protocone length .. i4 ees FY Bak Gag 74 8:8
Protocone breadth .. - Re OT ae ae 3
Protocone shape .. ay 60 55 55 46 34
Protocone length/Tooth fone abe eel!) R4. 41 33 38 46
2. ST. ARNAUD CEMETERY
H. sitifense (type specimen)
A-P length ie - “a 23
Transv. breadth... a fi 22°5
3. AIN EL HADJ BABA
R L
A-P length ays — di 225,20" 2), 20 20°2
Transv. breadth .. co ts 21 20°5 21 20°3
Protocone length .. Ef ne G2 6 ahs Dail 3
Protocone breadth .. xi me ma) iA. Br 7s
Protocone shape .. Le 51°6 66°6 52-2 47°5
Protocone length/Tooth fength Bi 27a QOEF Be" “aio

TABLE 8. Summary of measurements (mm.) of teeth of North African Pliocene Hipparion.

St. Arnaud Cemetery

The type specimens of H. sitifensis (sic) Pomel 1897 were recovered from
the St. Arnaud Cemetery. These two upper teeth, figured by Pomel (see also
Arambourg, 1956) are little hypsodont and rather small (although not quite
as small as the Mascara teeth). A broken calcaneum is also mentioned.

Lower teeth of a correspondingly small size and medium hypsodonty,
collected by Arambourg but as yet undescribed, have no ectostylid.

Ain el Hadj Baba

Upper isolated, but partially serial right (?) P4-M?, a left M?, lower
teeth, and limb bones were described by Thomas (1884) who referred them
to H. gracile (Kaup).

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

However, according to Arambourg (1956), the narrow and small number
of plications of the enamel, the double pli caballin, features of the protocone,
dimensions of the upper teeth and absence of the ectostylid on the lower teeth,
make the Ain el Hadj Baba specimens very similar to H. silifense types and
co-types, as well as to the Mascara specimens. Therefore he considered them to
be A. sitifense.

From the metapodial, it may be observed that the lateral digits were
still very strongly developed.

Utique (Northern Tunisia)

This area also includes the continental sands of the Ferryville region near
Bizerta (fig. 4).

Solignac (1927) indicated that the fragmented remains had been assigned
by Depéret to H. crassum Gervais, but Arambourg (1956) is inclined to refer
them to Stylohipparion.

St. Donat (Algeria)

Teeth of two maxillae, which Arambourg (1956) interprets as being
possibly referable to H. sitifense, were found at this site. Joly (1909) recognized
further teeth from this site and hastily referred them to H. gracile, but they
should probably be compared with Arambourg’s 1956 material and pooled
with A. sztifense.

PLEISTOCENE

North Africa

Oran: The teeth recovered by Pomel in the Oran area are among the
first discoveries of Hipparion in Africa and formed the basis of the first description.
Two sites were mentioned:

(i) ‘St. Pierre sandstone quarry, on the property of Mr. Brunie, in the
St. Charles district, in the east quarter of Oran, which is also the
type locality of Libytherium maurusium.’

Two lower teeth were described by Pomel (1897) as the type specimens of
EHipparion (?) libycum. The one, a left P; or P,, has an ectostylid while the other
is without an ectostylid, but the antero-external region is broken. The specimens
were successively referred, first by Van Hoepen (1932) to Stylohipparion steytlert
because of its similarity to P,; (Nas. Mus. C795) from Cornelia (vide infra, p. 347) ;
then by Joleaud (1933) to Libyhipparion libycum because of its fundamental
similarity to the du Bourg de Bozas Mission material from Omo, although it is
more elongated and somewhat larger; and later by Arambourg (1947, 1956) to
Stylohipparion libycum (or albertense).

A distal portion of a third metatarsal was also recovered from this deposit.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 341

(ii) Puits Kharouby.

Five upper molars were described by Pomel (1897) and referred to H.
massoesylium. However, Pomel himself was not too sure whether or not these
upper teeth belonged to those lower teeth which he had described under the
name Hipparion (?) libycum. Their reciprocal kinship was ascertained by Joleaud
(1933) who consequently referred H. massoesylium to his Libyhipparion libycum,
and by Arambourg (1947, 1956), who for the same reason, placed them in
Stylohipparion libycum (or albertense).

Ain fourdel (Constantine): The material consists of a M, and a lower
premolar, the additional stylids of which seem to have been overlooked by
Thomas (1884) who referred them to Hipparion gracile.

Joleaud (1933) emphasized their similarity to Pomel’s Oran specimens and
to the material discovered in the Orange Free State of South Africa. He proposed
that the Ain Jourdel material should be placed in a new species Stylohipparion
(?) thomas.

Arambourg (1947) can see no reason for not pooling Ain Jourdel (one
of the two teeth at least) and ‘H. massoesylium’ (= Joleaud’s Libyhipparion
libycum) with Stylohipparion libycum.

Beni Foudda (Constantine) : This site has been referred to as Ain Boucherit
by Arambourg.

A molar recovered here was attributed by Pomel (1897) to Hipparion
ambiguum but was referred by Arambourg (1947, 1956) to Stylohipparion libycum
(or albertense).

Wadi Natrun (Gart el Moluk Hill): Andrews (1902) described a left upper
premolar (?P*) of large dimensions (A—-P 29 mm.; transv. breadth 28 mm.),
but apparently it is little hypsodont (height 38 mm.). However, it is a rather
warn specimen.

When the specimen was discovered, very few African Hipparion had been
recovered, and the typical features of the few that had been recovered were
still largely unknown or as such unrecognized. The best comparative material
at that time was that from Oran: the enamel pattern of the specimen from
Wadi Natrun appeared definitely more complex and it differed in the absence
of isolation of its hypostyle. Therefore, in spite of its transversely more com-
pressed protocone, which explicitly suggested some close resemblance to

Hitpparion theobaldi (from the Siwaliks), the specimen was referred to the more
widespread H. gracile.

After the discovery of further material from Omo and Kaiso, Joleaud
(1933) emphasized the difference in the parastyle and mesostyle features from
Libyhipparion ethiopicum, and he stressed the resemblance with the Kaiso speci-
men, referring to it the Wadi Natrun premolar, which he called Hipparion
cf. albertensis (sic).

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

Central Africa

Koro Toro: The Hipparion material recovered from the ‘Villafranchian’
horizon of the Tchad has been provisionally referred by Coppens (1960) to
Stylohipparion. He is preparing a detailed description of these specimens, together
with the associated fauna. Therefore, it is not possible to comment on this
material at this stage.

East Africa

Omo: The first specimens were recovered by the du Bourg de Bozas
Mission (1903) and were described by Joleaud (1933). They consist of a few
isolated teeth, namely:

I incisor;

5 lower cheek teeth (1 right premolar, one left M3, two left molars
and one right molar) ;

1 fragmented upper right molar.

To this group Arambourg (1947) added a left M?! or M?, first interpreted
by Joleaud as Hippotigris.

A second series of specimens was recovered by Arambourg (1947) during
his 1932-33 expedition, and comprises 1 right P?; 1 left M%; 1 fragmented
right upper molar and 1 right M, or Mg.

The first of these groups was described by Joleaud (1933) who erected
the new genus and species Libyhipparion ethiopicum,’ generically distinct from
the South African specimens from Namaqualand and Cornelia on the one
hand, as well as from the Constantine material of Thomas, on the other, and
specifically separated from Pomel’s St. Charles teeth.

In a later reconsideration of these Omo specimens, to which the second
series of teeth from the same area was added, Arambourg (1947) rather empha-
sized (a) the close relationships between all the Ethiopian hipparionids; and
(b) their profound similarity to the East African (Kaiso, Serengeti pro parte,
and probably Olduvai) material and the Orange Free State discoveries.
Furthermore, Arambourg considered it reasonable to refer most of Pomel’s
and Thomas’ North African (St. Charles, Puits Kharouby and the Constantine)
material to S. albertense or possibly to S. libycum.

The characteristic features common to these specimens are:

(i) pronounced hypsodonty reaching 80 mm. in unworn teeth;
(ii) upper molars with very complicated enamel pattern and laterally
compressed and strongly elliptical protocone;
(iii) lower teeth with strongly developed additional stylids, rather large
oval or flattened and pointed ectostylid;
(iv) the presence of a ptychostylid; and
(v) ridged or pillar-like protostylid and well-built entostylid.

(1) With exception of left upper molar, considered by Joleaud as belonging to Eguus
(Hippotigris).

i
:
"

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 343

Therefore, there are no reasons for proposing any generic or specific
distinctions between these Pleistocene African Hipparion, which according to
the priority rules have been referred to Stylohipparion albertense.

Olduvai: Some of the first fossils identified in the Olduvai Gorge, when it
was first discovered by Kattwinkel in 1911, were the distal ends of 3 metatarsals
of hipparionids, which Hopwood (1937) referred to Stylohipparion cf. albertense
(Hopwood).

The dimensions of the lower articulations of metatarsal III were 40 x 30
mm. The broken metatarsal IJ] and metatarsal IV are stated to have had
‘originally’ almost the same lengths as metatarsal III, and to have had the
appearance of broadening distally.

Further specimens of teeth, recovered by Kattwinkel and Reck in 1913,
as well as an upper dental series collected by the British expedition, were
' referred to the same genus and species. The diagnosis proposed by Hopwood
(1937) for this Stylohipparion cf. albertense is:

Three-toed equid, with high-crowned molars of the Hipparion group.
Lower teeth are laterally compressed and display a strong, often laminated
ectostylid. Genotype: Stylohipparion hipkint Van Hoepen (1932).

Unpublished data

In recent years a large collection of equid material has been recovered
from the Olduvai Gorge by Dr. L. S. B. Leakey. All the specimens from Bed II
(and a few from Marsabit Road, Olorgesailie and Omo) are now being studied
by Dr. R. A. Stirton in California. Dr. Leakey kindly permitted the authors
to make a brief survey of this material. In order not to encroach on the final
description, only a few features relevant to this paper are mentioned:

(i) In this mixed sample there are 471 Equus teeth, of which 224 are
from the upper dentition. Of the 186 teeth of hipparionids, 100 are
upper. The vast majority of these are from Bed II, only a few deriving
from other sites.

(ii) This is then a reasonable sampling to ascertain the expected propor-
tions of upper and lower teeth. It will be indicated below that at
Langebaanweg there are approximately equivalent proportions of
uppers and lowers, but at the latter site there are almost no Equus
specimens.

(i11) The data is also useful to assess the situation at South Serengeti
(vide infra). In this respect it is interesting to note that in the Olduvai
collection, among the lower teeth, those of hipparionids have elong-
gated, strongly marked ectostylids, typical of Stylohipparion, while
those without ectostylids lack the typical features of Hipparion as well
as the protostylids of the Langebaanweg specimens. They are typically
Equus in type.

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

(iv) There is a great range of variation in the size of these Hipparion
teeth, and also in the length and shape of the protocone. On the
whole, the range seems smaller than that of the Langebaanweg
teeth, but the protocone is more elongated. In the Hipparion lower
(as well as in these Equus) teeth, there is a complete absence
of protostylids and hypostylids.

South Serengeti: A large collection of Hipparion has been made from various
localities in the South Serengeti, and was described by Dietrich (1942). A
complete list has never been published. However, from the available data, it
is possible to ascertain that the collection consists mostly of isolated and frag-
mented specimens, a few maxillary and mandibular fragments (among which
are symphysial portions with incisors), many loose upper and lower teeth,
both milk and permanent. There are also some rare postcranial specimens,
namely, a proximal fragment of a femur, a distal fragment of a tibia, two
third metatarsals, one third metacarpal and tarsals. Dietrich artificially
reassembled the isolated teeth into series, and most of the dental series figured
in his publication are reconstructed. From the scattered data and illustrations,

it is possible to establish an incomplete list of the South Serengeti dental

material (table 9).

Plate Number in

Description Origin and Coll. Number Dietrich (1942)

Milk Dentition

Upper r dP2-dP4 Vo 330 XIII 96
r dP2-dP3 Gadj 10 XIII 97
1 dP2-dP4 Gadj 2.39 XX 162
Lower r dP2-dP4 Vo 313 XIII 93b
r dP3-dP4 Marambu XIII oga
r dP4 Gadj 10-13.3.39 XIII 94
Permanent
Upper | P2—-M3 Vo, Gar. XIII 87
1 M2-Mg .. Vo 670 XIV 102
1 Po-M3°*.. Vo 330, Vo 670 XV 107, 108
1 P2-P4 Vo 670 XX 160
r Po-M3 .. Vo, Gar. XIII 88
r M2-Mg3 .. Vo 670 and Marumba XIV 103
Lower 1 M3-Pe2 Vo 313, 670 XV 106
1 P2-M3 Der., Gadj., Gar., Vo XV 105
1 Pe—-M3 Vo, Gar. XIV to1
1 Pe—-M3 Vo 330, Vo 670 XIII 90
1 Mr Garussi 2.39 XIV 99
1M Gar. River 200 XIII 95
r M3-P2 Vo 670, Gar., Olduvai Hill XVI 109
r P2—-M3 Vogel River XIII 89
r P2-M3 Vo 670, Gar., Vo XIII g1
r P3—-M3 Gadj 10 XIII 92
Symphysial region with incisors Vogel River 9-10-38 XVI 112

TaBLeE 9. Hipparion material mentioned by Dietrich (1942) and derived from
South Serengeti region

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 345

In contrast to the relatively unchanged nature of the postcranial skeleton
(vide infra) since Pliocene times, the advanced and progressive features of the
dentition of the South Serengeti hipparionids have been emphasized by Dietrich.
The teeth, both upper and lower, are strongly hypsodont (crown length:
g cm.). The upper molars show numerous enamel plications, and an elon-
gated, ungrooved protocone, completely isolated right down to the base of
the crown, and embedded in a thick cement layer. The incisors (see p. 350) are
longitudinally grooved on their anterior surface (in one case I, is missing).

It does not seem possible to distinguish different forms of hipparionids at
Serengeti on the basis of the upper teeth. On the contrary the lower teeth show
some constant features whereby it seems possible to distinguish two groups.*
A large group of teeth, all apparently recovered from the gray tuffs, does not
show any additional stylids while another group of some 50 (40 isolated teeth
and 2 incomplete dentitions) displays the ectostylids characteristic of most of
the African hipparionids then described. In these teeth the ectostylids are
sometimes very strongly developed, pillar-shaped or flattened and ridged.
Protostylids and hypostylids are also regularly present, and a very peculiar
entostylid (see p. 360) may be observed on one tooth (Garussi River 200, fig. 95
of Dietrich, 1942). ‘These teeth derive from deposits that seem to span over a
longer period of time than do those from which the teeth without stylids were
recovered: some of them probably occur in the gray tuffs, contemporaneously
with the first group, while others belong to a probably younger horizon.
However, as has been stated previously, the stratigraphy of the South Serengeti
is very poorly documented. Other than the presence of the stylids and the
average shorter total length of the dental series at a comparable stage of
wear (156 mm. v. 170 mm.), there seem to be no appreciable differences
between the two groups. However, the value that can be attached to the
assessment of the total length of the dental series in these specimens is doubtful
because (a) of the ‘reconstruction’ of most of Dietrich’s series, and (b) of the
variation of the total length due to wear.

In spite of the above distinction, Dietrich does not consider it necessary
to separate generically the South Serengeti material. ‘This attitude is reasonable
because

(1) lack of knowledge of a stratigraphical sequence rules out a clear
chronological sequence;

(2) too little was known of the skeleton, both cranial and postcranial ;

(3) the upper teeth showed no features whereby they could be separated
into two groups, and yet it is a statistical probability that, if there were two
species at the site, all the uppers could not have belonged only io one of these
species; and furthermore,

(4) the development of the ectostylids, ranging from absence through
mild formation to strong development, is an expression of dental adaptation
to environmental factors (vide infra, p. 352).

1 See also discussion on Arambourg’s concept of the non-validity of Hypsohipparion, p. 333.

6

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

On the basis of the data available then,! Dietrich considered that within
the Serengeti material there was an original and conservative African stock of
hipparionids for which he erected a new genus Hypsohipparion Dietrich, 1942,
with Hipparion albertense (Hopwood) from the Lake Albert Kaiso bone beds
as the type specimen. Into this genus and species he placed the Serengeti
material without stylids, as well as all the upper teeth and the postcranial
remains. ‘The diagnosis that he proposed was:

An advanced three-toed, short-snouted Hipparion of the dimensions
of a small caballus, the M?® of which reaches a crown height of g cm. The
protocone shows a tendency to become laminated, non-grooved and com-
pletely isolated from the protoconule. Teeth show no reduction of the main
cusps and styles. Plications are more marked than in all Pliocene species.
Lower teeth (deciduous premolars, permanent premolars and molars)
are (?)? all without ectostylids.

Dietrich believed that the group manifesting the stylids was displaying
the developmental trend of a selective pattern. He provisionally placed this
group in Stylohipparion as a practical measure, partially comparable to Stylo-
hipparion libycum, Stylohipparion ethiopicum, and Stylohipparion steytler1. However,
he maintained that the generic labelling had no strict taxonomic value because
of the scarcity of the material on which this was based. He did not see the
value of assigning his material to a new species.

Arambourg (1947) questioned the validity of the generic status of Hypso-
hipparion on the basis of a possible incorrect determination of what he considered
to be lower zebrine teeth (see p. 333). In later publications (especially 1956 and
1959) he seems to have definitely adopted this view permanently. In his opinion
the Serengeti material belongs partly to Stylohipparion and partly to E. zebra.
This opinion is discussed and considered unacceptable elsewhere in this paper.

Lake Eyasi: The few isolated teeth recovered from the Lake Eyasi shore
(west and north of Mumba Hill) are very much rolled and unsuitable for any
detailed study (vide supra, p. 287). They were referred to Hipparion sp. (Reck and
Kohl-Larsen, 1936).

Lake Albert (Uganda): From the east shore of the Lake, one incomplete
upper molar (B.M.N.H. M12615) was described by Hopwood (1926) as the
holotype of Hipparion albertensis (sic). The tooth is rather hypsodont, with a
complicated enamel pattern. Its antero-posterior diameter is 24-5 mm.

Dietrich (1942) used it as the genotype of his new genus Hypsohipparion
albertense, the proposed diagnosis of which is noted above.

South Africa

Namaqualand (Cape Province): Nine isolated mandibular teeth of the
same jaw, left P,-M, and M, and right P,-M, and M, (fragmentary) were
1 Through the kind collaboration of Dr. K. H. Fischer, Berlin, measurements of some of

the ‘Hypsohipparion’ teeth were made available to the authors: see table 20.
2 Dietrich includes this ‘query’ in his original diagnosis in German (p. 97).

ee

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 347

described by Haughton (1932), who erected a new genus and species WNoto-
hipparion namaquense. ‘The type specimen is S.A.M. 9982. Unfortunately a
diagnosis was not provided.

A small ectostylid and a protostylid are occasionally present. It is difficult
to comment on the height and relative hypsodonty of these teeth because the
crown is rather low, as the specimens are in a very advanced stage of wear
(table 10). The teeth possess a thick cement.

Cooke (1950) proposed a diagnosis for the genus Notohipparion Haughton:
‘rather low-crowned, heavily cemented hypsodont lower cheek teeth with an
extra antero-external cingulum fold or column, either isolated or fused with
the parastylid, present in all the permanent cheek teeth except the second
pre-molar, and a deep groove separating the strongly developed metaconid
and metastylid. The upper dentition is unknown’.

Because of the moderate hypsodonty and the weak development of
additional stylids, Dietrich (1942) considers Notohipparion to be more primitive
than Stylohipparion. This opinion had also been held by Van Hoepen (1932).

Arambourg (1947) compared these lower molars with the ‘single’ African
Pleistocene genus Stylohipparion but, probably because of the poor stratigraphy,
he did not revise its taxonomic status. In later comments on the African
Hipparion, he (1959) makes no further mention of these specimens.

P, Bs P, M, M, M,
A-P Length .. a big doi 20°5 27°0 26-0 ora) 24°0 29°0
Transv. breadth " a oto @ 17°0O 17°0 16:0 14°5 12°5
Height .. Be + ih AS Ohi I7°5 25°0 23°0 30°0 34.°0
Transv. breadth/A-—P length seat, "§aaee 63 65°4 69°6 60° 4 43
_ Height/A-P length .. As ms, AO?2 64°8 96°11 +100 125 Ty
Hypsodonty Index .. Ay bie, LO 97 67°9 69°5 48° 2 36°8

TABLE 10. Measurements (mm.) of teeth of ‘Notohipparion namaquense’ Haughton

Christiana (Cape Province): From this site an upper second (? third) left
molar (Archaeol. Surv. 113), considerably worn, undescribed and not illus-
trated, has been referred to Stylohipparion steytleri by Cooke (1950).

Transvaal australopithecine cave breccias: ‘Two lower teeth, recovered by
Broom in the breccia filling the Kromdraai Cave, have been referred to
Stylohipparion steytleri (Cooke, 1950). Hipparion teeth recovered from Bolt’s
Farm and Makapansgat Limeworks have been similarly referred (Cooke, 1963).
These specimens have not been described.

Cornelia (Uitzoek): Eleven isolated upper and lower teeth include a
group of five and another group of two teeth belonging to single individuals
(table 11):

(i) one right M! or M? (Nas. Mus. C558), one left M® (C555), one left
M,-—M, (C556),1 described by Van Hoepen (1930) as a new species

1 In the original publication (1930), C556 was stated to be composed of lower milk teeth
(DM, and DM,), and only in 1932 were they correctly acknowledged as permanent teeth.

348 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fiipparion steytleri, the type being C558 and the paratypes being
0555-550.

(i1) A series of five left lower teeth P;-M, (C795), recovered from the
type locality, were referred by Van Hoepen (1932) to the same
species.

(ii) A right M, (C797), in very early wear and therefore displaying
a completely uncharacteristic pattern, was used by Van Hoepen
(1932) to erect a new genus, namely, Stylohipparion, and a new species
hipkini, to which a left P, (C796), recovered from the same locality
and formation, was provisionally referred.

Van Hoepen (1932) subsequently referred steytleri to the new genus Stylo-
hipparion. He considered Stylohipparion to be ‘obviously the terminal stage of
the evolutionary line which originated from Hipparion with Notohipparion.’

Joleaud (1933), who compared steytlert, with Libyhipparion ethiopicum, and
hipkint with the Ain Jourdel specimen of Thomas, accepted the specific distinc-
tion between séeytlert and hipkini. Cooke (1950) considered this distinction to
be invalid. This opinion is strongly supported here, for both a priori and a
posteriori reasons: first of all, no upper teeth of hipkini have been recovered,
and Van Hoepen’s species is based on one single lower tooth (C797), which
constitutes a very inadequate basis for creating a new taxonomic unit.
Furthermore, the tooth is in a very early stage of wear, which is the case of
the obviously atypical, peculiar pattern of its occlusal surface. Thus the
‘difference’ between it and the Stylohipparion steytlert is not at a species level.

Van Hoepen provisionally referred specimen C796 to the new species,
but it is even less different from the known steytleri specimen. In any event,
being a P,, it does not show the typical features of a species.

On the other hand, and quite apart from this question of a specific distinc-
tion, the genus Stylohipparion Van Hoepen has been generally accepted by all
subsequent authors. Hopwood (1932), Dietrich (1942), Arambourg (1947,
19560, 1959), Cooke (1950) and Gromova (1952), among others, have considered
Stylohipparion as a valid unit (at least at a subgenus level), which expresses the
‘African’ trend of Pleistocene hipparionids. The following diagnoses have
been proposed :

Arambourg (1947): ‘“Tridactyl equid, with very hypsodont dentition.
Upper molars with compressed protocone of very elliptical section, wavy
parastyle; especially complicated enamel pattern, particularly around
the prefossette. Lower molars with broadly developed ectostylid, laterally
compressed, and close to the antero-external hypoconid pillar.’ (Direct
translation from the French.)

Cooke (1950): ‘High-crowned rather hypsodont lower cheek teeth
with a strongly developed isolated pillar external to the ectostylid, possess-
ing no external groove between hypoconid and hypoconulid, having no
protoconid and hypoconid and small rather widely separated metaconid

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 349

and metastylid. High-crowned upper cheek teeth with isolated oval

protocone and possessing a small flange on the antero-internal side of the

parastyle.’

We wish to point out that the ‘ectostylid’ referred to by Cooke is actually
the ptychostylid, while the ‘strongly developed isolated pillar’ is the ectostylid.

UPPER TEETH
St. steytlert (van Hoepen)

M! or M? M?
(C558) (C555)
A-P length A 22°O 21°0
Transv. breadth .. 22°0 18-0
Height .. 4 54°0 66-0
St. steytleri St. steytleri St. hipkini
LOWER TEETH P; P, M, M, M, M, M, BS
(C795) (C556) (C797) (C796)
me Acneth § .: 25°0 28:0 22°O 24°5 — 21°O 28°5 31°O
Transv. breadth 15°0 12°O 12-0 EL 5 12°O 12°5 15°0 14°5
Height .. ie 04°70 74°0 69:0 7a™6 2270 41°O 81-0 46°0

TABLE 11. Dimensions (mm.) of teeth of ‘Stylohipparion’ from the Orange Free State, South Africa.

From Uitzoek also comes an anterior portion of a lower jaw (Nas. Mus.
C679), containing four large first and second incisors, which are flattened
anteriorly and arranged almost in a straight line. Each incisor shows the
‘mark’ or the cement-filled depression in the enamel. The third incisor is small
and lies behind and in contact with the second incisor.

The above description (Cooke, 1950) is a slightly modified interpretation
(also suggested by L. H. Wells) of the original description by Van Hoepen
(1930), made, according to Cooke, so as to avoid the ‘startling supposition’ of
the complete absence of the third incisor. However, it should be pointed out
that specimen GADJ. 10 from South Serengeti shows complete absence of
not only I;, but also of the canines (Dietrich, 1942).

One of the special features of this specimen C679, besides the reduction
(? complete, i.e. absence) of I,, is that the lingual aspect of I, and of I, has
two surfaces, meeting at an angulated thick ridge. Each surface also has
longitudinal parallel ‘costae’.

A new genus and species Eurygnathohippus cornelianus van Hoepen (1930)
was created to include the above material. No measurements were published,
but from the illustrations and the cast it is clear that the jaw fragment belongs
to a very large skull.

Dietrich (1942)(1) made the suggestion that this mandibular symphysial

1 Dietrich incorrectly describes the specimen as belonging to a milk dentition (‘Milch-
vordergebiss’, p. 97).

59°

ANNALS OF THE SOUTH AFRICAN MUSEUM

region (for which no cheek teeth are known), should belong to Stylohipparion,
or at least to the advanced Hipparion of the Cornelia layers.

Ewer (1963) refers to Eurygnathohippus as a chalicothere. The basis for this
opinion is not clear to the authors.

NOTE ON Hipparion INCISORS

Eipparion incisors have rarely been discovered in Africa. In a recent,
comprehensive study of a large assemblage of Hipparion material from South
Aragon, Sondaar (1961) indicates a similar shortage there. The small number
of recorded specimens from Africa are:

(1) Upper incisors: the H. africanum type specimen (no. 141) from Oued el
Hammam presents a complete snout, and, from the same site, there are
also three maxillae with incisors. They have been described by Arambourg
(1959) without comment, because there is no comparative material.

(11) Lower incisors :

(a)
(5)
(¢)

The Libyhipparion ethiopicum (Joleaud, 1933) collection from the
Omo Valley contained one incisor (vide supra, p. 342).

Dietrich (1942) mentions some incisors from the South Serengeti
without going into any detailed description (vide supra, p. 344).

Eurygnathohippus cornelianus (Van Hoepen, 1930): an anterior portion
of a lower jaw. This may belong to an individual of the genus Hipparion
or Stylohipparion, as has been suggested already by Dietrich (1942)
and in a personal communication (1964) from Dr. L. S. B. Leakey
(who bases his view on material recently discovered at Olduvai).*

Each of these specimens raises difficult problems, which cannot be solved
until more complete material becomes available. A few of these problems

are:

(1)

The ‘Libyhipparion’ incisor does not show the sub-elliptical section of
Hipparion. It is completely subdivided into two clearly separated
‘marks’, no other example of which is known among Hipparion in the
literature. It has been suggested that this feature is an extreme mani-
festation of the enamel plications, typical of the African hipparionids.
It is impossible to state whether or not the feature is really exceptional.
It is not demonstrable in the other few available incisors.
Eurygnathohippus : The ‘costae’ and the blunt ridge on each specimen
are unique in this small. series. A possible explanation for these
features is that they are adaptations to browsing and strengthen the
teeth, set practically parallel to the horizontal symphysis.

The occasional reduction (? absence) of the third incisor is referred
to above.

1 Confirmed while in press. See Leakey, L.S.B.: Olduvai Gorge 1951-1961, 1. p. 26.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 351

A SUMMARY OF PUBLISHED POSTCRANIAL REMAINS OF AFRICAN Hipparion

The postcranial skeleton of African hipparionids is very poorly known.
The only described specimens are: |

Camp Berteaux: A femur fragment with the proximal epiphysis.

Oued el Hammam : Distal fragment of humerus (no. 159) ; radio-ulna (no. 123)
and fragments 13, 22 and 27; femur without epiphysis and a distal
fragment; distal fragment of tibia; pelvic fragment; two astragali;
one calcaneum; five metacarpals and five metatarsals, many with
lateral digits.

Ain el Hadj Baba: Metapodials with lateral digits.

Oran: One fragmentary calcaneum and one third metatarsal.

Omo: Right humerus.

Olduvai : Distal end of three metatarsals. There are also quite a number of
undescribed specimens. The authors examined a number of such
specimens recently collected at Olduvai Gorge. Data on the
metapodials is provided below.

South Serengeti: Proximal fragment of a femur; distal fragment of tibia;
two metatarsals III; one metacarpal III; tarsals.

Arambourg (1949) has probably described the largest collection known
from one site: it consists mainly of numerous complete metapodials (many of
them with lateral digits) which have been referred to Hipparion africanum.
From the description and the measurements, this North African Upper Miocene
Hipparion seems to have been about the same size as the smallest H. mediterraneum
of Pikermi. The extremities indicate that africanum was rather short and
heavily built. The lateral digits were more robust and strongly developed than
those of most other hipparionids, and they certainly were still functional.

From the Pliocene, only a few metapodials with very well developed
lateral digits are known from Ain el Hadj Baba. They have not been adequately
studied. It has been assumed that they belong to H. sitifense.

From the Pleistocene of East Africa a right humerus is known from Omo,
and a few fragmentary long bones and metapodials have been recovered at
Olduvai and South Serengeti. The material has not been described. Size and
morphology of the metatarsals do not seem to differ noticeably from H. gracile.
Dietrich (1942, p. 101) states that the South Serengeti metatarsal III ‘must

Breadth, distal end A—P diameter,

distal end
Olduvai (Hopwood 1937) .. sh 40 30
South Serengeti (Dietrich 1942) .. 37 35
H. elegans... i, ne Ae 29°6 25°71
H. moldavicum of a a 33°9 26°8

Hi. mediterraneum nr 4 ps 32°3

TABLE 12. Dimensions (mm.) of metatarsal III of Hipparion

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

have a length similar to H. gracile, i.e. 23-24 cm.’ The lower articulating
surface is 37 mm. broad and 35 mm. A-P, so that it is only possible to state
that the distal end is broader and thicker than in most of the European Hipparion
(table 12). The lateral metapodials reach almost to the distal articulating
facet and are rather strongly developed, but it is not possible to state firmly
whether or not they possessed three phalanges. Nothing is known about the
front extremities at present.

The only species with similar dimensions of the lower end of metatarsal
III is A. longipes with 40 and 33 mm. for the breadth and A—P dimensions
respectively. H. africanum has a distal breadth of 34:7 mm. It is difficult to
assess whether these greater dimensions of the East African Pleistocene forms
are indicative of larger overall dimensions of the animal, or whether they only
refer to more massive metapodials. Although there is no direct correllation
between the different structures, it may be mentioned that the larger size of
some of the South Serengeti molars may suggest that the first possibility is
not excluded. In general, too, Pleistocene mammals tend toward giantism.

On the basis of published material, it was not known whether or not the
African Pleistocene hipparionids were three-toed horses. There is some informa-
tion concerning the existence of metatarsal II and metatarsal IV at Olduvai,
of which it is stated that ‘they originally had practically the same length as
metatarsal III and that they are broadening distally’ (Hopwood, 1937).
However, no phalanges are mentioned.

Unpublished data

Examination of the Hipparion third metapodials from Olduvai Bed II,
now in Berkeley, California, reveals conclusively that the lateral digits were
robust, reached the distal ends of the metapodials, and were as well developed
as those of Hipparion from Pikermi. The metapodials II and IV are not repre-
sented by complete specimens or by the distal articular end. Measurements of
the fragments are considered to be of no value. Despite the lack of complete
lateral digits, the evidence of the articular grooves formed by them on the
third metapodials convincingly indicates that the East African Hipparion were
three-toed. Furthermore the data suggests that they possessed slightly longer,
but more massive, limbs than the Pikermi Hipparion (see also p. 363).

Table 13 indicates the dimensions of the available third metapodials
from Bed II, compared with two specimens from Pikermi.

It appears that the African hipparionids have hardly modified their
locomotor apparatus since the Pliocene.

THE ECOLOGICAL ADAPTATIONS OF Hipparion

Considerable plasticity exists in biological organisms. Factors such as
climate (with such variations as in temperature and humidity) considerably
affect water supply, food, animal and plant associations and the actual nature
of the environment, e.g. savanna, sand dunes, forests. The range of an intra-

ti ie

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 353

Proximal Distal

Metacarpal III Total Proximal breadth Distal breadth

length A-P (max.) A-P (max.)
1957. SHK II 935* .. 217 36 46 33 42
F. 3457 2: 206 37 44 34 245
1955. BK II 45 - 229 37 >45 36 48
MEAN . 215 37 45 34 45
Pikermi UC63402** 206 31 40 29 38

Metatarsal III

1957- SHK II 557* 253 38 48 35 45
1957- SHK II 729/730 260 37 47 36 45
1955. BK II 68 iA 257 40 46 35 45
1953. BK Il . 242 37 45 34. 44
wan7. otk Il 1177 .. 254 — — a3 >42
fgg. DK AL663 .. —_ 41 46 — oe
mons. BK ILig, .. — a — 38 48
neat, © 1 F 797 ae — — — 32 43

MEAN bad 253 38°6 46°4 36 44°7
Pikermi UC63415** .. 241 37 45 33 40

* These appear to belong to one individual.
** University of California, Berkeley (Dept. of Paleontology).
+ There is a unilateral pathological growth at the distal end.

TABLE 13. Dimensions (mm.) of third metapodials of Hipparion from Olduvai Bed II.
All the specimens, except the two labelled ‘Pikermi’, are from Olduvai.

species variation may be considerable and will allow extensive pliability in
adaptation to changing environmental conditions. Such modifications may not
change the genetic background or consequently, the taxonomic status. In
comparing different anatomical patterns, this concept must be borne in mind.

On the other hand, it is also obvious that the environment may favour,
either the greater adaptive potential of some individuals in a particular popula-
tion or mutations, so that selective forces may operate to promote different
races and eventually a new species. Therefore, in a survey of a biological
group which extends over a vast continent, it is essential to appreciate the
plasticity of the organism.

Such a study has been successfully developed for the family Equidae, and
the hipparionids in particular, because of the vast amount of available fossil
material, both in America and in Eurasia. Following other scholars, Gromova
(1952), in her revision of the genus Hipparion, has emphasized its variability
and evolutionary trends in relation to ecological factors.

The aims of this section are
(a) to summarize the main conclusions about the ecological and functional

significance of Hipparion characteristics;

(b) to summarize the available information concerning the African biotopes
occupied by Hipparion from Upper Miocene to Pleistocene times; and

(c) to draw possible conclusions about the migrations and evolution of
African Hipparion under the influence of these bionomical circumstances,

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

and thereby to contribute to the appreciation of the taxonomic status of
the African groups.

ECOLOGICAL AND FUNCTIONAL SIGNIFICANCE OF HIPPARION FEATURES

The overall dimensions of the Hipparion skeleton do not allow definite conclu-
sions about the biotope. It is known that the smallest races of extant Equidae
are to be found in the driest areas characterized by drier food and, in the
northern part of their dispersion, in areas of poor economic conditions. It is
probable that similar factors have influenced the Hipparion-associated fauna,
and that their general dimensions illustrate in some way the nature of the
biotope. However, larger and smaller forms (? species) have repeatedly been
recovered from the same deposit so that it appears that they co-existed in the
same climate, but they may have occupied, within the same area, slightly
different ecological niches. On the other hand, this is not a general rule because
exceptions have been documented. For example, at Pavlodar (on the right
bank of the Yrtych, Moldavia, U.S.S.R.) where it was possible to make inde-
pendent studies of the nature of the biotope at different ecological stations, a
species characterized by longer and more slender extremities and reduced
lateral digits, viz. H. longipes, lived in a drier habitat than the shorter-limbed
H. elegans did. Therefore, it is not possible to make any direct inferences about
the biotope from a mere consideration of the overall dimensions of the animal.
Various factors are probably involved which cannot be adequately isolated.

Relative dimensions of cheek teeth and incisors seem to be largely influenced by
the quality of the food: a drier, more steppe-type grass usually develops larger
teeth. However, the incisors and the cheek teeth may show differing reactions
and adaptability to this ecological feature. This has been demonstrated by
comparing species presenting large cheek teeth with species having small
cheek teeth, e.g. H. gratum and H. longipes, which show comparable incisors.
The relative size of the teeth in proportion to the overall dimensions of the
animal must also be considered, a smaller animal normally having relatively
larger teeth.

The development of the preorbital fossa, which has become an important
taxonomic consideration, is in the present state of our knowledge not related
to any ecological feature. Animals with large and with small preorbital fossae
are found together in the same xerophytic (H. proboscideum and H. matthew, at
Samos) and moist habitats (H. moldavicum at Taraklia, or H. theobaldi in the
Siwaliks). Preorbital fossae may even be missing (H. platygenys at Taraklia)
without any apparent ecological reason.

The isolated protocone, completely detached from the protoloph and the
protoconule right down to the base of the crown, is a constant feature of the
hipparionids. It obviously weakens the structure of the tooth. This has to be
compensated for, especially in the case of a hard and dry grass diet, by a
greater development of the cement in the anterior and the posterior valleys so
as to bind and strengthen these individual elements (Stirton, 1931).

ee ee eee Ce ee ee oe ee

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 355

Two elements are involved in shaping the protocone: antero-posterior
length and transverse breadth. It is known that, relative to the total dimension
of the tooth, the A—P length of the protocone is more constant than its transverse
breadth. Therefore, a protocone may look elongated, but this is only a reflection
of its narrowness. Thus it is advisable to express the functioning structure of
the protocone by means of two indices: 3
(a) the protocone length is expressed relative to the general mesiodistal

measurement of the tooth, i.e. the ‘length index’.

(6) ‘The so-called ‘shape index’ expresses the length of the protocone relative

to its breadth (table 14; figs. 12, 193).

These indices highlight the differences in adaptation between the African
and Eurasiatic Hipparion. |

The two indices for the African forms are compared to those for 7 Eurasiatic
Hipparion species. Available measurements of the total dental series have been
utilized. It may be noted from figure 12 that the length index of the African
Hipparion is constantly higher for each tooth than that of the corresponding
tooth of any of the Eurasiatic species. However, in both groups the length
index tends to increase in a mesiodistal direction. On the contrary, the shape
index (fig. 13) is constantly lower for the African forms, decreasing for both
African and Eurasiatic groups in a mesiodistal direction. Thus it may be
concluded from the two indices that the protocone of the African Hipparion is
both relatively longer (elongated) and narrower than in the Eurasiatic forms.

The shape of the protocone varies with progressive wear, the section
tending to become more oval and less elongated. But independently of these
modifications, for particular locations in the dental series at comparable
levels of attrition, there seems to be also an adaptive elongation or broadening
of the protocone which is linked with the type of food it has to deal with. It
is suggested that the efficiency of the protocone’s function in trituration lies
in the action of its buccal and lingual enamel crests, developed perpendicularly
to the lateral chewing movements of the mandible. It is obvious that the
efficiency is maximal for a narrow and elongated protocone, for here the
crests are at one and the same time the longest and the most perpendicular.
Thus they would favour a dry and harder bunch grass. This suggestion is
strengthened by the contemporaneity of the lengthening of the protocone and
the development of a typical xerophytic vegetation in America.

Enamel plications

It has been successfully demonstrated that the complexity of the enamel
pattern on the occlusal surface of Hipparion molars has developed synchronously
with the drier environment in Upper Miocene and Pliocene times. Like the
elongated protocone, the numerous enamel plications are oriented perpendicu-
larly to the movement of the jaws, and are increasing the triturating power of
the teeth for coping with harder grass. Although there seems to be a parallel
increase in the degree of plications with increasing toughness of food, there

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 357

H. africanum ©

H. sitifense :
Hypsohipparion :
Langebaanweg |

. elegans

H. moldavicum
. platygenys :
. periafricanum:
. gromovae
. concudense :
. koenigswaldi :

Fic. 12. Protocone length index in the various teeth of African and non-African Hipparion species.

3 58 ANNALS OF THE SOUTH AFRICAN MUSEUM

e@ H. africanum

m H. sitifense
Hypsohipparion
Langebaanweg

. elegans

. moldavicum

. platygenys

. periafricanum
. gromovae

. concudense

- koenigswaldi

dpoepono px

LB

Fic. 13. Protocone shape index in the various teeth of African and non-African Hipparion species.

Sa

a.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 359

must be a limit to the former. The quantity of available enamel is fairly constant
and probably limited, so that an increase in the complex pattern of the enamel
will result in a thinning out of the plications. Eventually this would produce
too weak a support for the triturating pressure required for hard grass. When
this point is reached, the plications again reduce in number and the protocone
lengthens. as a compensatory mechanism.

Therefore, it is not always possible to ascertain from the small number of
plications whether they represent a primitive condition corresponding to a
humid environment, or whether they are secondary, resulting from a reduction
under the influence of a hard grass food. Then other factors must be considered,
e.g. the shape of the protocone, to solve the problem.

A directly comparable situation is encountered in the development of
proboscidean teeth. Here an increase in the available grinding surface of the
enamel ridges (lamellae) could represent the increasing plications in the teeth
of the Equidae, especially as they are also oriented perpendicular to the grinding
movement. This parallelism is also demonstrated in the hypsodonty of these
two groups.

Hypsodonty

In the evolution of the Equidae most lines show a constant increase in
the crown height of the cheek teeth. However, from Merychippus, in Miocene
times, the progressive hypsodonty has advanced along two phyletic lines
(Hipparion and Pliohippus) at a faster tempo, again under the influence of a
general increase in aridity. The same observation has been made among
Bovidae. It seems obvious that it is once more an adaptive response of the
organism to the xerophytization of the diet.

The dietary changes involved in the evolution of Cenozoic hypsodont
horses probably corresponded to a shift from browsing on softer herbaceous
plants to grazing on harsh, siliceous grasses, which acted as abrasive agents on
the teeth. These changes are, in turn, a consequence of an increasing aridity
as has been clearly demonstrated for the Great Plains of America (Stirton, 1947).
The association of widespread sandy deposits and of bunch grass of the Stepidium
type, with the increasing aridity of this area is actually known to be contempo-
rary with the development of hypsodonty from Lower Miocene times onward.
This increased height of the crown has played an important role as a protective
modification in, the dentition. It is supposed, although impossible to prove
from the fossil record, that a mutation or other factors, favoured an increase
in activity of the odontoblasts and ameloblasts to lay down more dentine and
enamel when the tooth was still in the formative stage. The root closure (fusion)
may have been considerably retarded as well. Furthermore, it may be suggested
that dry climate has influenced the teeth by necessitating some protective
mechanism not only because of the harsh conditions of the grass, but also
because of the sand mixed with it as a foreign substance. Stirton (1947) states
that in the Great Plains area, a close inspection reveals much sand adhering

360 ANNALS OF THE SOUTH AFRICAN MUSEUM

to grass or lodged in between blades and stems, where it is blown by wind or
splashed after thunder showers.

The double knot in the lower cheek teeth of the hipparionids displays three
typical features:

(a) a primitive ‘stenonis’ type, inherited from Merychippus and generally —

retained throughout the evolutionary sequence among American species;
(b) a more progressive, typical ‘Hipparion’ type has been developed by most

European forms; and
(c) the ‘caballus’ type which was lately acquired, particularly in African

representatives.

It has been suggested that the double knot of the lower teeth was shaped
and typified under the influence of the protocone structure in the upper teeth,
with which it is functionally linked in occlusion. There is good reason for
developing a long, narrow metastylid parallel to an elongated, elliptical
protocone, as is found in most African Hipparion. Here, the shape of the metasty-
lid in particular, and consequently of the whole knot, conveys useful information
about diet and climate. However, the correlation is probably not a simple

one, because American Hipparion display an elongated protocone and a rounded -

‘stenonis’ type of metastylid simultaneously.

Total tooth structure

Upper teeth are rather compact. The outer enamel forms a fairly con-
tinuous and parallelipipedic structure, only grooved slightly by a small hypo-
glyph. The only weak structure which can be broken away from the tooth
under excessive strain is the protocone, separated from the rest of the tooth by
the internal depression. In the Langebaanweg material, this plane of fracture
is noted in Lg35, Lo42, Lo45, Lo56, S.A.M.11722 and S.A.M.11724 (pls. 1, 7,
8, 9g). It has been noted above that, with increasing dryness of the food, this
weakness is largely corrected by the development of a narrow protocone closely
adherent to the protoloph.

Lower teeth, on the other hand, are built on a very different pattern.
The many conids and stylids are much more individualized: they are contig-
uous but are separated from each other by rather deep depressions right down
to the base of the crown. These intervals are filled with cement, but it is softer
than the enamel and does not always supply sufficient compensatory protection,
as can be seen in many fossil teeth where the cement has disappeared. In drier
environments, the hard food influences this structure of the lower teeth,
demanding additional strengthening of the enamel-dentine pillars which are
embedded in their cement coating to prevent both fracturing and excessive
wear of the tooth. Support can be provided in two ways, either

(i) by reducing the depth of the internal and external depressions separating

protoconid from hypoconid, metaconid from metastylid, and metastylid ©

from entoconid, or
(ii) by additional stylids.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 361

Reduction of the depth of the depressions is commonly observed. As has
been stated previously, this occurs on the lingual surface by the development
of a ‘caballus’ type of double knot, while on the buccal surface it is secured by
a broader connection between the two halves of the ectolophid.

The alternate solution lies in the construction of additional pillars:
ectostylid and ptychostylid in particular, just in front of the external depression,
and protostylid and hypostylid on the antero- and postero-external corners.
Entostylids are occasionally developed in the postero-internal depression. In
one particular case (a ‘Stylohipparion’ Gar. Fluss no. 200, figured by Dietrich,
1942, pl. XIII, fig. 95), a strong additional pillar is seen between metastylid
and metaconid.

The higher frequency of these additional pillars in milk teeth seems to have
been necessitated by the lowness of the crown of these teeth and their softer
construction. This is the obvious way to prevent too rapid attrition which
cannot be compensated as there is no continuous growth of the tooth.

As suggested previously, enamel plications, shape and size of protocone,
structure of the double knot, increased hypsodonty and additional pillars are
many different methods of meeting the mechanical requirements of a tougher
diet. They do not all necessarily develop together at a parallel tempo. Some
of them are functionally linked but other elements develop compensatory roles
so that all need not be present at one time. Therefore, no single element can
give reliable information about climate, biotope and the quality of food, but an
assessment of the whole picture and of the reciprocal values of the different
features must be considered.

The extremities

Duerst (1926) has successfully demonstrated how a humid climate and a
soft grass develop a heavier type of equid with broader extremities (eurysome
type), while a leptosome type of a smaller, more slender animal with more
gracile extremities tends to be built in drier biotopes characterized by a
xerophyte vegetation. This seems to apply especially to metapodials which
become relatively longer.

The progressive reduction in size and the disappearance of the lateral
digits in equids, with the concentration of the whole body weight on the third
metapodial, has always been interpreted as an adaptation to increasing speed
in steppe surroundings. It has been repeatedly suggested that the high tempo of
evolution of Pliohippus towards monodactyly is an expression of its drier habi-
tats, while the lateral digits of African Hipparion are kept more or less functional
right into the Pleistocene, probably because of a more humid environment.

THE UPPER MIOCENE AND PLEISTOCENE BIOTOPES OCCUPIED BY AFRICAN
HIPPARIONIDS

Our knowledge concerning past climates and biotopical conditions in
those areas where African hipparionids have been recovered is still inadequate.

7 .

362 ANNALS OF THE SOUTH AFRICAN MUSEUM

The fragmentary information spans over a very long period of time (probably
10-12 million years) and is derived from a large continent where the climates
have certainly been variable. Consequently it is not always possible to draw
accurate conclusions from the collected data, nor is it possible often to correlate
data from one site with those of another.

Furthermore, this information is deduced from fossil assemblages of which
some are listed on the basis of debatable fragments, while others are wrongly
identified. For example, at Olduvai Gorge, recent and more extensive collections
will permit a better analysis.

Even where there is sufficient and accurately identified material, it
should be remembered that a fossil specimen is not necessarily found in the
typical habitat of the living form. It is well known that in periods of drought
or volcanic eruptions (which have played such an important role in the Ceno-
zoic sedimentation of East Africa, from Omo to Eyasi), animals have concen-
trated around swamps, temporary pools and tuffs, possibly fairly distant from
their normal biotope.

This biotope is often less restricted than one imagines, and the adaptation
of some organisms is quite fantastic. So, it seems futile to draw definite conclu-
sions from the presence of one or even a few animals. Real associations must
be considered to avoid misinterpretation of the presence of a species at a
particular site.

(a) In the Upper Miocene, the Oued el Hammam faunal assemblage
contained Palaeotragus, Samotherium, Damalavus, Gazella, Dicerorhinus, Hippopota-
mus and Hyaena. With slight generic differences, it corresponds very closely to
the fauna of the East African plateau of today, and indicates the typical
savanna with mimosa biotope of the tropics. According to Arambourg (1959),
the presence of Macaca and Cephalophus in the contemporaneous deposit at
Marceau suggests the proximity of forests. This is supported by the location
of the deposit in a higher and already tilted portion of the Tellian Atlas, and
by the lignite abundant in the deposit.

(6) The Lower and Middle Pleistocene faunal assemblages of the East
African Hipparion-bearing sites clearly indicate the same type of steppe or
plain and highland (mimosa) savanna environment, very similar to that of
the present day, ‘with a marked bias toward somewhat moister conditions in
most cases’ (Cooke, 1963). The main feature at Omo, Kaiso, Kanam, Olduvai
I-IV and Eyasi is undoubtedly that of an open savanna. No true forest associa-
tion is to be found in these areas, although forest strips and moist woodland
occurred occasionally, as is indicated, for example, by the presence of Wesotragus
moschatus. The cyclical and seasonal variation of the rainfall has provided
more humid conditions at times, and consequently the development of swamps
and (or) lakes in the lowlands, with very luxurious vegetation, are responsible
for the abundant Suidae, Bovidae, Giraffidae, Elephantidae and Equidae. A
swampy environment is explicitly suggested at Olduvai by Aonyx, and at
Omo by Kobus. A lake shore or grassland adjacent to water is evident both at

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 363

Omo and Olduvai (e.g. Hippopotamus, Omochoerus) where this is also reflected
by the nature of the deposit.

The presence of Oryx and Beatragus hunteri would suggest arid conditions
prevailing in the locality.

Although most of the fauna are suggestive of plains or a plateau, Tragelaphus
nakuae and Dzinopithecus brumpti indicate that a mountainous biotope was
probably part of the picture at Omo.

(c) In South Africa, forest and desert associations are similarly absent,
and the prevailing feature remains that of a steppe or a savanna fauna. The
Vaal River Gravels suggest a fauna of thornbush and grassland. The cave
breccia—probably less representative of a true biocoenose—contains both
open plains and dense bush-dwelling forms. It is believed that the rainfall
varied considerably during the Pleistocene, with oscillations from 20% to 40%
annually (Brain, 1958). The fossil forms represented there would have tolerated
such changes. In any case, even in the driest conditions (Taung) well-watered
bush valleys could be located at not great distances.

In conclusion, it may be stated in broad terms, that African Upper
Miocene and Pleistocene Hipparion sites seem to have been areas of (dry to)
moist savanna and grass steppe, or of open woodland with mixed savanna. It
is not possible to define with any more precision the environment proper to
Mipparion in this general biotope which extended over half of Africa for several
million years. It is probable that local circumstances and features have directed
the relationships and balance of the various species in particular areas: but as
a whole, and in spite of replacement of species, the African fauna has maintained
an essentially constant composition throughout late Cenozoic times, which
indicates that the broad ecological pattern did not vary greatly from the
Upper Miocene onwards. The main ecological feature, viz., the savanna, may
at times have been extensively altered by moister climates producing swampy
conditions and even patches of forest. Then later, it would be altered again
by a rather low rainfall. Nevertheless, drastic changes probably never occurred.
There seems little evidence of a true forest fauna or of conditions too dry to
sustain a normal steppe biocoenose (Cooke, 1963).

EFFECTS OF BIONOMICAL CONDITIONS ON THE EVOLUTION AND THE MIGRATION OF
AFRICAN HIPPARIONIDS

It has been generally accepted that the three-toed hypsodont Hipparion
were typical steppe forms. The savanna biotope is commonly emphasized as an
important equid environment of the late Tertiary in the Great Basin of the
United States, and the vast reduction of this habitat is strongly suggested as a
Suitable explanation for the extinction of Hipparion by the end of the Hemp-
hillian (Shotwell, 1961). The life span of Hipparion in North America, namely,
Clarendonian-Hemphillian, corresponds rather precisely to the extension of
the savanna or steppe biotopes in this area. The recent observations of Sondaar
(1961) strengthen this view: at Nombrevilla, in Spain, where Decennatherium,

364. ANNALS OF THE SOUTH AFRICAN MUSEUM

Aceratherium and Mastodon were recovered together with Hipparion, there is —
good evidence for a steppe environment. However, it has become clear that

fipparion is not restricted to such narrow ecological conditions. Thenius (1950)

interprets H. gracile as a forest-dwelling form, and Kurtén (1952), on the
basis of the Chinese ‘Pontian’ fauna, enforced the necessity of a reconsideration —

of this hypothesis. Because of the almost identical frequency (13°) of Hipparion
in both the gaudryi and dorcadoides faunas, characteristic, respectively, of the
‘forested’ or ‘southern’, and of the ‘steppe’ or ‘northern’ provinces (the classical
distinction of Schlosser, 1903), it was suggested that the Chinese Hipparion
thrived in various surroundings, or at least that the genus as such was not
exclusively a steppe form. Furthermore, it is not possible to make a direct
correlation between the geographical distribution of a particular species and
its biotope. Of the eleven different species recorded by Sefve (1927), some are
confined to one locality and others turn up in widely different areas; in some
instances, up to four different species have been recorded from the same fossil
pocket.

The adaptability of hipparionids explains their extraordinary diffusion

both in North America and in the ancient world, from Upper Miocene times ©
onwards. Ecological conditions influenced structural features in different and —

varying ways, often difficult to explain. A similar geographical distribution
and even the simultaneous presence of different fossil forms in one particular
pocket are compatible with slightly different cles niches or habitats in
a small vicinity.

Under favourable conditions, observations have often been made of the
association of more massive animals (with larger skull, shorter limbs, longer
lateral digits and strongly plicated enamel) with open forested areas, character-
ized by soft and wet ground and tender vegetation. Lighter forms (with slender
and elongated limbs, reduced lateral digits and simpler enamel pattern,
elongated and narrow protocone and increasing hypsodonty) are found in a
more xerophytic environment. Outside Africa, H. primigenium, theobaldt, crassum
are representatives of the former group; H. elegans, longipes, moldavicum,
proboscideum, matthew are typical of the latter.

In general, the various characteristics observed in the African Hipparion
are Clearly indicative of animals adapted to a rather xerophytic steppe environ-
ment, although this never seems to have been very severe, and was certainly
not constant everywhere. It has been stated above that nowhere in Africa
are Hipparion-bearing sites located in true desert areas, and none of the forms
recovered shows the extreme leptosome type displayed by some North American
forms, e.g. H. whitneyi.

Hipparion africanum, with its large skull, short and massive limbs with
lateral digits well developed, and rather less hypsodont teeth, is probably,
from the inadequate data available, the least adapted to a steppe environment.
However, the enamel plications and the narrow protocone show an orientation
in that direction. It is difficult to assess whether this moderate expression of

> — 7 2 <u * gg
On ee

’

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 365

the steppe features corresponds to the precocity (in time) of H. africanum or
reflects a humid and even partially forested environment.

The moderate transverse flattening of the protocone, the few and simple
plications, the absence of additional stylids, the reduced hypsodonty and the
small dimensions of the teeth of Hipparion sitifense, as well as the development
of the lateral digits, are to be interpreted in the same way, although the absence
of skeletal remains does not permit evaluation of the degree of massiveness.

The postcranial skeleton of the African Pleistocene forms is practically
unknown. However, quite a few metapodials have been recovered in the
Olduvai Gorge (Tanganyika). They are not particularly slender, and the
lateral digits are still well developed, but the absolute length of the third
metapodial is quite remarkable (metacarpal: 217-223 mm.; metatarsal:
242-260 mm.), being 5-9% longer than the corresponding elements from
Pikermi (table 13).

The typical features of the dentition are strongly marked hypsodonty, the
fairly complicated enamel pattern of the ‘mark’ walls, and the elongation of
the protocone, which may become very flattened. The extreme flattening
corresponds to a spectacular development of the additional stylids. It seems
clear that these features must be interpreted as a positive pattern of adaptation
to a steppe environment throughout Africa. However, the absence of any
extreme slenderness of the metapodials and the complicated enamel pattern
seem to strengthen the conclusion drawn from the faunal assemblages, viz.
that the xerophytic environments of the African Hipparion were never very
severe.

In the absence of sufficient cranial and postcranial remains, and because
of the vicarious role played by the different tooth structures in their adaptation,
it is unnecessary to comment any further on the special features of every single
group so as to define more precisely the particular environment of each.
Discussion is further limited by the fact that climate and biotope seem to have
been rather uniform over a great part of Africa, and have certainly not under-
gone drastic changes. Throughout the continent, African Hipparion reflect in
their known morphology the vastly extended steppe or savanna environment
with an absence of much aridity.

UNPUBLISHED MATERIAL FROM THE VAAL RIVER DEPOSITS, SOUTH AFRICA

Sydney-on-Vaal and Pniel
The exact location of these two specimens is not known (fig. 6). Cooke
(1949) describes other material from these sites which are in the northern
Cape Province.
Material:
MMK 431 —left M3
MMK 5225—left M, or M,
These specimens were diagnosed by the authors as belonging to Hipparion

366 ANNALS OF THE SOUTH AFRICAN MUSEUM

in the collection of equid material which was kindly sent on loan by the Director
of the McGregor Memorial Museum, Kimberley, South Africa.

MME 431 (pl. 1)

This is a left M? with fairly high crown, in an early stage of wear.

Half of the ectoloph: fairly deep, flattened arc, being angulated at the
parastyle and slightly angulated at the mesostyle. .

The parastyle: the lateral angle is chipped away at the occlusal surface
but near the base it is seen to be prominent. It is rounded and projecting, and
is separated by a fairly deep groove from the ridged anterior angle. The surface
is very oblique, almost in the same plane as the anterior surface.

The mesostyle is partly broken. It is fairly large and rounded, and it has
a deep groove.

The metastyle tends to be broad with a ridged lateral angle, a smaller
ridged posterior angle and a slightly concave surface, which is oblique. |

The protocone is isolated. It is very long, being about half the length of —

the tooth. Anteriorly and posteriorly, it is sharply angulated. Its inner surface
is flattened, and the lateral, outer surface is wavy, presenting an elongated __

cigar shape. |
The protoconule is short and has a flattened arc. Posteriorly it becomes
markedly angulated and turns laterally to join the base of the pli caballin.

The hypocone is elongated and flattened. Its two sides are almost touching. —
The hypoglyph is deep and circular, demarcating an angulated hypostyle.
The hypostyle is joined to the metastyle by a concave arc of enamel.
The pli caballin is long, with unequal sides almost touching the protocone. —

The prefossette: the anterior wall has a small plication and a very deep,
narrow pli protoloph which tends to separate the medial part of the prefossette.
The posterior wall has shallow plications and a deep narrow pli protoconule
which tends to separate the medial part of the fossette.

The postfossette: the anterior wall has an irregular shape, with one plication
and a very deep and narrow pli postfossette, which tends to separate off a
flattened portion of the medial part of the postfossette. The pli hypostyle is
duplicated. It is fairly deep, tending to isolate the medial part of the postfossette.
The anterolateral angle of the postfossette tends to form a rounded, almost __
isolated pillar. |

MME 5225 (pl. 1)

A left M, or M,, probably M,.

The tooth is partly fragmented, especially on its inner aspect.

The protoconid is partly broken mesially. It appears that the other wall is
flattened. The protoconid is shorter than the hypoconid. The metaflexid is
elongated, flattened and biconcave with rounded anterior and posterior ends.

The metaconid is broken away. Its outer wall appears to be a flattened arc.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 367

The parastylid is partly broken but its posterior wall turns sharply inwards
and almost touches the outer wall of the metaconid.

The metastylid: the outer wall is flattened in a mesiodistal direction. The
entoflexid is elongated, flattened and anvil-shaped.

The entoconid is rounded and pillar-like, being almost completely separated.

The hypoprotonocid groove is deep and wide, being rounded internally. It
has a narrow mouth laterally and a bulge posteriorly.

The ptychostylid is finger-like, projecting anterolaterally. The sides are
almost parallel and it has a narrow base.

The ectostylid groove is a marked indentation, broad and shallow.

The hypoconid has a flattened, elongated outer surface with posterior parts
tapering sharply inwards.

The ectostylid is an isolated triangular pillar, the outer wall being chipped
at the occlusal surface, but it can be seen to be flattened near the root. The
inner wall is flattened and meets the outer wall at a sharper angle. The two
walls are separated anteriorly by a somewhat irregularly concave base. The
inner wall of the enamel joins the metaconid.

The metastylid forms a distinctive broad arc near the base of the tooth where
it is intact.

MMK 5225 MMkK 431

A-P length .. ap «C8 235, A-P length 22% 3
Breadth — Breadth .. Qi
Max. A-P length Seman 8:1 Breadth/A—P 94°7
Max. A-P length hypoconid..~ 12 Protocone length A-P Pet
Max. A-P length metastylid. . 6-2 Prefossette A—P length 10°8
Max. A-P length entoconid .. 4°6 Prefossette breadth : 8°3
Max. A-P length entoflexid.. 11°8 Postfossette A—P length . 10°8
Max. A-P length metaflexid .. 6-8 Postfossette breadth se 7°0
Breadth hypoconid .. $4 5°6 Protocone A—P/total A—P ieee 56:2
Breadth entoconid .. gs 4°1 Crown height... ay -« 62,70
Crown height .. a .. Ca 40

TABLE 15. Dimensions (mm.) of MMK 431 and MMK 5225 from the Vaal River Deposits.

Taxonomic status

The above two teeth are referred to Hipparion Oe Aare dake libycum (see
p. 387-92 for diagnosis).

MATERIAL FROM LANGEBAANWEG, CAPE PROVINCE, SOUTH AFRICA

DESCRIPTION OF HIPPARION TEETH

Milk dentition

Material:

S.A.M. 1 eee DM?
S.A.M. 11718—right DM?

368 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAAD ryan m(pl, re)

A right DM? in medium stage of wear with a thick layer of cement
particularly on the inner side.

Half of the ectoloph: it is shallow and has a slight arc. It is sharply angulated
at the mesostyle and slopes up into the parastyle.

Parastyle: the lateral angle is chipped but it is seen to project markedly
in a lateral direction near the base of the tooth. The surface is oblique and not
quite in the same plane as the anterior surface of the tooth. There is a marked
groove. The anterior angle is rounded and forms a ridge.

The mesostyle is stumpy, projecting, and flanging at the surface. A fairly
marked overlap exists anteriorly. The surface is rounded, not grooved. The
sides tend to be parallel.

The metastyle is slight and rounded.

The posterior ectoloph has a vertical bulge at the centre of its surface.

The protocone is oval in a mesiodistal direction, with the medial side slightly
more flattened than the other side. It is distinctly separated from the other
cones by a thick layer of cement.

The protoconule is an elongated, flattened arc with a fringed effect.

The protoglyph is deep and wide with an irregular base.

The hypocone is elongated, with a rounded medial side and a flattened
lateral side, tending to come to a point posteriorly. Anteriorly it is almost
completely isolated by a very deep indentation of the hypoglyph. Opposite,
there is an indentation tending to separate it from the metaconule. The
metaconule also has a fringed appearance.

The hypoglyph has a broad mouth. Lateral to the mouth of the hypoglyph,
the enamel shows a V-shaped indentation.

The pli caballin is long, narrow and arched, tending to touch the protocone.
At the base of the pli anteriorly, there is a trace of a duplication of the pli.

The prefossette: the anterior wall shows two deep plications and a deeper,
narrow pli protoloph which tends to isolate the medial part of the prefossette.
The posterior wall of the prefossette shows numerous plications which are
markedly complex. One of the plications adjacent to the pli protoconule is
large and has become isolated, forming the typical island lateral to the base of
the pli caballin, and wedged between the medial portion of the pre- and
postfossette. The pli protoconule is deep and narrow, almost touching the
medial wall of the paracone. The pli prefossette is deep and complicated with
a bifid base, tending to isolate the posterolateral angle of the prefossette.

The postfossette: the anterior wall has numerous plications that are not as
complicated as the posterior wall of the prefossette. The pli postfossette is
deep and narrow, tending to isolate the medial part of the postfossette. The
posterior wall of the postfossette shows numerous plications, the most lateral
one being rather deep. The pli hypostyle is deep and angulated, tending to
isolate the medial wall of the postfossette.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 369

S.A.M.11718 (pl. 2)

This is a right DM? of the same individual as 11717.

The anterior portion of the tooth is broken away but the remainder of the
tooth shows the same features as 11717 with the following differences:

(a) Protocone is more elongated and more oval.

(6) Hypocone is more oval.

(c) Pli caballin is bifid medially.

(d) Pli protoconule is more complicated, and is duplicated.

Dimensions SAM 10717 Hypsohipparion
(DM®) (DP*—DP* ..)
A-P rt re 31°9 28-34
Transv. breadth .. 26-2 23-26
ment .. os QI 28-34

(medium wear) (early wear)

TABLE 16. Dimensions (mm.) of Langebaanweg milk
molar compared with ‘Hypsohipparion’.

The features which are observable on the two Langebaanweg milk molars
are identical to the description given by Dietrich (1942) for the Hypsohipparion
material (19 individual teeth and 2 maxillary fragments). The isolation of the
hypocone and the shape of the protocone of Dietrich’s (1942) specimen illus-
trated in his figure 162 is typical of the Langebaanweg milk teeth. The
dimensions of the latter fall into the range of Dietrich’s specimens (table 15).

Lower permanent dentition
Material:

P,: Left: L947, L1465p (pl. 6).
Right: Logg (pl. 5).

Peers: Lett: L937, L943 (pl..6).
Right: Lg41, L946, Lg56, Li451a, L1465a (pls. 5, 6,

8,19)

M, or M,: Left: Lg44, Lg52, Lo59, L1448 (pls. 6, 8).
Right: L948, Lg54 (pl. 7).

M;: Left: L938, Lo49, L1465B (pls. 8, 9).
Right: L1465F (pl. 2).

One specimen, L1465c, a M, or Mg, is excluded because of its size and
its fragmented appearance which does not permit recognition of any typical
hipparionid features.

The following specimens seem to belong to single individuals:

1. L937, Logg.
2. Lo41, L943, L944, L948.
3. L954, L957.
4. L938, Lg59.

370 ANNALS OF THE SOUTH AFRICAN MUSEUM

Classification of specimens according to wear:

A. Unworn or slight wear: Lo49.

B. Early to medium wear: L947, L1465p, Lo39, L937, L943, Lg41, Lo46,
Li451a, L1465a, L944, Lo52, Lo59, L1448, Lo48, Lg38, L14658,
Li465F.

C. Late wear: L954, L957.

General description

All the Langebaanweg specimens obviously belong to one group, viz.
ENipparion (Hipparion) albertense baardi subsp. nov. The dimensions are presented
in table 17. The following features are distinctive:

1. Entoconid: Usually in P, it is more or less flattened in a bucco-lingual
direction. It tends to be more rounded or quadrangular in shape in P,, P,,
M, and M,. However, in early wear a slight indentation of the enamel produces ©
a ridged effect on its lingual surface mesially.

2. Metaconid: It has a slight tendency to a bilobed formation due to a —
fairly marked indentation on the mesiolingual aspect, which is only recognizable ;
in avery early stage of wear.

3. Double knot: This is of the ‘caballus’ type which is usually found in
African Hipparion. |

4. Metastylid: It has a triangular shape with the right angle found distally ©
on the lingual side of the entoflexid.

5. Metaflexid: It constantly shows unique invagination on both its —
anterior and its posterior walls. |

6. Entoflexid: ‘The buccal wall shows varying degrees of waviness —
(plications).

7. Internal depression: This valley, lying between the metaconid and ©
metastylid, is very broad, shallow and irregular in P,, while in P,—P, it is ©
broad but slightly deeper giving it a U-shaped appearance. |

8. Ectostylid: This is constantly absent, but on Lg52 there is a small
elongated thickening of the enamel at the crown-root junction. However this —
‘bud’ does not arise from the cingulum but extends from the posterior root onto —
the crown just above the junction (pl. 8).

9. Ptychostylid: Present on P,, and occasionally it is very well-developed. |
In P,—P, there are irregular indentations on the bucco-distal aspect of the | :
protoconid, i.e. in the valley (external depression) opposite the ptychostylid. |
Inconstantly, slight indentations are seen on the wall of the hypoconid in the
depression. | |

In M,—M, the ptychostylid and the indentations are not constantly | :
present. In M, these features are both absent. |

10. Protostylid: It may appear as a laminated ridge or as an isolated |
pillar. Usually, when present, it extends along the whole height of the crown. _

373

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372 ANNALS OF THE SOUTH AFRICAN MUSEUM

It can always be observed as a prominent ridge along the mesio-buccal border
of the tooth. It is constantly present on P,—P,, and always present on M,—M,.

11. Entostylid: One entostylid has been observed (Lg46) in the entoflexid
on the bucco-mesial aspect of the entoconid. It presents as a distinct flattened
enamel island (pl. 8).

12. Talonid: Bilobed.

Upper permanent dentition
Material:

PF or P*: Left: L934, L936, Lo42, L1463, 114678.) ee ee
(ols; To, 5, 7):
Right: Lo55, L1467c, S.A.M.11722, S.A.M.11724
(pls. I, 3, 7> Q).
M! or M?: = Left: Lo45, L953, Lo58, L1467B, L1467p, L1467Hn,
».A.M.11719 (pls: 3,4, OF
Right: Lg40/950, L951, L956, L1459 (pls. 1, 2, 4).
M*: Right: L935 (pl. 9).
? premolar: Left: S.A.M.11723 (embedded in plaster for sectioning).
Cheek teeth: ?: L1785 (fragment).

The following specimens probably belong to single individuals:

1. Lo35, S.A.M.11722, S.A.M.11724.
ia a7o) oe inter 7 To:

. L942, L945, Lo56.

. Lg40/Lg950, Lg51.

- L934, L953, L955, L1459.

. L14678, L1467c, L1467p, L1467E.

DM oO B CO ND

A completely unworn tooth, L1467A, recovered from the same site as
L1467B-E and H (viz. Baard’s Quarry), does not present the typical features
of hipparionids. It shows the same type of fossilization as the other L1467
specimens, but, because of its exceptional dimensions (ca. 32 X 28 mm.), it is
provisionally excluded.

The teeth are classified according to their degree of wear as follows:

(a) completely unworn or just erupting: S.A.M.11716.

(b) early wear: L1463, S.A.M.11719.

(c) moderate wear: L935, Lo40/950, Lo42, Lo45, Lo51, Lo56, L1467B-£,
SA Vito) SAME IT eA,

(d) advanced wear: L934, L936, L953, L955, L958, L1459, L1467H.

a

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE

General dimensions (mm.) of P?—M?:

1. A-P length: range of variation: 21-8-30°5

Mean: 26-6

2. ‘Transverse breadth: range of variation: 23°5-30°3

Mean: 26-9

3. Breadth/length index: range of variation: go-7—-111°4

4. Hypsodonty:

Means

All P?—-M?
Unworn teeth
Early wear
Moderate wear

Mean: 99°1

Absolute
crown height
(mm)

49°7

70
63-69:2
51°8

* The ratio between crown height and breadth.

Hypsodonty
Index*

(%)
49°6
36-4
42°3
51°6

373

Height/length
Index

(%)
193
249

233
201

To indicate the relationships between the Langebaanweg dimensions and
those of other African Hipparion, the following data has been extracted from
table 18 and summarized (table 19):

A-P length

Transv. breadth ..

Breadth/length

Height (unworn and early
wear) .. i sie

Hypsodonty index (un-
worn and early wear)

Height/length index (un-
worn and early wear) ..

Hi. africanum AH. sitifense

19-23
18—22°5

23-27
20° 5-26
97°2
48—60
38° 3-41°3

223-240

“f1ypsohipparion’
albertense
(South Serengeti)
25-31
24°5-29°5

95°1

70

249

Langebaanweg

26°6
26°9
99"I
67°4
39°3

238

TABLE 19. Comparison of dimensions (mm.) of Langebaanweg upper teeth with summary of
data of other African Hipparion.

From these figures, it is clear that the Langebaanweg Hipparion has
hypsodont upper teeth, but this hypsodonty has not reached the same degree
as it has in the South Serengeti specimens.

Description of selected individual teeth:

Two specimens, an unworn left premolar (S.A.M.11716) and a left M?
or M? (S.A.M.11719) in early wear, have been selected for detailed

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 375

description. They display the characteristic features of the other Langebaan-
weg specimens. ‘The typical features of the whole group will be outlined
below (see p. 376).
S.A.M.11716 (pl. 5)

A left P4, unworn.

Occlusal surface: Paracone and metacone are V-shaped and they are
curved towards the medial side. The protoconule and metaconule are less
angulated than the lateral cone and tend to curve less towards the medial
side. The paracone and the metacone tend to overhang the fossettes.

Protocone: ‘The cement is split off near the apex of the tooth. It is lozenge-
shaped, completely isolated and the apex tends to curve in a buccal direction.

Shape: The tooth has medium hypsodonty. It has a gentle convexity from
root to apex in a lingual direction. The styles stand out as clear costae, forming
deep hollows between them. One-third of the distance from the occlusal surface,
the hypoglyph becomes increasingly deep so that in the upper half of the tooth
the hypocone becomes well demarcated.

Appearance of the tooth, sectioned just above the middle :

The halves of the ectoloph are deep and arc-shaped with a slight angularity
at the mesostyle.

The parastyle is prominent. Most of the prominence is taken up by the
lateral angle which is rounded. The surface, flattened but with a slight groove,
is markedly oblique and almost in the same plane as the anterior (mesial)
surface of the tooth.

The mesostyle is prominent, and, because its rounded surface is flanged, it
overlaps the buccal depressions on both sides.

The metastyle is very slight (almost negligible), its rounded ridge decreasing
in prominence from the occlusal surface to the root.

The protocone is slightly elongated and oval. Completely separated from the
protoconule.

The protoconule is a flattened arc, angulating fairly sharply toward the pli
caballin.

The hypocone is elongated posteriorly, rather flattened. There is no tendency
to isolation. There is a deep V-shaped hypoglyph, and the hypostyle is
angulated.

The pli caballin is bifid, the posterior process just touching the protocone.

The prefossette: There are plications which are so deep and complex that
the centre of the prefossette is very narrowed and reduced. The anterior wall
is plicated and the antero-lateral angle is knob-shaped, tending to be separated
from the prefossette. The pli protoloph is deep and narrow, touching the very
deep and narrow pli protoconule so that the medial portion of the prefossette
is an almost isolated triangle. The plications on the posterior part of the pre-

376 ANNALS OF THE SOUTH AFRICAN MUSEUM

fossette are very deep and complex. One of the more medial ones tends to
flange out to form an almost isolated irregular pillar just lateral to the base of
the pli caballin and wedged between the most medial part of the prefossette
and postfossette.

The postfossette: ‘The plications on the anterior wall are also deep and
complicated but not as numerous as on the posterior wall of the prefossette.
The pli postfossette is deep and irregular-shaped, almost touching the base of
the pli hypostyle, thus tending to isolate an oval portion of the medial part of
the postfossette. Just lateral to the pli postfossette, the plications tend to isolate
a small rounded pillar.

S.A.M.11719 (pl. 3)

A left M? or M?.
It is slightly higher crowned than S.A.M.11716, and slightly less curved.

Half of the ectoloph: The paracone is deep and quadrangular-shaped,

angulated at the mesostyle and parastyle.

The parastyle is prominent and narrow. The sides tend to be parallel,
slightly widening at the base. ‘

The metastyle is partly broken. It forms a slight ridge.

The protocone is elongated, oval and angulated anteriorly and posteriorly.
It is completely isolated. .

The protoconule forms a broad flattened arc, slightly angulated toward the
pli caballin. 4

The pli caballin tends to be duplicated with the anterior tongue longer and —
almost touching the protocone.

The hypocone is rather short, tending to be separated from the metaconule.
There is a very deep hypoglyph that is very broad-mouthed.

The prefossette: There are a few anterior plications, continuous with a
deep narrow pli protoloph which tends to isolate the medial wall of the fossette.
The pli protoconule is shallow. The plications tend to be absent. However, —
the original plications have become isolated to form an irregular circular —
island (prefossette loop) just lateral to the pli caballin and wedged between ~
medial parts of the pre- and postfossette.

The postfossette: The plications tend to be worn away but it can be seen ~
that they were complex anteriorly. The pli postfossette is very deep and touching ~
the medial wall of the metacone. The pli hypostyle is deep, angulated and —
wide, and it has almost isolated the medial part of the postfossette. |

Typical features of the upper teeth:

1. Protocone: The characteristic isolated protocone of Hipparion is observed —

in all the complete molars mentioned above. On L935, Lo42, L945, Lo56, —
S.A.M.11722 and S.A.M.11724, as well as on L1785, the lingual portion of —
the tooth has been broken away along the pre- and postprotoconal grooves and ~

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 377

does not allow any observations of the protocone. However, there is no reason
to suppose that the protocone in these teeth displayed different features, and
it is as umed that the description proposed for the other teeth applies to these
specimens as well.

The protocone is elongated in a mesiodistal direction, and clearly shows
angulation in those teeth not worn or in early stages of wear. It becomes more
elliptical or oval in the later stages of attrition. Thus the shape index (breadth/
length) varies from 36-4 to 58-9, with an absolute mean of 48-9 for all the
teeth and an average of 48-3 for those teeth which have not reached an advanced
stage of wear (table 14).

The length index ranges from 42-7 to 28-7, with a mean of 35:1 for all
the teeth, of 34-7 for teeth in early wear, and of 35-7 for those in advanced
wear. This smaller range of variation indicates that the actual length of the
protocone does not change as much with wear as does the breadth, and that
the apparent elongation of the protocone of the unworn teeth is more due to
its narrowness than to its actual length.

Thus it is clear that the Langebaanweg specimens display this typical
feature of African hipparionids (see p. 355). In so far as the few available speci-
mens in varying stages of attrition from other African sites permit comparison,
there seems to be a great similarity in the shape and dimensions of the protocone
between H. africanum and the Langebaanweg Hipparion. The ‘African’ character
is even more strongly expressed in ‘Hypsohipparion’ albertense (tables 14, 20;
figs. 12, 13).

2. The hypocone is commonly angulated, tending to isolate in early wear.

Later it becomes more rounded, and the isolation from the metaloph is no

longer discernible, the anterior groove tending to disappear. Parallel variations
in the shape of the hypoglyph are observed with progressive attrition.

3. The marks are constantly closed.

4. The plications are numerous, profound and often bifurcated, especially
in early or moderate stages of wear (e.g. S.A.M.11716, Lg40/950, Lg51).
The enamel pattern is most complex on the posterior wall of the prefossette
and on the anterior wall of the postfossette. The plications have been counted
according to the method suggested by Gromova (1952), and the average
plication formula for the Langebaanweg specimens has been established:

Kee at 505) (075-5) — (On)
(1-2)

There is a constant isolation of a prefossette loop in the lingual-distal angle
of the prefossette, being bilobed and large in early stages of wear, and rather
triangular and arrow-shaped in advanced wear (e.g. specimens L934, L951,
S.A.M.11716; best example is the latter on pl. 5).

The pli caballin is constant, often double, in early and moderate stages

of wear. It is not observed in very advanced wear.
In all these features there is not a clear basis of distinction between the

8

ANNALS OF THE SOUTH AFRICAN MUSEUM

378

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 379

Langebaanweg material and the ‘Hypsohipparion’ upper teeth from Serengeti,
the plication formula of which is found to be very similar:

(0-4-5) —(2-5) — (0-3) —(o-2)
(0-3)
POSTCRANIAL EQUID REMAINS

At Langebaanweg seven phalanges have been recovered. Because of
their equid structure it could not be excluded a prior: that they belonged to
Hipparion represented by the dental remains at the sites. It must be pointed
out that a few teeth of Equus (to be described in another publication) have
been recovered at these sites. However, they only constitute 10% of the total
number of equid remains collected thus far. Consequently, because of the
lack of comparable Hipparion remains from Africa, the Langebaanweg specimens
are hesitantly included in this paper, more as a documentation for comparison
by others than a positive contribution to the reconstruction of the Hipparion

limb.

Material:
First phalanges: L1456, L1462a, L1462B, L1462c (pls. 10, 11).
Second phalanges: L1449, L1462p (pl. 12).
Third phalanges: L1444 (pl. 13).

All the absolute dimensions namely, length, diameter of the proximal
and distal ends and of the median shaft are usually considerably larger than
those (after Gromova, 1952) of the three species of Hipparion compared, i.e.
elegans, moldavicum and longipes (table 21). There is virtually no overlap between
the two series, although the highest figures for H. longipes (the largest of the
three species) are sometimes very close to the lowest figures for Langebaanweg
(table 22). Sondaar (1961) mentions one first phalanx of the forelimb third
digit with an exceptional length of 69:8 mm. In Europe the highest mean is
known from Eppelsheim, H. primigenium, with a length of 64-3 mm. and a
proximal extremity breadth of 41-3 mm.

A comparison of the dimensions of the proximal end of the first phalanges
from Langebaanweg with the breadth of the distal extremity of metatarsal ITI
from Olduvai and South Serengeti (tables 12, 13) makes it obvious that the
Langebaanweg specimens require a much broader metatarsal for articulation.
For H. longipes the respective breadths of the distal end of metatarsal III and
the proximal end of the first phalanx are 40 mm. and 42 mm. The mean
distal breadth of the known specimens of metatarsal III from East Africa is
44:7, the maximum being 48 mm., the widths of the proximal end of the first
phalanx at Langebaanweg are 50-56 mm. Therefore, either the Langebaanweg
specimens belong to a slightly more massive limb of an Hipparion or else they
belong to Equus.

Compared with Equus zebra and burchelli, the Langebaanweg specimens are
larger in a number of dimensions, but there is a greater amount of overlapping

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 381

and the dimensions are closer together than in the comparisons with the
Hipparion above (figs. 14, 15; tables 22, 23, 24). The difference between the
Langebaanweg and the Equus proximal phalanges average 6%.

The situation is less clear for the relative dimensions. The lowest index for
Langebaanweg is constantly smaller than the highest for burchelli, and very
often even overlaps H. longipes. ‘The relatively smaller indices indicate that in
the Langebaanweg specimens the length versus the breadth and the breadth
versus the A-P diameter are somewhat greater than in Hipparion and burchelli.
In these limited comparisons it seems that, to some extent, the Langebaanweg
architecture differs from these groups.

It cannot be denied that the Langebaanweg measurements fit more
satisfactorily within the range of Equus, but more material and comparisons
are needed before this discussion can be taken further.

First phalanx, III digit

A Oo &® eG& Axx x
B @O dw AA xx
B/A 0 ® Wy Ax vA x
C OCOD wRA Axx x x
C/A Ch Ye@xdCA x x
C/B rx a7 y © GOx ©
D CHO xe x
© (Fore)
E COO dew Lx x © (Hind) H. elegans
Seas, H. moldavicum
E/A 0 00 ver x A x
3 (Fore) H. longipes
(Hind)
F CRE HWA xx x A(F
ore)
: As (Hind) E. burchelli
G CXS) We @/Ax x x V (Fore) ¢
¥ (Hind) at
G/A COYOO CeLtOxA x x x Langebaanweg sp.

10 20 30 40 50 60 70 80 90

Abs. dimensions and Indices scale

Fic. 14. Absolute dimensions and indices scale for the first phalanx of the third digit in various
species of Hipparion and Equus compared with the Langebaanweg specimens.

8A

382 ANNALS OF THE SOUTH AFRICAN MUSEUM

Second phalanx, III digit

A § (inal H. elegans aX Wadd xx
Fore) .
B § (fina) H. moldavicum CXS Dorie. =
; ihere H. longipes
Cc COX) éeyvy LAK x ———ae
2B (herd) E. burchelli losin
Vv (Fore) @O* Oe x x 7 > Tae
@ (Hind) E. zebra C/A
D x Langebaanweg sp. COOOVT eLAxx
E @OQ 67 FAxA
) ® «x vy 64 FJO4E
F- CKO © xxi
0 Ox 064 vA ¥ . F/E
G KO ey TAL
OCiIO xe x y AT A GA
Abs. dimensions 10 20 30 40 50 60

Indices scale 50 60 70 80 90 100 110

Fic. 15. Absolute dimensions and indices scale for the second phalanx of the third digit in various
species of Hipparion and Equus compared with the Langebaanweg specimens.

ORIGIN AND DIFFERENTIATION OF AFRICAN HIPPARIONIDS

In recent years, it has become evident that Africa was more a true and
independent centre of evolution and dispersion than a refuge for ‘Pontian’
Eurasiatic animal forms. Discoveries at different levels of the history of the
mammals and in different regions have supported this concept proposed
originally by Pilgrim (1941), and later vindicated by, inter alios, Arambourg
(1959) and Cooke (1963).

Several independent observations have definitely established the original
role played by Africa in the development of late Tertiary mammals in general,
and have illustrated its contribution to the establishment of the ‘Pontian’
fauna in particular:

(i) The endemic evolutionary differentiation of the Proboscidea and
Hyracoidea from the Oligocene onwards;

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 383

A. ABSOLUTE DIMENSIONS

H. longipes Equus burchelli Langebaanweg
(largest) (largest) (smallest)
PHALANX I
Total length va 54 = 69°5 75-0 70°2
Breadth prox. end ek * 42 50 50
Diameter prox. end :f ns 31°7 35 36°8
Diameter prox. artic. surf. ns 28-2 27°3
Breadth distal end BS oe 34°8 40 41°2
Diameter distal end m, A 21°5 22 23-2
Breadth median shaft .. zs 30°3 32 34°1
PHALANX II
Maximum length a5 rt 42 40 47°2
Length median ant. surface... 34°5 34 34°1
Breadth prox. end hy th 39°5 46 47°8
Diameter prox. end “ie “gs 28°3 29 31
Breadth distal end <n By 39°4 43 Ane
Diameter distal end e ie 23 29 24°1
Minim. breadth shaft .. + 32°9 41-5 38° 4
PHALANX III H. moldavicum
Anterior length .. “i + 54°1 47°5 61
Maximum breadth ‘fe fa 54 59 72
B. INDICES
PHALANX I H. longipes
Prox. breadth/length .. wi 63°2 Fi~3 66-2
Prox. diam./length ws ty 48-9 50 47°4
Prox. diam./breadth .. bY 80-9 70 68
Distal breadth/length .. dy 53°5 af 53°7
Breadth med. shaft/length ns 43°6 45°7 44°3
PHALANX II
Prox. breadth/length .. ei 98 TS. 101
Distal breadth/length .. +3 ee} 107 86°3
Distal diam./breadth .. - 65:7 72:5 57°8
Minim. breadth shaft/length .. Gz* 7 104 o1°3

TABLE 22. Comparison of dimensions (mm.) and indices of Langebaanweg smallest phalanges
with the largest of H. longipes and E. burchelli. The italicized figures indicate those dimensions in
which the Langebaanweg specimens are smaller.

H. medit.
PHALANX III Langebaanweg (Pikermi) WH. elegans H.moldav. E. burchelli
ant = post ant _—— post ant —— post
Length ant. surface .. 61 50 A528 Age 2 JIG “bdet » 4Ou. Ages
Maximal breadth a 72 55 AG 3) "AA" 2 54. 43°4 59 58°5
Index B/L af 118 110 IOI°L 93°6 103 FOO) 120.) 122

TABLE 23. Dimensions (mm.) of phalanx III of Langebaanweg and of some Eurasiatic species
and E£. burchellz.

384. ANNALS OF THE SOUTH AFRICAN MUSEUM

Hipparion Equus Langebaanweg

— — ——— — ——————“~“ | ———————“@“ |\ qq“ | ——_ | _______._...

elegans | moldav.| cabs.\. | zebra

Index of length
Phat Li ant. 135°9| 145°1 | 116°4 | 110 127 123
3d digit/Phal. II | post.140°1| 157°3 | 118°8 | 116-9

Index of length ant. 78°7| 86:9 | 64°2 |) 57°6|Li144q4 | L144q4 | L1444 | Li444
III/I | | — | ——_—
post. 85 gi*2 66°4 | 61°3 | L1462a | L1462B | Lr1462e | L1456

85°3 75°8 77°3 79°3

TABLE 24. Comparison of relative lengths of phalanges.
(In the case of Langebaanweg indices have been calculated from the possible associations.)

(11) the presence in the Lower Miocene of East Africa and in the Upper
Miocene deposits of North Africa of a complete specialized faunal assemblage
with typical African characteristics before parallel diversification took place
in Europe;

(ii1) the constant specific differences between comparable Pontian
Eurasiatic and African forms; and

(iv) the African pre-Pleistocene differentiation of Bovidae with the evi-
dence of the local origin of Cephalophini, Neotragini and probably the
Alcelaphini (Wells, 1957).

It is suggested that, because of the stability of its climate, its geography
and its vegetation, Africa has provided a suitable habitat throughout the
Tertiary. Furthermore, groups have evolved and later have developed parallel
forms. Elsewhere, these have become extinct, but survival was favoured in
Africa. Even in modern times this seems to be the case, e.g. the Proboscidea,
Giraffidae, Artiodactyla, and Anthropomorpha. The surviving species, far
from being newcomers and refugees from outside, are local representatives
of a fauna which has disappeared elsewhere.

Arambourg (1952) and Cooke (1960), inter alios, demonstrated that the
Plio-Pleistocene forms of elephants may have differentiated within the continent
of Africa, paralleling the steps of progress observed elsewhere, but not necessarily
linked to Eurasia by periodic invasions as was previously thought.

Analogous circumstances may have influenced the origin and the diffe-
rentiation of the hipparionids. There is abundant evidence that Hipparion
evolved in North America from some Merychippus stock in Miocene times.
Migration probably took place at a rather rapid pace, Hipparion being found
on both sides of the Mediterranean basin before the end of the Miocene.
Hipparion has been recovered from brackish water deposits dating from the
Sarmatian in the area of Sebastopol (Borissiak, 1914) and Istanbul (Chaput
and Nafiz, 1934; Chaput and Gillet, 1938). It is also known from the Upper

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 385

Tortonian Beds in the Rhone Valley (Denizot, 1939). Villalta and Crusafont-
Pairo (1946a, b, c; 1947; 1948) have described Hipparion from the Vallés—
Penedés area in Catalonia, in continental and brackish water deposits inter-
calated in sediments ranging from the Burdigalian to the Tortonian, with
Hipparion appearing in the Upper horizon (Vallesian, parallel to the Meotian
of Eastern Europe, i.e. Upper Sarmatian). In the basin of Teruel, Hipparion
was recently found to appear in Los Algezares in some gypsum intercalation
of the middle light-red clayish deposit overlying the Los Monotos series,
dated as the Vindobonian (Sondaar, 1961).

Parallel to these occurrences, and probably very close to the last one in
the chronological sequence, hipparionids have been recorded in the Upper
Miocene of the Maghreb, i.e. H. africanum (Arambourg, 1959) from Oued el
Hammam and Marceau.

It is still not known which Atlantic or Pacific bridge the hipparionids used
to invade the Old World. The only reasonable assumption which has been
proposed is that the migration route probably lay very much to the north.
The absence of any important migration of Camelidae and Antilocapridae
from America, or of hyaenas, antelopes and giraffes to America, provide good
evidence in support of this high latitude of the migratory passage (Simpson,
1947). On the other hand, and as a confirmation, the most ancient Old World
Hipparion seem to have been much better adapted to a humid and forested
biotope of temperate, or even cold, climates than most of the American forms,
which evidence a drier and steppe environment. Rather than indulging in
some fantasy that a direct connection between America and Africa existed by
way of a hypothetical chain of islands across the North Atlantic (Joleaud,
191g9a, b), it seems more reasonable to accept the normal Bering route which
certainly meets the climatic requirements of the migrants.

Furthermore, in the present state of our knowledge, it is neither possible
to formulate any conclusion about those species of Hipparion responsible for
the origin of the African group, nor to reconstruct the phyletic sequence of
African Hipparion. The fact that H. africanum is found contemporaneous with, or
possibly prior to, every other European form and that it already shows typical
_ African differentiation, refutes a European origin of the group from any of
the known ‘Pontian’ species. On the other hand, and for the same reasons,
there seems at present to be no possible way of establishing an African origin
for the European hipparionids. However, the metapodials of H. africanum are
rather strong and the lateral digits are well-developed, which must be inter-
preted as ‘primitive’ features, less specialized than in most of the early Eurasiatic
forms. But other characters, e.g. the articulation of the external cuneiform
with the second metatarsal only, a typical mechanical adaptation to tridactyly,
show more primitiveness in the relatively contemporaneous H. catalaunicum
(Pirlot, 1956). It seems that here there is evident some expression of an inde-
pendent mosiac pattern of evolution in different (Eurasiatic and African)
groups. Although they obviously descend from a common ancestral stock, they

386 ANNALS OF THE SOUTH AFRICAN MUSEUM

have been precociously isolated on different continents and have developed _
their independent stages of evolution.

For several reasons it is not possible to depict accurately the actual evolu- _
tionary development of the African Hipparion. As has been stated previously,
the fossil record is rather poor. At more than thirty sites where Pleistocene
hipparionids have been recovered, the available material consists almost
exclusively of isolated teeth, and virtually nothing is known of the skeleton.
Furthermore, because of the plasticity of the group, and of the fluctuating
character of many features, which seem to be highly adaptive to ecological
requirements, it would appear hazardous to reconstruct artificially an ortho-
genetic series. Gromova (1952) states that the diversity among Old World
hipparionids usually remains at the level of a particular genus and does not
exceed specific differences. On the basis of our present knowledge, this typifies
the African representatives of the group: overall dimensions, proportions,
size of the teeth, plications of the enamel, development of stylids, elongation
of the protocone, preorbital fossae and development of lateral digits. Most
of these features are readily adaptive and susceptible to functional interpretation
under climatic and ecological circumstances.

On the other hand, it cannot be denied that parallel to the ecological
adaptation which has been emphasized previously, some general and fairly
continuous trends have characterized the evolution of the hipparionids through-
out the world. From Merychippus onwards, in late Cenozoic times, the various
groups have shown a permanent tendency to

(a) increased hypsodont dentition;

b) lengthening and flattening of the protocone;

c) reduction of the external depression in the lower teeth;
development of additional stylids; and

e) evolution of the double knot in three different but constant directions,

i.e. towards a ‘stenonis’? type in America, towards a ‘“Hipparion’ type

in Eurasia, and towards a ‘caballus’ type in Pleistocene Africa.

There is insufficient evidence to state that the reduction of lateral digits
and the elongation of the third metapodials have obeyed a clearly directed
evolutionary trend. The enamel pattern has not been controlled by any
constant progressive influence, but it has only responded to the changing
ecological conditions.

There is little doubt that these trends have been developing on more or
less parallel and independent pathways and at different and varying speeds
along the several lines, with the resulting mosaic type of differential radiating
evolution.

For the reasons stated earlier in this paper, the degree of relationship
between Hipparion africanum and the Pleistocene stock of African Hipparion
is not obvious. There is a great gap in our record, and this is not sufficiently
bridged by the fragmentary remains of H. sitifense. Therefore it is difficult to
appreciate to what extent the distance between H. africanum, H. sitifense and

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 387

the Pleistocene forms illustrates the chronological or the phyletic separation.

All Pleistocene African Hipparion show a marked progression in the
expression of trends (a), (b) and (d) above; and together with the special
orientation of trend (e), this vindicates their common origin and their genetic
homogeneity.

It is suggested that the “African features’ and the more stable biotope
are responsible for the prolonged presence of Hipparion in Africa during the
Pleistocene. However, the differential expression of these trends along various
lines of evolution allowed for some differentiation during this period.

A basic group, represented by ‘Hypsohipparion’ and the Langebaanweg
specimens, is characterized by a greater conservatism in the building of addi-
tional stylids: the ectostylid is constantly absent. The flattening of the protocone
and the hypsodonty are strongly manifested in ‘Hypsohipparion’, but are less
marked at Langebaanweg. This basic group seems to be restricted to the
Lower Pleistocene (the basal grey tuffs of Serengeti and the archaic forms at
Langebaanweg).

‘Stylohipparion’ is more advanced in the expression of additional stylids:
the constancy of the ectostylid indicates a group subjected to a particularly
clear trend of evolution (‘Entwicklungswucht’ of Dietrich, 1942) in the process of
building a new Hipparion form. This successful group has rapidly extended
throughout the continent, being found in Lower and Middle Pleistocene
deposits from the Maghreb to the northern Cape Province, and possibly
differentiated on a geographical basis. The oldest record derives from North
Africa. It is not impossible that the group migrated eastward and then south-
ward, and, after being widespread throughout Africa, became extinct first in
North Africa during the Lower Pleistocene, then in East Africa where it is
still found in the Middle Pleistocene deposits.

TAXONOMIC STATUS OF THE AFRICAN HIPPARIONIDS
Family Equidae Gray 1821

Sub-family Equinae Steinmann & Déderlein 1890

Elongated face with completely closed orbits. Molar teeth, at first mode-
rately, then later strongly hypsodont, with closed valleys, the external portions
of which become isolated in the shape of pits, called fossettes or marks. Abundant
cement layer on the outer surface and in all the hollows of the tooth. Complete
homeodonty, the largest tooth being P, of a triangular prismatic shape,
pointed forwards. Ulna is first fused with the radius; however, later in the
development, it is reduced to its proximal portion (olecranon process), which
has the appearance of a mere apophysis of the radius. Limbs are tridactyl in
primitive forms, with complete lateral digits although they no longer reach
the ground, except during galloping, when they become functional. In advanced
forms, however, limbs are monodactyl, and the lateral metapodials are reduced

388 ANNALS OF THE SOUTH AFRICAN MUSEUM

to small splints, eventually only persisting in their proximal portion.
Unguligrade.

HIPPARION de Christol 1832

Face is relatively short, with profound lacrimal fossa. Prismatic hypsodont
molars, of moderate height, slightly curved, with more strongly plicated
enamel pattern than in Equus. The protocone pedicle has narrowed, isolating
the protocone, giving the appearance of a column which is attached to the
protoconule only at its base. On its occlusal surface, the protocone shows an
island of dentine, circled with enamel, of a rounded or oval shape. The enamel
of the fossettes is rather strongly plicated. Upper and lower incisors show a
typical mark. Metapodial moderate to long. Persistent tridactyly; the lateral
digits are strongly developed and more closely related to the third metapodial
than in Merychippus.

Pomel (1897) included in his definition that the lateral metapodials are
not very different from those of Equus, but they broaden distally into an articular
head; there are three phalanges, the most distal not reaching the ground.

Hipparion africanum Arambourg 1959

Hipparion sp. Arambourg, 1951: 2464, Arambourg, 1954: 295.
Hipparion africanum Arambourg, 1959: 75, Pl. 10, 11, 12, 13, 16, fig. 7.

‘Hipparion with skull of great dimensions, but with limbs of moderate
size and heavy extremities. Face and snout elongated; nasal aperture long
and broad; orbits far back; preorbital fossae long, simple, distant from orbit.
Dental series of moderate size: P?-M%=141 to 154 mm. Upper cheek teeth
with strongly plicated enamel; compressed protocone, elliptical or lenticular.
Cingular formations developed on lower milk teeth. Limbs are relatively
short, with strong metapodials, in which the lateral digits are still
well-developed.’ (Translated from Arambourg, 1959, p- 95.) .

Hipparion sitifense Pomel 1897

Hipparion sitifensis (sice) Pomel, 1897: 14, Pl. 1, figs. 11-15, pl. 2, figs. g-10.
Hipparion sitifense Pomel, Arambourg, 1956: 817, pl. 26, figs. 1-5a.
Hipparion gracile [Non] Kaup, Thomas, 1884: 10, pl. 2, figs. 1-3.

Hipparion crassum [Non] Gervais, ‘Thomas, 1884: 10.

Hipparion characterized by its small teeth (A—P and transverse dimensions
of P3—-M2, ca. 19 mm.), little hypsodont and markedly curved in the median
plane. The upper teeth have a flattened protocone, completely independent
from the protoloph right down to the base of the tooth. The enamel plications
are few and simple. The lower teeth possess no ectostylid. The lateral digits
are well-developed.

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 389

Hipparion (Stylohipparion) libycum Pomel 1897

Hipparion ? libycum Pomel, 1897: 8, pl. I, figs. 1-7, pl. 2, figs. 11-12.

Hipparion massoesylium Pomel, 1897: 11, pl. 1, figs. 8-9.

Hipparium? (sic) ambiguum Pomel, 1897: 15, pl. 2, figs. 2-4.

Hipparion crassum Gervais, Solignac, 1927: 756.

Hipparion steytleri van Hoepen, 1930: 21, figs. 14-19.

Eurygnathohippus cornelianus van Hoepen,? 1930: 23, figs. 20-22.

Stylohipparion hipkini van Hoepen, 1932: 31, figs. 14-20.

Stylohipparion steytlert (van Hoepen), van Hoepen, 1932: 33, figs. 21-23.

Notohipparion namaquense Haughton,’ 1932: 421, text-fig. 5.

Libyhipparion ethiopicum Joleaud, 1933: 7, pl. 1, figs. 1-4, 6-8, 10-12.

Equus (Hippotigris) sp., (pro parte) —Joleaud, 1933: 25, pl. 1, figs. g and 13.

Libyhipparion steytlert (van Hoepen), Joleaud, 1933: 12.

Stylohipparion cf. albertense (Hopwood), Hopwood, 1937: 130.

Stylohipparion, Notohipparion, Libyhipparion, Dietrich, 1942: 98, pl. 13, figs. g1-95, pl. 14,
fig. 99 pro parte, pl. 15, fig. 105, pl. 16, fig. 109.

Stylohipparion albertense (Hopwood), Arambourg, 1947: 303, pl. 10, fig. 3, pl. 11, figs. 3-5.

Diagnosis :

Three-toed equid with rather hypsodont cheek teeth with a strongly
developed ectostylid, particularly constant on P, and M,. Typically the ecto-
stylid is broad in the centre and narrowed at each end, the greater length
being mesiodistally and antero-lingually orientated. The upper teeth show the
typical ‘African’ features, namely, complex enamel plications of the ‘mark’
walls, and the narrow elongated protocone. |

Hipparion (Hipparion) albertense serengetense, subsp. nov.
Hipparion albertensis (sic) Hopwood, 1926: 17, fig. 4.
Hypsohipparion albertense (Hopwood), Dietrich, 1942: 97, pl. 4, fig. 39, pl. 13, figs. 87-90,
93b, 96, 97; pl. 14, figs. ro1—104, pl. 15, figs. 106-108, pl. 16, figs. 110, 112; pl. 20, fig. 160.
Equus (Hippotigris), Arambourg, 1947: 306.
Stylohipparion albertense (Hopwood) (pro parte) Arambourg, 1947: 306.
Diagnosis :

Three-toed equid with very hypsodont teeth (the M? of which reaches a
crown height of 80-90 mm.). The lower teeth are characterized by the constant
absence of the ectostylid. The protocone is very elongated, the cement is very
thick, the enamel plications being complex. The dental series is short, average
length being 156 mm.

Eipparion (Hipparion) albertense baardi, subsp. nov.

Type specimen: Lg46 (pl. 8) in the S.A. Museum, Cape Town.
Paratypes: Lo51, S.A.M.11717 (pls. 4, 2) in the S.A. Museum, Cape Tega
Type site: Baard’s Quarry, Langebaanweg, C.P.

1 It must be a misprint. Everywhere else, except in the title, Pomel writes Hipparion
ambiguum.

2 This is tentatively included here on the basis of our belief that this symphysial fragment
is referable to other Stylohipparion specimens from this area.

3 This may prove to be a transitional form between the typical Hipparion and the more
progressive. Hipparion (Stylohipparion).

390 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis :

Three-toed equid with rather hypsodont teeth, about 70 mm. crown height.
The lower teeth are characterized by the constant absence of the ectostylid
and by a tendency to form other additional stylids, especially protostylid
extending along the total height of the crown. The protocone is elliptical but
less elongated than in serengetense, enamel is thin and plications are complex.

DISCUSSION

1. THE AFRICAN NON-PLEISTOCENE HIPPARIONIDS constitute
one genus (Hipparion) and two species (africanum and sitifense). The lack of
marked affinity to any of the Pleistocene forms which constitute a homogeneous
unity precludes any consideration of this group in terms of the subgenera
proposed for the Pleistocene forms.

2. THE AFRICAN PLEISTOCENE HIPPARIONIDS constitute a
certain unity marked by the characteristics of the genus Hipparion. Furthermore
they have in common a number of ‘African features’ which distinguish them
from the non-African forms. Nevertheless these different features do not
necessitate generic distinction.

Within the group there exist variables of differing degrees of importance.

The most obvious differentiating characteristic is the presence or absence
of ectostylids. The evolutionary trend and ecological adaptation of this feature
have been discussed (pp. 331, 332, 333) and it is considered that it constitutes a
differentiation at the subgeneric but not generic level. The statistical constancy
of absence or presence of this feature within groups otherwise identical indicates
that consideration must be accorded on both a non-generic and a non-specific
basis. Consequently the African Pleistocene Hipparion are subdivided into
two subgenera, viz.

HMipparion (Hipparion) de Christol 1832, and
Hipparion (Stylohipparion) van Hoepen 1932.

3. THE CONSIDERATION OF THE SPECIES of Hipparion (Stylo-
hipparion) is beset with the obvious difficulty of the paucity of the material
available. Previous workers, faced with the same problem, have either cautiously
avoided speciation or unjustifiably proposed species that are void, e.g.
Van Hoepen, 1932—steytleri, hipkini. Arambourg (1947, 1956, 1959) refers to
libycum only when he is discussing the North African Pleistocene material;
otherwise he generalizes to the extent of using the generic (subgeneric) name
Stylohipparion without species determination. We concur with this cautious
attitude. We recognize that material referable to this subgenus has been
recovered from three different geographical areas, namely, the Maghreb,
East Africa (incl. Omo) and South Africa. Joleaud (1933), also aware of the
geographical distribution, recognized in his new genus Libyhipparion three

HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 391

LAANGEBAANWEG HYPSOHIPPARION

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Fic. 16. Diagrammatic superimposition of an upper and a lower molar tooth of the South
Serengeti ‘Hypsohipparion’, and the Langebaanweg Hipparion.

species, viz. libycum, ethiopicum and steytleri. However, despite the differences of
the localities, the wide range of variation within the small amount of available
material does not provide the evidence for species separation.

Comparison of the various ‘diagnoses’ provided by Hopwood (1937),
Arambourg (1947), and Cooke (1950), and reference to descriptions by Pomel
(1897) and Joleaud (1933) indicate quite clearly that no true, clear-cut diagnosis
is possible. Therefore, it is proposed to extend Arambourg’s generalized usage
of libycum to cover all the material included in this subgenus, but with the
awareness that subsequent discoveries may bring to light information to
highlight the differences of locality on a subspecies level.

392 ANNALS OF THE SOUTH AFRICAN MUSEUM ~

4. A better situation exists with reference to Hipparion (Hipparion) which
includes a portion of the material from South Serengeti (i.e. ‘Hypsohipparion’
Dietrich) and the Langebaanweg specimens.

The dimensions and the main features (fig. 16) of the teeth from both areas
are almost identical. However, there are differences in those features which
emphasize the ‘African’ character of the dentition; namely, the hypsodonty,
the elongation and narrowing of the protocone (tables, 16, 19, 20; figs. 12, 13).
For both of these characters the Serengeti material displays a more progressive
stage. Furthermore the latter also exhibits a thicker cement. These differences,
although constant, do not constitute sufficient grounds for species separation.
Therefore the species albertense is retained for both groups. Taking into considera-
tion the ecological adaptation of these features, it is proposed to distinguish
these two groups on a subspecies level. The Serengeti “Hypsohipparion’ group,
erected by Dietrich, is now considered to be Hipparion (Hipparion) albertense
serengetense, subsp. nov. The reason for dropping the genus Hypsohipparion has
been discussed above. The Langebaanweg group constitutes the subspecies
Hipparion (Hipparion) albertense baardi, subsp. nov. The name baardi is chosen
because the first specimens were recovered from Baard’s Quarry, owned by
Mr. J. Baard.

It seems that these two subspecies constitute the earliest Pleistocene forms.
The material (to be described by Dr. Stirton) from the base of Olduvai
Bed I should confirm or correct this suggestion.

ACKNOWLEDGEMENTS

This study was supported in part by the U.S. Public Health Service,
National Institutes of Health grant no. GM 10113-02(3, 4) and USPHS
General Research Support Grant 1-So1—FR-—05367-01. In addition, generous
assistance was obtained from

The Wenner-Gren Foundation for Anthropological Research Inc.,
New York; .

Fonds National de la Recherche Scientifique (Brussels) ;

Fondation Universitaire de Belgique;

The Boise Fund, University of Oxford; and

The Dr. Wallace C. and Clara A. Abbott Memorial Fund of the
University of Chicago.

We are indebted to the African Metals Corporation (AMCOR) for
permission to work at the Langebaanweg sites, and we are grateful to Messrs.
Glathaar, Krumm, Muller and De Bruyn and Dr. Boardman for their co-opera-
tion. Mr. John Baard, owner of the farm ‘Langberg’, has kindly allowed
excavations. In particular we are thankful to Mr. Robin Warren, a chemist
of AMCOR, for his continuous efforts to recover material and for his enthusias-
tic assistance. Mr. Q. B. Hendey, research assistant to one of us (R.S.) at the
South African Museum, Cape Town, has been extremely helpful. Some of the
information on the geology of the Langebaanweg sites results from discussions

ya

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HIPPARION FROM LANGEBAANWEG, CAPE PROVINCE 393

with Mr. R. R. Inskeep, Dr. A. Fuller and Mr. D. Needham of the University
of Cape Town, and Mr. Robin Warren.

Dr. R. Bigalke, Director of the McGregor Memorial Museum, Kimberley
has been most co-operative in sending requested material for study. Professor R.A.
Stirton, Museum of Paleontology, University of California at Berkeley, provided
facilities for the study of some of the East African material there, which Dr.
L. S. B. Leakey allowed us to examine. Dr. Stirton kindly permitted us to
publish our observations on the material.

Dr. K. H. Fischer, Institut fir Palaontologie und Museum der Math.-
Naturwissenschaftlichen Fakultat der Humboldt-Universitat zu Berlin
kindly took some measurements (at the authors’ request) on the original
Serengeti material.

Mrs. M. A. Norris patiently typed the manuscript.

SUMMARY

The recent discoveries of hipparionid material at the Langebaanweg
fossil sites (Cape Province, South Africa) necessitated a revision and interpreta-
tion of similar material recovered elsewhere in Africa. At the same time new
data on material from the Vaal River Gravels, Olduvai Gorge and South
Serengeti are added. A review of the geological and faunal data described
from approximately 40 sites in Africa leads to a tentative chronological correla-
tion. Consideration of the anatomical features (mostly of the dentition) provides
opportunity for a discussion of the ecological adaptation and evolutionary
trends. The problems of the origins and migrations of African hipparionids
are dealt with.

It is concluded that the African hipparionids belong to a single genus,
Hipparion. The Miocene forms remain limited to H. africanum. The Pliocene is
represented by H. sitifense, while the Pleistocene forms are referable to two
subgenera, viz. Stylohipparion and Hipparion.

The Serengeti ‘Hypsohipparion’ group, is referred to Hipparion (Hipparion)
albertense serengetense subsp. nov.

The Langebaanweg material forms a unified group and is referred to
Hipparion (Hipparion) albertense baardi subsp. nov.

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.
$
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|
5
;
4
«¢

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Ann. S. Afr. Mus., Vol. XLVIII Plate VII

D ———__.

Occlusal aspect. Scale: 1 cm. A—MMK 431. B—MMK 5225. C—L1467c. D—L1463.
E— L956.

Ann. S. Afr. Mus., Vol. XLVIII

®eclusal aspect. Scale: 1 cm.

A—S.A.M. 11717.

B—S.A.M. 11718.

D—Lo34. E—L1465r.

Plate VIII

C—Lg40 | 950.

Ann. S. Afr. Mus., Vol. XLVIII Plate IX

D

Occlusal aspect. Scale: 1 cm. A—S.A.M. 11719. B—Lg53. C—1L1467p. D—L955.

Ann. S. Afr. Mus., Vol. XLVIII Plate X

Occlusal aspect. Scale: 1 cm. A—1L1467H. B—L14678. C—L1459. D—Lg51. E—Lo58.

Ann. S. Afr. Mus., Vol. XLVIII Plate XI

A—D: occlusal aspect. Scale: 1 cm. A—L1467z. B—Lo39. C—L1451a. D—S.A.M. 11716.
E—S.A.M. 11716: section across middle of crown.

> ia

Ann. S. Afr. Mus., Vol. XLVIII Plate XII

Occlusal aspect. Scale: 1 cm. A—-Lg943. B—Lg41. C—L1465p. D—Lo47. E—Lo44.

Ann. S. Afr. Mus., Vol. XLVIII

Occlusal aspect. Scale: 1 cm. A—L94

Plate XII

8. B—Lo42. C—Lg54. D—Lg36. E—S.A.M. 11724.

Ann. S. Afr. Mus., Vol. XLVIII Plate XIV

Occlusal aspect. Scale: 1 cm. A—Lg946. B—Log52. C—L1465a. D—Lo59. E—L1448.
F—Lo38.

Ann. S. Afr. Mus., Vol. XLVIII Plate XV

es FE

Occlusal aspect. Scale: 1 cm. A—Lo45. B—L14658. C—S.A.M. 11722. D—Lg35.
E—Lo49. F—L957.

=g

Plate XV1

Ann. S. Afr. Mus., Vol. XLVIII

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Ann. S. Afr. Mus., Vol. XLVIII Plate XVI

F

A, B, C—L1462p: lateral, anterior and proximal surfaces respectively.
F—L1462c; anterior, lateral and proximal surfaces respectively.

Amn. o. Afr. Mus., Vol. XLVIII Plate XVII

A, B, C—L1449: lateral, anterior and proximal surfaces respectively.
D, E, F—1L1462p: proximal, anterior and lateral surfaces respectively.

Ann. S. Afr. Mus., Vol. XLVIIT

A, B, C—1L1444: lateral, anterior and inferior surfaces respectively.

Plate XIX

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TaBLE oF CoNTENTs and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. x 7 in. (7$ in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

Smiru, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name

of publisher. Thus:

SmitH, C. D. 1954. South African Plonias. In Brown, X. Y. Marine faunas. 2nd ed. 3: ibis
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopied by the XV International
congress of zoology, London, Fuly 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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iH, BARRY : LIDRARY,

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ON THE EPIPTERYGOID—ALISPHENOTD=y,
TRANSITION IN THERAPSIDA

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ON THE EPIPTERYGOID—ALISPHENOID TRANSITION
IN THERAPSIDA
By
T. H. Barry

South African Museum, Cape Town
(With 22 figures in the text)

CONTENTS

PAGE
Introduction . ‘ ; - 399
Material . : ‘ . 407
The palatoquadrate of Pristerodon . 402
Phylogeny ! P : - 405
Placodermi : : F . 405
Crossopterygii . : ‘ . 406
Labyrinthodontia , : $107
Cotylosauria. ; . . 408
Pelycosauria . ‘ , » XO
Therapsida ‘ - ; ABS
Triconodonta . ; ; ae
Living reptiles and mammals :) Cag
Conclusions . ; , 2 421
Acknowledgements. p - 423
Summary . : : ; -) 424
References : : ‘ « 424

INTRODUCTION

Mainly as a result of the researches of Gaupp (1902), Allis (1919), Broom
(1907, 1909 and 1914), Fuchs (1912) and Gregory and Noble (1924) it is
now generally accepted that the reptilian epipterygoid and mammalian
alisphenoid, although superficially different in many respects, are homologous.
_ structures. The views of these workers have received support from embryologists,
comparative anatomists and palaeontologists alike over the last few decades
and the theory has now virtually become axiomatic.

In modern reptiles there is a marked degree of variation in the development
_ of the epipterygoid, ranging from the primitive, rather solid, construction of
the bone in Sphenodon, to the absence or vestigial development of the bone in
adult Ophidia and Crocodilia with the Lacertilia displaying a thin columnar
structure. The relationship of the branches of the trigeminal nerve to the
_ bone, remains constant throughout the class, the profundus emerging anterior
_ to the epipterygoid and the maxillary and mandibular branches posterior to
the bone.

399

Ann. S. Afr. Mus. 48 (17), 1965, 399-426, 22 figs.

400 ANNALS OF THE SOUTH AFRICAN MUSEUM

In mammals the development of the alisphenoid as a relatively broad and
flattened bone is fairly constant except in Echidna where it appears to be
absent. In mammals, however, the relationship of the branches of the trigeminal
nerve to the alisphenoid varies. This ranges from the ‘reptilian’ type, where
the bone lies between the profundus and maxillary branches, found in Didelphis,
through stages where first the maxillary and then also the mandibular branches
pierce the bone, to where the profundus and maxillary branches both emerge
anterior to the alisphenoid.

The palaeontological evidence for the homology. of the reptilian epiptery-
goid and the mammalian alisphenoid presented in the early years (mainly by
Broom) naturally reflects the inadequate state of knowledge of the early
reptile fossils at the time. Although seldom stated explicitly there was a strong
tendency to see in the lacertilian condition the basic type from which the
mammalian condition could be derived. |

As far back as 1907 Broom stated that palaeontological evidence supported
the view that the alisphenoid of mammals had evolved from the reptilian
epipterygoid concluding: :

‘In the very primitive reptiles, of which Procolophon may be taken as a
type, we have a lizard-like columella cranii. In the early types, which
have specialized along the mammalian line, such as the Therocephalians,
we still have a columella cranii. In the even more mammal-like ano-
modonts, such as Dicynodon and Oudenodon, there is a columella cranii, but
no alisphenoid. When we come to examine the Cynodonts—those remark-
able reptiles, so very nearly related to the Mammals as to be regarded by
many as the group ancestral to them-—we find a broad flattened bone,
which most anatomists would not hesistate to call the alisphenoid, and
yet there can be little doubt, it is only the columella cranii of the anomodont
flattened out’ (p. 114).

In 1914. Broom offered more direct evidence to substantiate his view
stating:

‘A short Seinen: is pee in Diadectes sm a ions liarickiee pderorlls
-cranii is met with in Procolophon. A short epipterygoid occurs in the Pely-

. .cosaur Dimetrodon, In Anomodonts the epipterygoid is present as a long
___slender'rod—in-some species rounded and in others much flattened . . .
and its lower end is considerably expanded _antero-posteriorly,-giving it a

. long suture with the. pterygoid. In Gorgonopsians. the epipterygoid is a

. -long, much flattened red which inferiorly has probably -similar relations
_to the pterygoid as.seen in. the Anomodonts. In the -Therocephalians the
epipterygoid is.only satisfactorily known in Scylacosaurus. Here, ...itisa
relatively short, flattened structure with a very wide base which lies along
the pterygoid. In the Cynodonts, at least in the higher forms as exemplified

_ . by. Diademodon and Gynognathus, in the region-occupied by the epipterygoid
in the Therocephalians there is a very much larger bone which there can

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA 401

be little doubt is also an epipterygoid development. The upper part is
greatly expanded antero-posteriorly and forms much of the cranial wall.
The lower portion of the bone is so much more developed than in the
Therocephalian that it completely replaces the posterior part of the
pterygoid, . . . and extends outwards as far as the quadrate. There can be
little doubt that while this bone is homologous with the epipterygoid of
the lower forms, it is also the homologue of the mammalian alisphenoid’

(p- 30).
In the next thirty years very little advance was made. In 1944 Olson still

describes the structural changes undergone by the epipterygoid in the transition
from the primitive reptilian condition to that of mammals as follows:

‘In primitive reptiles the bone is composed of a restricted basal portion
and a slender ascending ramus, the columella cranii. In somewhat more
advanced forms, the basal portion is expanded into a posterior quadrate
ramus and an anterior pterygoid process. At this stage the ascending
ramus remains a thin rod. In the advanced mammal-like reptiles, the
ascending ramus is much expanded and has attained intimate association
with the periotic behind and the parietal above. The quadrate and
pterygoid processes are somewhat elongated’ (p. 110).

In the twenty years that have elapsed since Olson wrote the foregoing
descriptions of the palatoquadrate of all the principal groups of reptiles and of
the forms ancestral to the reptiles have become available. A restatement of the
course of the evolution of the epipterygoid-alisphenoid has therefore become
necessary. In the following pages a short review of the present state of our
knowledge of this element is given. Although the epipterygoid is a distinct
element in the early tetrapod skull its evolution cannot be discussed in isolation
since both the number of palatoquadrate ossifications and the extent of these
elements vary. An account of the palatoquadrate as a whole will therefore be
given and the evolutionary trends evident in its ossifications discussed.

My interest in the transformation of the epipterygoid (and quadrate
in so far as it affected the epipterygoid) originated while studying serially
ground sections of the skull of the anomodont Pristerodon buffaloensis. This
technique revealed important features of the palatoquadrate complex which
would not have been readily seen in material cleaned in the normal way. As
some of these features contradict vital points in the evolutionary sequences as
interpreted by Broom and Olson, an account of the Pristerodon palatoquadrate
will be presented first and will be followed by a general review of the trans-
formation of the palatoquadrate.

MATERIAL

A skull of Pristerodon buffaloensis Toerien, No. B.P.I. 339, was made available
for sectioning by Dr. A. S. Brink, Assistant Director of the Bernard Price

Z

402 ANNALS OF THE SOUTH AFRICAN MUSEUM

Institute for Palaeontological Research, Johannesburg. It was discovered
in 1956 by Mr. James W. Kitching of the same Institute in Cistecephalus Zone
strata on the farm Kirkvors, today known as De Hoop, approximately two
miles north-west of Murraysburg, Cape Province. Pristerodon belongs to the
family Endothiodontidae of the Sub-order Anomodontia (Sub-class Synapsida;
Order Therapsida).

THE PALATOQUADRATE OF PRISTERODON

The ossified palatoquadrate of Pristerodon buffaloensis is represented by the
separated epipterygoid and the quadrate (figs. 1 and 4). Viewed laterally the
epipterygoid is L-shaped, with both the vertical and the horizontal limbs well
developed. The vertical limb (or columella cranii) is relatively thin and extends
upward and slightly forward to meet a ventrally extending ridge originating

PILA ANTOTICA

PARABASISPHENOID

Cae a eA EPIPTERYGOID

PTERYGOID

QUADRATOJUGAL

QUADRATE RAMUS PTERYGOID

fe
QUADRATE RAMUS EPIPTERYGOID iy
QUADRATE

Fic. 1. Pristerodon buffaloensis. Lateral view of skull reconstructed from serial sections.
Squamosal cut away to show palatoquadrate complex.

PILA ANTOTICA
AUDITORY CAPSULE

ASCENDING PROCESS ~

PALATOQUADRATE BAR

PARS QUADRATA

Fic. 2. Tropiocolotes tripolitanus. Lateral view of skull of 15°2 mm.
embryo. (After Kamal, 1960.)

a

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA 403

trom the under-surface of the parietal. The posteriorly directed horizontal
portion is thickest near the angle of the bone, tapering posteriorly to become
rod-shaped (quadrate ramus of the epipterygoid). The entire base of the
epipterygoid is applied to the dorsal and/or dorso-lateral surface of the quadrate
ramus of the pterygoid. The quadrate rami of both the pterygoid and the
epipterygoid are directed postero-laterally, towards the inner surface of the
quadrate.

The quadrate is relatively large, has the shape of an inverted Y in cross-
section, the upper leg lying in a deep antero-posteriorly directed ventral
pocket of the squamosal. It is suturally attached to two other elements only, the
quadratojugal on the outside and the stapes to which it is attached through a
short horizontal ridge projecting medially from the inner condyle. Antero-
dorsally to this ridge there is a longitudinal groove in the vertical face of the
quadrate, extending backwards from the anterior border for approximately
one-third of the length of the vertical plate (fig. 4). Throughout its length
this groove is seen to follow remarkably closely a course parallel to the posterior
portion of the quadrate ramus of the pterygoid, the latter bone terminating
close to, but free from the inner surface of the quadrate, approximately half-way
along its length. A distinct rounded bulge terminates the groove in the left
quadrate. As the groove follows what could have been the course of a posterior
extension of the horizontal limb or quadrate ramus of the epipterygoid it
would seem reasonable to assume that the groove housed a cartilaginous rod
connecting the quadrate with the epipterygoid.

The occurrence of a solid link between the quadrate and epipterygoid in
an adult Pristerodon, recalls conditions found in the developmental stages of
the skull in many recent reptiles and in developmental stages of Sphenodon
(fig. 3) as well as in the adult. The complex, as reconstructed in Pristerodon,

ASCENDING
PROCESS

\ PTERYGOID

PALATOQUADRATE BAR

Fic. 3. Sphenodon punctatus. Embryo. Lateral view of skull. (After Howes
& Swinnerton, 1901.)

404 ANNALS OF THE SOUTH AFRICAN MUSEUM

shows a remarkable resemblance to the palatoquadrate of Zonurus (Condylus),
as featured by Broom (1925), and of the 15-2 mm. developmental stage of the
Egyptian lizard Tropzocolotes tripolitanus (fig. 2), described by Kamal (1960).
Similarly the relationship between the horizontal limb and the quadrate
ramus of the pterygoid in Pristerodon is mirrored by conditions found in Sphenodon,
the base of the palatoquadrate cartilage occupying the dorsal and dorso-lateral
surface of the quadrate ramus of the pterygoid in both forms.

In this connection it is important also to consider the relationship of the
nerves to the complex, and especially that of the trigeminal nerve to the
epipterygoid. In recent reptiles the nerve emerges from the braincase through
the incisura prootica. The profundus branch then passes medially to the
epipterygoid while the maxillary and mandibular rami pass behind the epiptery-
goid. In Pristerodon there are no indications of the routes followed by the
presumed branches of the trigeminal nerve but judging by the nerve foramina
and grooves found in some early gnathostomes and tetrapods it would seem
indicated that present-day relationships had an early origin.

QUADRATE RAMUS
OF EPIPTERYGOID

LONGITUDINAL
GROOVE

ASCENDING
RAMUS OF
EPIPTERYGOID

BULGE

POSITION OF QUADRATE

QUADRATE RAMUS OF PTERYGOID

AREA OF ATTACHMENT
OF STAPES

Fic. 4. Pristerodon buffaloensis. Medial view of palato-
quadrate complex. Left side.

In the phylogenetic review that follows, it will be seen that the basic
form of the palatoquadrate is retained from the early gnathostome condition
up to the early ‘stem reptile’ stage. The number of ossifications in the palato-
quadrate would seem to vary from stage to stage and even within stages and
could possibly be of minor evolutionary importance, especially as cartilaginous
inter-connections are indicated. If this is so it might explain the apparent
inconsistency of an otic process on the epipterygoid ossification in certain
labyrinthodonts while in other labyrinthodonts and later forms it forms part
of the quadrate ossification. After the attainment of the basic reptile stage, or
possibly even already during the labyrinthodont stage the palatoquadrate

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA 405

begins to undergo evolutionary change which will have far reaching effects.
The development is channelled into two main streams; one leading towards
the expansion of the epipterygoid and reduction of the quadrate, as exemplified
in those trends showing mammalian affinities, and another resulting in the
retention of the quadrate and reduction of the epipterygoid in trends showing
reptilian affinities.

PHYLOGENY

The first adequately known vertebrates are agnathous forms found in
the Late Silurian. Jawed vertebrates have as yet not been discovered in the
Silurian but the variety of these forms in the Lower Devonian indicates they
must have been undergoing development in the Silurian (Romer, 1955).

PLACODERMI

The early Devonian gnathostomes are predominantly placoderms.
Although widely varied in appearance, the jaw apparatus is usually of a
relatively primitive type. Amongst them the acanthodians are generally
regarded as the earliest and most archaic and their morphology could, therefore,
throw light on the problems connected with the formation and evolution of
the primitive jaws.

According to Watson (1937) the acanthodian palatoquadrate is large
and superficially divisable into two elements, a short horizontal suborbital or
palatal portion and a large vertical postorbital or paraotic portion. In the
genera Climatius and Cheiracanthus the palatoquadrate is ossified as a single
unit, but in Mesacanthus and Ischnacanthus the palatal and paraotic portions
are ossified independently. In Acanthodes (fig. 5), the last surviving member of
the group, and therefore possibly specialized, the palatoquadrate is ossified
as three separate structures, but it seems certain that these bones, in life,
comprised parts of a single palatoquadrate. Anteriorly the paraotic plate
ends in a vertical border, which forms the hind margin of the orbit. From the
top of this border the bone curves downwards posteriorly to end in the thickened
quadrate condyle. Behind the vertical border the paraotic plate is perforated
by a foramen, extremely large in Cheiracanthus, which could possibly have
served for the exit of the maxillary and mandibular branches of the trigeminal.
The palatal portion of the palatoquadrate terminates anteriorly at a point in
line with the middle of the orbit. This is some distance behind the front end
of Meckel’s cartilage and would appear to indicate that the palatoquadrate
was continued forward as cartilage.

The palatoquadrate does not seem to have contact with the neurocranium
in Climatius, but in Mesacanthus, Cheiracanthus and Acanthodes, the paraotic
flange bears an otic process which articulates with the skull behind the post-
orbital process. The palatal part of the bone has a basal articulation.

3

406 ANNALS OF THE SOUTH AFRICAN MUSEUM

OTIC PROCESS POSTERIOR BONE

ANTERIOR (PONE... OF .PALATORUABRATE IN PALATOQUADRATE
IN PALATOQUADRATE

my

°
fi

of w *.
alin

\
x

? CARTILAGINOUS earn rue

PALATOOUAPRATS VAL Jo cee

—[S= ———— —=

Fic. 5. Acanthodes sp. Reconstruction of the skull from specimens from the Lebach
ironstones. Complete except for squamation. (After Watson, 1937.)

According to Watson (1937) the palatoquadrate in arthrodires is directly
comparable with that occurring in several acanthodians. In Pholidosteus the
palatoquadrate has two ossifications, an anterior which articulates with the
ventral part of the neurocranium just behind the olfactory capsules, and a
posterior quadrate.

CROSSOPTERYGII

In the Crossopterygii the processes and future subdivisions of the
palatoquadrate become more apparent.

In Eusthenopteron the palatoquadrate is a single ossification with, according
to Jarvik (1954), five distinguishable parts; an anterior pars autopalatina,
articulating with the ethmoidal region and the anterior neural endocranium;
a basal process articulating with the basipterygoid process; an antero-dorsally
directed ascending process which articulates with the suprapterygoid process
of the neural endocranium; a paratemporal process, which articulates with
the anterior end of the otic shelf and a thickened pars quadrata posteriorly.
A thin film of bone, dotted with large fenestrae, makes up the rest of the
palatoquadrate (fig. 6).

Jarvik states that the profundus passed medially to the ascending process
while the maxillary and mandibular branches of the trigeminal probably
passed through the notch between the ascending and paratemporal processes.

Osteolepis macrolepidotus similarly possesses a completely ossified palato- —

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA 407

RATE

ae cae ea See

- a i = ey By

WS SSS 7y

ECTO- Jj ON ee i

PTERYGOID a
PTERYGOID

Fic. 6. Eusthenopteron foordi. Lateral view of skull with dermal bones
removed. (After Jarvik, 1954.)

quadrate (Watson, 1954). But in Megalichthyes (Watson, 1925), the palato-
quadrate is ossified as a continuous series of bones. Of these the anterior one,
which extends into the basal process, represents the epipterygoid, the posterior
the quadrate.

LABYRINTHODONTIA

The Coal Measure embolomerous anthracosaurs Palaeogyrinus decorus and
Eogyrinus altheyi (Panchen, 1964) both display well developed palatoquadrates.
In Palaeogyrinus (fig. 7) a large facet forms the anterior termination of a massive
buttress, shaped like the mouth of a horn, which tapers forward as the palatal
ramus. The latter appears to have rested on the horizontal palatal ramus of
the pterygoid, but its lower edge cannot be made out. In mesial view the
columella cranii appears as a slender rod extending dorsally until it expands
to form an anteriorly directed process and a second vertical process. Panchen
believes that the anteriorly directed process, the dorso-mesial surface of which
is grooved, formed the dorsal edge of a notch for the profundus and that the
second process probably divided the maxillary and mandibular branches of the
trigeminal nerve. In lateral view the columella is continuous with a sheet of
bone covering the whole of the upper half of the quadrate ramus of the pterygoid
and which runs from the level of the cranial base to the quadrate condyle. A
similar expansion is present in Edops (Romer and Whittier, 1942). Panchen
believes that the epipterygoid may have extended as cartilage further down
the quadrate ramus of the pterygoid and there seems also to have been a
broad process projecting downward and backward from the level of the basis
cranil. There is also a complex articulation for the basipterygoid process of
the basisphenoid formed by both epipterygoid and pterygoid.

The quadrate is small in Palaeogyrinus but is a much more extensive
ossification in Edops.

Panchen states that the considerable ossification in the cartilaginous
palatoquadrate shown by these forms, and probably by Eogyrinus must be

408 ANNALS OF THE SOUTH AFRICAN MUSEUM

SPHENETHMOID PARIETAL

7:
4:

\

(a fi{(7
fe [Ce

Ae
hae

a a >
SNe <1) ae ae
x ~~ ~ ZZ o%

SS
£2
“a \ N77 PTERYGOID
\

Fic. 7. Palaeogyrinus. Lateral view of suspensorial region as though
sectioned through a plane parallel to the quadrate ramus. (After
Panchen, 1964.)

interpreted as a primitive condition and that a progressive reduction of the
epipterygoid ossification is to be seen in later labyrinthodonts.

Although Triassic labyrinthodonts are not on the phylogenetic line
leading to reptiles, the structure of the palatoquadrate is nevertheless of
interest. It consists of two parts, a horizontal cartilaginous anterior part and
a mainly vertical ossified posterior part (Sushkin, 1899; Watson, 1919 and
1926, and Save-Séderbergh, 1936). The vertical part had, at least in Lyrocephalus,
basal, ascending and otic processes. Save-Séderbergh maintains that all
these processes were embodied in the epipterygoid in the most completely
ossified individuals, but that in Aphaneramma and Platystega the ascending
process only was included in the epipterygoid. The epipterygoid was joined to
the quadrate ossification by means of a cartilaginous link which rested in a
groove of the ramus of the pterygoid.

We may therefore assume that the palatoquadrate survived in laby-
rinthodonts complete from the quadrate to at least the front end of
the epipterygoid.

COTYLOSAURIA

Whether or not the seymouriamorphs are considered amphibians or
reptiles, the skull of primitive forms such as Seymouria and Kollassia show that
they are not far removed from the anthracosaurian amphibians from which
the seymouriamorphs and more advanced reptiles have been derived (Romer,
1956). The epipterygoid, which is not well known and seems to be slow to

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA 409

ossify, extends as a rod-like structure to the skull roof. The primitive quadrate,
when well ossified, ran forward to meet the epipterygoid. When the quadrate
is less fully ossified, a cartilaginous area may have bridged the gap between
the quadrate and the epipterygoid (fig. 8). It is presumed that the tendency
for the epipterygoid to remain unossified is a degenerate feature (Romer, 1956).

The earliest known reptiles date from the Carboniferous but knowledge
of their anatomy is fragmentary. Abundant forms appear in the Upper
Carboniferous and Lower Permian but as these include not only primitive
forms but also more advanced types, it is certain that a very important section
of the early history of the reptiles is still unknown.

In diadectids both epipterygoid and quadrate are greatly developed.
Diadectes itself features a quadrate of considerable height and width but little
length (Romer, 1956). In the South African pareiasaurs both epipterygoid and
quadrate are plate-like. The epipterygoid, which has a slender ascending
process, furthermore stands transversely on the quadrate ramus of the pterygoid
near its articulation with the basipterygoid process (Boonstra, 1934). According
to Romer (1956) the margins of the base of the epipterygoid in pareiasaurians
are unfinished, indicating a cartilaginous connection with the quadrate which
has a similarly unfinished surface facing towards the epipterygoid.

The captorhinomorphs Limnosceles, Captorhinus and Labidosaurus from the
Lower Permian of the American Southwest are among the oldest cotylosaurs

EPIPTERYGOID

*BASI-
. OCCIPITAL

PTERYGOID Mei. a

CARTILAGINOUS PORTION OF
PALATOQUADRATE RECONSTRUCTED QUADRATE

Fic. 8. Seymouria. Lateral view of the skull with dermal bones removed.
Cartilaginous portion of palatoquadrate reconstructed. (After Romer,
1956.)

410 ANNALS OF THE SOUTH AFRICAN MUSEUM

of which we have detailed knowledge. Here the quadrate also consists of a
vertical sheet of bone with an essentially flat outer surface and an epipterygoid,
separated from the quadrate but presumably attached to it in life by cartilage,
with a rod-like ascending process (fig. 9).

WII’ NYMR

EPIPTERYGOID

Z

a ae ae -——.
wea = ~~

7? CARTILAGINOUS
PALATOQUADRATE

27 PTERYGOID QUADRATE

Fic. 9. Captorhinus. Lateral view of the skull with dermal bones removed.
Cartilaginous portion of palatoquadrate tentatively indicated. (After
Romer, 1956.)

Although we have no knowledge of the epipterygoid in the aberrant
early Permian form Bolosaurus (Watson, 1954), the fact that the inner part of
the quadrate continues forward as a ramus, on the outer surface of the pterygoid,
would seem to indicate that the palatoquadrate was basically the same as the
general cotylosaurian type.

The Lower Triassic form Procolophon belongs to the last group of survivors
of the cotylosaurs. In this form the epipterygoid has an expanded footplate, the
anterior extension of which is more fully developed than the posterior portion.
From the posterior portion of the footplate a slender ascending ramus extends
dorsally. The quadrate is large and has a process extending forward which
partially replaces the quadrate ramus of the pterygoid. In Procolophon the
latter is reduced in length as well as in height.

PELYCOSAURIA

If, as is generally accepted, the Pelycosauria represent an early stage in
mammalian history and that they possess ‘. . . many archaic features which

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA A411

illustrate the structure of the primitive reptilian stock’ (Romer & Price,
1940: 1), then it should not be surprising to find that this group still displays
a palatoquadrate complex in which the epipterygoid and quadrate portions
are linked. This condition would merely represent the retention in this group
of the basic gnathostome condition of a unified palatoquadrate as exemplified
in the placoderms, and retained, with modifications, in the crossopterygians,
labyrinthodonts and probably also the early stem reptiles.

The epipterygoid has a wide base closely applied to the lateral surface of
the anterior portion of the quadrate ramus of the pterygoid (fig. 10). Anteriorly
a slender extension curves down along a groove on the dorsal surface of the
palatal ramus of the pterygoid. A continuation of this groove in some cases
beyond the anterior end of the bone suggests a further cartilaginous extension
of the palatoquadrate. The slender rod-like portion of the epipterygoid gains
contact, in Dimetrodon, with the anterior surface of the paroccipital process.

EPIPTERYGOID

FOOTPLATE OF %
EPIPTERYGOID PTERYGOID

Fic. 10. Dimetrodon limbatus. Lateral view of skull with dermal bones
cut off to show palatoquadrate. (After Romer & Price, 1940.)

The quadrate is essentially a triangular plate varying greatly in extent in
the few forms where it is adequately known. In some cases in which it is
well developed it appears to gain contact with the epipterygoid along the
dorsal margin of the internal surface of the pterygoid (fig. 10), while in others
in which the bone is more restricted in size, it appears to have been continued
forward as cartilage.

! THERAPSIDA
Dinocephalia

Of the slightly more advanced mammal-like reptiles, the therapsids, the
Dinocephalia are generally regarded as the oldest and in many respects the
most primitive suborder. In them, according to Boonstra (personal communica-
tion), the epipterygoid is small or reduced except in the titanosuchid genus

412 ANNALS OF THE SOUTH AFRICAN MUSEUM

Anteosaurus. In the latter the bone extends right up to the skull roof, has an
expanded waist and a relatively long anterior footplate. In Jonkeria (titano-
suchid) the footplate is also extended anteriorly but dorsally the epipterygoid
only reaches halfway to the roofing bones while the footplate as well as the
dorsal ramus are reduced in the tapinocephalid Struthiocephalus. In none of
the Dinocephalia do we find evidence of a posterior extension to the footplate
which could have formed a link between the epipterygoid and the quadrate.
According to Watson (1914) the Dinocephalia, as a group, has a very special
importance because alone amongst the therapsids it retained a large quadrate.
This feature as well as the absence of the quadrate ramus of the epipterygoid
indicates affinities with the reptilian rather than the mammalian lines of
evolution.

Anomodontia

The anomodonts are generally regarded to be an aberrant group of
mammal-like reptiles with but weak mammalian affinities. This is borne out
by the structure of the palatoquadrate complex in this group. As in the Dino-
cephalia the anomodonts Kannemeyeria erithrea (Case, 1934), Dicynodon kolbe
(Broom, 1932), D. sollast (Watson, 1948), Lystrosaurus murray: (Broom, 1932),
Daptocephalus leoniceps (Ewer, 1961), Kingoria nowacki (Cox, 1959), Dicynodon
grimbeeki and Pristerodon buffaloensis, all display an epipterygoid with a long,
thin dorsally projecting columellar portion, reminiscent of recent reptiles
(see later). However, in contrast to the latter the base of the epipterygoid is
expanded, to varying degrees, in all of the above, extending for some distance
along the upper edge of the quadrate ramus of the pterygoid. In Kannemeyeria
the base extends from a point close to the quadrate to beyond the front of the
interpterygoid space.

In the anomodonts investigated, there is no direct contact between the
epipterygoid and the quadrate (fig. 11), but there is a distinct probability
that a cartilaginous link existed in life in some of these forms, as has been shown

PTERYGOID ° QUADRATO-
JUGAL

EPIPTERYGOID
QUADRATE

Fic. 11. Dicynodon sp. Lateral view of skull.

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA 413

for Pristerodon. However, the general tendency within the group is for the
bar, linking the epipterygoid to the quadrate portion, to become reduced;
an evolutionary trend, as will be shown later, that was present also in the line
that culminated in modern reptiles.

Gorgonopsia

In all known Gorgonopsia the epipterygoid is high and relatively narrow,
with a well-developed footplate. In ‘Lycaenodon’ (fig. 12), Scymnognathus, Lepto-
trachelus, Cynariops (Boonstra, 1934a) and Aelurognathus (Haughton, 1924), the
footplate has a long tapering anterior extension, with a short posterior extension
ending some distance away from the quadrate. Boonstra found no evidence
in this group of a widening of the vertical portion of the epipterygoid.

EPIPTERYGOID
PROOTIC

FOOT PLATES

_——<

Fic. 12. ‘Lycaenodon’. Lateral view of skull with dermal bones cut away.
(After Boonstra, 19344.)

Therocephalia

In the early Tapinocephalus Zone forms such as the pristerognathids
Scylacosaurus and Scymnosaurus (Boonstra, 1934b, 1954), and most of the primitive
therocephalians, the vertical portion of the epipterygoid is still relatively
narrow, but in Glanosuchus macrops (Boonstra, 1954) the epipterygoid has
become dumb-bell-shaped, although still not much widened. The base has
no noteworthy posterior process. However, in the lycosuchid Trochosaurus
major (Boonstra, 1934b) occurring in the same zone, the epipterygoid has

414 ANNALS OF THE SOUTH AFRICAN MUSEUM

developed into a large, broad plate which obscures the lateral opening into
the pituitary fossa and the foramina for the Vth and VIth nerves. The epiptery-
goid here is more than twice as broad as in any known gorgonopsian and has
the appearance of a cynodont epipterygoid. It differs from that known in
Scymnosaurus and Scylacosaurus in having an expanded upper end in addition
to a wide footplate, a condition reaching its greatest development in the later
whaitsids (Boonstra, 1934).

Of the Cistecephalus Zone Therocephalia, Euchambersia mirabilis (Boonstra,
1936) also shows the dorsally and ventrally expanded condition (fig. 13) but
the shaft of the bone does not show much expansion. In the scaloposaurid
Ictidosuchops intermedius (Crompton, 1955), however, the shaft is a broad one,
and ends in a slightly more expanded dorsal portion which fits into a shallow
depression in the antero-dorsal wall of the prootic, consequently forming
part of the true lateral wall of the braincase. The anterior extension of the
footplate is long, but the posterior one is short. At the juncture of the posterior
extension and the ascending ramus there is a shallow notch which most probably
housed the maxillary or both the maxillary and mandibular branches of the
trigeminal nerve (Crompton, 1955).

EPIPTERYGOID

QUADRATE RAMUS OF PTERYGOID

Fic. 13. Euchambersia mirabilis. Lateral view of skull with dermal bones
cut away. (After Boonstra, 1936.)

Fic. 14. Aneugomphius ictidoceps. Lateral view of skull with outlines of
palatoquadrate bones indicated. (After Brink, 1956.)

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA 415

In Theriognathus microps (Boonstra, 1934), Notosollasia laticeps (Boonstra,
1934) and Aneugomphius ictidoceps (Brink, 1956), as in all known whaitsids, the
epipterygoid is broad and flat with greatly expanded dorsal and ventral ends
(fig. 14). The base is long and in most forms posteriorly overlaps the prootic
obscuring the incisura prootica. The hind wall of the epipterygoid in Aneugomphius
shows two deep notches, probably for the exit of the maxillary and mandibular
branches of the trigeminal nerve.

Cynodontia

Brink (1960) states that the epipterygoids of the primitive cynodont
Scalopocynodon gracilis are *. . . Cynodont-like in being broadly expanded, but
Therocephalian-like in their feeble overlap of the prootics’ (p. 145). The
anterior extension of the footplate is long and broad while the remainder of
the ventral margin follows the dorsal margin of the quadrate ramus of the
pterygoid (fig. 15). Brink made the interesting observation that the anterior
margin of the epipterygoid is very thick. This may well support the view that
this portion of the vertical limb is the more stable and is older phylogenetically
than the thinner, posterior section of the limb. The quadrate is in the shape of a
wedge lodged loosely in a cavity anteriorly in the squamosal. The anterior
margin is straight, blunt and vertical, while the posterior margin is sharp
and broadly curved. The quadrate is a much smaller bone than the epipterygoid
and is well separated from the latter.

In Thrinaxodon lorhinus (Parrington, 1946) the epipterygoid is greatly
expanded and dorsally is broadly in contact with the prootic. The quadrate
ramus of the epipterygoid joins the prootic, but does not reach the quadrate,
leaving a large foramen between the epipterygoid and prootic for the maxillary
and mandibular rami. The quadrate, which is about one-third the size of
the epipterygoid, has a wide, dumb-bell-shaped articulating surface and a
thin, slightly curved dorsal process which medially is greatly strengthened by
a pillar-like development which continues to the apex of the bone.

In an unidentified cynodont, which compares closely with Trirachodon,
Parrington (1946) found two foramina between the epipterygoid and prootic.
A groove comparable with that leading into the single foramen in Thrinaxodon,
leads into the lower, larger foramen from below and behind. The dorsal
smaller foramen is probably new. Parrington states that the significance of this
development lies in the possibility that the maxillary left the skull by the
upper foramen, which is apparently mainly enclosed by the epipterygoid, and
is equivalent to the foramen rotundum, and the mandibular by the lower,
larger foramen, the equivalent of the foramen ovale. The size of both foramina
indicates that blood vessels accompanied the nerves.

In Diademodon mastacus (Brink, 1955) the foramen for the trigeminal is
also divided into two, although incompletely in this species (fig. 16). The
foramen lies slightly above and behind the pituitary fossa. The epipterygoid
itself covers the anterior half of the prootic. The footplate is extensive and

416 ANNALS OF THE SOUTH AFRICAN MUSEUM

extends forward as a very thin tapering process while it reaches the quadrate
with its posterior extension.

In the cynognathids, the thin plate-like epipterygoid overlaps the anterior
superior process of the prootic. The footplate of the base is extensive. It sends
back a posterior extension which occupies the position of the quadrate ramus
of the pterygoid and which reaches the relatively reduced quadrate.

QUADRATE RAMUS
OF PTERYGOID

Fic. 15. Scalopocynodon gracilis. Lateral view of skull with dermal bones
cut away. (After Brink, 1960.)

EPIPTERYGOID

QUADRATE RAMUS OF PTERYGOID

Fic. 16. Diademodon mastacus. Lateral view of skull with dermal bones
cut away. (After Brink, 1955.)

'\ EPIPTERYGOID

eS See
5 ee Se
ic = = ——
Hae \ -— \\ \\ Se NOTCH
“¢ , SS sya yt sl Pg ele \\ Av FOR V2 & V3
\

(Ee: ==. geal \ a Va
SS Li ak
PTERYGOID QUADRATE RAMUS

OF EPIPTERYGOID

Fic. 17. Diarthrognathus broomi. Lateral view of skull. (After Crompton,
1958.)

EPIPTERYGOID —ALISPHENOID TRANSITION IN THERAPSIDA 417

Ictidosauria

In Diarthrognathus broom: the dorsal portion of the epipterygoid is extremely
broad and overlaps the prootic above the prootic incisure. Here the posterior
edge of the epipterygoid is slightly indented for the maxillary and mandibular
branches of the trigeminal nerve (fig. 17). Crompton (1958) states that although
it is difficult to ascertain, it would appear that the quadrate ramus of the
epipterygoid meets the antero-medial edge of the quadrate. The quadrate is
much reduced, S-shaped in occipital view, and the articular surface is markedly
concave. This latter feature, according to Crompton, is of great significance as
in all known therapsids, including Bzenotherium and Oligokyphus, the articular
face of the quadrate is either flat or convex, but never concave. It is also
significant that the mammalian malleus (articular) possesses a convex articular
surface which meets a convex articular face in the incus (quadrate).

In Bienotherium (Hopson, 1964) the epipterygoid and the anterior part of
the prootic together form the side wall of the brain case (fig. 18). The quadrate
ramus of the epipterygoid is more vertically orientated and deeper than in

EPIPTERYGOID FRONTAL PARIETAL

eRe eg PROOTIC

ORBITAL
FISSURE

Fic. 18. Bienotherium yunnenense. Lateral view of braincase reconstructed
from serial sections. (After Hopson, 1964.)

418 ANNALS OF THE SOUTH AFRICAN MUSEUM

the cynodonts and could have reached the quadrate. The ascending ramus is
broad and high with a posterior border which is overlapped laterally, above
the trigeminal foramen, by the prootic. This latter is the reverse of the usual
reptilian condition in which the prootic lies medial to the epipterygoid, and
is ascribed by Hopson to the development of a forward extension of the anterior
portion of the prootic in such a way that it lies lateral to the cavum epiptericum.
In Bienotherium both this extension—the ventro-lateral flange —and the antero-
dorsal border of the prootic extend well forward of the prootic incisure, an
observation which has led Hopson to conclude that the prootic component of
the trigeminal foramen in this form is not strictly homologous with the prootic
incisure in cynodonts, because the former is merely a notch in the anterior
border of the ventro-lateral flange. This conclusion seems to be supported by
the fact that the trigeminal foramen pierces the side wall of the skull anterior
to the depression of the semilunar ganglion. It also indicates that the maxillary
and mandibular branches extend in an anterior direction through the cavum
epiptericum in Bzenotherium, whereas normally, in those forms with broadened
epipterygoids, they pass in a posterior direction.

The trigeminal foramen is slightly constricted at mid-length presumably
indicating the incipient subdivision of the single trigeminal foramen into the
separate foramina rotundum and ovale of mammals.

TRICONODONTA

In a group of Mesozoic mammals, the Triconodonta, the brain case has
an essentially reptilian structure. According to Kermack (1963) Triconodon
mordax and Trioracodon ferox still possessed a cavum epiptericum lying outside
the ossified lateral wall of the braincase (formed by the petrosal) and as in
Morganucodon and possibly all pre-Cretaceous mammals, the alisphenoid
formed the lateral boundary of the cavum. The ramus profundus passed out
through the anterior end of the cavum epiptericum while the maxillary and
mandibular nerves passed out through two foramina formed in the posterior
edge of the alisphenoid.

According to Kermack, the semilunar ganglion in Morganucodon lay inside
the primary wall of the braincase and the mandibular branch of the trigeminal
nerve passed though the foramen pseudovale in the anterior lamina of the
petrosal. The lamina extends much further forward in Morganucodon than it
does in cynodonts. The maxillary branch of the fifth nerve presumably left
the braincase, passing through the notch at the front end of the anterior
lamina. After crossing the cavum epiptericum both branches would have left
it by passing behind or through the alisphenoid in some way reminiscent of
cynodonts.

In basic construction the braincase of the triconodonts differs from that
of an advanced therapsid only in the narrower cavum epiptericum in the
former, a difference due to the relatively larger size of the brain in mammals.

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA 419

LiviING REPTILES AND MAMMALS

Amongst recent reptiles Sphenodon has virtually retained its larval palato-
quadrate in the adult (compare figs 3 and 1g) but in the lizards Lacerta
(Gaupp, 1906), Cordylus, Eremias and Mabuia (Broom, 1903), Tropiocolotes
(Kamal, 1960), Varanus (Frazetta, 1962), Tupznambis (Jollie, 1960) and many
others, and in the chelonians Emys (Kunkel, 1912) and Chrysemys (Shaner,
1926), the palatoquadrate complex undergoes considerable changes during
ontogeny. Ossifications within the complex culminate in the formation of
widely separated epipterygoid and quadrate portions, while the rest of the
palatoquadrate is resorbed. The quadrate is retained as a comparatively
strong element while the epipterygoid is whittled down to a narrow rod-like
structure (fig. 20). The base of the bone is not expanded. It has, however, a
cartilaginous epiphysis, fitting into the fossa pterygoidei which may be linked
with the quadrate through a strand of connective tissue, both epiphysis and
connecting strand probably representing remnants of the palatoquadrate
cartilage.

EPIPTERYGOID PROOTIC

QUADRATE

Fic. 19. Sphenodon punctatus. Lateral view of the skull with dermal
bones cut off to show palatoquadrate.

Pp QUADRATE
TERYGOID SHOBTIC

QUADRATE RAMUS OF PTERYGOID
Fic. 20. Tupinambis nigropunctatus. Lateral view of skull. (After Jollie,
1960.)

420 ANNALS OF THE SOUTH AFRICAN MUSEUM

Further reduction of the epipterygoid has taken place in the lizards
Agama, Lyrwcephalus and Calotes (Ramaswami, 1946), Ophioceps and Anniella
(Jollie, 1960), Phrynocephalus (Siebenrock, 1895), Physignatus, Chlamydosaurus
and Amphibolurus (Beddard, 1905), where the epipterygoid is very short.
However, in the Chamaeleontidae, Dibamidae (Boulenger, 1887) and
apparently most Amphisbaenidae (known only in Trogonophis, Bellairs, 1950),
the epipterygoid is absent. In the Chelonia the epipterygoid is still present
but very small (Parker, 1880) while it is much reduced or vestigial in Ophidia
and Crocodilia, although the embryos of crocodiles still show the ascending
process (Parker, 1883; Shiino, 1914). In birds it appears to be absent (Goodrich
1930), the quadrate being the only part of the palatoquadrate to be retained
in the adult. :

Where present the epipterygoid lies lateral to the lateral head vein,
postero-lateral to the profundus and antero-medial to the maxillary and
mandibular branches and the orbital artery.

In mammals the alisphenoid appears to be lost in Echidna. In this feature,
as will be seen later, Echidna probably shows reptilian affinities. In Dasyurus
the dorsal end of the ala temporalis fuses with the orbital cartilage, but in all
other mammals, as far as is known, the ala temporalis ends freely (De Beer,
1937). In Didelphis the alisphenoid lies between the profundus and maxillary
rami, as in reptiles. In Trichosurus, Mus, Mustela and many others (De Beer,

ALISPHENOID / =
ORBITAL FISSURE FORAMEN OVALE
FORAMEN ROTUNDUM

Fic. 21. Felis domestica. Lateral view of skull with jugal arch cut away.

1937) the alisphenoid is pierced by the maxillary ramus (foramen rotundum),
while in Felis (fig. 21), and the majority of mammals the mandibular ramus
too may pass through it (foramen ovale). In the fourth group the maxillary
emerges freely in front of the alisphenoid (De Beer, 1937).

Edinger and Kitts (1954) state that the mandibular branch of the trige-
minal appears to have been variable in its relations to the alisphenoid in
living mammals as the foramen ovale is absent in some of them. Several

EPIPTERYGOID —ALISPHENOID TRANSITION IN THERAPSIDA 421

genera of extant Perissodactyla and Artiodactyla lack the foramen but in
both these orders a separate foramen ovale was the usual condition in the
early Tertiary. Comparison of fossil and recent material suggests a similar
trend in certain families of the Rodentia and Insectivora. They relate the
formation of the foramen ovale with an earlier palaeoneurological development
when the mandibular nerve came to branch off the trigeminal stem intra-
cranially, during or near the origin of mammals. It is only in some later
forms that the nerve used the foramen lacerum for its passage from the brain-
case. This probably indicates that its absence is a secondary condition and
possibly reveals an evolutionary trend within the mammals to abolish the
separate exit of the mandibular nerve.

(CONCLUSIONS

The evolutionary history of the palatoquadrate has been followed, as
far as the availability of information on fossil material permits, from the
early gnathostome condition to that in recent reptiles and mammals. It has
been shown that the primitive palatoquadrate was a solid structure, consisting
mostly of several ossifications, with four main processes for its attachment to
the neurocranium. These are the otic, ascending, basal and pterygoid processes.
In tetrapods the ossifications are reduced to two.

During its evolutionary history the palatoquadrate shows a tendency to
become reduced in size, the pterygoid process shortens considerably while the
area between the otic and ascending processes becomes deeply excavated
thereby accentuating and demarcating the epipterygoid and quadrate portions.
The demarcation of these two elements becomes more pronounced as evolution
progresses.

Conditions in recent and fossil forms show that the palatoquadrate of
both recent reptiles and mammals are deducable from the early gnathostome
condition and that they evolved through the placoderm, crossopterygian,
labyrinthodont and early stem reptile stages. In the groups leading to or
showing affinities with the lines leading to modern reptiles, the trend is towards
the progressive reduction of that part of the palatoquadrate anterior to the
quadrate. In most reptiles the epipterygoid is the only part of this area to be
retained but in some recent forms even this too disappears. In those lines
leading towards mammals the opposite occurred and regression of the quadrate
took place while the epipterygoid expanded (fig. 22).

From the foregoing it is clear that the phylogenetic and ontogenetic
evidence do not support the view that the epipterygoid in the primitive reptile
possessed a restricted base. The latter condition is a specialized reptilian one
as is proved by conditions obtaining in adult forms of many recent reptiles.
It is a stage in the gradual reduction of the epipterygoid in this group. Olson’s
(1944) views on the evolutionary development of the epipterygoid, must then
differ from those given here.

422 ANNALS OF THE SOUTH AFRICAN MUSEUM

S PLACODERM STAGE

TD CROSSOPTERYGIAN STAGE
\
LABYRINTHODONT STAGE

etd COTYLOSAURIAN STAGE

LINES SHOWING LINES SHOWING

REPTILIAN AFFINITIES MAMMALIAN AFFINITIES
a

‘
EVOLUTIONARY | EVOLUTIONARY

TREND

Za sae

ee 7 ares
an. 4 A Ao ie

GORGONOPSIAN
PROCOLOPHONIO

DICYNODONT
Ki
'
LIZARD !

SPHENODON

CYNODONT

MAMMAL
CHAMELEON

Fic. 22. Schematic presentation of probable evolutionary trend in development of palatoquadra
leading towards recent reptiles and mammals.

EPIPTERYGOID—ALISPHENOID TRANSITION IN THERAPSIDA 423

Although the therapsids show a marked degree of variability in the struc-
_ ture of the epipterygoid, we find that the groups which show mammalian
affinities have also retained certain basic features of the early tetrapod palato-
_ quadrate complex. The most noticeable of these is the retention of the extensive

base of the epipterygoid, indicating that a considerable portion of the base of
_ the palatoquadrate anterior to the quadrate, of the early labyrinthodont and
stem reptile, had been retained.

Two further features are closely linked with the development of the
alisphenoid in mammals. Firstly the expansion posteriorly of the rod-like
ascending process of the epipterygoid and secondly, the increase in size of
the brain in forms approaching the transitional stage.

Backward extension of the ascending process would result in the maxillary
and mandibular branches of the trigeminal nerve being pushed backward by
the epipterygoid as the latter progresses past the incisura prootica through
which the trigeminal emerges.

The increase in size of the brain in mammals and the consequent lateral
expansion of the braincase, could conceivably result in the nerves becoming
trapped between the posteriorly extending epipterygoid and the expanding
braincase. Close contact between the epipterygoid and braincase could force the
nerves to seek an exit through the epipterygoid. Whether this results in the
formation of a notch only, a foramen or two foramina depends largely on the
separate development of the epipterygoid and prootic, the way these bones
make contact and where this contact is made. In this way the maxillary ramus
can emerge through the foramen rotundum while the mandibular emerges
farther back through its own foramen ovale or merely through a notch in the
posterior portion.

If the expansion of the braincase is continued anteriorly the cavum
epiptericum would be obliterated, as it has been in mammals. This could
mean that the pila antotica, which forms the inner wall of the cavum in reptiles,
could have been pushed against the alisphenoid and as the pila will then
virtually have become redundant, there is no need for it to develop in mam-
mals. The possibility that this could have happened is strengthened by the
fact that isolated cartilaginous fragments of the pila antotica have been found
in many higher mammals, e.g. Lepus (Voit, 1909), Homo (Macklin, 1914),
Felis (Terry, 1917), Didelphys (Toplitz, 1920), Halicore (Matthes, 1921) and
Tarsius (Henckel, 1927).

ACKNOWLEDGEMENTS

I wish to record my gratitude to the South African Council for Scientific
_ and Industrial Research for a research grant during the tenure of which most
of this work was carried out. I should like to thank Professor E. Jarvik, Profes-
sor E. Stensié and the staff of the Palaeontological Institute of the Swedish
Museum of Natural History for their kindness, co-operation and assistance

424 ANNALS OF THE SOUTH AFRICAN MUSEUM

while working in the Swedish Museum of Natural History, Stockholm. My
thanks are also due to Dr. A. S. Brink, Dr. M. E. Malan and Dr. L. D. Boonstra
for reading the manuscript and making valuable suggestions, and Mrs. I.
Rudner for copying some of the drawings used in this paper.

The trustees of the South African Museum are grateful to the Council
for Scientific and Industrial Research for a grant to publish this paper.

SUMMARY

The serial sectioning of the skull of the anomodont Pristerodon buffaloensis
has produced valuable evidence regarding the structure of the palatoquadrate
complex and the evolution of the therapsid epipterygoid. A short review is
given of the structure of the palatoquadrate in the most important known
fossil vertebrate groups.

It is emphasized that the mammalian alisphenoid should not be looked
upon as a transformation of the rod-like epipterygoid, as found in more spe-
cialized true reptiles as Broom and others implied, but that both conditions
are derived from a common ancestral type.

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Ramaswaml, L. S. 1946. The chondrocranium of Calotes versicolor (Daud.) with a description of
the osteocranium of a just-hatched young. Quart. 7. micr. Sci. 87: 237-297.

Romer, A. S. & Price, L. W. 1940. Review of the Pelycosauria. Spec. Pap. geol. Soc. Amer. 28:
i-x, 1-538.

Romer, A. S. 1955. Vertebrate paleontology. Chicago: University Press.

Romer, A. S. 1956. Osteology of the reptiles. Chicago: University Press.

SAVE-SODERBERGH, G. 1936. On the morphology of the Triassic stegocephalians from Spitz-
bergen and the interpretation of the endocranium in Labyrinthodontia. K. svenska Vetensk.
Akad. Handl. (3) 16 (1): 1-181.

SHANER, R. F. 1926. The development of the skull of the turtle, with remarks on fossil reptile
skulls. Anat. Rec. 32: 343-367.

Suno, K. 1914. Studien zur Kenntnis des Wirbeltierképfes. 1. Das Chondrocranium von
Crocodilus mit Beriicksichtigung der Gehirnnerven und der K6épfgefasse. Anat. Hefte
Abt 1. 50: 253-382.

426 ANNALS OF THE SOUTH AFRICAN MUSEUM

SIEBENROCK, F. 1895. Das Skelet der Agamidae. S.B. Akad. Wiss. Wien. Abt. 1. 104: 1089-1196.

Susuxin, P. O. 1899. Schadel von Tinunculus. Nouv. Mem. Soc. Nat. Moscou. 16: 1-163.

TerRRY, R. J. 1917. The primordial cranium of the cat. 7. Morph. 29: 281-433.

T6puitz, C. 1920. Bau und Entwicklung des Knorpelschadels von Didelphis marsupialis. Zoologica,
Stuttgart. 70.

Vortr, M. 1909. Das Primordialkranium des Kaninchens unter Beriicksichtigung der Deck-
knochen. Anat. Hefte. Abt. 1. 38: 425-616.

Watson, D. M, S. 1919. The structure, evolution and origin of the Amphibia— The ‘Orders’
Rachitomi and Stereospondyli. Phil. Trans. (B) 209: 1-73.

Watson, D. M.S. 1926. The evolution and origin of the Amphibia. Phil. Trans. (B) 214: 189-257.

Watson, D. M. S. 1937. The acanthodian fishes. Phil. Trans. (B) 228: 49-146.

Watson, D. M. S. 1948. Dicynodon and its allies. Proc. zool. Soc. Lond. 118: 823-877.

Watson, D. M. S. 1954. On Bolosaurus and the origin and classification of reptiles. Bull. Mus.
comp. Zool. Harv. 111: 297-449.

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TaBLe or Contents and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 4# in. X 7 in. (7$ in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

SmitH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmitH, C. D. 1954. South African Plonias. In Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).
Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:

Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.

When another species has been called by the same name:

[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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N. A. H. MILLARD

_ THE HYDROZOA OF THE SOUTH AND WEST
> GOASTS OF SOUTH AFRIGA. PART III.

THE GYMNOBLASTEA AND SMALL FAMILIES OF
; CALYPTOBLASTEA

MUS. COMP. ZOOL.
LIBRARY

AUG 12 1966

June 1966 Junie UNIVERSITY.
Volume 48 Band
Part 118 Deel

ANNALS OF THE SOUTH AFRICAN MUSEUM

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ji ——

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Sat —CLoyer Towns

THE HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH
AFRICA. PART III. THE GYMNOBLASTEA AND SMALL FAMILIES

OF CALYPTOBLASTEA Petree’ .
WiUS, COMP. eM Na fe

LIBRA
ne ARY
€ »
N. A. H. Mitiarp AUG l2 1O86
Koology Department, University of Cape Town MARY a, O,
UNIVERSIT
(With 15 text-figures and 1 plate) ny
CONTENTS
PAGE
Introduction : : ‘ ‘ = OT
Station List . ; ‘ : : . 428
List of Species. : ‘ ‘ - ‘433
Tubulariidae : ; ‘ ; . 434
Myriothelidae . : ; : . 435
Corynidae . : , ‘ : . 440
Solanderiidae ; 3 ; ' - 444
Bougainvilliidae . : ‘ : - 49
Clavidae_. ‘ ; ; ; » 452
Eudendriidae : 3 : : - 454
Hydractiniidae . : ; é 5. RG
Pandeidae . : : ; s AGI
Aequoreidae ; . ; ‘ =) 461
Lovenellidae : ‘ ; : . 464
Haleciidae . P : : : . 464
Campanulariidae . : : é gaze
Summary . ; : : : 2+ 404
Acknowledgements : ; : . 484
References . ‘ . 5 ; -,) 404
INTRODUCTION

This paper represents the third and final part of a systematic account of
_ the hydroids from that part of the South African coast stretching from South
_ West Africa on the west to the southern border of Natal on the east. For
details on the origin of the collections the reader is referred to the introduction
to part I of the series (Millard, 1962).
: When examining Gymnoblast hydroids one must bear in mind that a
_ considerable quantity of the more delicate material brought up by dredging
_ becomes destroyed by rough handling or is so badly damaged that identification
beyond the genus level is impossible. Moreover most genera require the presence
_ of gonophores, usually female, for identification to species level. This applies
. particularly to such genera as Eudendrium and Tubularia. It is felt that no useful

‘

Ann. S. Afr. Mus. 48 (18), 1966: 427-487, 15 figs., 1 pl.

427

a

428 ANNALS OF THE SOUTH AFRICAN MUSEUM

purpose would be served by including dubious records and, at the risk of
giving a false impression of the abundance, poorly preserved material and
sterile material of doubtful specific identification has generally been omitted.

The author wishes to express thanks to the following: the South African
Museum for permission to examine the collection of the s.s. Pieter Faure, the
Zoology Department of the British Museum of Natural History for accommoda-
tion and permission to examine their collection in 1960, Dr. W. Engelhardt
of the Munich Museum and Dr. J. S. Pringle of the Natal Museum for the
loan of type material, Dr. M. E. Thiel of the Hamburg Museum for identifica-
tion of certain medusae and finally all past and present members of the Zoology
and Oceanography Departments of the University of Cape Town who have
helped to accumulate material in the now very extensive University collection.

Type specimens of new species have been deposited in the South African
Museum and have been given a Museum registered number in addition to the
University catalogue number.

STATION LIST

A. Littoral material from Oudekraal on the west coast of the Cape Peninsula.
Position: 33°58°5'S/18°22-2’E.

Date

A 40 16.1.34
A 116-122 = 15,.3.34
A 139 17-3-34
A 353

A 383 13.5-34
A 384 25.83.34
A 405 18.10.34

AFR. Material dredged by the government research vessel, R.S. Africana.

Date Position Depth (m.) Bottom
AFR 736 17.8.47 30°4.2°4.'S/15°59°2E 201 co gn S, Sh
AFR 801 7.10.47 32°34°4.S/17°52°2'E 71 gn M,R
AFR 866 9.1.48 34°36°8’S/19°16-4’E 38 R,S
AFR 967 23.3.48 35°07'S/20°4.9’E gI £S
AFR 985 5.4.48 34°4.7°4’S/20°19’E © 80 gn M
AFR 994 19 4.48 34.°34°5'S/21°22°5'E 68 co S, Sh
AFR 0002 6.6.49 33°10'S/17°57'E 73

B. Littoral material from Lambert’s Bay on the west coast. Date: July 1938.
Position: 32°05'5/18°18’E.

BMR. Bushman’s River Estuary, south coast, on sand and muddy banks.
Date: 14.9.50. Position: 33°41'S/26°40’E. Depth: 2—4°5 m.

BRE. Breede River Estuary, south coast, littoral. Date: 3.2.52. Position:
34°25,'S/20°51°5’E. |

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 429

CP. Littoral material from various localities on the west coast of the Cape

Peninsula.
Date Locality Position
CP 325 20.9.48 Sea Point 33°55°2'S/18°22°6’E
CP 327 16.12.48 Blaauwberg Strand 33°48'S/18°27-5’E
CP 331 8.8.49 Kommetje 34°08°5’S/18°19°4’E
CP 336 12.5.4.9 Oudekraal 33°58°5'S/18°22-2’E
CP 378 3.6.52 Cape Town docks (on 33°54°5'S/18°25°5’E
Squalus fernandinus )
CP 646 5.12.60 Oudekraal 33°58°5'S/18°22-2’E

CPR. Material from various localities in the Cape Province.

Date Locality Position Depth (m.)
CPR 9 30.4.50 Glentana Strand 34.°04’S/22°20’E littoral
CPR 46 20.6.59 Umgazi Bay 31°43’S/29°26’E 27

HAM. Keiskama River Estuary, Hamburg, on the south coast. Date: 9.1.50.
Position: 33°17'S/27°32’E.

HB. Littoral material from Hondeklip Bay on the west coast. Date: 8.2.40.
Position: 30°19’S/17°16’E.

_ KNY. Knysna Estuary on the south coast. Position: 34°05’'S/23°04’E (average).

Date Depth (m.) Bottom
KNY 30 16.7.47 5-7 M,S
KNY 70 15.7.47 2-6 S
KNY 164 9.7.50 0-3 M
KNY 165 9.7.50 I-13 S
KNY 176 11.7.50 Floating buoy
KNY 212 7.7.60 O-1 Ss
KNY 270 14.2.64 o-I M

L. Littoral material from East London on the south coast. Position: 33°01’S/
27°54'E.

Date
L 44 6.3.37
L 56-177 —.7-37

LAM. Dredged in Lambert’s Bay, west coast.

Date Position Depth (m.) Bottom

LAM. 24 16.1.57 32°04°6’S/18°18-15’E v7 R

LAM 30 19.1.57 32°05°1'S/18°17-7’E 20 R

LAM 35 19.1.57 32°05°5'S/18°17-7’E 27 R, Sh
LAM 46 22.13.57 32°04°4’S/18°17-7’E 23 R

LAM 50 23.1.57 32°08'5’S/18°17°7’E 16 R,S
LAM 52 Q1.1.57 32°04.7'S/18°18-2’E 17 Ss

LAM 59 23.1.57 32°09’S/18°18’E 16 R,S

LB. Langebaan Lagoon, west coast.

Date Position Depth (m.) Bottom
LB 127 26.4.48 33°05°6’S/18°01-6’E Littoral R
LB 166 15.7.46 33°09'0'S/18°03°4’E

7
LB 266 3.5.51 33°07°4’S/18°02+1’E Littoral N)

430 ANNALS OF THE SOUTH AFRICAN MUSEUM

Date Position Depth (m.) Bottom
LB 296 5.5.51 33°04'S/18°00’E Ship’s hull
LB 314 4.5.51 33°05°7'S/18°01°5’E Littoral R
LB 371 7.5.53 33°05°7'S/18°01'5’E Littoral R
LB 378 7.553 33°05°9'S/18°01°9’E O-1 Wooden piling
LB 380 7.5.53 33°06-7'S/18°01:0’E 4 S, Sh
LB 386 8.5.53 33°06°5'S/18°02’E Ship’s hull
LB 398 24.7.53 33°07°1'S/18°02:9’E O-1 S
LB 403 5.12.53 33°10'S/18°03°5’E Littoral M
LB 542 4.5.60 r Littoral
LB 556 22 7,08 33°07°3'S/18°03-0’E Littoral Wooden piling

LIZ. Dredged in Algoa Bay, Port Elizabeth, south coast.

Date Position Depth (m.) Bottom
LIZ 2 5.4.54 33°55°7.9/24°37°2 E 9
LIZ 3 5-4..54 33°56°1'S/25°40’E 17 S
LIZ 7 6.4.54 33°58-1'S/25°38-9’E 9 R, St
LIZ 11 6.4.54 33°57°2'S/25°38-0’E 9 Clay, R
LIZ 13 6.4.54 33°58°2'S/25°38-8’E 7 S
LIZ 16 7.4.54 33°58°4'S/25°40°5’E 14 St
LIZ 24-25 11.4.54 34.°00°4'S/25°44°5’E 39 co S, Sh
LIZ 40 11.4.54 34.°00°8’S/25°42°4’E 6 R

LU. Littoral material from Luderitz Bay, South West Africa.

Date Position
LU 59 24.2.63 26°38’S/15°05'5’E
LU 113-118 = 22.2..63 26°38’S/15°09°3’E

MB. Dredged in Mossel Bay, south coast.

Date Position Depth (m.) Bottom
MB 8-12 12.1.56 34.°04°3'S/22°13°9'E 19 R
MB 19 13.1.56 34.°08-7'S/22°07-2’E 13 R, S, Sh
MB 25 13.1.56 34°09'°3'S/22°10°1’E 29 S
MB 37 16.1.56 34.°09°3'5/22°10°:0’E 31 N)
MB 47 17.1.56 34°11°3S/22°10:0'E 10 R
MB 52 17.1.56 34.°11°0'S/22°09°9’E 14 RS
MB 55 17.1.56 34.°10°7'S/22°09'6’E 9 R
MB 60 18.1.56 34.°04°3'S/22°14:2’E 18 R, co S, Sh
MB 64 18.1.56 34.°04°8/S/22°13°1’E 26 R, co S, Sh
MB 69 19.1.56 34.°08-6'S/22°07°3’E 13 RSs ses
MB 70 19.1.56 34.°08-9’S/22°07-9’E 18 S
MB 81 20.1.56 34°06-2’S/22°10-9’E 27 M
MB 84 21.1.56 34°11°4’S/22°10°1’E 29 R
MB 88 18.1.56 34.°04°8/S/22°13-1’E 26 R, co S, Sh

OLF. Olifant’s River Estuary, west coast. Date: 23.1.55. Position: 31°42’S/
18°15’E.

PP. Littoral material from Paternoster, west coast. Date: 24.9.57. Position:
32°43'S/17°55 E.

SAMH. Material from the collection of the South African Museum. Specimens
147-352 and 361-383 were dredged by the s.s. Pieter Faure. Their positions
were given in the original records as compass bearings off salient points

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 431

on the coast, and were probably not very accurate. These have been
converted into latitude and longitude and are given to the nearest minute.

SAMH 147
SAMH 157-162
SAMH 166-170
SAMH 174-177
SAMH 179-180
SAMH 189
SAMH 202-203
SAMH 214
SAMH 221-227
SAMH 230-233
SAMH 235
SAMH 239-247
SAMH 250-253
SAMH 269-274
SAMH 283
SAMH 296
SAMH 315
SAMH 327-328
SAMH 335-338
SAMH 341-343
SAMH 351-352
SAMH 357
SAMH 361
SAMH 380-383
SAMH 404
SAMH 405-408

Date
23.6.1898
15.7.1898
11.11.1898
19.11.1898
22.12.1898
28.12.1898
7-3-1899
24.3.1899
19.6.1899
20.9.1899
5-7-1900
11.10.1900
15.7.1901
17.7.19OI
25.7.1901
13.8.1901
10.9.1901I
22.9.1904
4.10.1904
19.10.1904
22.8.1905
19.6.1914
11.11.1898
15.3.1899
—.4.1962
—.7.1962

Position

South of Mossel Bay
34°08’S/22°16’E
33°49'S/25°56’E
33°45'S/26°44E
32°52’S/28°12’E
33°09’S/28°03’E
33°59'S/25°51E
33°50'S/26°35’E
34°26'S/21°42’E
34°15'S/22°10°5’E
34°27'S/20°58’E
34°08'S/22°59°5’E
33°13°5'S/27°58’E
33°07'S/27°47°5'E
32°50’S/28°18-5’E
32°47'S/28°28’E
33°54'S/26°51’E
34°12’S/22°15°5’E
34°12’S/22°15°5’E
34°15°5'S/22°14’E
33°52’S/26°09’E
33°55'S/18°27’E
33°49'S/25°56’E
33°47'5/26°19’E
28°30’S/16°10’E
28°30’S/16°10’E

SB. Saldanha Bay, west coast.

SB 132
SB 153-168
SB 174
SB 178
SB 196
SB 231
SB 235
SB 267
SB 269

Date
26.3.53
—9-57
27-4-59
28.4.59
1.5.59
4.5.60
4.5.60
25.4.62
25.4.62

Position
33°04'S/17°59°3’E
33°02°5S/18°02’E
33°02°8’S/18°00-6’E
33°03°6’S/18°00-4’E
33°04°4'S/17°56-4’E
33°00°6’S/17°59°6’E
33°04°1'S/17°59°7'E
33°02’S/17°57°2’E
33°02°1'S/17°58’E

SCD. Dredged off the south coast.

SCD 5

SCD 20
SCD 22
SCD 26
SCD 29
SCD 37
SCD 50
SCD 52
SCD 56
SCD 60
SCD 61
SCD 75

SCD 79-81

Date
19.4.58
26.5.58
26.5.58
23.5.58
22.6.58
19.5.58
18.5.58
20.8.58
19.8.58
16.8.58
15.89.58

- 16.7.59

16.7.59

Position
34°15'S/25°05’E
34°07°3'S/23°23'8’E
34°26-7’S/23°26-0’E
33°47'S/26°04’E
33°38°6'S/26°54°7’E
32°15°2'S/28°57-7'E
31°38°8S/29°34°4’E
34°01°S/25°45°5'E
33°37'S/26°56-6’E
33°02S/27°56-2’E
32°17°7'S/28°54°5'E
32°33'S/28°38’E
32°43'S/28°28’E

Depth (m.)

73-78°5

86
24-27
gI

Depth (m.)
8

Littoral
15
15
35

Depth (m.)

Bottom

St

brk Sh
brk Sh, St
f£S

£S

M

M

S, Sh, St

Bottom

432

SCD 84-85
SCD 94
SCD 111
SCD 112
SCD 113
SCD 114
SCD 117-118
SCD 119
SCD 126
SCD 129-133
SCD 154
SCD 169
SCD 175
SCD 179
SCD 188
SCD 190
SCD 206
SCD 215
SCD 239
SCD 258
SCD 265
SCD 276
SCD 281-283
SCD 284
SCD 312
SCD 314
SCD 328
SCD 330
SCD 333
SCD 347
SCD 354
SCD 387
SCD 394

SH. Cape Town docks, from pylons and cable below tug jetty.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Date
17-7-59
20.7.59
23-7-59
20.7.59
26.11.59
26.11.59
14.2.60
14.2.60
3.6.60
3.6.60
25.11.60
24.11.60
30.11.60
24.11.60
30.11.60
29.11.60
30.11.60
25.11.60
29.11.60
14.7.61
14.7.61
14.7.61
11.2.62
6.2.62
9.2.62
9.2.62
10.2.62
11.2.62
11.2.62
12.2.62
11.10.62
8.12.62
2.12.62

Position
33°03'5/27°55E
33°55'5'9/25°51'E
34°35'S/21°11’E
33°55°5'9/25°51'E
34°24'S/21°45’E
34°29'S/21°49°5'E
34.°24'S/21°46’E
34°33'S/21°52’E
34°26°5'S/21°48’E
34.°48'S/22°06’E
34°03'S/25°59'E
33°58-9/S/25°41°4’E
34°20'S/23°31’E
33°58°9/S/25°41°4’E
34°10'S/23°32’E
34.°05'8'S/23°23°2’E
34°51'S/23°41E
34°03 'S/25°58’E
34.°02:0'8/23°28-4’E
33°53°8'S/25°42°5'E
33°48'S/25°47'E
33°53°8'S/25°42'5'E
34°04'S/23°23’E
33°01'S/27°55'E
33°58'S/25°4.7'E
33°58'S/25°43’E
34°43'S/25°40’E
34°03'5'S/23°23E
34°02'S/23°27’E
34°10'S/22°15’E
32°08’S/29°12’E
34°04°4'S/23°25'6’E
33°50°7'S/25°47°5/E

Date
SH 433 14.4.61
SH 436 3.4.62

Depth (m.)

Floating buoy
{S$
M
M

R
R,S

STJ. St. John’s River Estuary, eastern coast of Cape Province. Date: 20.1.50
Position: 31°37'S/29°37’E.

Estuary,

south coast.

Date:

SWD. Dredged off the coast of South West Africa.

SUN. Sunday’s River
33°42'S/25°53E.
Date
SWD 12 10.2.63
SWD 39 12.2.63
SWD 42 13.2.63

Position
26°35'S/15°o1’E
26°37°5'S/15°04'5'E
26°38:0’S/15°00°2’E

TB. Dredged from Table Bay, Cape Town.

TB 6
TB 7-8

Date
4.8.46
25.10.46

Position
33°49°5/S/18°27°5E
33°52°5'S/18°27°5’E

Depth (m.)
71
40
7%

Depth (m.)
13
17

7.1.50. Position:

Bottom

ann

Bottom
S, Sh
S Sh

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 433

Date Position Depth (m.) Bottom
TB9 25.10.46 33°52'S/18°28’E 15 S
TB 13 4.8.46 33°49°5'S/18°27°5’E 13 S, Sh
TB 14 25.10.46 33°52°5'9/18°27°5’E L7 S, Sh
TB 15 11.2.47 33°4.7°5'S/18°24°3’E 19 S, Sh, St
TB 16 25.10.46 33°52’S/18°28’E 15 N)
EB 17 26.6.47 33°52°7'9/18°28-7’E 9 S, St
TB 19 a 7A7 33°51°2'S/18°27-3/E 23 R,S
TB 21 15.12.57 33°48-6’S/18°24°6’E 15 S, Sh, St.
TRA. Material collected by commercial trawlers.

Date Position Depth (m.) Bottom
TRA 32 Q.11.47 34°49’S/20°21°5’E c. QI
TRA 33 20.7.49 34°55'S/21°10’E c. gO R,S
TRA 35 21.1.50 34°34’S/20°50’E 70 M,S
TRA 38 —.7.50 34.°30'S/20°56’E 73 M,S
TRA 42 —.7.51 34°30'S/20°55’E Cc. 70 M, St
TRA 56 28.11.52 34°40'S/21°35’E 73 RS
TRA 57 27.11.52 34°24'5/21°55’E 59
TRA 59 26.11.52 34°28'S/21°45’E 70 S, St
TRA 86 233.53 32°41°7’S/17°58°5E 9 N)
TRA 92 —.1.54 35°03’S/21°50’E 110 R,S
TRA 99 18.1.56 34°25'5'S/21°50°2’E 60 S
TRA 150 6.3.58 34.°42°2’S/20°25:0’E gl M
TRA 156 15.10.58 34°12’S/18°22’E 22 Cable
TRA 159 6.7.58 33°56’S/25°36’E Turtle

WCD. Dredged

off the west coast of the Cape Province.

Date Position Depth (m.) Bottom
WCD 7 24.3-59 34° 09°3'S/18°17'5’E 43 R
WCD 12 24.3.59 34°09°4'S/18°16°5’E 75 R
WCD 18 29.4.59 33°05'6'S/17°54°5E 64 kh M
WCD 20 30.4.59 33°07°5'S/17°52°5’E 86 R
WCD 25 1.5.59 33°06°5'S/17°55°4’E 86 en M
WCD 81 15.9.49 34°05’S/18°21’E II S
WCD 125 23.4.62 33°08’S/17°46’E 157 gen M
WCD 134 25.4.62 33°07°3'S/17°57°5/E 26 S
WCD 145 29.8.63 33°50°3'S/18°23°2’E 15 R
WCD 156 22.10.63 34.°01°7'S/18°14°7’E 100 R
WCD 158-160 21.10.63 33°55°8'S/18°21°3’E 37 R
WCD 164 7.10.63 32°52’S/18°25’E 29 R

LisT OF SPECIES

Family Tubulariidae
Tubularia solitaria Warren, 1906. Tubularia warrent Ewer, 1953.

Family Myriothelidae
Monocoryne minor n. sp. Myriothela tentaculata n. sp.

Myriothela capensis Manton, 1940.

Family Corynidae

Bicorona elegans, n.g., n.sp. Staurocladia vallentini (Browne, 1902).

Sarsia eximia (Allman, 1859).

4

434 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Solanderiidae

Solanderia procumbens (Carter, 1873).

Family Bougainvilliidae

Bimeria vestita Wright, 1859.

Bougainvillia macloviana (Lesson, 1836).

Bougainvillia sp.

Dicoryne conferta (Alder, 1856).
Rhizorhagium robustum (Warren, 1907).

Family Clavidae

Clava sp.

Merona cornucopiae (Norman, 1864).

Family Eudendriidae

Eudendrium ?capillare Alder, 1856.
Eudendrium ?carneum Clarke, 1882.

Eudendrium deciduum Millard, 1957.
Eudendrium ramosum (Linn., 1758).

Family Hydractiniidae

HAydractinia altispina Millard, 1955.
— Hydractinia kaffraria Millard, 1955.

Hydrocorella africana Stechow, 1921.
Podocoryne carnea M. Sars, 1846.

Family Pandeidae

Leuckartiara octona (Fleming, 1823).

Family Aequoreidae

Aequorea africana n. sp.

Family Lovenellidae

Lovenella chiquitita Millard, 1957.

Family Halectidae

Halecium beanii (Johnston, 1838).
Halecium delicatulum Coughtrey, 1876.

Halecium dichotomum Allman, 1888.

Halecium halecinum (Linn., 1758).

Halecium ?muricatum (Ellis & Sol.,
1786).

Halecium tenellum Hincks, 1861.

Family Campanulariidae

Campanularia hincksi Alder, 1856.
Campanularia integra MacGill., 1842.
Campanularia laminacarpa n. sp.
Campanularia ?mollis (Stechow, 1919).
Campanularia morgansi Millard, 1957.

Clytia hemisphaerica (Linn , 1767).
Clytia hummelincki (Leloup, 1935).
Clytia paulensis (Vanh6ffen, 1910).
Obelia dichotoma (Linn., 1758).
Obelia geniculata (Linn., 1758).

Family Tubulariidae

Tubularia solitaria Warren, 1906.

Tubularia solitaria Warren, 1906: 83, pl. 10, 11. Millard, 1957: 179.
Records. West coast: A 122. LB 166 (recorded by Day, 1959). SB 153U.

South coast: L 172.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 435

Tubularia warrent Ewer, 1953
Tubularia warreni Ewer, 1953: 351, fig. 1-4. Millard, 19594: 299. Millard, 1959): 240.

Records. West coast: LB 296A, 386A (reported by Day, 1959). South
coast: CPR 9G. KNY 176A (reported by Day, Millard and Harrison, 1952,
as T. ?betheris).

Description. This material conforms to previous descriptions of the species
with the exception of the CPR sample which differs in its paler coloration.
The general effect, when alive, was a creamy colour, with pink tinges in the
manubrium and gonophores. It is obviously a young colony, reaching a
maximum height of 1-75 cm., and the largest hydranths measuring only about
2:5 mm. in length and 1 mm. in basal diameter. The blastostyles are unbranched
and few in number (5-9) though fully mature gonophores are present. Apart
from the colour there is nothing to distinguish it from TJ. warreni.

Family Myriothelidae
Monocoryne minor n. sp.
Fig. 1
Material. The holotype, SCD 215Q, is a single specimen detached from
its substratum and the only specimen known. Most of the colour has gone,
though signs of a dark-red pigmentation are visible on the gonophores. South
African Museum registered number: SAMH 410.

Description. Polyp measuring approximately 5 mm. in length, though
somewhat contracted and curled up. Basal part of body (1 mm.) covered in
transparent perisarc which is continued proximally in a number of threads and
filaments, some of which appear to be filled with living coenosarc. Remainder
of body cylindrical, bearing about 110 capitate tentacles. Tentacles solitary
or arranged in groups of 2, 3, or 4 united at their bases. Length of tentacles
variable: sometimes the middle one of a group of three is the largest as in
M. gigantea, but sometimes all members of the group are equal in length and
sometimes the first of a group of four is longest.

Eleven gonophores, apparently male, borne irregularly on the body of
the polyp, the largest about 0-3 mm. in diameter. Gonophores pear-shaped,
not distinctly demarcated from pedicel.

Nematocysts. At least three kinds visible in smears of preserved material:

(1) Desmonemes, the most abundant type. Capsule oval, thread with
about 2 complete coils, on which a spiral ridging is visible. Size
variable: 9:0-13°5 x 6:3-10°8 p.

(ii) Stenoteles, fairly common. Capsule egg-shaped, with shaft occupying

about half length. 15:3-18-0 & 13:0-15'3 mp.

(iii) Undetermined heteronemes, rare. Capsule elongated, with shaft

occupying about 2 length. 16-2-18-9 x 6-3—7°6 p.

Remarks. As there is only one specimen available no sections were cut and

accurate measurements could not be made without damage. A few detached

436 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 1. Monocoryne minor n. sp.

A. The whole animal (holotype).
B. A typical group of 4 tentacles.
C-F. Nematocysts. (C, stenotele. D, undetermined heteroneme. E—F, desmonemes).

tentacles were used for examination of nematocysts. The specimen is fairly
well preserved, but there is no knowing how many tentacles may have been
broken off during handling, or whether the base was originally attached to a
firm substratum. Since the tip of the body is curled over, the mouth could not
be observed. »

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 437

The arrangement of the tentacles differs from that in the only two known
_ species of Monocoryne, namely M. gigantea (Bonnevie, 1898) and M. bracteata
(Fraser, 1941), in both of which the middle one of a group is the largest. It is
also much smaller than either of them. The gonophores are not seated in the
axils of the tentacles but distributed at random over the body.

Myriothela capensis Manton, 1940

Mpyriothela capensis Manton, 1940: 276, pl. 1 (figs. 12, 13), pl. 3 (fig. 27), figs. 7, 8b, 9. Millard,
1957: 186.

Records. West coast: CP 331 (one almost mature male specimen attached
to weed). LAM 52F (one mature female specimen attached to weed).
LU 118Y (two mature male specimens torn from their base and two young
specimens on crustacean appendage).

Description. The material from Luderitz Bay (LU 118Y) had retained its
colour after about four months in spirit. The whole distal end of the body
was a vivid splash of colour—the capita of the body tentacles and tips of the
gonophores bright magenta, shading to pink on the stalks and colourless on the
base of the polyp. Specimens preserved for a longer period are pale pink or
completely colourless.

Largest specimen 1-6 cm. in length. Structure and nematocysts as described
by Manton. One mature female specimen with actinulae larvae (one gonophore
on opening contained three larvae). Female gonophores reaching o-g x 1-0
mm.

Myriothela tentaculata n. sp.
Fig. 23
Material. WCD 7Q: 5 specimens from west coast attached to encrusting
polyzoan, two of them rather badly damaged and the others in various states
of preservation. The best preserved was selected as the holotype and another
was used for sectioning. South African Museum registered number of holotype:

SAMH 411.

Description of holotype. Total length 2-5 cm. No colour left in specimen except
for a circle of dark-red spots round the distal end of each gonophore.

Basal region of hydranth (8 mm.) naked and drawn out into an irregular
shape, bearing about g chitinoid, flattened, adhesive discs attached to the
substratum.

Above this a single whorl of 17 long, tapering blastostyles reaching a
maximum length of about 2 cm. These are generally somewhat coiled and bear
a superficial resemblance to the tentacles of an octopus. They are the most
distinctive feature of the species. Blastostyle unbranched, bearing 4-6 gono-
phores in the proximal 5 mm., of which the most distal is the oldest and the
others in various stages of development. Capitate tentacles present amongst
the gonophores and others (about 25) scattered irregularly over the distal
region of the blastostyle. Tentacles rather poorly developed and resembling
boot-buttons. Gonophores male, spherical, reaching a diameter of about

438 ANNALS OF THE SOUTH AFRICAN MUSEUM

2 mm., each with a very short pedicel and 8-10 pigmented spots around distal
end.

Distal region of hydranth (about % of length) covered with densely packed
capitate tentacles and bearing a terminal mouth. Tentacles well-developed,
with slender stalks and large capituli, but more poorly developed in the region
of the blastostyles.

Nematocysts. Four types distinguishable from smears and sections:

(i) ‘Heteronemes’ (fig. 2D), elongated and banana-shaped, with an
axial body which stains redin Mallory’s stain and a fine much-coiled
thread. 44°1-45°9 x 8-1 p.

(11) Desmonemes of two sizes (fig. 2E-H), the larger 17-1-18-0 x 12-6—
14°4 pw, the smaller 10-2-10°8 x 7-2—7-7 w. In both, part of the thread
stains red in Mallory’s stain, the rest remains unstained. It bears a
distinct spiral marking which may indicate the presence of barbs.
The larger form contains a thread in 3 complete longitudinal coils
and the smaller a thread in 2 coils.

(iii) Steroteles (fig. 2C). Oval capsules containing a butt which stains red
in Mallory’s stain and increases in diameter towards the base. The
thread is coiled transversely in the lower half of the capsule and forms
an opaque mass in this region. A few were found exploded and showed
a characteristic butt, though the thread and spines had been broken
off. 14°4. X 10° p.

(iv) Atrichous isorhizas (fig. 2B). Elongated capsules with the contents
difficult to discern, but apparently containing a long tangled thread
which stains faintly blue in Mallory’s stain. 15-3-19°8 Xx 4:0-7°2 p.

Mistology. The hydranth body wall (fig. 2M) is very similar in structure to
that of M. capensis. It is 110-180 p thick, with a fairly thin layer of mesogloea
10-20 pw thick from which slender lamellae bearing muscle fibres project into
the ectoderm. The lamellae arise at intervals of approximately 14 » and are
40-80 p» deep. From the endoderm arise villi anything up to 1 mm. in depth,
each containing a very thin supporting lamella of mesogloea. The apical cells
of the villi contain dark-staining granules, but the remainder are clear and
vacuolated. 85 villi were counted in one section. In the region of the mouth the
granular cells are more numerous and many cells contain droplets of a yellowish
material. |

The body tentacles (fig. 2K) are exactly like those of M. capensis as
described by Manton (1940), with an apical pad of mesogloea fibrils about
50 » in thickness and a central cavity which passes into the stalk without
constriction. The endoderm of the stalk is separated from that of the hydranth
body by a thin layer of mesogloea. The capita of the tentacles are richly armed
with nematocysts of all 4 types, of which desmonemes are the most numerous.

The blastostyle is the most distinctive structure in the species. In the
proximal region where the gonophores arise the endoderm bears villi which

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 439

20

Bea
SS EEE

<<

\

zh

i

PX

Fic. 2. Myriothela tentaculata n. sp.

A. The whole animal (holotype).

B-H. Nematocysts. (B, atrichous isorhiza. C, stenotele. D, ‘heteroneme’. E and F, large
desmoneme in side view and end-on view. G and H, small desmoneme in end-on view and
side view.)

t.s. blastostyle in proximal region with young male gonophore arising from it.

l.s. body tentacle.

ls. blastostyle tentacle.

t.s. through body-wall of hydranth in distal region including the origin of 2 tentacles.

(e, ectoderm. enb, endoderm of bell. m, mesogloea. ml, mesogloeal lamella. mp, apical pad
of mesogloeal fibrils. n, nematocysts. se, subumbrella ectoderm. sp, spermatogenic cells.
v, endodermal villi).

Sr AS

440 ANNALS OF THE SOUTH AFRICAN MUSEUM

project into the central cavity and fill it almost completely (fig. 2J). The cells
of the villt are densely packed with granules which stain red in Mallory’s
stain, and many of the granules float freely in the interstices between the villi.
The mesogloea bears lamellae about 20 » deep on its ectodermal surface.
The ectoderm contains many developing nematocysts which are particularly
abundant around the origins of the tentacles. In the slender distal part of the
blastostyle ectoderm and mesoderm are normal and well-preserved, but the
endoderm appears to be largely senile. Although remains of villi and their
mesogloeal lamellae can be recognised, most of the internal cavity is filled with
dead cells and fragments of cells most of which have an opaque yellowish
coloration.

The blastostyle tentacles (fig. 2L) are similar to those of the body, but each
has a much shorter stalk and a thinner pad of mesogloeal fibrils (about 20 p
thick in maximum). The lumen is very narrow and the endoderm separated
from that of the blastostyle by a basal lamella of mesogloea. The superficial
layer of the capitum is densely packed with nematocysts, including
‘heteronemes’ and atrichs, while desmonemes and stenoteles appear to be
entirely absent.

The structure and development of the gonophore is similar to that in
other species (Briggs, 1929, 1931; Manton, 1940). Only male gonophores are
present and the largest one sectioned is 1-24 mm. in diameter with a layer of
spermatogenic cells 0-24 mm. thick around the spadix. It is apparently almost
mature, for a velar invagination is present in the exumbrella ectoderm though
not yet perforate. The endoderm of the spadix bears rounded villi which
project into the central cavity which is continuous with that of the blastostyle.

Remarks. Of the six southern hemisphere species listed by Manton, (p. 288)
this species is most closely related to M. capensis. It differs from it in its larger
size, its adhesive structures, which are less obviously tentacular, and its long
and tendril-like blastostyles.

The nature of the nematocysts supports the establishment of a new species.
All are larger than the corresponding types in M. capensis, the banana-shaped
‘heteronemes’ being enormous, about 4 times those of M. capensis and over
twice those of M. penola and M. cocksi. The presence of atrichs in the hydranth
is unusual since in other species this type is confined to the actinula. The term
‘heteroneme’ has been used following Manton, although the only evidence
that this type possesses a distinct butt is that of Allman (1876), and needs
confirmation. No haplonemes could be identified in this material, and if
present cannot be distinguished from stenoteles.

Family Corynidae
Genus Bicorona nov. gen.
Diagnosis. Corynidae with firm perisarc and upright, monopodially
branched stems. Hydranths with two whorls of capitate tentacles. Gonophores
in the form of fixed sporosacs borne on the body of the hydranth.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 441

Bicorona elegans nov. sp.
Fig. 3

Types and records. Holotype: SB 161X (South African Museum registered
number: SAMH 412), from Saldanha Bay on the west coast. Other records:
A 116, 139, 353 (west coast); L 44, 56 (south coast).

Description of holotype. A luxuriant colony of upright branching stems
reaching a maximum height of 5-8 cm., growing in the lower intertidal region
of the shore.

Hydrorhiza embedded in sponge, branching and giving rise to upright
stems. Stem unfascicled, with monopodial growth and terminal hydranths.
Branches arising alternately, either bearing one terminal hydranth, or
rebranching in a manner similar to the stem. Stem and branches covered
with firm perisarc which is closely annulated throughout except for a smooth
area on the origin of each branch. Hard part of perisarc terminating just
below hydranth, but continued as a gelatinous layer onto the base of the
hydranth.

Hydranth Tubularia-like, with two whorls of capitate tentacles. Aboral
tentacles 10-21 in number in mature hydranths, usually with long and short
tentacles alternating. Oral tentacles short, 4-7 in number. ‘Neck’ region of
hydranth (below aboral tentacles) with indistinct longitudinal striations.

Gonophores in form of fixed sporosacs, borne on approximately 7 short
blastostyles which form a whorl just distal to the aboral tentacles. Gonophores
oval to spherical, 2-4 to a blastostyle, the oldest one terminal.

Histology (fig. 3K). Stem normal, with no endodermal canals, terminating
in a swollen region in the last perisarcal segment.

‘Neck’ region of hydranth with longitudinally ridged ectoderm, surrounded
by thick gelatinous sheath continuous with the perisarc of the stem.

Hydranth expanding at level of aboral tentacles, with spacious enteron.
Ectoderm of hydranth normal. Endoderm specialized: immediately below
aboral tentacles thickened and granular but with no indication of parenchyma,
in the region between the two whorls of tentacles folded inward around sup-
porting strands of mesogloea in the nature of endodermal villi, in region of
hypostome thickened.

Tentacles solid, with central core of ‘chordal’ endoderm. Aboral tentacles
with no ‘supporting lamella’ of mesogloea, and endoderm continuous with
that of hydranth. Oral tentacles attached obliquely to hydranth body, with a
‘supporting lamella’ of mesogloea which may be incomplete in its aboral
region.

Blastostyle hollow and tubular. Gonophores with a hollow spadix, without
subumbrella cavity or radial canals. Female gonophore containing many
small eggs arranged in a single series around the spadix (counts give numbers
varying froma 29 to 84). Eggs 0:05-0:10 mm. in diameter. Male gonophore
containing a thick mass of spermatogenic cells.

Nematocysts (fig. 3H, J) all stenoteles, of 2 sizes: large ones reaching

5

442 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 3. Bicorona elegans n. sp.

A. Two complete stems from the holotype.

B-G. Stages in the development of the hydranth (B-F from A 139, G a mature hydranth
from the holotype).

Hand J. Small, and large, stenotele from A 116.

K. l.s. hydranth with female gonophores from L 44.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 443

26 X 17p, small ones reaching 13:5 <X 7 (measurements from preserved
material, undischarged).
Measurements (mm., preserved).
(NV.B. the holotype material is well expanded, A 116 less so.)
holotype A 116
Stem, diameter ; : . 0°21-0°35 0°26-0°43
Hydranth, length from end of ‘hard’ perisarc to

hypostome . ; . 0°96-1'95 1°29-1°73
Gonophores, length : i ¢ feaehing. . 0-47 0°50
maximum diameter ; a MERGING w\0-4.4: 0-36

Remarks. The material chosen as the holotype is a well-developed, mature
colony, with presumably the maximum number of tentacles on the polyps.
One of the other samples (A 139) contains shorter stems with younger, well-
relaxed polyps, permitting observations on the origin of the tentacles (fig.
3B-F).

In the young polyp bud, two whorls of capitate tentacles appear simul-
taneously, 4-7 oral and approximately 7 aboral. These are at first short and
knob-like, but they lengthen as development proceeds, especially the aboral
ones. The number of oral tentacles is not increased, but before long a new
series of aboral tentacles appears, about 7 in number, alternating with those
of the first series and at a slightly Jower level on the polyp. Polyps at this stage
of development (fig. 3F) thus appear to have three whorls of tentacles, one
oral and two closely alternating aboral. When the second series of aboral
tentacles is about half the length of the first, young blastostyles make their
appearance, one in the axil of each aboral tentacle of the first series. The oldest
polyps in this particular sample have reached this stage, but the appearance
of a third series of aboral tentacles would bring the number up to about 21
(the maximum number observed in the holotype). In the fully mature polyps
of the holotype and other samples, however, the aboral tentacles are arranged
in a single whor] and at one level (fig. 3G). Whether this is due to further
growth in circumference of the polyp or to differences in the state of contraction
can only be determined from living material.

Annandale (1915) established a new genus Dzucyclocoryne for a species
previously described in 1907 as Syncoryne filamentata, in which the generic
diagnosis is the presence of two whorls cf capitate tentacles. But D. filamentata
bears free medusae and thus, following the practice adopted in these papers,
cannot be included in the same genus as forms with fixed sporosacs. There are
also other differences from the present species (e.g. unbranched stems, smooth
perisarc, different position of gonophores) which make the establishment of a
new genus for this material desirable, viz. Bicorona.

Dicyclocoryne and Bicorona appear to be more closely related to the Corynidae
than to any other family of Capitata, and this is supported by the presence
of only one category of nematocyst. Although stenoteles and desmonemes have

444. ANNALS OF THE SOUTH AFRICAN MUSEUM

been reported from medusae of this family, only stenoteles (large and small)
are known from the hydroids (Russell, 1938, and Millard, 1959b, for Sarsia
exumia). In the Pennariidae on the other hand, with which Bicorona also has
certain affinities, 3 or 4 different categories of nematocyst occur (Weill, 1934,
and Millard, 1959a).

Sarsia eximia (Allman, 1859)
Coryne sp.: Millard, 1957: 179.
Sarsia eximia. Russell, 1953: 50, pl. 2 (fig. 3), figs. 17a, 18A, B. Millard, 19596: 241. Kramp,

1959: 79, fig. 15. Kramp, 1961: 27.

Records. West coast: LB 556A. LU 113D. SH 433D, 436A. South coast:
CPR oF. KNY 165E (recorded by Day, Millard and Harrison, 1952, as
Syncoryne Peximia).

Remarks. Living material from Table Bay docks (SH 436A) and Langebaan
_ (LB 556A) was kept in the laboratory where it released medusae. At liberation
these measured 0-9 mm. in depth and 0-8 mm. in diameter, with a hypostome
of 0-3 mm. in length. After 3 days the size had increased to 1:1 mm. depth and
I°2 mm. diameter, with a hypostome of 0-4 mm. At this stage the teniacles
reached a length of about 1-8 mm. when extended and bore about 12 clusters
of nematocysts. The structure was similar in every way to that of S. eximia.
The medusa of S. exzmia has not previously been recorded from South Africa.

Staurocladia vallentint (Browne, 1902)

Cnidonema capensis Gilchrist, 1919: 509, pl. 30.
Staurocladia vallentini: Browne and Kramp, 1939: 274, pl. 14 (figs. 3-4), pl. 15 (fig. 4), pl. 19

(fig. 2).
Cnidonema vallentini: Ralph, 1947: 414, pl. 35 (figs. 1-6).

Records. West coast: LB 266A, 398A (recorded by Day, 1959, as Cnzdonema
vallentint). A 40 (recorded by Bright, 1938, as Eleutheria valleniint).

Description. Material from Langebaan (LB) consists of numerous young
medusae found crawling on weed (Gracilaria confervoides), most of them under-
going asexual reproduction. Medusae reaching a maximum diameter of about
I mm., and bearing 20-30 tentacles. Aboral branch of tentacle with 4-5
nematocyst batteries, of which one is terminal and the rest dorsal. No gonads
present.

Material from Oudekraal (A) includes two larger medusae, reaching a
diameter of 3 mm. and bearing gonads. Specimens rather damaged, but
apparently bearing over 40 tentacles. Nematocyst batteries as in LB material.

Hydranths not as yet found.

Family: Solanderiidae

Solanderia procumbens (Carter, 1873)
Fig. 4 and Plate I
Ceratella procumbens Carter, 1873: 10.

Ceratella spinosa Carter, 1873: 12.
Solanderia atrorubens: Marshall, 1892: 12, pl. 5, pl. 7 (figs. 2-4).

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 445

Solanderia procumbens: Vervoort, 1962: 535.
Solanderia spinosa: Vervoort, 1962: 535.
Non Dehitella atrorubens Gray, 1868.

General. The material of this species is divisible into two forms on general
appearance (here designated as form A and form B), which do not seem to
be specifically distinct, but are kept separate in case they should later prove
to be so. In the description which follows the skeleton of the largest specimen
of each (WCD 158F and E respectively) is described and the soft parts of the
best preserved (SAMH 247).

Records
Form A: West coast: WCD 158F. South coast: SAMH 247. TRA 590A.
Form B: West coast: WCD 158E. South coast: SCD 154A.

Description, form A. The largest colony (Plate IA, left) a magnificent
fan-shaped specimen 33 cm. high with a spread of 38 cm. Hydrorhiza 5 cm.
broad, and flattened below for attachment. Main stem flattened in the plane
of branching and expanding from a base 2-4 cm. wide and 1-2 cm. thick to a
broad leather-like blade 5 cm. wide and 0:5 cm. thick; beyond this giving rise
to a number of large branches, which in their turn branch and rebranch.
Branching in one plane, with a tendency for the branches to bend to one side
and give off more sub-branches on this side, suggesting the influence of a
strong current during growth. Method of branching variable, often dichoto-
mous, sometimes alternate and sometimes unilateral. Smaller branches quite
round in section, comparatively thick and stumpy (about 2 mm. in diameter),
with rounded tips.

Fibrous meshwork of main stem reticulate, with rounded, rectangular,
or hexagonal apertures between the trabeculae, but no marked longitudinal
arrangement. Trabeculae of unworn areas bearing numerous spiny structures
0-3-0°4 mm. in height, some of them flattened and spatulate and resembling
abortive hydrophores with one or two supporting ribs (fig. 4D).

Fibrous meshwork of branches arranged in a more definite fashion, in
which the longitudinal trabeculae predominate. Groups of trabeculae often
raised up into prominent longitudinal ridges surmounted by blade-like crests
parallel to the long axis of the branch (Plate IB, lower right corner). Spines
reduced to tubercles on the branches and absent on the smallest subdivisions.

Hydrophores numerous and scattered irregularly over surface of branches.
A typical one consisting of a scoop-shaped bracket supported on its lower
surface by longitudinal ribs continuous with the trabeculae of the branch
(fig. 4B). Ribs normally not reaching margin, which is smooth and rounded.
Hydrophores, however, subject to much variation, some being bilobed (fig. 4C),
some divided into two halves (one on each side of the hydranth) and some
represented by a laterally seated hemihydrophore only. Badly eroded hydro-
phores with margin worn down between the supporting ribs (fig. 4H). Normal
hydrophores approximately 0-5 mm. in length, and 0-5 mm. in basal width
tapering to 0:3 mm. at margin.

— 446 ANNALS OF THE SOUTH AFRICAN MUSEUM ©

Hydranths arising from coenosarc above hydrophores and, rarely, directly
from the stem with no protective supports whatever; bearing approximately
15 capitate tentacles (but difficult to count due to poor preservation).

Gonophores arising directly from coenosarc of stem, spherical, shortly
stalked. Immature male gonophores present on two specimens, the largest
measuring 0°34 mm. in diameter and 0-32 mm. in length; containing several
layers of spermatogenic cells around a central spadix, and 4 distinct radial
canals.

Colour : stem and larger branches dark brown, occasionally with a purplish
tinge, shading to yellowish-brown on the smaller branches.

Form B. Colonies smaller than form A, the largest reaching 19:5 cm. in
height and 15-5 cm. in spread, with more slender branches and lighter coloration
(Plate 1A, right). Main stem rounded in section. Hydrophores on smaller
branches restricted to two sides, but irregularly scattered on larger ones. The
‘double’ type of hydrophore predominant (fig. 4E, G), bilobed or single ones
rare (fig. 4F).

Nematocysts. Three types present in smears of preserved material:

(i) Large oval stenoteles (fig. 4R, S$). Fairly common. Undischarged
capsules with a broad, central shaft and a long thread coiled trans-
versely in the lower half. Discharged capsules with a broad, slightly
tapering butt armed with 3 large spines and several spiral rows of
smaller spines. Thread broken off in all examples seen.
15°3-24°3 X 11°7-18°0 pw

(ii) Small oval heteronemes (fig. 4P). Abundant. Undischarged capsules
with a central shaft and a thread which appears to be irregularly
coiled. A single discharged and rather distorted capsule bore a short,
swollen butt and a number of spines. 8-1-8-7 x 6°3 p

(i11) Elongate-oval capsules of unknown category (fig. 4Q). Rare. Only

discharged capsules seen with no sign of butt or thread. 15-3 X 54 u

Remarks. It is felt that form A and form B represent different growth-forms
of the same species, form B including younger colonies which have possibly
grown more rapidly. Their main claim to distinction is the nature of the
hydrophores which are predominantly ‘single’ in form A and ‘double’ in form
B, although both types can be found in all colonies. There is no difference in
the nematocysts. The ‘double’ hydrophore is in some ways similar to the
‘spines’ which flank the hydranth in S. secunda (Inaba). However, diagrams
by Stechow (1909, pl. 4, fig. 7) and Vervoort (1962, figs. 2b, 7) show that the
latter are little larger than the area of one mesh of the skeleton, whereas in
the present material they approximate to the area of 4 meshes.

Carter’s dried type material of Ceratella procumbens was available for com-
parison in the British Museum. It includes one specimen from the Cape of
Good Hope (reg. no. 1867.3.22.1) and three from Natal (reg. no. 1872.8.1.1).
The largest is one of the latter and measures 26-5 cm. in height with a spread

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 44.7

S

Fic. 4. Solanderia procumbens (Carter).

A-C. Hydrophores from form A, WCD 158F (A and B single, C bilobed).

D. A portion of the older part of the stem in WCD 158F showing spines.

E-G. MHydrophores from form B, SCD 154A (F single, E and G double).

H. A typical eroded hydrophore from False Bay.

J-N. Hydrophores from Carter’s type material (J and K single, L double, M and N eroded).
P-S. Nematocysts (P, a small heteroneme. Q, undetermined type. R and S, large stenotele).

448 ANNALS OF THE SOUTH AFRICAN MUSEUM

of 11 cm. The main stem is 1-8 cm. wide and 0-6 cm. thick. This is evidently
the specimen whose measurements were quoted by Carter in 1873. I select it
as the lectotype.

In spite of the fact that the type material is battered and obviously beach-
worn the resemblance to form A described above is unmistakable. Here too
the hydrophores are predominantly single (fig. 4J, K) though bilobed and
double ones also occur (fig. 4L). In the older and badly worn parts the
hydrophores are eroded so that the supporting ribs project and give a
spinous appearance to the margin resembling that in S. fusca (Gray)
(fig. 4M, N).

S. spinosa (Carter, 1873), is here considered to be a synonym for
S. procumbens. The dried holotype of this species from Port Natal is also present
in the British Museum (reg. no. 1872.8.1.17). It is less beach-worn than the
type material of S. procumbens and this would account for the minor differences
said to distinguish the species. Its general appearance is similar to form A
described above though most of the hydrophores are of the ‘bilobed’ type.
The ‘spines’, which are the main specific character, are present only on the
older part of the colony as in the present material, and are comparatively
delicate structures which would easily be rubbed off by sand erosion. This
would account for their absence in the type material of S. procumbens.

Another species which possesses well-developed hydrophores is S. fusca
(Gray, 1868), from Australia, described and figured by Spencer (1892). Gray’s
type material is present in the British Museum (reg. no. 1884.12.6.15—-16; the
first of these designated as lectotype by Vervoort (1962: 533). The most
obvious characteristic of this species is the nature of the hydrophores which
have spiny margins due to the projection of the supporting ribs beyond the
edge. That this effect is not in this case the result of wear is evident from the
fact that spiny hydrophores are present in the youngest and thinnest branches
and, according to Spencer, also in the living animal.

S. atrorubens (Gray, 1868), is closely related to S. fusca and possibly a
synonym, although Vervoort, 1962, considers that it can be distinguished by
its method of growth. An examination of the type material in the British
Museum, probably from Australia (of which no. 1962.4.14.1 has since been
designated as lectotype by Vervoort, 1962, p. 535), showed that it has the
same spiny hydrophores.

The fact that hydrophores of S. procumbens may acquire a spiny appearance
as the result of erosion has probably led to confusion between this species and
S. fusca and S. atrorubens. Marshall’s record of S. atrorubens from Port Natal can
be included in the synonymy of S. procumbens as his figures show that the edges
of the hydrophores are smooth. Other species recorded from South Africa,
including S. labyrinthica (Hyatt, 1877), and S. rugosa Marshall, 1892, and also
Brazier’s record (1887) of S. atrorubens are insufficiently described but are
possibly all synonyms of S. procumbens.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 449

Family Bougainvilliidae
Bimeria vestita Wright, 1859
Fig. 5 A-F
Bimeria vestita: Hincks, 1868: 103, pl. 15 (fig. 2). Allman, 1872: 297, pl. 12 (fig. 1-3). Hamond,

1957: 297, figs. 3-4.
Leuckartiara vestita forma nana: Vervoort, 1946a: 294.

Records. South coast: MB 88L. SAMH 158, 177, 233, 235, 327, 337; 343;
361. SCD 85T, 118M, 347F.

Description. Colonies epizootic on other hydroids, and reaching a height
of 3-8 mm. Stem upright, bearing from 1 to 6 or 7 alternately arranged
hydranth pedicels. Pedicels occasionally rebranching. Perisarc annulated at
base of stem, on origin of pedicels and at other irregular intervals, covered
throughout with adherent particles. Stem and pedicels narrower at base than
at distal end. Perisarc continued over the hydranth, sheathing the tentacles
for part of their length and covering the greater part of the hypostome. Tentacles
varying in number from about 10 to 16.

Gonophores borne on the stem and hydranth pedicels, each on a short
annulated pedicel, completely covered by a thick coat of gelatinous perisarc
male and female on separate colonies. Male sporosac elongated-oval, with
branching spadix. Female sporosac oval to spherical, bearing a single terminal
ovum, which develops in sztu into a planula larva (fig. 5 B—D).

Measurements (mm., preserved).

Stem, diameter . : : ; « 0°04-0:09
Pseudohydrotheca, cid, ! ; . O°21-0°49
diameter. : ; . O*12—0°30

Gonophore, female, length (without as : © reachang 0-16
diameter . . ‘teaching: 0-16

male, length (without periunite) . reaching 0°33
diameter. ; . . reaching O12

Remarks. These colonies are very similar to those described by Hincks,
Allman and Hamond, although they are not so richly branched as some of
Allman’s specimens. The size of the hydranths and the thickness of the stem
appears to be less than in most descriptions, though it corresponds well with
Hamond’s material.

The only previous record of this species from southern Africa is that of
Vervoort, 19464, from Inhaca in Portuguese East Africa. Female gonophores
are apparently described here for the first time.

Bougainvillia macloviana (Lesson, 1836)

Perigonimus maclovianus: Vanh6ffen, 1910: 284, fig. 10.
Bougainvillia macloviana: Jaderholm, 1923: 3. Millard, 1959): 242, fig. 1 A-C. Vannucci & Rees,
1961: 69.

Records. West coast: SB 178D.
Description. Colony epizootic on other hydroids. Stems only 2 mm. in

6

450 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 5.

A-F. Bimeria vestita Wright. A, a typical stem. B—D, stages in the development of the female
-gonophore (C and D drawn without the perisarcal covering). E, a contracted hydranth.
F, a male gonophore. (A—E from SAMH 361, F from SAMH 177).
G. Clava sp. Two hydranths sketched from living material (CP 646A).
H-J. Merona cornucopiae (Norman) from SCD 119R. H, part of the colony showing hydranths,
a single blastostyle and nematothecae. J, three nematothecae on a larger scale.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 451

height, slender, flexuous, branching irregularly, bearing medusa buds, of which
the oldest has its tentacles unfurled.

Remarks. In its general appearance this colony strongly resembles
Vanh6offen’s description of the species and also the material previously described
(Millard, 1959). The identification is strongly supported by the discovery
of abundant mature medusae of this species from the same area in Saldanha Bay
(identification by M. E. Thiel of the Zoologisches Museum, Hamburg). It is
interesting also that the two ships on whose hulls this species was recorded in
19590 both came from Saldanha Bay.

Bougainvillia sp.
Records. West coast: LB 542A. South coast: KNY 165D (reported by Day,
Millard and Harrison, 1952, as B.ramosa). SAMH 170, 245. SCD 190A,
281G. TRA 33Z, 150F.

Description. Stems slender, upright, weakly fascicled at the base in the larger
colonies and branching profusely in a more-or-less alternate fashion. Maximum
height 5:3 cm. Larger stems generally smooth, smaller branches wrinkled or
corrugated, particularly over the origins. Smaller colonies unfascicled and less
profusely branched. Medusa buds with 4 unbranched oral tentacles and 4
marginal bulbs, each with 2 black ocelli and 2 marginal tentacles.

Remarks. It is felt that this material could not be assigned to a species with
any certainty. The general growth-form is very similar in all the colonies,
though some are obviously older than others. Well-developed medusa buds
may be present on stems of only 3 mm. in height.

The stems are stiffer and more profusely branched than those of
B. macloviana, yet not so sturdy as those described as B. ramosa (Millard, 19596)
from ships’ hulls.

The material might well be included in B. ramosa forma musca Allman,
1864, yet no mature medusae of B. ramosa have so far been recorded from this
country, whereas medusae of B. macloviana do occur on the west coast (see
above).

Dicoryne conferta (Alder, 1856).
Eudendrium confertum Alder, 1856: 354, pl. 12 (figs. 5-8)
Dicoryne conferta: Allman, 1872: 226, 293, pl. 8. Jaderholm, 1909: 47, pl. 3 (fig. 6).
Records. South coast: SCD 133C.

Description. A dense colony growing on a gastropod shell occupied by a
hermit, reaching a maximum height of 2:5 cm. Hydrorhiza reticulate. Stem
unfascicled, branching irregularly, increasing slightly in diameter towards
distal end. Perisarc roughly corrugated throughout, but more definitely
annulated on origin of stem, terminating below hydranth when the latter is
expanded and covering the base of the body as a ‘pseudohydrotheca’ when
contracted. Hydranth long, with a single whorl of tentacles near distal end.

‘Blastostyles’ in the form of tentacle-less hydranths borne on stem and

452 ANNALS OF THE SOUTH AFRICAN MUSEUM

hydrorhiza. Male gonophores present, borne on lower region of blastostyle
either as a tight cluster or distributed along its length. The two tentacles of the
swimming sporosac clearly visible within the perisarcal covering of the
gonophore.

Nematocysts all small, 5:4 x 2°7 p

Remarks. This appears to be the first record of the species from the southern
hemisphere. The appearance of the colony is very characteristic and closely
resembles that illustrated by Jaderholm in 1909.

Rhizorhagium robustum (Warren, 1907)
Parawrightia robusta Warren, 1907: 187, pl. 33 (figs. 1-5), figs. 1-4.
Records. South coast: L 177.

Description. A sterile colony growing on a coralline alga. Most stems bear
a single terminal hydranth, but many 2 or 3. In some cases a number of stolons
or stems (it is impossible to determine which) are twisted together simulating
a fascicled stem which is quite free from the substratum and reaches a maximum
height of 17 mm. Perisarc well developed and forming a very distinct
‘pseudohydrotheca’ over the base of the hydranth. Tentacles 18-22.

Family Clavidae

Clava sp.
Fig. 5G.
Records. West coast: CP 646A.

Description. Colony growing on a stone just below low tide level. Hydrorhiza
reticulate, penetrating into calcareous matter on stone, covered with a thin
layer of perisarc which forms a very low collar round the base of each hydranth.

Hydranths reaching a maximum height of 7 mm., creamy pink in colour
when alive, with 22 to 30 scattered filiform tentacles which reach a maximum
length of 1 mm. when fully extended. |

Gonophores absent.

Nematocysts of two kinds:

(i) Microbasic euryteles, 0-67—0-72 < 0°22 p.
(ii) Desmonemes, 0°45 X 0°32 uw

Remarks. This species cannot be definitely identified in the absence of
gonophores. The trophosome is similar in size and all other characters to
C. multicornis (Forskal, 1775), forma genuina Broch, 1916. The sizes of the two
categories of nematocyst are also close to those given by Weill, (1934) for
C. squamata (= C.. multicornis).

Merona cornucopiae (Norman, 1864)
Fig. 5 H, J.
Merona cornucopiae: Rees, 1956: 499, figs. 1-3.

Records. South coast: LIZ 25P. SCD 110R.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 453

Description. Colonies growing on the bivalve Crassatella capensis Lamy. Of the
two samples recorded above the first consists of a single bivalve bearing a male
colony of about 160 hydranths. The second consists of three bivalves bearing
colonies of between 30 and 50 hydranths each, two of these colonies bearing
female gonophores. In each case the colony is restricted to one corner of the
shell.

Hydrorhiza in the form of an open reticulum at the margins of the colony,
but consolidated into a mat in the denser regions.

Hydranths with 16 to 20 scattered filiform tentacles, surrounded by sturdy
perisarcal tubes into which they can be completely retracted. Tubes slender at
base, expanding distally, often regenerated.

Gonophores borne in clusters on blastostyles which arise separately from
hydrorhiza, male and female on separate colonies. Blastostyle surrounded at
base by short collar of perisarc.

The hydrorhiza of one colony only (SCD 119R, male) bearing numerous
minute nematothecae. Nematotheca in the form of an asymmetrical perisarcal
funnel borne on a long and slender pedicel. Containing a cluster of large
nematocysts.

Measurements (mm.)
SCDi1r9R LIZ 25P

Perisarcal tube, height . . 1°83-4°25 2°15-2°72
maximum diameter : . 0°30—0°38 0:32-0:35
Blastostyle, total height . a EAT 0°63
Gonophore, length ; : . OF17-0'25 O°1G—0°35
maximum diameter . O°12-0°20 O°12—-0°21
Nematotheca, height. . 0:03-0:06
maximum diameter : ; . 0°07—0:09
lenethof pedicel — . : . 0°22—0°54

Nematocysts. At least two types present:

(1) Microbasic euryteles. Elongated capsules measuring 16:2 x 4°5 uw
undischarged. Abundant in nematophores, scarce on hydranth
tentacles. A single discharged and rather damaged capsule observed.

(ii) ?Desmonemes. Ovoid capsules measuring 7:2 X 2°7-3:°6 w undis-
charged. Abundant on hydranth tentacles, scarce in nematophores.

Remarks. This species has so far been reported only from northern seas.
It is a new record for South Africa and its presence here is surprising. Even
more so is the discovery of nematophores on the hydrorhiza There can be
no doubt about the identification of the species thanks to the detailed description
of living material by Rees and the characteristic habitat of the animal.

The nematophores, when first noticed, were thought to be the hydrothecae
of an epizootic species, but the microscope showed that they arose from the
same hydrorhiza as the clavid ‘host’, a fact which was convincingly proved
when the same nematocysts of identical measurements were found in both. A

454 ANNALS OF THE SOUTH AFRICAN MUSEUM

noteworthy point is that nematophores occurred in only one colony (male)
out of four which were similar in every way other than sex.

Family Eudendriidae
Eudendrium annulatum Norman, 1864.

Remarks. The only record of this species from South Africa is that of
Ritchie (1909) from the entrance to Saldanha Bay, and that a doubtful one.
In 1960 the opportunity offered to examine two slides of Ritchie’s material
from Saldanha Bay and to compare them with Norman’s preserved type
material of E. annulatum. In Ritchie’s material the stem is not so densely
annulated as in the usual conception of EF. annulatum, although in the type
material of the latter smooth areas do occur in some parts of the larger branches.
More important is the fact that in Ritchie’s material bifurcating spadices could
be distinctly seen on some of the female gonophores, whereas in E. annulatum
the spadices are unbranched (Broch, 1916, p. 62). Ritchie’s record of E. annula-
tum from South Africa should thus be discarded, and the material is provisionally
placed in E. carneum (see p. 455).

Eudendrium ?capillare Alder, 1856.

Eudendrium capillare Alder, 1856: 355, pl. 12 (figs. 9-12). Broch, 1916: 62. Stechow, 19254:
202. Leloup, 1952: 124, fig. 63. Picard, 1955: 183.

Eudendrium parvum Warren, 1908: 272, pl. 45 (figs. 1-4), fig. 1.

Eudendrium ?parvum: Millard, 1959a: 305, fig. 1G, H.

Records. South coast: SCD 154K. SH 433A.

Description. SCD 154K: Stems unbranched or sparsely branched. Perisarc
annulated at origin of stem and branches and often at other irregular intervals.
Old female gonophores present on ‘blastostyles’ which arise from stem or
hydrorhiza. Pedicel of blastostyle corrugated, bearing 3-5 gonophores or their
empty capsules irregularly distributed near distal end. Gonophores covered
by transparent capsule which has a warty appearance possibly due to shrinkage.
Soft parts too badly preserved for nematocyst examination or tentacle counts.

SH 433A: Living material kept in laboratory for several weeks. Stems
unfascicled, but profusely branched and reaching a maximum height of 1-7 cm.
Perisarc annulated on origins of stems and branches and at other irregular
intervals. Hydranths orange-pink in colour with white tentacles and hypostome;
with 23-28 tentacles held alternately elevated and depressed. Various stages
of female gonophores present on ‘blastostyles’ which arise from the stem or its
branches. Pedicel of blastostyle corrugated. Young blastostyle with fully-formed
hydranth and a ring of gonophores around its base; each gonophore with an
unbranched spadix arching over a single egg. Older blastostyles with the
hydranth showing signs of reduction and the gonophores more irregularly
distributed, each gonophore without a spadix and bearing a single embryo
surrounded by a transparent capsule. Nematocysts of two types: large isorhizas,
27 X 115 pw, and small heteronemes, 8 x 2-4 u (undischarged).

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 455

Remarks. Warren’s E. parvum is now considered to be a synonym for the
cosmopolitan F. capillare. This conclusion is based on examination of —

(i) material of E. capillare from Marseilles bearing female gonophores
and kindly supplied by J. Picard in 1958.

(ii) material of E. parvum from Knysna deposited by Warren in the
British Museum (slides 22.3.6.104-107). This is obviously not the
holotype, which came from Park Rynie, but is labelled co-type.
Both male and female gonophores present.

) Warren in 1908 summarised the differences between the two species. One
of these was the presence of 3-chambered male gonophores in E. parvum as
against two in EL. capillare. But the basal chamber in EF. parvum is very small
indeed and its development may well be a matter of degree. Moreover, Alder
in his original description of E. capillare mentions 2- or 3-chambered gonophores
(‘two or three capsules in linear series on each pedicle’). :

Another difference was the presence of a terminal tubercle on the male
gonophore in E. parvum. This appears to be a variable character. It is present
in only some of the gonophores in Warren’s material from Knysna, and is
apparently present or absent in FE. capillare (Stechow, 19252).

Further, Warren mentioned the extension of the perisarc over the base
of the hydranth in E. parvum. This perisarc is very delicate and may well have
been missed in earlier descriptions of E. capillare. It is visible in Picard’s material
of the latter from Marseilles.

_ Although Warren did not describe female gonophores in the holotype of
E. parvum, they are present in his material from Knysna and are exactly like
those of E. capillare.

Finally, the mention of small nematocysts only in E. parvum by Warren,
1908, and Millard, 1959a, is in agreement with Picard’s statement (1955) that
only small microbasic euryteles occur in E. capillare.

_ The presence of both large and small nematocysts in the present material
(SH 433A) is the only reason for the query in the identification. One is loath
to create a new species on the basis of nematocysts only, but one is forced to the
conclusion that either there are two species with exactly similar female gono-
phores or that LE. capillare has the potentiality of producing two kinds of
nematocysts, which is not always realised.

Eudendrium ?carneum Clarke, 1882

?Eudendrium annulatum : Ritchie, 1909: 70.
Eudendrium carneum: Vannucci, 1954: 101, pl. 1 (figs. 1-9), pl. 2 (fig. 8), pl. 4 (figs. 2-5). Millard
1959a@: 302, fig. 1A-F.

Records. West coast: A 118. CP 336A.

Description. Fascicled, branching stems reaching a maximum height of
about 5 cm. Colonies more heavily annulated than is usual for the species,
the groups of annulations on the main stem more common and more extensive,
often with about 15 rings; hydranth pedicels usually completely annulated,

456 ANNALS OF THE SOUTH AFRICAN MUSEUM

though some with smooth areas. Hydranths with 15 to 24 tentacles.

Young female gonophores with bifurcating spadices. Male gonophores
2-chambered.

Nematocysts of two types: large isorhizas, 20 X gp, and small
heteronemes, 7 X 4 pL.

Remarks. This material appears to be a smaller and more closely annulated
form of E. carneum. The young female gonophores are exactly the same, though
completely mature ones with basket-shaped capsules were not present. The
male gonophores are 2-chambered, whereas material from the east coast had a
minimum of 3 chambers (Millard, 19592). However, Vannucci (1954) has
described 2-chambered male gonophores in the same species.

The nematocysts are similar to those of E. carneum, though both types are
slightly smaller. The presence of large nematocysts excludes the material from
E. racemosum which also has a forked spadix in the female gonophore.

This material is very similar to that reported from Saldanha Bay by
Ritchie (1909) as E. annulatum(?). Ritchie’s material also has bifurcating
spadices. (See also p. 454.)

Eudendrium deciduum Millard, 1957
Eudendrium deciduum Millard, 1957: 184, fig. 2.
Records. South coast: LIZ 7R. MB 10R, 47N.

Description. No large nematocysts have been observed in this species.
Small heteronemes (probably microbasic euryteles) similar to those of E. carneum
(Millard, 1959a, fig. 1B) present, measuring 6-3-8-1 x 2-7—3-6 w undischarged.

Eudendrium ramosum (Linn., 1758)

Eudendrium ramosum: Hincks, 1868: 82, pl. 13. Stechow, 1923a: 83. Weill, 1934: 388, fig. 237.
Leloup, 1952: 127, fig. 64.

Records. West coast: AFR ooo2K. South coast: SCD37R (dubious
identification).

Description. The first specimen (AFR o002E) has long, graceful stems
reaching 17°5 cm. Main stem fascicled, branches unfascicled and given off in a
pinnate fashion either in one plane or twisted into a spiral. Female gonophores
present, with unbranched spadix. Gonophore-bearing hydranths with reduced
tentacles.

The second sample (SCD 37R) includes portions of a colony with a more
bushy and stiff appearance and with both main stem and principle branches
fascicled. Female gonophores present, with unbranched spadix.

Nematocysts (from AFR o002E)

(i) Small heteronemes, 7:2 X 2°7 p, present on both body and tentacles.
(ii) Large isorhizas, 18-0-18-9 < 7:2—-7:6 » present on body only.

Remarks. The first of these specimens agrees well with published descriptions

of E. ramosum and there can be little doubt as to the identification. The appear-

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 457

ance and measurements of the two categories of nematocyst in the undischarged
state are also close enough to those given for the species by Weill (1934), the
small heteronemes probably being microbasic euryteles.

The identification of the second specimen is more doubtful due to the
different appearance of the colony and the fact that the hydranths were too
poorly preserved for nematocyst examination.

This is the first record of the species from South Africa.

Family Hydractiniidae

Hydractinia altispina Millard, 1955
Hydractinia altispina Millard, 1955: 215, fig. 1. Millard, 1957: 179.
Records. West coast: B62, 92 (reported by Millard, 1955). LAM 50W,
59C. LU 59Z. SAMH 407. SB 153T.

Hydractinia kaffraria Millard, 1955
Fig. 6
Hydractinia kaffraria Millard, 1955: 217, fig. 2. Millard, 1959a: 307.

Records. South coast: BMR 23L. BRE 111A. HAM 3Q. KNY 30P, 70E,
164, 270J. SUN 3N. (All reported by Millard, 1955.)

Remarks on living material. Specimens from Knysna Estuary were kept alive
on their host snails for a period of two weeks in the laboratory in January, 1956,
when the release of the sexual products from mature sporosacs was observed.

The ripe male and female sporosacs have the structure of degenerate
medusae, with radial and circular canals and rudimentary marginal tentacles,
but the sexual products are discharged while still attached to the gonozooids
and there is no active medusoid generation.

The male sporosacs when mature are practically spherical and measure
approximately 0-65 mm. in length and 0-63 mm. in diameter. The four radial
canals are distinctly visible and around the opening are 4-6 rudimentary but
distinct marginal tentacles. The cavity is filled with spermatogenic cells
attached to the central spadix. While still attached to the gonozooid irregular
powerful contractions of the whole bell expel the active spermatozoa through
the aperture. A cloud of massed spermatozoa surrounds the sporosac for some
time until finally dispersed by the movements of the hydroids and their host.
Once the sporosacs are empty they fall off and lie loose on the bottom, but
show no further activity. At this stage they are more oval than spherical, the
length being greater than the diameter.

The mature female sporosacs are larger than the male, measuring
approximately 0-90 mm. in length and 0-86 mm. in diameter. Radial canals
are visible but the marginal tentacles are very indistinct and not so well
developed as in the male. The ripe eggs separate from the spadix and lie loose
the cavity of the bell. Fertilization apparently occurs inside the bell, since
eggs contain either one or two nuclei immediately after discharge. The area

4.58 ANNALS OF THE SOUTH AFRICAN MUSEUM

a ee Roe: eee Se

Fic. 6. Hydractinia kaffraria Millard.

A. A female sporosac in the process of releasing eggs.
B. A male sporosac.
(Both from living material.)

around the aperture of the sporosac performs irregular contractions which
constrict the opening, but no complete contractions of the bell have been
observed. These may possibly occur sporadically in the natural condition.
Under the microscope the pressure of the coverslip is sufficient to squeeze out
the eggs one by one, and the final fillip is provided by the contraction of the
margin. The diameter of the eggs is greater than that of the aperture, but their
elasticity allows them to be ejected with ease. Occasionally the sporosacs fall
off the gonozooids before all the eggs are discharged and continue to perform
gentle contractions of the margin, though no swimming movements have been
observed.
Hydrocorella africana Stechow, 1921
Fig. 7

HAydrocorella africana Stechow, 1925): 409. Millard, 1957: 183.

Records. West coast: A 405. AFR 801Q. CP 646B. HB 4D. SAMH 408.
WCD 12], 25C. South coast: AFR 866R, 967.0.V, 985E, 994M. SCD 114Q,
133B, 175U, 206R. TRA 33.00.D, 42K, 56C, 99E.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 459

Description. Common on shells of gastropods and hermits. Has been observed
on Turbo sarmaticus Linn., Argobuccinum argus (Gmelin), Fusus verruculatus Lam.,
Nassa speciosa A. Adams, and on shells occupied by the hermits Clibanarius sp.,
Pagurus arrosor (Herbst), Diogenes costatus Hend. and Eupagurus placens Stebb.

Skeleton as described by Stechow (1925) with two grades of calcareous
ridged processes; the larger 5 mm. or more in height and covered with naked
coenosarc bearing gastrozooids and gonozooids; the smaller about 0-5-1 mm.
in height, scattered amongst the hydranths. Young colonies with smaller
processes only.

Living gastrozooids reaching 3 mm. in length when extended, with 5-12
extensile tentacles, of which 1 or 2 are usually much longer than the others.

Gonozooids reduced, reaching about 0-5 mm. in length in living material,
with about 6 rudimentary, knob-like tentacles, and each bearing 3 or 4 spherical
sporosacs in various stages of development. Male and female sporosacs on
separate colonies. Male sporosacs bearing the spermatogenic cells around a
central, hollow spadix, reaching 0-33 mm. in length and 0-33 mm. in maximum
diameter. Female sporosacs containing a single central egg surrounded by a
number of blind, hollow outgrowths from the basal spadix, reaching 0-36 mm.
in length and 0-56 mm. in maximum diameter. Developing gonozooid often
enclosed by curved laminar outgrowths of the skeleton, which form a sort of
basket-work imprisoning it.

Colour: skeleton and spines white, hydranths creamy white, female
sporosacs orange.

Nematocysts of two kinds: microbasic euryteles, 0-72 < 0:27 4, and
desmonemes, 0°45 X 0:27 pL.

Details of female sporosacs. Sections were cut to elucidate the rather unusual
structure of the female reproductive bodies.

Female gonozooids are normal in structure with the exception of the
tentacles which are reduced to knobs. In each gonozooid a number of young
ova are present in the endoderm of the central region of the body. The youngest
sporosac is nothing more than a bulge in the ectoderm into which one of the
enlarging ova has been pushed together with a few endoderm cells. In one
case such a bulge contains two ova, one large and one small. At a slightly later
stage the sporosac is well-defined and contains a single large ovum seated
on a low evagination of the gonozooid endoderm, which presumably represents
a reduced spadix. A few endoderm cells are flattened against the periphery
of the egg. There is no entocodon development and the ectoderm is
single-layered.

After this the ovum enlarges enormously and becomes packed with yolky
material. At the same time from the basal spadix several (usually 4) hollow,
finger-shaped processes grow out around the ovum until they partially surround
it. These may represent radial canals, but they do not communicate distally
and there is no sign of a circular canal or any other medusoid structure. This
is the oldest stage seen in sections, but presumably fertilization occurs in situ,

460 ANNALS OF THE SOUTH AFRICAN MUSEUM

z4ng

n
Ve San

Serer

Pb cune Ce
DNS ean eg

Fic. 7. Hydrocorella africana Stechow.

A colony growing on a shell occupied by a hermit.

A section through colony and shell on a larger scale, showing many small processes, one
large one, and contracted hydranths.

Expanded gastrozooids sketched from a living colony.

Gonozooids bearing sporosacs, two female and one male.

Planula larva after release.

ts. 3 female sporosacs at different stages.

ls. female gonozooid and sporosac. Young eggs visible in endoderm of gonozooid.

an 4

.

MOO

Qs

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 461

for on several occasions gourd-shaped planulae have been found imprisoned
by the over-arching extensions of the skeleton.

Remarks. This species was described by Stechow (19250) with particular
emphasis on the skeleton, but it has never been illustrated. Stechow mentioned
male sporosacs only and failed to observe the gonozooids.

Podocoryne carnea M. Sars, 1846

Podocoryne inermis Allman, 1876: 255, pl. 10 (figs. 4-5).
Hydractinia carnea: Vervoort, 1946): 126, fig. 49. Millard, 1957: 181.

Records. West coast: LB 380C, 403C. SB 132N, 174M, 231Y, 267V,
2690B. TB 13, 14, 15, 21K. WCD 134C. South coast: KNY 212G. LIZ 3B,
eA 250). SCD 26K, 111C, 113E, 126L, 239G, 2585, 281E, 330F, 333F.
All on shells of the gastropod Nassa (Hinia) speciosa A. Adams.

Remarks. One of these colonies (SCD 26E) is without spines and would
correspond to the spineless form found in Europe (‘Podocoryne inermis’ of Allman).

Family Pandeidae

Leuckartiara octona (Fleming, 1823)
Leuckartiara octona: Rees, 1938: 12, figs. 3-5. Millard, 1957: 182.

Records. South coast: LIZ 3A. SCD 20J, 26D, 94B, 258T, 281F, 314C,
333G.

Remarks. To avoid misidentification, only those specimens with medusa
buds are included in the above records. SCD 281F was growing on the shell
of Nassa speciosa A. Adams, SCD 314C on Nassa analogica Sow. and all others on
Bullia annulata (Lam.). |

Family Aequoreidae

Aequorea africana n. sp.
Fig. 8
Holotype: MB 70G from Mossel Bay on the south coast. South African
Museum catalogue number: SAMH 413.

Description. A colony with well-extended hydranths growing on an empty
snail-shell.

Stem unbranched or branching sympodially up to three times, reaching a
maximum height of 0-2 cm. Stem and branches annulated or corrugated,
increasing in diameter from base to distal end, bearing terminal hydrothecae.
Shorter stems closely and distinctly annulated throughout, longer ones closely
annulated in basal region, irregularly corrugated in more distal part.

Hydrotheca thin and membranous except for base which is somewhat
stouter and remains as a saucer-shaped Halecium-like structure in damaged or
regenerated specimens (fig. 8A), with distal region irregularly creased and
folded longitudinally to form an operculum below which the hydranth can be

462 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 8. Aequorea africana n. sp.

A. An empty stem surmounted by a saucer-shaped structure (the remains of the hydrotheca).
B-E. Hydrothecae containing hydranths in various stages of expansion, E with a gonotheca.
F and G. Gonothecae containing medusa buds, F arising from hydrorhiza, G from the stem.
H. An expanded hydranth showing the web between the tentacle-bases.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 463

withdrawn. Hydranth long and extensile, with conical hypostome and 11-17
tentacles united by a web at their bases (fig. 8H). Tentacles often moniliform
in appearance.

Gonophores borne on stem or hydrorhiza, each on an annulated
pedicel which increases in diameter towards the distal end. Gonotheca
not sharply demarcated from pedicel, pear-shaped, containing one young
medusa-bud.

Measurements (mm.)

Stem, length ; : . Or18—2°05
maximum diameter ; . 0°045-0:07
Hydrotheca, length : ’ » 0°32-0°55
maximum diameter : ; ; Obi O16
Hydranth, length to hypostome (preserved) . reaching 0:72
Gonotheca, length, with pedicel : : : ; . 0°29-0°40
maximum diameter : . OF12-O'15

Remarks. This species is closely related to several members of the ‘Campanu-
linid’ group of hydroids, in particular to the following:

(i) Campomma hinckst (Hartlaub, 1897). This species was linked by
Hartlaub with the medusa Eucheilota maculata WHartlaub, 1894,
probably erroneously according to Rees (1939, p. 442) and Russell
(1953, P- 313). Campomma hincksi is the sole representative of its
genus, which is retained provisionally only until such time as the
medusa can be definitely placed. Both C. hinckst and E. maculata are
known only from Europe. Aequorea africana strongly resembles Leloup’s

material of C’. hincks: from Belgium (1952, fig. 70).

(ii

_

Campanulina paracuminata Rees, 1938. This is almost certainly the
hydroid of Aequorea aequorea (Forskal, 1775) (= Aequorea forskalea
Péron and Lesueur, 1809; see Russell, 1953, p. 350, and Huvé, 1952,
p. 36). The medusa is known from Atlantic coasts including South
Africa (Kramp, 1961).

(iii) Campanulina acuminata (Alder, 1857). This is probably the hydroid
of Aequorea vitrina Gosse, 1853 (see Russell, 1953, p. 354) or of Aequorea
pensilis (Haeckel, 1879) (see Huvé, 1952, p. 37). A. vitrina occurs
in N.W. Europe and A. pensilis in the English Channel and
in the Pacific and Indian Oceans including South Africa (Kramp,
1961).

The present material may well be Campanulina paracuminata, whose medusa

is known from South Africa and material of which was examined in the British
Museum in 1960; but as there are slight structural differences (e.g. in the nature
of the operculum and the number of tentacles) and as there are other South
African species of Aequorea with unknown hydroids (e.g. A. coerulescens (Brandt,
1838) and A. macrodactyla (Brandt, 1838)), a separate species has been
established as a temporary measure.

464 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Lovenellidae

Lovenella chiquitita Millard, 1957
Lovenella chiquitita Millard, 1957: 198, fig. 7. Millard, 1959b: 250, fig. 3.

Records. West coast: A 384D. SWD 39G. TRA 156C.

Remarks. ‘The placing of this species in the genus Lovenella is a temporary
measure only and is based on the morphology of the hydroid generation,
which shows close affinity to Lovenella clausa (Lovén, 1836).

The medusa, in its absence of cirri at the time of liberation, is more
closely related to Phialella. That it is not P. quadrata (Forbes, 1848) was estab-
lished by comparison with material of the hydroid generation of the latter in
the British Museum in 1960, for which privilege the author is indebted to
Dr. W. J. Rees. It might well be P. falklandica Browne, 1902, the medusa of
which has been found in South Africa at Saldanha Bay (identification by
Dr. M. E. Thiel, Zoologisches Museum, Hamburg), yet the youngest known
medusae of this species (0-8—-1-0 mm. diameter) have ‘four perradial tentacles,
and the four interradial tentacles are just beginning to develop and are visible
as four minute bulbs’ (Brown and Kramp, 1939: 298), suggesting that they are
liberated with only 4 tentacles and not 8 as in the present species.

Family Haleciidae

Halecium beanu (Johnston, 1838)
Fig. 9 A-F
Halecium beanii: Hincks, 1868: 224, pl. 43 (fig. 2). Broch, 1918: 38, fig. 13. Millard, 1957: 188.

Millard, 1958: 168. Ralph, 1958: 332, fig. 10 a, b, e-k. Vervoort, 1959: 224, fig. 6.

Records. West coast: CP 336C. SB 196 M. SWD 12D, 42F. TB 17A, 21B.
WCD 125U, 145V. South coast: CPR 46L. LIZ 7S. MB 47T, 60Q. SAMH 157,
214, 273, 328, 335, 341. SCD 37M, 85M, 154C, 387G, 394B.

Description. Stiff, shrubby colonies, many of them epizootic on other
hydroids, the largest reaching a height of 7-4 cm.

Remarks. Since no criterion can be found for distinguishing between
sterile colonies of H. beanit and H. halecinum, only samples containing female
gonophores have been included above. The species is certainly more common
than is indicated by the records, as a further 26 sterile samples are present in
the collection, all, or most of which, probably belong to the same species.

Attempts have been made by various authors to distinguish H. beani from
H. halecinum on the basis of the angle of the hydrothecal margin and the shape
of the basal part of the hydrophore, but, as the accompanying diagrams (fig.
g A-E, all chosen from fertile specimens) will show, both characters are so
variable that no reliability can be placed upon them.

Halecium delicatulum Coughtrey, 1876
Fig. 10L
Halecium parvulum: Millard, 1957: 189, fig. 4A. Vervoort, 1959: 227, fig. 7.

Halecium parvulum, var. magnum Millard, 1957: 190, fig. 4 B—O.
Halecium delicatulum: Ralph, 1958: 334 (synonymy), figs. 11e, h-n, 12 a-p.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 465

Fic. 9. Halecium beanii (Johnston) (A-F), and H. halecinum (Linn.) (G-L).
A-E and G-K, hydrophores from various colonies.
F and L, gonothecae.

Records. West coast: AFR 736Z. CP 327C. LAM 30P, 35A. TBo, 17C,
21C. WCD 18T, 156A, 160P. South coast: LIZ 16F. MB 19Q, 84E, 88J.

SAMH 179, 189, 242, 250, 381. SOD 5F, 22A, 29H, 37N, 52J, 85K, 154B,
179H. TRA 92M.

466 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks. It has been pointed out by Ralph (1958) that the large variety
(var. magnum Millard) described in 1957 has similar dimensions to the type
material of H. flexile Allman, 1888, which is now recognized as a synonym for
H, paroulum and H. delicatulum. Since so great a variation of size is possible
it is not justifiable to retain a subspecies on this character alone.

Halecium dichotomum Allman, 1888.
Fig. 1o A-K

Halecium dichotomum Allman, 1888: 13, pl. 6. Billard, 1910: 4. Stechow, 19256: 419. Millard,
1957: 188 (excluding the male gonophores).

Records. West coast: WCD 20G, 164C. South coast: LIZ 7W. MB 8V,
12W. SAMH 162, 227, 243, 274, 352. SCD 5G, 20J, 37P, 81P, 85N, 154D,
1751, 1885, 239F, 265G, 333C, 387F, 394A. TRA 38], 92N.

Description. Colonies very variable in appearance and growth-form. Stem
fascicled, branching with the typical dichotomy described by Allman and
generally in all planes, usually geniculate, with a tendency for annulation in
the region of the nodes. Primary hydrophores usually sessile, secondary hydro-
phores usually symmetrical and annulated in the basal region. Hydrotheca
low and wide, with walls flared outwards, over 0:17 mm. in diameter at the
margin.

Among the variety of colonies examined two extremes of growth-form
occur:

(i) Large upright colonies reaching 11-12 cm. in height, with strongly

fascicled stems and branching mainly in one plane. Main stem more
or less straight and dichotomy not obvious. Larger stems and branches
stiff in appearance and up to 2 mm. thick, though unable to support
themselves out of fluid. Smaller branches graceful and flexuous.
Stem and branches with long internodes with no annulation other
than a shallow constriction near the base. Secondary hydrophores
scarce. Closer examination shows that the typical dichotomy is in
fact present in this form, but that one limb is always short (the branch)
and the other long, contributing to the axis of the stem. The latter
limb is enveloped by peripheral tubes in fascicled regions, so obscuring
the dichotomy (fig. 10 A, B).
Low, scrubby colonies, often epizootic, reaching a height of 1-2 cm.
Stem usually weakly fascicled and strongly geniculate. Branching
profuse and in all planes, and stolonisation common, resulting in a
tangled mat which may cover large areas of the substratum and is
very easily recognised. The typical dichotomy is common in this form
and very obvious (fig. 10 C). The internodes of the stem and branches
tend to be shorter and more annulated, and secondary hydrophores
are abundant and often closely annulated in their basal regions.

In two specimens (MB 12W and WCD 164C) of the low, scrubby form,
the colony is epizootic on a dead polyzoan and the hydrorhiza is provided with

(ii

ee

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 467

\ :
) 0 :

a
a a eel

Fic. 10. Halecium dichotomum Allman (A-K) and H. delicatulum Coughtrey (L).

A and B. Portions of stem from the distal and proximal ends respectively of the tall form, to
show the incorporation of one limb of the dichotomy into a main stem. Peripheral tubes
teased apart in B. (TRA 92N.)

Cand D. Portions of stem from TRA 38J, showing typical dichotomy in C and a unilateral
branch in D.

E and F. Female and male gonophores.

G-J. Details of secondary hydrophores.

K. Part of the hydrorhiza to show root-like structures (MB 12W).

L. Part of a branching stem in H. delicatulum for comparison.

468 ANNALS OF THE SOUTH AFRICAN MUSEUM

unusual rootlike projections which enter the pores of the host and anchor the
colony (fig. 10 K). Similar ‘roots’ have been observed on other mounted
specimens.

Female gonophores as described by Allman. Male gonophores on separate
colonies, slender, elongated and often curved, tapering distally to a small
rounded aperture, closely annulated throughout or smooth in distal region
(fig. 10 F). Reaching 1-24 mm. in height and 0:36 mm. in maximum diameter.

Remarks. The variation in the growth-form of this species is remarkable,
and at first glance the two extremes might be taken for different species.
However, female gonophores have been found in both, and intermediate
types exist. Comparison with Allman’s type material in the British Museum
(slide no. 88.11.13.9) confirms the identification, and shows that the hydro-
phores are similarly variable. Primary hydrophores, though usually sessile,
may be quite long. The hydrotheca may be strongly flared out, weakly flared
at the extreme distal edge only (the commonest type), or the everted part
may be quite worn off.

This variation may introduce difficulties in the identification of sterile
colonies and possible confusion with species such as H. delicatulum, though to one
familiar with the species no difficulty arises. Useful diagnostic characters are
the normally sessile primary hydrophores, the typical dichotomy in which 2 or
even 3 internodes arise from a previous one at the same level, and the character-
istic curved apophysis by which each internode arises from its predecessor.
Dichotomy may also occur in H. delicatulum, but the two limbs usually arise
at different levels and the apophyses are not curved (fig. 10 L). A. dichotomum
may be distinguished from JH. tenellum by its fascicled stem and larger
hydrothecae.

With the abundant material available, it was possible to clear up the
confusion which previously existed over the nature of the male gonophores
of this species. A re-examination of the material described from False Bay
(Millard, 1957: 188) showed that the smooth gonophores from sample FAL 78Z
belonged in fact to an epizootic male colony of H. beanii inextricably tangled
with a female colony of H. dichotomum. The annulated gonophores without
lateral openings from sample FAL 64N and thought to be female, were in
fact male.

Halecium halecinum (Linn., 1758)
Fig. 9 G—L
Halecium halecinum: Hincks, 1868: 221, pl. 42. Broch, 1918: 36, fig. 11. Vervoort, 1946): 158,
figs. 63-64. Vervoort, 1959: 225.
Records. South coast: SCD 85P.

Description. A small epizootic, yet fertile, colony, reaching a maximum
height of 0-6 cm. Stems stunted and branching irregularly, but fascicled at base.
Internode length about twice width at distal end.

Secondary hydrophores very variable, most are asymmetrical, curving

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 469

towards abcauline side and with an obliquely set aperture. Some are quite
symmetrical in the basal region, though asymmetrical more distally. In only
a few is the aperture perpendicular to the axis.

Female gonothecae banana-shaped with concave adcauline side and
terminal aperture. Containing a single row of 3-4 large eggs.

Remarks. This species is possibly more abundant than is indicated by this
single record, as without gonophores it is impossible to distinguish from
H. beanu. This is the only fertile colony observed and the first fertile, and
therefore unquestionable, record from South Africa.

Halecium ?muricatum (Ell. & Sol., 1786)
Fig. 11 A, B
Halecium muricatum: Hincks, 1868: 223, pl. 13 (fig. 1). Broch, 1918: 43, fig. 17. Vervoort, 19460:
163, fig. 67.
Records. West coast: TB 19A. South coast: SCD 56U.

Description. Fascicled stems reaching a maximum height of 4-7 cm.,
branching in an irregularly pinnate fashion, straight for most of length though
sometimes weakly geniculate in distal regions, fairly rigid and able to support
themselves out of fluid, though more graceful in appearance than in H. beanit.
Nodes oblique and sloping alternately to left and right.

Primary hydrophore long; borne on broad apophysis at distal end of
each internode, from which it is separated by a distinct groove; usually asym-
metrical with adcauline side more convex than abcauline; with a well-marked
pseudodiaphragm in distal region below hydrotheca and usually a second one
in proximal region. Pseudodiaphragm better developed on adcauline side.
Secondary hydrophores similar, though not so long and usually with only one
pseudodiaphragm. Hydrotheca relatively deep, margin everted, more so on
adcauline side.

Gonophores absent.

Measurements (mm.)

TB19A SCD 56U

Stem, internode length . . 0°70-0°86 0:61-0:77
diameter across node ! . O'19-0'24 0°12—-0°28
Hydrotheca, diameter at margin. ; 5 . 0°24-0°31 0°24—0°29
depth (diaphragm to margin) . . 0°09-0°14 0:08-0-12

Remarks. The identification of this species must remain dubious until the
gonophores have been discovered. The stem and hydrophores are remarkably
similar to those illustrated by Broch (1918) for H. muricatum, yet H. muricatum
is an arctic species and has not been reported from the Southern Hemisphere.

Another closely related species is H. filicula Allman, 1877, from the Gulf
Stream and N. Atlantic Ocean. This species has smaller dimensions (cf.
Billard, 1906: 163) and differs in the presence of 2 annulations at the base of
the hydrophore and in the absence of a pseudodiaphragm. The gonothecae

470 ANNALS OF THE SOUTH AFRIGAN MUSEUM

(Billard, 1906, fig. 2) appear to be similar to those of H. muricatum, and the

two species may prove to be conspecific.

0'S5mm.

Fic. 11. Halecium ?muricatum (Ell. & Sol.) (A-B) and H. tenellum Hincks (C-F).

A and B. Portions of stem from TB 19A and SCD 56U respectively.
Cand D. Female gonothecae in side and front views respectively, D with 2 larvae (SCD 37Q)
E. Part of colony and female gonotheca with several larvae (SCD 60B).

F. Male gonotheca from SAMH 315.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 471

Halecium tenellum Hincks, 1861
Fig. 11 C-F
Halecium tenellum: Millard, 1957: 193, fig. 5. Hamond, 1957: 307, fig. 14. Ralph, 1958: 340.
Vervoort, 1959: 229, fig. 8. Naumov, 1960: 454, fig. 344.
Records. South coast: MB 8X. SAMH 315. SCD 37Q, 60B, 394C.

Description. A number of colonies, mostly epizootic, with unfascicled
stems reaching a maximum height of 0-4 cm. Stems geniculate and often
branching in a dichotomous manner, usually with many athecate internodes
as previously figured (Millard, 1957), but often of more normal appearance
as figured by Vervoort (1959) and Hamond (1957).

Male and female gonophores present, on separate colonies. Male
gonothecae compressed, broad and bluntly rounded at distal end when young,
pointed at distal end when mature, reaching a maximum of 1:07 mm. in
length and o-60 mm. in diameter. Female gonothecae borne on stem or
hydrorhiza, compressed, pear-shaped in anterior view with bluntly rounded
distal end, and a circular, terminal aperture blocked by a small papilla which
is shed on the escape of the contents, reaching a maximum of 1:07 mm. in
length, 0-51 mm. in diameter and 0-30 mm. in thickness; with no hydranths;
containing 2—7 eggs on a branching blastostyle, which develop into planulae
in situ.

Remarks. Vervoort was correct in assuming that the male gorophores
previously described (Millard, 1957) were young ones, as larger mature ones
in the process of shedding their contents have now come to light.

Family Campanulariidae

Campanularia hincksiw Alder, 1856
Fig. 12 A-D
Campanularia Hincksii Alder, 1856: 360, pl. 13 (fig. 9). Hincks, 1868: 162, pl. 24 (fig. 3), fig. 18
Billard, 1906: 172, figs. 4-5 (incl. var. grandis).
Campanularia hincksii: Vervoort, 1959: 311, fig. 55a.

Records. South coast: SAMH 283. SCD 354G.

Description. Pedicels unbranched and smooth, except for occasional
regeneration lines, with a single spherule at distal end.

Hydrotheca inverted cone-shaped, with 8-12 broad, marginal teeth in
which the distal end is either hollowed out to form two points or (more rarely)
truncated. Hydrotheca polygonal in end-on view, the angles between the
teeth forming longitudinal striations visible in side view, which may continue
to the base or peter out half-way down. Diaphragm in form of annular thecal
thickening.

Female gonotheca with very short stalk, elongated, broad near base and
narrowing slightly to truncated distal end, with about 8 low, rounded annula-
tions. Only 2 gonothecae seen, one empty and one almost spent yet still
containing 2 eggs.

4.72 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm.) SAMH 283 SCD 354G
Pedicel length ; : : : . 2:26-4:19 3°70-6°81
maximum diameter ; : . 0°08.0°13 O*12-0°15
Hydrotheca, length : . 085-124 1°20—-1°55
maximum diameter : ! . 0°47-0°77 0:60-0:88
length/diameter t : . 1:36-1:97 1°48-2-27
Gonotheca, length . ; ; 1°52
maximum diameter O64

Remarks. This is the first record of the species from South Africa. The
measurements correspond well with Billard’s var. grandis, but the material
resembles the typical form more in the double nature of the marginal teeth.
These teeth appear to become truncated as they wear down with age.

Campanularia integra MacGillivray, 1842
Fig. 13 A-D
Campanularia caliculata: Warren, 1908: 338, fig. 19.
Campanularia integra: Billard, 1907: 340. Jaderholm, 1917: 4. Broch, 1918: 159 (synonymy).
Millard, 1957: 193. Millard, 1958: 171.
Clytia compressa: Vanhoffen, 1910: 303, fig. 24.
?Campanularia gracilis: Stechow, 1925): 423, fig. 6.
Orthopyxis caliculata: Ralph, 1957: 838, fig. 6 a—f.

Records. South coast: CPR gE. LIZ 7U, 13C. MB 52L, 55J. SCD 84X,
179K.

Description. Colonies creeping on weeds and other hydroids, particularly
Lytocarpus filamentosus.

Remarks. In the identification of this species I have followed Broch (1918)
who considers C. integra, C’.. caliculata and C. compressa as synonymous, although
this opinion is not accepted by all recent authors. The difference between
these species is supposed to lie in the gonotheca, which is said to be spirally
annulated in C’.. integra, smooth and round in section in C. caliculata, and smooth
and compressed in C. compressa. Broch claims to have found intergrading forms
between C. integra and C. caliculata, while the South African material shows
intergrading forms between C’. caliculata and C. compressa. Here the gonotheca
is usually compressed and smooth, but may also be round in section, and is
sometimes roughly corrugated, though not distinctly annulated.

Medusa systematists distinguish the ‘medusa’ of C. caliculata (Agastra mira
Hartlaub, 1897) from that of C. compressa (Agastra rubra Behner, 1914) on
minor points such as the arrangement of eggs. Of the two, the South African
material resembles more Agastra rubra, as the eggs are large and comparatively
few in number.

Campanularia laminacarpa n.sp.
Fig. 12 E-K
?Campanularia tincta: Jaderholm, 1923: 6
?Campanularia africana: Stechow, 19256: 420, 421 (material from Agulhas Bank, station 105).

Non Campanularia tincta: Warren, 1908: 337, fig. 18.
Non Campanularia africana Stechow, 1923): 104.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 473

Fic. 12. Campanularia hincksii Alder (A—D) and C. laminacarpa n. sp. (E-K).
A-C, hydrothecae, and D, gonotheca, from SAMH 283.
E-G, hydrothecae from TRA 38P, and K, from SCD 84U.
H. Male gonophore from SCD 84U.
J. Female gonophore from the holotype, TRA 32A.

474 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype: TRA 32A, a female colony from the Agulhas Bank growing on
Thyroscyphus aequalis Warren. South African Museum registered number:
SAMH 414.

Other records: South coast: SAMH 174, 269, 351. SCD 37V, 61F, 79M,
84U, 112A, 117L, 154E, 169Y, 265E, 394F. TRA 35Z, 38P, 56U, g2B.

Description. Colony stolonic and epizootic on the Sertulariid Thyroscyphus
aequalis. Hydrothecal pedicel upright, unbranched, smooth or corrugated, with
a spherule of smaller diameter at the distal end.

Hydrotheca tubular or slightly expanding to margin, smooth, with length
about twice diameter at margin, with a diaphragm in the form of a well-
developed annular thickening of the perisarc, with 10-14 rounded marginal
teeth.

Gonotheca erect, flat, smooth, generally held in a plane at right angles
to the stem of the host, broadening to distal end which is abruptly truncated,
with a wide, operculate aperture. Male and female similar. Female containing
a single sporosac bearing numerous small eggs. Male with a single sporosac
with the structure of a degenerate medusa, spermatogenic cells arranged in
4 longitudinal bands with indications of 4 radial canals.

Measurements (mm.)

Hydrothecal pedicel, length . . : . O-ZI-1°34
maximum diameter ; . P . 0:06-0'11
Hydrotheca, height : : : : ‘ : . 0°36-0°71
diameter at margin . : , : ; ; .0°225-0°34
diameter/height é : ; : . 0°46-0°74
Gonotheca, height . ; : ‘ ; ; . I*53-2-01
maximum diameter ‘ . O-81—1-20

Remarks. This species has previously been confused with Campanularia
africana Stechow, 19236. The discovery of gonothecae, which are larger and
quite different in appearance from those of the latter species, necessitates the
establishment of a new species. Unfortunately the two species have identical
trophosomes and there is no method of distinguishing sterile material.

Campanularia africana was originally described from Park Rynie, Natal, as
C. tincta by Warren in 1908 (p. 337, fig. 18). It has cylindrical to ovate gonothe-
cae, narrowing towards the distal end and then everted to form a short, circular
collar around the aperture. Although there are variations in shape the
gonothecae are never flattened, but always round or nearly so in section. I
have examined Warren’s material, which was kindly loaned to me by the
Director of the Natal Museum, and found his diagrams to be a faithful represen-
tation of the structure. The gonothecae contain a number of large planulae.
It was to this material that Stechow (1923) gave the new name of Campanularia
africana. The species presumably also includes Pennycuik’s material from
Australia (1959: 169), which possessed male gonothecae, one with ‘5 shallow
annulations’.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 475

Fic. 13. Campanularia integra MacGill. (A—D) and C. ?mollis (Stechow) (E-J).

A and B, Gonothecae, and C and D, hydrothecae from CP 258.
E-J, various hydrothecae from CP 646C showing variation in thickness of walls.

476 ANNALS OF THE SOUTH AFRICAN MUSEUM

Stechow (19255) stated that he had examined material loaned to him by
Warren. This was obviously not the type material, and was not recorded by
Warren, for it came from Algoa Bay, and it was growing on Thyroscyphus
regularis (= T. aequalis) and presumably unfertile. This material was probably
C. laminacarpa, as was Stechow’s Valdivia material (unfertile) reported in the
same paper, at any rate that growing on T. aequalis. C’. laminacarpa possibly
only occurs on YT. aequalis as I could only identify gonothecae in material
growing on this host, and I have only quoted records (above) of material
growing on Thyroscyphus. Of these, 7 have gonothecae of the flattened type.

Several other sterile colonies present in this collection, and growing on
other species of hydroids, cannot be identified with certainty, nor can unfertile
material in the literature.

Campanularia ?mollis (Stechow, 1919)
Fig. 13 E-J

Clytia mollis Stechow, 1919: 44, fig. L.
Campanularia mollis: Picard, 1951: 344, fig. 3.
Orthopyxis mollis: Ralph, 1957: 840, fig. 7e-k.

Records. West coast: CP 646C.

Description. A flourishing colony growing on weed. Pedicel generally
smooth, with a single small spherule at distal end, often with regeneration nodes.
Hydrotheca cylindrical, with length 2-3 times height, with 7—9 clearly defined
marginal teeth separated by broad and shallow bays, diaphragm in form of
annular thecal thickening. Marginal teeth sometimes bifurcated at the tip.
Margin often reduplicated.

Perisarc of pedicels very thick, that of hydrothecae variable, sometimes
thick throughout, sometimes thin for the most part though always thickened
at margin and near base.

Gonothecae absent.

Measurements (mm., without reduplications).

Pedicel, length . 0°58—1°87
maximum diameter ! . O'10—0°17
Hydrotheca, length : : : . 0°57—-0°80
diameter at margin . : : : . 0°:26-0°33
length/diameter : ; . 1*94-2°93

Remarks. The identification of this species must remain doubtful in the
absence of gonothecae, as there are several closely related species with similar
trophosomes. It is close to C’. mollis though differing from previous descriptions
of the species in the larger hydrothecae, better defined marginal teeth and
smooth pedicels. A few specimens do, however, show indications of a spiral
twisting at the base of the pedicel. The species has not been recorded from
South Africa before.

ee

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 477

Campanularia morgansi Millard, 1957
Campanularia morgansi Millard, 1957: 195, fig. 6. Millard, 1958: 171.
Records. South coast: LAM 30M. MB 47Y. SAMH 202, 221, 230, 206.
SCD 37W, 84W, 117K, 354F. West coast: WCD 12H.

Genus Clytia Lamouroux, 1812

Remarks on type species. The genus Clytia was established by Lamouroux in

1812 (p. 184) for 3 species, namely:
Sertularia volubilis
Sertularia syringa Linn., 1767
Sertularia verticillata Linn., 1758

Of these, S. syringa has been transferred to Calicella Hincks, 1859, and
S. verticillata is generally considered as a species of Campanularia but has recently
been declared the type species of a new genus Verticillina by Naumov (1960).

In the literature 2 species have been confused under the specific name of
volubilis :

(i) The name Sertularia volubilis was originally established by Linnaeus
in 1758 (p. 811) for Ellis’s ‘small climbing Coralline with bell-shaped
cups’ described and figured by the latter in 1755 (p. 24, pl. 14, fig.
a, A). But Ellis’s description was very inadequate and the material
might be either Campanularia or Clytia. However, the name has been
retained for a well-known species of Campanularia with fixed sporosacs
and a smooth gonotheca which has been declared by Naumov in
1960 to be the type species of this genus (i.e. Campanularia volubilis
(Linn., 1758)).

(ii) In 1786 Ellis and Solander (p. 51, pl. 4, fig. e, f, E, F) described
and figured under the name of Sertularia volubilis material with an
annulated gonotheca which is unmistakably recognisable as the
species later described by Alder in 1856 as Campanularia johnstont.
Ellis and Solander added the corollary that ‘there are different
varieties and sizes of this twining bell-shaped Coralline’. It is to this
description that Lamouroux referred when he created the genus
Clytia.

So far as I can determine no type species has been established for the genus
Clytta and I therefore select Sertularia volubilis Ellis & Solander, 1786, non
Sertularia volubilis Linn., 1758.

However, the specific name volubilis is obviously untenable. The name
johnston Alder, 1856, is antedated by the name of the medusa of the same species,
namely Medusa hemisphaerica bestowed by Linnaeus in 1767 on the ‘Medusa
hemisphaerica’ of Gronovius, 1760 (the latter name being part of a Latin descrip-
tion and thus not valid). The same medusa was later placed in the genus
Phialidium Leuckart, 1856, by which name it is now generally known, but which
is antedated by Clytia Lamouroux, 1812.

478 ANNALS OF THE SOUTH AFRICAN MUSEUM

To sum up, the name of the type species of the genus Clytia becomes
Clytia hemisphaerica (Linn., 1767).
Synonyms: Medusa hemisphaerica Linn., 1767
Phialidium hemisphaericum (Linn., 1767)
Sertularia volubilis Ellis & Solander, 1786, non Linnaeus, 1758.
Campanularia johnstoni Alder, 1856

Clytia hemisphaerica (Linn., 1767)
Fig. 14 A-F
Laomedea gracilis: M. Sars, 1857: 160, pl. 2 (figs. 1-3, 5).

Campanularia johnstont Alder, 1856: 359, pl. 13 (fig. 8). Vervoort, 1959: 312.
Clytia raridentata: Vanhoffen, 1910: got, fig. 22. Fraser, 1944: 145, pl. 26 (fig 118). Millard,

1957* 197-
Campanularia raridentata: Stechow, 1919: 58, fig. Q.
Thaumantias raridentata: Stechow, 1923a: 107, fig. M. Stechow, 19256: 426.
ila atl Stechow, 1925): 431, figs. g-10. Millard, 1957: 196. Millard, 1958: 172, fig. 3B,
Clytia cae : Ralph, 1957: 820, 823, figs. 1h-u, 2, 3a~f. Millard, 1958: 172, fig. 3A, D, F.
Laomedea (Phialidium) pelagica: Vervoort, 1959: 313, fig. 558, ¢.

Records. West coast: SB 178C. TB 7, 16. South coast: LIZ 7T, 11K, 40K.
MB 8N, 52K, 60N, 64P, 69B, 81W. SAMH 161, 166, 176, 180, 203, 239, 253,
383. SCD 5L, 37X, 50N, 52U, 56V, 61G, 75H, 79P, 84V, 129E, 179D, 265],
283T, 284E, 330C, 387L, 394G. STJ 31M. TRA 57D, g2W, 150B.

Description. Numerous colonies growing on weeds and other hydroids.
Stems solitary or occasionally sparsely branched, annulated at top and bottom
and occasionally throughout. |

Hydrotheca variable in size, with 8-15 marginal teeth. Teeth acute,
sharp or rounded, but always covering a smaller area than the bays between
them, often asymmetrical and leaning towards one side. Diaphragm distinct,
variable in thickness, but always clearly demarcated from the hydrothecal
wall.

Gonotheca generally smooth, with truncated distal end, but sometimes
with I or 2 irregular corrugations; none of these specimens with distinct
annulations. |

Remarks. Ralph has demonstrated for Clytia johnstoni in New Zealand a
variation in size, shape of marginal teeth and degree of annulation on the
gonotheca which can be correlated with latitude. While specimens from the
southern region correspond to the classical conception of C. johnstoni, with
annulated gonothecae and broad marginal teeth, those from the northern
region have smooth gonothecae and acute marginal teeth and thus cover the
form known as C. gracilis. We thus have no alternative but to combine the two
species. So far as is known there is no difference between the medusae. The
range of variation also covers material from South Africa recorded as
C.. raridentata, which is here considered a synonym. See also remarks on p. 477.

Since the latitude on the south coast of Africa is the same as that of the
north end of New Zealand, it might be expected that the material in these

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 479

Fic. 14. Clytia hemisphaerica (Linn.) (A-F), and C. hummelincki (Leloup) (G-L).

A-D. Various hydrothecae showing variation in total size and shape of marginal teeth: A
with triangular teeth, B and D with asymmetrical teeth and C with very slender teeth.

E and F. Gonothecae, corrugated type (a smooth one shown in A).

G. A hydrotheca containing a hydranth, and a gonotheca.

Hand J. Empty hydrothecae.

K and L. Gonothecae containing young medusae. :

(A from TB 16, B from TRA 57D, C from Natal, D-F from False Bay, G-L from SCD 328H.)

480 ANNALS OF THE SOUTH AFRICAN MUSEUM

two regions would be similar, and that the variation in South Africa would
continue from where it left off in New Zealand as one moves up the east coast
into warmer waters. This was tested out in the material available and
measurements of hydrothecal size and pedicel length are given below.

Hydrotheca: height (mm.) Pedicel: length (mm.)
Range Mean Range Mean
West coast o:6-1:0 0:8 0°7—3°4 2°4
(32-34° S) (16) (16)
South coast 0°4—0'9 0-6 0°5-5°7 1°9
(34-35° 8) (40) (39)
East coast 0-4-0°8 0-6 1-6—3°8 2°5
(32-28° 5) (32) (20)
Portuguese
East Africa 0°3—-0°6 O°5 0°4-1°6 I‘O
(27-23° S) (23) (18)

These figures show that the hydrothecal height from the south coast
corresponds well with that from a similar latitude in New Zealand (Ralph
gives measurements of 0:40-0:81 mm. between 40° S and 34° S) and that there
is a clear decrease in size from the west coast (where the water is colder than
the south coast) round the south and up the east coast. A similar trend in
pedicel length is not so clear, although pedicels from Portuguese East Africa
are certainly shorter than elsewhere. No tendency for a change in the proportions
of the hydrotheca was seen.

The marginal teeth are always acute in South African material as in
northern New Zealand, and on the east coast there is a tendency for the bays
between them to become larger and the teeth narrower (fig. 14C).

With one exception the gonothecae are quite smooth or with a few irregular
corrugations, as in those illustrated by Ralph from northern New Zealand.
This is as might be expected, and it is difficult to account for the single sample
where all gonothecae are distinctly annulated (6-8 rings) described from
Portuguese East Africa as C. johnstoni (Millard, 1958, fig. 3D).

The variable nature of the gonotheca explains the discrepancy which
exists in the literature over the nature of the gonotheca of Clytia (Thaumantias)
raridentata (cp. Fraser, 1944, and Stechow, 1923).

The nature of the diaphragm allows for no differentiation between species,
and, although variable in thickness, it is always distinct from the hydrothecal
wall and thus different from the type characteristic of the genus Campanularia.

Clytia hummelincki (Leloup, 1935)
Fig. 14 G—-L

Laomedea hummelincki Leloup, 1935: 19, fig. 7.
Records. South coast: SCD 328H.

Description. A rich colony growing on the surface of Lepas sp. taken from a
buoy. Colony stolonic, stem unbranched, long, closely annulated in basal

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 481

region and often for short areas at other irregular intervals, otherwise smooth,
with a single, rather flattened ‘spherule’ at distal end.

Hydrotheca in the shape of an inverted cone and usually with straight
sides, expanding evenly to margin, with height approximately equal to
maximum diameter. Margin untoothed. Diaphragm very delicate, usually
oblique. Hydranth with 15-29 tentacles, completely retractable into
hydrotheca.

Gonotheca arising separately from hydrorhiza on a short pedicel of 2-4
segments, elongated and expanding to distal end which is truncated, containing
I or 2 medusa buds. Oldest medusa deep, with manubrium, 4 radial canals
and 4 tentacle bulbs visible.

Measurements (mm.)

Pedicel, length : : ; . 1°92—4°73
maximum diameter ; : d . 0°09-0'15
Hydrotheca, length ; , : ; ; . 0°25-0°38
diameter at margin . : : . 0°22—0°42
length/diameter ’ . 0°70—1°36
Gonotheca, length . ; . 0°73-1:26
maximum diameter ; , ; . 0°28-0°45

Remarks. This material agrees entirely with Leloup’s description of
Laomedea hummelincki from the West Indies, except that these pedicels are
somewhat longer and stouter. The nature of the diaphragm and the presence
of medusa buds in the gonotheca (observed here for the first time) shows the
species to be a Clytia. This is only the second record of this rare species, and a
new record for South Africa.

Clytia paulensis (Vanhéffen, 1910)
Fig. 15

Campanularia paulensis Vanh6ffen, 1910: 298, fig. 19 a, b.
Clytia paulensis: Stechow, 1919: 45, 155. Stechow, 1923a: 110, fig. N. Stechow, 1925): 428,

fig. 7. Stechow, 19254: 211.
?Clytia ulvae Stechow, 1919: 47, fig. N. Stechow, 1925): 428.

Records. South coast: LIZ 11M. SAMH 336. SCD 79N, 154G, 258W,
276U, 333D, 387M. TRA 38K.

Description. Several colonies growing on the stems of other hydroids. Stem
unbranched, or giving rise to 1 or 2 secondary pedicels in a sympodial manner, ©
closely annulated at base, in distal region and at irregular intervals between.

Hydrotheca very thin and fragile, with 7-10 double marginal teeth.
Margin bowed out between teeth giving an undulating outline in end-on view
and sometimes the effect of longitudinal striations in the upper part of the
hydrotheca. Depth of hydrotheca 1? to 3} times diameter at margin.

Gonotheca as described by Stechow: smooth and fragile, with annulated
pedicel and truncated distal end, containing 1-3 medusa buds.

482 ANNALS OF THE SOUTH AFRICAN MUSEUM

A
E
e
5 C
B D E

re

Fic. 15. Clytia paulensis (Vanh6ffen).

A-E and J. Hydrothecae.
F. Gonotheca.

Gand H. MHydrothecae drawn from Stechow’s slide of Clytia ulvae from Marseilles, 1910.

Measurements (mm.)

SAMH

336
Pedicel length } - O°57-1'°14
maximum diameter 0:04-0:06
Hydrotheca, length . 0°35-0°58

diameter at margin. 0°16-0°33

length/diameter . 1°76-2°47
Gonotheca, length : —_

maximum diameter —

TRA
38K
0°48-1-82
0:04-0:06
0°41-0°72
0°19-0°31
1:90-2:68
0°715
0°385

SCD
2538W
0°52—-0°94
0:04-0:05
0°45-0°69
0°15—0°28
2°17—-3°07
0°66-—1-00
0°30—0:36

SCD SCD Clytia ulvae

333D 154G (Marseilles)
O*QI-1°13. 0°85-1'00 0-22—0-90
0°045-0°05 07035 0:03-0°035
0°53-0°68 0°42—-0°50 0°35-0°44
O°2I-0°25 O°15-O0°16 0O-12—0°14
2°20-3°10 2°80—-3°13 2°71-3°23

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 483

Remarks. Measurements taken from different localities show that the size
and proportions of the hydrotheca are very variable. Vanhoffen’s material
_from the Antarctic is well within range, while some of Stechow’s material
from South Africa (1925)) and Australia (1925a) is a little smaller, though
the proportions are similar.

It is highly probable that Clytia ulvae Stechow, 1919, is a synonym for
C.. paulensis. | have been able to examine a prepared slide of Stechow’s material
of C. ulvae from Marseilles kindly loaned by the Munich Museum and find
that the marginal teeth are in fact double (fig. 15G, H), though the bays
between members of a pair are almost as large as those between pairs. However,
the depth of the former bays is variable within a single colony of C. paulensis
so that the small teeth may be 4 to 2 the size of the large double teeth. I have
included measurements of Stechow’s material (taken by myself) for comparison.
These show that the hydrothecae are slightly narrower and the proportion
of length/diameter in consequence slightly greater, though the material might
well fit near the end of a series of changing proportions arranged as above.

Obelia dichotoma (Linn., 1758)

Obelia dichotoma: Millard, 1952: 420, 426, 433, fig. 3. Millard, 1957: 198. Millard, 1958: 174.
Millard, 1959): 250.

Obelia dubia: Vanhoffen, 1910: 307, fig. 27. Nutting, 1915: 77, pl. 19 (fig. 1). Stechow, 1925):
435. Fraser, 1937: 86, pl. 17 (fig. 87).

Campanularia obtusidens Jaderholm, 1904: 2, pl. 1 (fig. 1).

Records. West coast: CP 378. LAM 46P. LB 378F. OLF 21C. SAMH 404,
405. I'B 8. South coast: KNY 165C (recorded by Day, Millard and Harrison,
1952). LIZ 2G, 11J. MB 37D, 81V. SAMH 147, 160, 223, 338, 342. SCD 112G,
258V, 281D, 312C. TRA 38L, 42W.

Remarks. Obelia dubia is considered to be a synonym for O. dichotoma. The
South African material shows all grades of hydrothecal types from the typical
O. dichotoma form where the marginal teeth are usually not distinct to that of
O. dubia as illustrated by Jaderholm (1904) and Vanhéffen (1910), where
there are distinct marginal teeth and indications of longitudinal striations. It
is impossible to draw a dividing line between them. The hydrothecal pedicel
is extremely variable in length and the diaphragm may be straight or oblique
within the same colony.

The gonothecae of O. dubia were illustrated by Fraser (1937) and are said
to be ‘almost smooth, or provided with broad, shallow undulations’. In the
South African material the gonothecae are usually smooth, but examples
are also present (with the typical dichotoma hydrothecae) where the gonothecae
are corrugated, approaching very closely the type illustrated by Fraser.

Obelia geniculata (Linn., 1758)
Obelia geniculata: Millard, 1957: 198. Millard, 1959): 250.
Records. West coast: A 383 (reported by Bright, 1938). CP 325. LAM 24H.
LB 127, 314G, 371B. PP 1V. SAMH 357. SB 168G, 235J. TB 6A. TRA 86P.
WCD 81G. South coast: TRA 42V.

484 ANNALS OF THE SOUTH AFRICAN MUSEUM

SUMMARY

A total of 43 species of hydroids is recorded, including 25 Gymnoblasts
and 18 Calyptoblasts. Of these 5 are new species, namely Monocoryne minor,
Myriothela tentaculata, Bicorona elegans, Aequorea africana and Campanularia
laminacarpa, and 8 are new records for the Republic of South Africa.

The new genus Bicorona is established for a Corynid species with 2 whorls
of tentacles and fixed sporosacs.

The status of the Campanularian genus Clytia is discussed and a type species
selected.

ACKNOWLEDGEMENTS

The Trustees of the South African Museum acknowledge gratefully the
receipt of grants from the University of Cape Town and from the Council for
Scientific and Industrial Research towards the cost of publication. The author
wishes to acknowledge the receipt of a grant from the Fourcade Bequest of the
University of Cape Town for the purchase of a microscope.

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486 ANNALS OF THE SOUTH AFRICAN MUSEUM

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HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 487

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WEILL, R. 1934. Contribution a l’étude des Cnidaires et de leurs nématocystes. I. Recherches
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1-347; 11: 349-701.

Ann. S. Afr. Mus., Vol. XLVIII Plate 1

Solanderia procumbens (Carter).
A. Whole colony of form A on left (WCD 158F) and form B on right (WCD 158E).

B. Details of the skeleton in form A (WCD 158F). An old stem shown on left with reticulate

meshwork, and a younger stem on right with more marked longitudinal trabeculae and
many hydrophores.

(Photos: Dr. G. 7. Broekhuysen)

a
2
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-
-
-
ea

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hex

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TaBLE or CoNnTENTs and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 4? in. x 7 in. (7$ in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers)
first and last, preceded by a colon (without prefix ‘p’). Thus:

Smrru, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmirH, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).
Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:

Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.

When another species has been called by the same name:

[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

ny

Ne A. HH: MILLARD

HYDROIDS OF
THE VEMA SEAMOUNT

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UNIVERSITY.
HYDROIDS OF THE VEMA SEAMOUNT

By
N. A. H. MiLiarp,
Koology Department, University of Cape Town

(With 1 figure in the text)

CONTENTS

PAGE
Peivoduetion §; « te. «© ‘s) 52) 480
Systematic account Os. ee ea
Discussion ‘ : } : , 404
Summary he pe Pe eae es
Acknowledgements : : : - 495
References : ; f : ; ~ 495

INTRODUCTION

The material recorded in this paper is part of a collection of benthic
fauna made by the Department of Oceanography of the University of Cape
Town and the Division of Sea Fisheries, Cape Town, in November, 1964. The
Vema Seamount is situated in 31° 38’ South Latitude and 08° 20’ East Longi-
tude, approximately 550 miles off the west coast of South Africa, and is
surmounted by a plateau of about five square miles with a mean depth of 73 m.
The hydroid material was obtained from a rocky bottom in depths varying
from 42 to 61 m. and was part of a mixed community containing also sea-weeds,
sponges, ascidians and holothurians. Further details of the expedition are
reported by Simpson and Heydorn (1965). It is of interest that from a cursory
examination of the fauna these authors remark on an apparent affinity with
Tristan da Cunha. This was particularly evident in the fish, and the common
rock-lobster (Jasus tristant), which is now being exploited commercially, is
similar to that occurring in the Tristan area.

The hydroid collection is the property of the South African Museum and
will be lodged there.

Family Haleciidae
Halecium beani (Johnston, 1838)

Three infertile samples, two from 54 m. and one from 61 m. Stems reaching
a maximum height of 8-6 cm.

489

Ann. S. Afr. Mus. 48 (19), 1966: 489-496, 1 fig.

490 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hydrodendron caciniformis (Ritchie, 1907)
Fig. 1
Ophiodes caciniformis Ritchie, 1907: 500, pl. 23 (fig. 11, 12), pl. 24 (fig. 1).
Hydrodendron caciniformis: Millard, 1957: 186, fig. 3. Ralph, 1958: 342, fig. 13b, c, 14a.
Phylactotheca caciniformis: Pennycuik, 1959: 174.
Ophiodissa caciniformis : Vervoort, 1959: 218, fig. 1, 2.

One sample from 54 m. Stems reaching a maximum height of 0-9 cm.,
most of them unfascicled, though a few weakly fascicled at base. Structure of
stem and hydrophore similar to the South African material, though dimensions
on the whole slightly smaller.

Gonophores borne in numbers on hydrorhiza. Elongated barrel-shaped,
with short pedicel and truncated distal end, lightly annulated. Reaching
1°35 mm. in length and 0-58 mm. in maximum diameter. Male and female
similar in appearance and distinguishable only under the microscope. Male

Ear eee eS SE ee Imm.

Fic. 1. Hydrodendron caciniformis

A. Portion of stem. B. Male gonophore. Cand D. Female gonophores.

HYDROIDS OF THE VEMA SEAMOUNT 491

generally slightly smaller than female. Blastostyle consisting of an elongated
spadix bearing a single layer of rather flattened eggs in the female and a mass
of spermatogenic cells in the male, the whole enveloped by a diffuse layer of
tissue rich in large stenotele nematocysts. No hydranths present.

Although gonothecae were described by Bale (1919), for Ophiodes australis,
which Ralph suspects to be a synonym, they were not illustrated, and this is
the first certain record of gonophores for H.. caciniformis.

I cannot agree with Pennycuik that Ophiodissa expansa Fraser, 1948, from
the Pacific coast of North America is a synonym. Not only does Fraser illustrate
the nematotheca of this species without the everted rim which is so typical of
H. caciniformis, but the gonophore is different. In O. expansa it is borne on the
stem, is much more strongly annulated (‘very strongly crested rugosities’ :
Fraser, 1948: 227) and has a narrower aperture.

Family Campanulariidae
Obelia geniculata (Linnaeus, 1758)
One young infertile colony growing on weed from 54 m. Maximum height
of stem 0-5 cm.
Family Lafoeidae

Lafoea fruticosa M. Sars, 1851

One small infertile colony from 54 m. Maximum height of stem 1:0 cm.

Family Sertulariidae
Amphisbetia bidens (Bale, 1884)

One infertile colony from 54 m. Maximum height of stem 3-4 cm.

Amphisbetia minima (d’Arcy Thompson, 1879)
One colony, with gonophores, from 54 m. Maximum height of stem 0-6 cm.

Dynamena quadridentata nodosa Hargitt, 1908
Two infertile samples growing on weed, both from 61 m. Maximum height
of stem 0-6 cm.

Parascyphus simplex (Lamouroux, 1816)
Thyroscyphus tridentatus: Ritchie, 1909: 74, fig. 1.
ar alae simplex: Splettstdsser, 1929: 126. Totton, 1930: 179, fig. 29. Ralph, 1961: 755,

g. Ib.

Two small infertile colonies from 42-50 and 54 m. Stems reaching a
maximum height of 1-0 and 0-8 cm. respectively. Stem usually unbranched,
but in some cases giving off one, or even two, lateral branches. Structure of
stem and hydrothecae as in previous descriptions.

This species has not been recorded from South Africa, but has been
reported from Gough Island by Ritchie. Apart from this there are several
records from Australasia and one from Great Britain,

492 ANNALS OF THE SOUTH AFRICAN MUSEUM

Salacia articulata (Pallas, 1766)
One colony, with gonophores, from 54 m. Maximum height of stem
2°3 cm.
Sertularella arbuscula (Lamouroux, 1816)

Two very typical, though infertile, colonies from 54 and 61 m. Maximum
height of stem 6-8 cm.

Sertularella flabellum (Allman, 1886)
Two infertile colonies from 61 and 54 m. Maximum height of stem 2-0 cm.

Sertularella mediterranea Hartlaub, 1901

One infertile colony from 42-50 m., consisting of stems reaching a maxi-
mum height of 2:3 cm., most of them with one lateral branch. Also a fragment
from 54 m.

Sertularella megista Stechow, 1923
One infertile colony from 54 m. Maximum height of stem 3-9 cm.

Sertularia distans gracilis Hassall, 1848

Four infertile colonies growing on weed, two from 54 m. and two from
61 m. Maximum height of stem 0-4 cm.

Also a colony with rather larger dimensions and of more doubtful identi-
fication from 61 m. The stems in this sample reach a maximum height of
1:0 cm. The hydrothecae are larger than the normal material found on the
South African coast (Millard, 1957, 1958, 1964), the internodes longer, and the
typical basal hinge-joints are absent. One of the stems has a branch arising
from the posterior surface.

Sertularia marginata (Kirchenpauer, 1864)

One infertile colony growing on weed from 54 m. Both simple and branched
forms present. Maximum height of branching stem 1-7 cm.

Family Plumulariidae (subfamily Halopterinae)

Antennella quadriaurita Ritchie, 1909

Antenella quadriaurita Ritchie, 1909: 92, fig. 9.
Antennella quadriaurita: Stechow, 1919: 113.

Three infertile samples from 42-50, 54 and 61 m. and one fertile sample
from 42-50 m.

Stems reaching a maximum height of 2-2 cm., normally solitary, but some-
times clustered together at base, and sometimes giving off up to four subsidiary
branches. These branches arise quite irregularly, usually from the posterior
surface of the basal athecate region, and successive branches never form the
main axis of the stem as in Monostaechas faurei Millard, 1958. Stem with alternate

HYDROIDS OF THE VEMA SEAMOUNT 493

thecate and athecate internodes, of which the athecate ones are longer in the
basal region and the thecate in the distal region. Athecate internodes generally
bearing two nematothecae each, but sometimes one, and only rarely three.
Hydrotheca with depth and marginal diameter approximately equal.
Other details as described by Ritchie.

Gonothecae (not previously described) borne on thecate internodes just
below hydrothecae, pear-shaped, with truncated distal end and wide spherical
aperture, with two nematothecae on basal region (probably female). Pedicel
of two segments.

A. quadriaurita is known only from Gough Island and Havana. There is
little to distinguish this species from A. africana Broch, 1914, other than the
number of nematothecae borne on the athecate internodes. For the type
material of A. quadriaurita from Gough Island Ritchie gives two to four, but
generally three, and for the Havana material Stechow gives two. In A. africana
there is normally only one, but occasionally two, and very rarely three. As
shown in the following table, material from the Vema Seamount is inter-
mediate between that from Gough Island and A. africana from South Africa.
A. quadriaurita is retained as a separate species solely on the fact that most of the
athecate internodes have more than one nematotheca, though it is likely that
more material from the Atlantic will show a completely intergrading series,
in which case it will be necessary to sink A. africana in the synonymy of A.
quadriaurita. The gonophores of the two species are identical.

Number of Number of
nematothecae per I 2 3 4 internodes
athecate internode examined

Gough Island od -— 40% 55% 5% 20
(from Ritchie) |

Vema Seamount... 23% 7a 2%, — go

South Africa .. NaeoisG4 70 575% B22 % aa 506

Halopteris constricta Totton, 1930

Two samples, both from 54 m., one bearing gonophores. Maximum height
of stem 0-8 cm.

Family Plumulariidae (subfamily Plumulariinae)

Plumularia pulchella Bale, 1882

Three samples, all from 54 m. The most luxurious colony has stems
reaching a maximum height of 1-0 cm. and bears gonophores. Some of its stems
bear one to three irregular lateral branches.

Plumularia setacea (Linnaeus, 1758)
Six samples, none of them luxurious, two from 42—50 m., three from 54 m.

494 ANNALS OF THE SOUTH AFRICAN MUSEUM

and one from 61 m. Two samples bear gonophores, and the maximum height
of the stem is 1:7 cm.

Plumularia spinulosa Bale, 1882

Three samples, one from 42-50 m. and two from 54 m. Maximum height
of stem 0-5 cm. Young gonophores present in one sample.

Family Plumulariidae (subfamily Kirchenpauerinae)

Pycnotheca mirabilis (Allman, 1883)
One infertile sample from 42-50 m. Maximum height of stem 3-1 cm.

Family Plumulariidae (subfamily Aglaopheniinae)

Aglaophenia pluma pluma (Linnaeus, 1758)

Six samples, one from 42-50 m., three from 54 m. and two from 61 m.
Four of these colonies fertile and both male and female corbulae present.
Stems reaching a maximum height of 2-4 cm. This material is of more delicate
build than that found in South Africa, and in many cases the internodal septa
and intrathecal septum are poorly developed or absent. The median nemato-
theca is always short as shown by Broch (1933, fig. 18a).

Lytocarpus filamentosus (Lamarck, 1816)

Five samples, all infertile, one from 42-50 m., three from 54 m. and one
from 61 m. Maximum height of stem g:0 cm.

Discussion

Of the 23 hydroid species here recorded, 21 also occur in South Africa
and it appears that in the case of the hydroid fauna at any rate the two areas
have close affinities with one another. Of the remaining two (Parascyphus
simplex and Antennella quadriaurita) both have been reported from Gough Island
in the South Atlantic by Ritchie, 1909. The hydroid fauna of the Tristan group
of islands is, however, very poorly known, and it is probable that further
investigation would show more species in common with the Vema Seamount.

Further analysis shows that of the total 23 species 10 are cosmopolitan,
namely:

Halecium beaniu

Obelia geniculata

Lafoea fruticosa

Amphisbetia minima (south of the Mediterranean)
Dynamena quadridentata

Sertularella mediterranea

Sertularia distans gracilis

Sertularia marginata

HYDROIDS OF THE VEMA SEAMOUNT 495

Plumularia setacea

Aglaophenia pluma pluma

With these should probably be included Parascyphus simplex and Hydro-
dendron caciniformis, which have a peculiar scattered distribution, the former
being known from Australasia, Gough Island and the west coast of Scotland,
and the latter from the Cape Verde Islands, Mediterranean, West Indies,
tropical West Africa, Australasia and South Africa.

Seven species have an Indo-Pacific distribution, namely:

Amphisbetia bidens: Australia, Madagascar, South Africa.

Sertularella arbuscula: Indian Ocean, Australia, South Africa.

Halopteris constricta: New Zealand, South Africa.

Plumularia pulchella: Australasia, South Africa.

Plumularia spinulosa: Australasia, Japan, South Africa.

Pycnotheca mirabilis: Australasia, Japan, west coast of North America, South
Africa.

Lytocarpus filamentosus: Australia, Madagascar, South Africa.

Three species are so far known only from South Africa, namely:

Salacia articulata

Sertularella flabellum

Sertularella megista

One species has an Atlantic distribution, namely:

Antennella quadriaurita: Gough Island, Havana.

SUMMARY

A total of 23 species of hydroids is recorded from the Vema Seamount.
Of these the gonophores of Hydrodendron caciniformis and Antennella quadriaurita
are described for the first time.

The distribution of the species in the rest of the world is discussed. It is
concluded that the affinities of the hydroid fauna are mainly with South Africa.

ACKNOWLEDGEMENTS

The Trustees of the South African Museum gratefully acknowledge the
receipt of a grant from the Council for Scientific and Industrial Research
towards the costs of publication.

REFERENCES

Bate, W. 1919. Further notes on Australian hydroids, IV. Proc. roy. Soc. Vict. 31: 327-361.

Brocu, H. 1933. Zur Kenntnis der adriatischen Hydroidenfauna von Split. Skr. norske
Vidensk-Akad., Mat.-natur. Kl. 1933 (4): 1-115.

Fraser, C. McL. 1948. Hydroids of the Allan Hancock Pacific expeditions since March,
1938. Allan Hancock Pacif. Exped. (1) 4: 179-335.

Mitiarp, N. A. H. 1957. The Hydrozoa of False Bay, South Africa. Ann. S. Afr. Mus., 43:
173-243.

496 ANNALS OF THE SOUTH AFRICAN MUSEUM

Mitiarp, N. A. H. 1958. Hydrozoa from the coasts of Natal and Portuguese East Africa.
Part I. Calyptoblastea. Ann. S. Afr. Mus. 44: 165-226.

Miuuarp, N. A. H. 1964. The Hydrozoa of the south and west coasts of South Africa. Part II.
The Lafoeidae, Syntheciidae and Sertulariidae. Ann. S. Afr. Mus. 48: 1-56.

Pennycurk, P. R. 1959. Faunistic records from Queensland. Part V. Marine and brackish
water hydroids. Pap. Dep. Zool. Univ. Qd. 1: 141-210.

Raupu, P. M. 1958. New Zealand thecate hydroids. Part II. Families Lafoeidae, Lineo-
lariidae, Haleciidae and Syntheciidae. Trans. roy. Soc. N.Z. 85: 301-356.

RaupH, P. M. 1961. New Zealand thecate hydroids. Part III. Family Sertulariidae. Trans.
roy. Soc. N.Z. 88: 749-838.

RitcutE, J. 1907. On collections of the Cape Verde Islands marine fauna, made by Cyril
Crossland, M.A.(Cantab.), B.Sc.(Lond.), F.Z.S., of St. Andrews University, July to
September, 1904. Proc. zool. Soc. Lond. 1907: 488-514.

RITCHIE, J. 1909. Supplementary report on the hydroids of the Scottish National Antarctic
Expedition. Trans. roy. Soc. Edinb. 477: 65-101.

Simpson, E. S. W. & Heyporn, A. E. F. 1965. Vema Seamount. Nature 207: 249-251.

SPLETTSTOsSER, W. 1929. Beitrage zur Kenntnis der Sertulariiden. Thyroscyphus Allm., Cnidos-
cyphus nov. gen., Parascyphus Ritchie. Zool. Fb. 58: 1-134.

StecHow, E. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und
anderer Gebiete. <ool 7b. System. 42: 1-172.

Totton, A. K. 1930. Coelenterata. Part V. Hydroida. Nat. Hist. Rep. Terra Nova Exped. 5:
131-252.

VERVoORT, W. 1959. The Hydroida of the tropical west coast of Africa. Atlantide Rep., no. 5:
211-325.

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TABLE OF CONTENTs and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. = 7 in. (73 in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

SmiTH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmiTH, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 ( = natalensis West).

PotA., HULEEY

THE VALIDITY OF RAJA RHIZACANTHUS REGAN
mee RAvASPULLOPUNCTATA SMITH, BASED ON
A STUDY OF THE CLASPER

November 1966 November
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UNIVERSITY.

THE VALIDITY OF RAJA RHIZACANTHUS REGAN
AND RAJA PULLOPUNCTATA SMITH, BASED ON A
STUDY OF THE CLASPER

By
Pe ely ELEY
South African Museum, Cape Town
(With 8 figures in the text)

CONTENTS

PAGE
Introduction Tee ees Reece Pe! tr:
The Claspers of Myxopterygia. . . 498
Clasper of R. rhizacanthus
(i) the external anatomy . : - 499
(i) ime intermal anafomy..,,.. +. . 560
Clasper of R. pullopunctata
(i) the external anatomy . ‘ . 505
Gi the internal anatomy 00.) 505
Discussion . ; ‘ ‘ ‘ ‘ . 508
memiowredrements, . 2) 2) «= / 3» BL3
RAE kk ee we lw at
PpUERRiCes ait his ON io ELS URE ASE Os Al
INTRODUCTION

The present classification of South African skates is more or less tentative,
for this difficult genus shows great variability in its morphological characters,
which vary with age and/or sex in the individuals within the species. This
high intraspecific variability probably accounts for the present inclusion of
95-100 species in the genus Raja. Several South African species have, as yet,
proved to be indistinguishable from European species from the corresponding
latitudinal belt in the northern hemisphere (Bigelow & Schroeder, 1953), and
it would appear that these pairs represent cases of ‘masked bipolarity’ (Ekman,
1953). But in very few cases have actual comparisons of the specimens from
the two regions been made, although Norman (1935) has given some brief
references.

Raja alba, Lacépéde, from the N.E. Atlantic is reported here under the
same name, but it appears to be cosmopolitan in distribution, being recorded
from Morocco (Pietschmann, 1906), West Africa (Fowler, 1936) and Angola
(da Franca, 1959).

ANS)

Ann. S. Afr. Mus. 48 (20), 1966: 497-514, 8 figs.

498 ANNALS OF THE SOUTH AFRICAN MUSEUM

Norman expressed some doubt about the identification of R. batis from
South Africa but the material he examined included a single stuffed specimen,
on which accurate measurements were impossible. This South African species
is now thought to be distinct from R. batis Linnaeus, and Smith (1964), on the
basis of the shape, the presence of a single median spine and the general
markings, has reclassified this skate as R. pullopunctata, although he points out
that only juvenile material was examined.

Raja rhizacanthus Regan, has been thought to be identical with the Euro-
pean thorn-back skate R. clavata Linné, but this has been a matter of speculation.
Barnard (1925) and Fowler (1940) have included R. rhizacanthus as a synonym
of R. clavata, but Norman regarded the two as distinct species on the basis of
spination and the position of the vent. This has been followed by Smith (1961).

It is the purpose of this paper to compare and contrast certain morpho-
logical characters, in the hope that the validity of these latter two South
African species may be determined with greater certainty.

Since skates show such great variations in external features, a more
critical method of examination needs to be employed. Leigh-Sharpe (1920-
1926) has pointed out that in the genus Raja, individual species can be recog-
nized by the morphological structures of their claspers. He has extended this
further and has postulated the existence of pseudogenera (termed Alpha-,
Beta- and Gamma-raia, etc.), recognizable by the similar presence or absence of
the clasper elements. Ishiyama (1958) has shown that in Japanese rajids,
although intraspecific variations in the clasper can be recognized, the ‘external
and internal structures of the clasper are species specific without exception’
(p. 224). On this basis he has also been able to postulate phyletic, generic and
inter-specific relationships, and has concluded that ‘the characters in the male
external organ (claspers) of the Japanese rajids seem to give the most reliable
basis for the systematics of this group of fish’. (p. 243.)

Hence it would appear that since the anatomy of the clasper is valid for
the purposes of identification, it should also be valid for the purposes of com-
parison of geographically separated species. However, ‘genitalic differences
must be evaluated just like other characters’ (Mayr et al., 1953, p. 109), and so
differences in the external and measurable characters of the specimens must
also be taken into account.

THE CLASPERS OR MyYXOPTERYGIA

Comprehensive anatomical studies have been carried out on the claspers
of European elasmobranchs by Petri (1878), Jungersen (1899) and Huber
(1901). Two points of view were held as to the function of the claspers: that
they were ‘holders’ (analogous to hands) was originally postulated, but they
were later recognized as organs of intromission. Extensive study on the external |
anatomy and the function of the clasper in the various elasmobranch groups
has been made by Leigh-Sharpe, and further work on the function of the organ
has been performed by Friedman (1935).

RAJA RHIZACANTHUS REGAN AND RAJA PULLOPUNCTATA SMITH 499

The nomenclature used in this paper is mainly in accordance with that of

Leigh-Sharpe and Ishiyama, together with that of the above-mentioned
authors. I Ag

THE CLASPER OF R. RHIZACANTHUS REGAN

The claspers from seven adult specimens of R. rhizacanthus, trawled off
Cape Columbine and in False Bay, were examined.

apopyle

hypopyle

slit
Signal

pseudorhipidion shield

Sw 0-01

pseudosiphon

sentinel
* sentina
| 2:0cms |
spike

Fic. 1. Raja rhizacanthus

A, external view of right clasper from dorsal side; B, lateral view of right clasper, opened to
show structural features of glans.

(i) The external anatomy of the clasper (fig. 1 A, B)

The clasper of R. rhizacanthus is a moderately long, strongly built structure,
the outer surface of which is entirely naked. Proximally the clasper is cylin-
drical, but from about 1/3 its length from the distal end, it becomes broader
and dorso-ventrally flattened, especially on its ventral surface. It tapers
gradually towards its distal end and terminates in a broadly rounded, spatulate

500 ANNALS OF THE SOUTH AFRICAN MUSEUM

tip. The ventral surface is pale and the dorsal surface tones with the general
coloration of the body.

The apopyle (fig. 1 A) is situated dorsally, some little distance behind the
vent, and the appendix groove, arising from this point, runs across to the outer
margin of the organ, and continues along the distal two-thirds of the lateral
edge almost to the tip of the clasper. In some cases, due to the skin stretching
across the dorsal terminal 1 cartilage, the groove may be seen from the dorsal
side. Proximally the groove is bordered on by the two ‘scroll-like’ dorsal and
ventral marginal cartilages, so that the edges of the groove closely approximate
to form a tube, running from the apopyle to the hypopyle. Distally, from about
half the clasper length, fleshy lips border the groove, and during erection,
brought about by the powerful adductor muscles, and by vascular dilation
(Friedman), these lips are opened and the structures of the clasper glans
protrude (fig. 1 B).

Two pouches are present on the dorsal surface of the concavity of the
clasper glans (fig. 1 B), the smaller, more proximal slit and the larger, distal
pseudosiphon. The median hypopyle is bordered dorsally by the pseudo-
rhipidion, a plate-like structure employed as a splash-plate for ejaculating
spermatozoa and distinguished from the rhipidion of Leigh-Sharpe in not
being fan-shaped or pitted. Posterior to the hypopyle, and at about the same
level as the pseudorhipidion, is a fleshy pad, the szgnal, which is capable of
rotation about the longitudinal axis of the clasper. The sentinel is strongly
developed and in older specimens its knife-like edge is exposed. There is a deep
sentina situated distally, its inner border raised into a curved spike. The ventral
shield is narrow and not well developed. The lining of the concavity is naked,
scale and spine being absent.

All these above-mentioned structures are associated with the cartilages
of the terminal group.

(ii) The internal anatomy of the clasper (figs. 2, 3)

The skeleton of the clasper is a continuation, in the median axis, of the
basipterygium of the fin, and is composed of two groups of cartilages, the basal
group (including the basipterygium) and the terminal group.

(a) The basal group (fig. 2)

Ishiyama has pointed out, that in Japanese rajids, the basal group is not
species specific, and, although White (1937) has proved that the number of
proximal segments and other cartilages in the basal group tend to vary in large
groups of the elasmobranchs, the basal group may only be used in phyletic
considerations among the batoid fishes. Examination of the basal group of
R. rhizacanthus seems to support this view, since the same seven cartilages are
present as in the Japanese rajids (see Ishiyama, fig. 7).

The basal group in R. rhizacanthus consists of the basipterygium, and
b, and b, cartilages, the Beta-cartilage, and the axial cartilage, which extends

RAJA RHIZACANTHUS REGAN AND RAJA PULLOPUNCTATA SMITH 501

basipterygium

j ES Fy \ Wk fi
of Hy ‘ SS in rays
b, cartilage

beta cartilage

dorsal marginal
axial

4:0cms

Fic. 2. Raja rhizacanthus. Cartilages of the basal group of the
clasper.

to the terminal part of the clasper, and is covered by the dorsal and ventral
marginal cartilages (fig. 2). The basipterygium is a short, stout structure, to
which are attached in a groove on its lateral margin, the last 6 fin rays of the
pelvic fin. Distally, the basipterygium is elongated on its dorsal surface to
_ form a protuberance which overlaps the b, cartilage. The proximal end of the
b, cartilage is visible from the dorsal side, but the Beta-cartilage, situated
dorsally to the b, and b, cartilages, completely obscures the b, cartilage and
the distal end of b,. The Beta-cartilage is more or less flat, serving as a region
of attachment for the m. flexor internus, but is slightly thickened and upraised
at its proximal end, forming a shelf.

The distal ends of the b, and Beta-cartilage are superimposed so that both
of these elements play a part in the joint with the axial and marginal cartilages.

‘The long axial cartilage is cyclindrical at its proximal end, but it becomes
dorso-ventrally flattened distally and assumes a spatulate appearance. It is
tightly enclosed by the calcified dorsal and ventral marginal cartilages and is
only free at its spatulate tip, although it is visible dorsally. The dorsal marginal
cartilage attaches tightly to the axial, starting where the axial is connected to
the b,/Beta joint, and runs along the outer lateral margin of the axial for about
half its length. The dorsal marginal cartilage is somewhat broadly pointed
anteriorly but it becomes expanded distally, its outer lateral edge forming the
dorsal edge of the ‘scrolled’ groove of the clasper. The ventral marginal carti-
lage, unlike the dorsal marginal, starts posteriorly to the axial and is a flat,
tongue-like structure. Its outer lateral margin forms the ventral lip of the

502 ANNALS OF THE SOUTH AFRICAN MUSEUM

‘scroll’ and medially it is attached to the axial, overlying it distally. The ventral
marginal cartilage extends farther down the length of the clasper than the
dorsal marginal cartilage. Attached distally to the axial and to the marginal
cartilages, are the elements which comprise the terminal group of cartilages.

(b) The terminal group

The elements which comprise the terminal group of cartilages are asso-
ciated with the structures of the glans, and may be divided into three types:
ventral terminal, dorsal terminal and accessory cartilages.

1. Ventral terminal (figs. 3 A, B, G)

The ventral terminal cartilage is attached to a groove in the distal end of
the ventral marginal cartilage. It has a J-like shape, being by far the largest
cartilage of the terminal group. It is strongly curved medially, and runs around
the axial cartilage, so that it can be seen from the dorsal side. The upright
portion of the ventral terminal cartilage is fairly well developed and there is a
small shelf on its dorsal side, which is seen as the shield of the clasper glans.
Distally, the ventral terminal is thick and somewhat pointed.

2. Dorsal terminal 1 (figs. 3 C, G)

The dorsal terminal 1 cartilage (‘cover-piece’ of Jungersen) is situated on
the dorsal side of the clasper, partially covering the dorsal terminal 2 cartilage,
to which it is attached along the length of its longest margin. It is flatly rounded
and curved proximally, so that it is elongated on its outer lateral margin. From
this point, there is a shelf, running in a curve along the anterior margin, which
serves as a point of insertion for the aponeurosis of the m. dilatator, which conse-
quently covers the whole area of the cartilage. Distally, the cartilage is flat
and its distal end is attached, together with the dorsal terminal 2 cartilage, to
the axial cartilage. The dorsal terminal 1 cartilage is associated with the
development of the pseudosiphon.

3. Dorsal terminal 2 (figs. 3 D, G)

This curved element is inserted between the dorsal terminal 1 and the
dorsal marginal/axial cartilages. It is firmly attached to the distal region of the
dorsal marginal cartilage, and by means of a ligament (which forms the shelf
between the slit and pseudosiphon) it is attached at its distal end to the outer
lateral margin of the axial cartilage, at about 2/3 the length of that cartilage
from its anterior end. The dorsal terminal 2 cartilage is strongly curved, and
raised along the side of longest curvature, to form a flat surface ‘articulating’
with the dorsal terminal 1 cartilage. The dorsal terminal 2 cartilage is asso-
ciated with the slit. \

4. Accessory cartilages 1 and 2 (figs. 3 E, G)

These two cartilages are strongly united and are attached to the distal
end of the ventral marginal cartilage. The accessory cartilage 1 is the S-shaped

RAJA RHIZACANTHUS REGAN AND RAJA PULLOPUNCTATA SMITH 503

axial

dorsal marginal

dorsal terminal 1

dorsal terminal 2

accessory 3

SUID 0-01

accessory 1

ventral terminal

accessory 2

| 2:0 cms |

E

Fic. 3. Raja rhizacanthus. Cartilages of the terminal group of the clasper.

A, ventral terminal (ventral view); B, ventral terminal (dorsal view); C, dorsal terminal 1

(dorsal view); D, dorsal terminal 2 (dorsal view); E, accessory cartilages 1 and 2 (dorsal

(view); F, accessory cartilages 3 and 4 (ventral view); G, terminal cartilages in situ from
dorsal side.

element, and its knife-edged protruberance is the sentinel. Accessory cartilage 2
is elongate in the longitudinal axis of the clasper, and its distal end is recurved
and hook-like, forming the spike of the clasper glans.

5. Accessory cartilages 3 and 4 (fig. 3 F, G)

Accessory cartilages 3 and 4 form the signal of the glans and are situated
on the ventral side of the clasper, between the upright portion of the ventral
terminal cartilage and the accessories 1 and 2, The smaller proximal cartilage,

504 ANNALS OF THE SOUTH AFRICAN MUSEUM

accessory 4 is attached in a groove formed between the accessory cartilages
1 and 2 and the ventral marginal cartilage; accessory 3 is movable on accessory 4
and its distal end is bluntly rounded.

Musculature

Only a brief mention of the musculature of R. rhizacanthus needs be made
for it is identical with that of R. clavata as given by Jungersen.

M. dilatator

Proximally this consists of a single muscle mass, but distally it seems to be
split into dorsal and ventral muscles. Its origin is on the axial, b, and b, carti-
lages, and it inserts as an aponeurosis on the shelf of the dorsal terminal 1
cartilage.

The m. compressor sacci, m. adductor, m. flexor (not extensor as given in Junger-
sen), and muscles of the fin are the same as for R. clavata.

hypopyle

SUD 0-f
-
=.
ae
2.
re}
a

pseudosiphon

shield

spike

Fic. 4. Raja pullopunctata

A, external view of right clasper from dorsal side; B, lateral view of right clasper, opened to
show structural features of the glans.

RAJA RHIZACANTHUS REGAN AND RAJA PULLOPUNCTATA SMITH 505

THE CLASPER OF R. PULLOPUNCTATA SMITH

The claspers from three adult specimens of R. pullopunctata, trawled West
of Cape Town and in Algoa Bay were examined.

(i) The external anatomy of the clasper (fig. 4. A, B)

The clasper of R. pullopunctata is long and stout; its cyclindrical body is
dorso-ventrally flattened distally, and slightly truncate, giving a spatulate
appearance. The glans is much broader than the rest of the clasper, due to the
strong development of the shield, which can be clearly seen from the dorsal
side. The claspers are entirely naked, scale and spine being absent. The apopyle
(fig. 4 A) opens some distance away from the vent, on the dorsal side.

The rhipidion (fig. 4 B) is well developed and fan-shaped and it is attached
along the inner ventral wall of the clasper, from the region of the hypopyle to
about half-way along the length of the glans. Its lower surface is pitted with
cavities of irregular outline, and this spongy tissue is said by Leigh-Sharpe to
be erectile and to act as a splash-plate for the ejaculating spermatozoa.

R. pullopunctata may be placed in the pseudogenus Gamma-raia, for there
is a reduction and crowding of the structural features of the glans. As in all
Gamma-raia, the signal is absent, while the shield is very well developed and
extends along the whole ventral surface of the glans, from the level of the
hypopyle almost to the distal extremity of the clasper. The distal region of the
shield is covered by pleated epithelia, but this is so thin at the outer lateral
edge, that the knife-like appearance of the shield is not unlike the sentinel of
R. rhizacanthus.

Both the slit and the pseudosiphon are present in the dorsal wall of the
concavity of the glans. In R. pullopunctata the pseudosiphon is larger and more
distally placed than the slit, and these two cavities are separated by a prominent
cartilaginous shelf. The pseudosiphon is well separated from the hypopyle by
the elongated dorsal marginal cartilage.

Medially placed is a large, single, tongue-like structure, which examination
of the terminal cartilages reveals as the spike (cf. Leigh-Sharpe). Although
covered by a thick layer of epithelial tissue, the calcified cartilage can be easily
felt. The small accessory 1 cartilage is not greatly developed, and the sentinel
is not a recognizable element in the external appearance of the clasper.

(ii) The internal anatomy of the clasper (fig. 5)

The basipterygium, b, and b, cartilages and the Beta-cartilage are the
same as in R. rhizacanthus. The axial cartilage is long, cyclindrical at its proximal
end, but narrower and dorso-ventrally flattened distally. The distal end of the
axial is more heavily calcified than the proximal region, and is slightly curved
outwards, terminating in a bluntly rounded point. The dorsal marginal carti-
lage starts more proximally than the ventral marginal cartilage, and runs for
about 2/3 the length of the axial, to which it is tightly bonded for its entire
length. The dorsal marginal cartilage is convex at its proximal end, but becomes

506 ANNALS OF THE SOUTH AFRICAN MUSEUM

axial

dorsal marginal

ventral marginal

dorsal terminal 2

ventral terminal

Fic. 5. Raja pullopunctata. Cartilages of the terminal group of the clasper.

A, ventral terminal (dorsal view) ; B, ventral terminal (ventral view) ; C, dorsal terminal 1

(dorsal view); D, dorsal terminal 2 (dorsal view); E, accessory 1 (ventral view); F,

accessory 2 (dorsal view); G, terminal cartilages in situ from dorsal side, but with dorsal
terminal 1 removed.

flatter distally and is extended in a small point, forming the shelf between the
pseudosiphon and the hypopyle. The ventral marginal cartilage is flat and
slightly expanded distally, so that it more or less overlaps the axial cartilage,
to which it is united. Its distal end is truncate.

1. Ventral terminal cartilage (figs. 5 A, B, G)

As in R. rhizacanthus, the ventral terminal is J-shaped, but its distal end is
not so strongly developed. The cartilage is attached to the inner surface of the
‘scroll-like’ ventral marginal cartilage. It is locked to this cartilage by means of

RAJA RHIZACANTHUS REGAN AND RAJA PULLOPUNCTATA SMITH 507

a protuberance, situated ventrally about 1/3 of the length of the ventral
terminal from its proximal end. The protuberance engages with a corre-
sponding point (the minute ‘sentinel’) of the accessory cartilage 1, hooking
the two elements together. The ventral terminal cartilage does not extend to
the tip of the clasper. The outer lateral edge of this cartilage is well developed
and calcified, forming a convex lamina, the shield of the clasper glans. Distally,
the apex of the ventral terminal is grooved, through which the distal regions
of the axial and dorsal terminal 2 cartilages pass, before becoming dorso-
ventrally flattened. The upright limb of the J-like ventral terminal is raised
into a ridge on its dorsal surface.

2. Dorsal terminal 1 (fig. 5 C)

This cartilage is situated on the dorsal surface of the clasper, and is
attached along its outer edge to the dorsal terminal 2 cartilage. The dorsal
terminal 1 cartilage is ‘shield-like’ in appearance and its proximal end is
elongated into an obtuse point, to which the dorsal bundle of the m. dilatator
attaches. The inner distal region of this cartilage is so curved that it wraps
around the axial stem and can be seen on the ventral side, where it slightly
overlaps the upright limb of the ventral terminal cartilage. Distally, the dorsal
terminal 1 is attached by connective tissue to the axial, dorsal terminal 2 and
accessory cartilages.

3. Dorsal terminal 2 (fig. 5 D)

The dorsal terminal 2 cartilage is attached to the inner distal end of the
dorsal marginal cartilage, so that the point and connective tissue, which forms the
shelf between the pseudosiphon and hypopyle, lie behind the dorsal terminal 1.
The dorsal terminal 2 cartilage is a long, thin, single cartilaginous element,
curving and then re-curving, so that its pointed distal end is attached to the
axial cartilage (fig. 5 G). Proximally, the dorsal terminal 2 is strongly calcified
and has a short blunt protuberance on its outer lateral margin, while distally
calcification is not so marked, the soft cartilage becoming dorso-ventrally
flattened. About half-way along its length, on the inner edge, there is.a liga-
ment (forming the shelf between the pseudosiphon and the slit), which arches
and inserts on the axial cartilage about 1/5 the length of this cartilage from

its distal end (fig. 5 G).

4. Accessory cartilage 1 (fig. 5 E)

This minute V-shaped cartilage is attached to the distal end of the ventral
marginal cartilage. The longer of its two arms runs forward along the outer
lateral edge of the ventral marginal, while the shorter traverses the distal end
of the ventral marginal cartilage. The base of the V is slightly pointed (the
minute ‘sentinel’) and is curved inwards, forming the point on which the
ventral terminal engages. It must be noted that this cartilage does not manifest
itself as an external structural element in the glans.

508 ANNALS OF THE SOUTH AFRICAN MUSEUM

5. Accessory cartilage 2 (fig. 5 F)

The accessory cartilage 2 resembles an inverted Y. Its cyclindrical proxi-
mal end is attached to the inner distal end of the ventral marginal cartilage.
The longer limb of the accessory is tightly bonded to the axial, so that distally
these two cartilages resemble a single element. The shorter arm of the cartilage
is simple in shape and projects laterally outwards. Its terminal end is slightly
upturned and forms the spike of the clasper glans.

Musculature

The musculature of R. pullopunctata was found to be identical with the
description of the musculature of R. clavata given by Jungersen, and like
R. batis, the m. dilatator is in two bundles.

(a) Dorsal m. dilatator

This muscle has its origin on the proximal region of the axial cartilage
and its insertion on the dorsal and lateral regions of the dorsal terminal 1
cartilage. The muscle is not strictly dorsal in position but more medio-dorsal,
and is much larger than the ventral m. dilatator.

(b) Ventral m. dilatator

The muscle is entirely separate from its dorsal counterpart and has its
origin on the ventral proximal region of the axial cartilage and the distal end
of the b, cartilage. It inserts as connective tissue across the surface of the ventral
terminal cartilage, although some fibres attach to the tip of the axial cartilage.

The m. compressor sacct, m. adductor, m. flexer and the muscles of the fin are
the same as for R. clavata.

DIscussION

Two pairs of claspers from specimens of R. batis Linnaeus, caught S.W.
of Ireland, and a pair of claspers from R. clavata Linnaeus, caught at Lowestoft,
were examined for purposes of comparison.

Although there is a marked difference in the size of the clawiee of R.
pullopunctata and R. batis, the general external appearance of the claspers is
similar. In both, the clasper is long, stout and naked, becoming dorso-ventrally
flattened and spatulate at its terminal end. The outer lateral edge of the shield
can be seen from the dorsal side in both species, but in R. bats it is more promi-
nent. This may be due to differences in the size of the claspers. However,
examination of the clasper glans reveals that the structural features in R.
pullopunctata and R. batis are different (figs. 4 B, 6).

The pseudosiphon and slit are present on the inner dorsal surface of the
glans in both species, but their formation is somewhat different, although they
are associated with the degree of development of the dorsal terminal cartilages
in both cases. In R. pullopunctata there is a marked shelf-like ridge separating

RAJA RHIZACANTHUS REGAN AND RAJA PULLOPUNCTATA SMITH 509

shield
slit

~hypopyle

rhipidion

sentinel

pseudosiphon spike

Fic. 6. Raja batis. Lateral view of right clasper, opened to show
structural features of the glans.

the pseudosiphon from the slit; this shelf is found to be wanting in R. batis.
The pseudosiphon is similarly situated in the two species, but in R. pullopunctata
the pseudosiphon resembles a concavity rather than the simple pouch found in
R. batis. The slit is more proximally situated in R. pullopunctata, occurring about
half-way along the length of the glans (about one-third the length in R. batis).
The hypopyle seems to be more distally placed in R. batis, but this may be due
to the heavier development of the shield. The rhipidion is pitted and fan-
shaped in both species, occurring along the inner ventral wall of the hypopyle,
and extending about half-way along the length of the glans in R. pullopunctata
and about two-thirds the length in R. batis. The rhipidion is more compact in
R. batis. The prominent shield extends along the whole inner ventral surface of
the glans in both species, but it is thinner and covered with pleated epithelia

510 ANNALS OF THE SOUTH AFRICAN MUSEUM

along its entire length in &. batis, while in R. pullopunctata the broader shield
is only pleated at its distal region.

The most noticeable difference in the structure of the claspers is in the
presence of the enlarged sentinel in R. batis, which extends distally and fits
together with the spike. In R. pullopunctata, the sentinel is reduced and lies under
the rhipidion, so that it is not seen when the glans is opened, and these two
elements are not expanded terminally as they are in R. batis.

As has been pointed out, the visible external characters of the clasper are

dorsal terminal 2

axial

dorsal terminal 3

wane
heise:

Fic. 7. Raja batis. Cartilages of the terminal group.

A, dorsal terminal 1 (dorsal view); B, ventral terminal (dorsal view) ; C, accessory 1
(dorsal view); D, accessory 2 (dorsal view); E, dorsal terminal 2 and 3 in position.

RAJA RHIZACANTHUS REGAN AND RAJA PULLOPUNCTATA SMITH By. ee

closely associated with the structure and number of the internal cartilages,
and therefore differences in the structure of the glans will be shown by differ-
ences in the terminal cartilages. In R. pullopunctata there are a total of five
cartilages in the terminal group (2 dorsal terminals, 1 ventral terminal and
2 accessory terminals), while in R. batis six terminal cartilages are present (3
dorsal terminals, 1 ventral terminal and 2 accessory terminals) (figs. 5, 7).

Although there is a great difference in the size of these elements, their
differences in shape and their position may be examined. The dorsal terminal 1
cartilage is similarly situated in both R. batis and R. pullopunctata, occurring on
the dorsal side of the clasper, and it is flat and plate-like, being slightly convex
distally. In both species, the median edge is bent in such a manner, that the
cartilage wraps round the axial stem and may be seen on the ventral side.
Distally, the cartilage is pointed in the two species, but in R. pullopunctata, the
proximal edge is elongated into a point for the attachment of the aponeurosis
of the m. dilatator, while in R. batis (fig. 7), the proximal edge is almost straight
and somewhat more convex. The dorsal terminal 2 cartilage is attached to the
distal end of the dorsal marginal cartilage in both species, and it is interesting
to note that in R. pullopunctata, the dorsal terminal 2 extends to the tip of the
axial cartilage, while in R. batis, the crescent-shaped dorsal terminal 2 (fig.
7 D, E) is attached to about half way along the axial and that the dorsal
terminal 3 extends to the tip of the clasper, i.e. these two cartilages in R. batis
have been reduced to a single cartilage in R. pullopunctata, although there is
linking between the dorsal terminal 2 and the axial in both cases. The terminal
end of the axial cartilage is similarly curved (laterally outwards) in the two
species, and the attachment of the dorsal terminal 2 of R. pullopunctata and the
dorsal terminal 3 of R. batis (fig. 7 E) is similar.

Although both R. pullopunctata and R. batis possess two accessory cartilages,
the accessory cartilage 1 (forming the sentinel) is reduced to a small point in
R. pullopunctata. In R. batis this cartilage (fig. 7 C) is well developed and extends
down towards the spike, and fits closely to it. The ventral terminal cartilages
are similarly situated in both R. batis and R. pullopunctata and these elements
are similar in shape. Both possess the ‘hooking’ protuberance on the inner
ventral margin, the enlarged convex outer lateral edge and the raised dorsal
ridge. However in R. batis (fig. 7 B) the distal end of the convex edge projects
slightly from the main body of the limb, and the distal end of the ventral
terminal is not so markedly grooved as in R. pullopunctata.

It may therefore be concluded that, in the structure of the glans and in
the number and arrangement of the internal cartilaginous elements, the
claspers of R. pullopunctata and R. batis are different.

Besides differences noted in the structure of the claspers of the two species,
some differences can also be seen in the external characters. Smith (1964)
points out that there is a difference in shape between the two, but examination
of later and adult stages of R. pullopunctata reveals that the shape of the disc
varies and that the pectorals and anterior margins of the disc become more

512 ANNALS OF THE SOUTH AFRICAN MUSEUM

concave, as they do in R. batis. The author is unable to detect any appreciable
difference in shape.

Examination of specimens of comparable size indicates that the inter-
orbital is narrower in &. pullopunctata than in R. batis, and that there is little
difference in the comparative size of the eye.

The most noticeable difference is the presence of the large, single, median,
nuchal spine in R. pullopunctata, which is found both in juveniles and adults
(although sometimes worn down). Except about the eyes, the upper surface of
the disc is entirely devoid of other large spines, as it is in R. batis.

Therefore Smith’s proposal that R. pullopunctata and R. batis are distinct
species as based on the general external morphology, is confirmed on the basis
of the clasper structure. Recently, two large ‘black-bellied’ skates, lacking
signs of a median nuchal spine, were obtained in trawls off Cape Columbine
in 250 fathoms. ‘The external characters of the clasper of the mature male were
seen to be identical with those of R. batis from European waters. Therefore
R. batis Linnaeus also may occur off South Africa; however, further investi-
gation on these specimens is necessary.

It is interesting to note that a ‘black-bellied’ skate from the Falkland
Islands, R. flavirostris Philippi 1892, greatly resembles the South African
R. pullopunctata in possessing a single, large, median, nuchal spine. But as yet,
clasper material is unavailable for the purposes of comparison.

The author was unable to find a single difference, both in the external
structure of the glans and in the number and arrangement of the internal
cartilaginous elements, between the claspers of R. clavata and R. rhizacanthus.
The claspers in these two species are identical.

Although Norman points out that the South African species is closely
related to the European species, he considers that the two are distinct on the
basis of the degree of spination in adults, the position of the vent and the shape
of the snout. Preliminary examination of some two hundred specimens of
R. rhizacanthus has shown that the shape of the snout varies considerably in
local populations, as does the degree of spination, and the position of the vent.

Therefore bearing this in mind, and in view of the fact that the claspers
are identical, the author agrees with Barnard and Fowler that R. rhizacanthus
specimens should be referred to R. clavata and that R. rhizacanthus therefore
becomes a synonym of R. clavata. This is further supported by the fact that
masked bipolarity is already acknowledged in R. alba, so that it is not unlikely
that distribution of this type also occurs in R. clavata.

It may be seen from fig. 8 that the distribution of R. clavata ranges from
Arctic waters to the tropical waters of the Indian Ocean, having been recorded
from greater extremes than R. alba. However, through most of their recorded
range both species occur in the same waters. As yet, these two species have never
been recorded from the equatorial waters of West Africa, but equatorial
submergence probably occurs, allowing gene flow from northern hemisphere
to southern hemisphere and vice versa.

RAJA RHIZACANTHUS REGAN AND RAJA PULLOPUNCTATA SMITH 513

AX, 4 4

ar a

ooo A
A- @
A:.-
@
* wt
, A
C

@r oe

. alba te
AR. clavata er

Fic. 8. Map showing the masked bipolar distribution of R. alba and R. clavata.

ACKNOWLEDGEMENTS

I am grateful to Messrs. M. J. Holden and C. H. Humphries of the
Fisheries Laboratory at Lowestoft for the R. clavata and R. batis material and
to Mr. S. X. Kannemeyer for assistance in collection of the South African
skates. My thanks are due to Dr. N. F. Paterson for checking the manuscript
and for many helpful recommendations.

The Trustees of the South African Museum acknowledge gratefully the

receipt of a grant from the Council for Scientific and Industrial Research
towards the cost of publication.

514: ANNALS OF THE SOUTH AFRICAN MUSEUM

SUMMARY

After detailed studies of the anatomy of the claspers of the males, it is
considered that Raja rhizacanthus Regan is synonymous with R. clavata Linnaeus
and that R. pullopunctata Smith is distinct from R. batis Linnaeus, but that
R. batts Linnaeus may also occur in South African waters.

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New Mer McGraw-Hill.

Norman, J. R. 1935. Coast fishes Pt. I. The South Atlantic. ‘Discovery’ Rep. 12: 3-58.

PIETSCHMANN, V. 1906. Ichthyol. ergebnisse einer reise nach Island. Anniln. naturh. Mus. Wien.
21: 72-148.

Petri, K. R. 1878. Die Copulations-organe der Plagiostomen. Z. wiss. Zool. 30: 288-335.

Situ, J. L. B. 1961. The sea fishes of southern Africa. Cape Town: C.N.A.

SmirH, J. L. B. 1964. Fishes collected by Dr. Th. Mortenson off the coast of South Africa in
1929, with an account of the Bom Cruriraja in South Africa. Vidensk. Meddr dansk naturh.
Foren. 126: 283-300.

Wuirte, E. G. 1937. Interrelationships of the elasmobranchs with a key to the order Galea.
Bull. Am. Mus. nat. Hist. 74: 87-117.

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TaBLE or CONTENTs and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. = 7 in. (74 in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

SmitH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmiTH, C. D. 1954. South African Plonias. In Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, Fuly 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 ( = natalensis West).

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MUS. COMP. Zool,
LIBRARY

HARDING & SMITH
MAY 26 1967

ARD

SOME SOUTH AFRIGANA®:
FRESH WATER COPEPODA

April 1967 April

Volume 48 Band

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ANNALS OF THE SOUTH AFRICAN MUSEUM
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SOME SOUTH AFRICAN FRESH WATER COPEPODA

MUS. COMP. ZOOL.

By

: LIBRARY
J. P. Harpine anp W. A. SMITH MAY 28 1967
British Museum (Natural History), London
HARVARD
UNIVERSITY

(With 9g figures in the text)

CONTENTS

PAGE
Introduction hr aed GREG
Systematic account... Pac: ts
pomuaary 6. 8 See . 520
Acknowledgement gj LOE PY ined pea
Relerences, |. : ; : . 521

INTRODUCTION

The material consisted of 26 tubes from the western Cape Province, 45
tubes from the Jukskei—Crocodile system, 39 tubes from the Vaal River, and
32 tubes from other parts of the Transvaal. The material was collected by
Dr. A. D. Harrison and his colleagues.

The collection comprised 9 species of Diaptomidae and 11 species of
Cyclopoida. Unidentified Harpacticoida were also present in some of the
samples. The following species were identified:

CYCLOPOIDA

. Macrocyclops albidus (Jurine) sub.-sp. oligolasius Kiefer
. Eucyclops serrulatus (Fischer)

. Eucyclops gibson (Brady)

. Tropocyclops prasinus Fischer

. Paracyclops fimbriatus (Fischer)
. Paracyclops finitimus Kiefer

. Acanthocyclops vernalis (Fischer)
. Microcyclops varicans (Sars)

. Mesocyclops leuckarti (Claus)

. Thermocyclops oblongatus (Sars)
. Thermocyclops emini (Mrazek)

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. 9. Afr. Mus. 48 (21), 1967: 515-521, 9 figs.

O05)

516 ANNALS OF THE SOUTH AFRICAN MUSEUM

DIAPTOMIDAE

12. Lovenula barnardi Sars

13. Lovenula falcifera (Lovén)

14. Paradiaptomus lamellatus Sars

15. Metadiaptomus transvaalensis Methuen
16. Metadiaptomus capensis (Sars)

17. Metadiaptomus colonialis (Douwe)

18. Metadiaptomus purcelli (Sars)

19. Tropodiaptomus spectabilis (Kiefer)

20. Eudiaptomus sp.

The names used for the Cyclopoida are in accordance with Kiefer’s
monograph (Kiefer, 1929a). This gives a full synonymy and here we give
synonyms only with reference to other works on South African copepods.

Macrocyclops albidus (Jurine) sub.-sp. oligolasius Kiefer

The species M. albidus is a cosmopolitan one but specimens in Africa south
of the Sahara if examined sufficiently closely are usually found to belong to the
subspecies oligolasius which was first described (Kiefer, 1928a) from the Linjanti
River where it flows into the Zambezi. Hutchinson et al. (1932) record it from
the south-eastern Transvaal. Sars (1927) did not find any representatives of
the genus Macrocyclops in his survey of the Entomostraca of the Cape Province
but two of the present samples from the western Cape Province contained
M. albidus oligolasius, both from a vlei at Betty’s Bay in brown and acid water.
It was also present in the Vaal River, the Olifants Vléi near Johannesburg,
the Klein Jukskei1 River and from the Great Usutu River in the eastern

Transvaal.
®

Eucyclops serrulatus (S. Fischer)

This cosmopolitan species was found in all regions sampled in the present
survey. It is rather surprising that so common a species was not found in the
Cape Province by Sars (1927). Several of the present samples contained it.
Hutchinson et al. (1932) record it from south-eastern Transvaal and it was
present in many of the present samples from the Transvaal.

Eucyclops gibson (Brady)

Cyclops gibsoni Brady 1904: 123, pl. 6, figs. 1-10.
Cyclops longistylis Brady 1910: 242. (Nigeria.)
Afrocyclops gibsoni (Brady), Sars 1927: 121, pl. 13, figs. 1-18.
Eucyclops gibsoni (Brady), Hutchinson et al. 1932.

This is a species widely distributed in Africa south of the Sahara.

Found in the western Cape Province in the Krom River, Stellenbosch,
and under the Blouberg road bridge, Milnerton, and in the Transvaal at the
Amato Dam, Benoni, and at the Pretoria sewage works.

Brady’s type of longistylis from Nigeria is in the BM(NH) on a slide and
appears to be this species as Sars suggested.

SOME SOUTH AFRICAN FRESH WATER COPEPODA 517

Tropocyclops prasinus Fischer

Leptocyclops prasinus (Fischer) Sars 1927: 119, pl. 12, figs. 11-20.
Eucyclops prasinus (Fischer) Hutchinson e¢ al. 1932.

This is a very widely distributed probably cosmopolitan species, but was
represented in the present collections only by a single male from the Crocodile—
Jukskei system. It is a very small species and may have passed through the nets.
It is recorded from the Cape (Sars, 1927) and the Transvaal (Hutchinson
@ Gl..5. 132). |
Paracyclops fimbriatus (Fischer)

This species is reported from the whole of the Palaearctic and Nearctic
regions and most parts of Africa. It was found in four of the samples from the
Vaal River, two of which contained the related form Paracyclops finitimus as well.

Paracyclops finitimus Kiefer
Platycyclops poppei (non Rehberg) Sars 1927: 127, pl. 14, figs. 5-17.

This species was first described from Morocco (Kiefer, 1928b: 99) and has
been found in New Zealand. Sars had already recorded it under the synonym
given above from the Cape Peninsula.

It was found in three samples from the Cape Province: the Krom River at
Stellenbosch and the Berg River and in nineteen from the Transvaal: at
Benoni, in the Crocodile—Jukskei System, in the Klipspruit near Johannesburg,
in the oxidation pond of the Pretoria sewage works and in the Vaal.

Acanthocyclops vernalis (Fischer)

This species is widely distributed in the Northern Hemisphere and has
been recorded from South America and Kerguelen. In the present collection
it is represented by a single adult female from the Berg River.

Microcyclops varicans (Sars)

Owing to uncertainties of identification the precise distribution of this
species is not known; but there are records from all principal land masses. I
know of no record for South Africa south of the Vaal River; but Hutchinson
et al. (1932) record it from the Transvaal. It was present in three tubes from
the Crocodile—Jukskei System in marginal vegetation and from two stations
on the Vaal River.

Mesocyclops leuckartt (Claus)

This cosmopolitan species was found in twenty-four of the samples: from
the western Cape Province, from the Vaal River and from the Transvaal.

Thermocyclops oblongatus (Sars)

(Figs. 1-9)
Mesocyclops oblongatus Sars 1927: 114, pl. 11, figs. I-15.
(Thermocyclops) infrequens Kiefer 1929b: 315, fig. 10.
Sars (1927) described T. oblongatus from the Cape Province. Unfortunately
his figures are not very accurate, in particular showing a peculiarly shaped

518 ANNALS OF THE SOUTH AFRICAN MUSEUM

receptaculum and a fifth leg with the outer terminal seta less than half the
length of the inner. Kiefer (1929b) described T. infrequens and many specimens
of the present collection from both the Transvaal and the Cape agree with
Kiefer’s description and figures, but none with Sars. We have examined Sars’s
original material and find that the receptaculum and fifth leg are the same as
in Kiefer’s species and as these were the only features by which the two species
were thought to be distinguishable we conclude that oblongatus and infrequens are
two names for a single species. There are two tubes of Sars’s material, one which
had at some time been allowed to dry up was from a small pond at Salt River,
collected by Dr. Purcell and referred to in Sars’s paper, and the other of speci-
mens in good condition from an unspecified locality. Both tubes appeared to
contain the same species and figs. 1-4 are from a specimen from the second tube.
Figs. 5-9 are of corresponding parts of specimens from the Transvaal in the
present collection. The species is very close to T. schuurmanae Kiefer, the main
difference being that the rounded prominence on the lamella of leg 4 (figs.
2 and 6) are larger, closer together and bear more spinules. Kiefer’s original
specimens of 7. schuurmanae were from the Transvaal (Kiefer, 1928a: 15) and
Hutchinson ef al. (1932) also record it from several Transvaal localities and
from a Cape Vlei. However, apart from a few T. emini, all the Thermocyclops in
the present collection are 7. oblongatus, not T. schuurmanae. The localities are:
the Zeekoe Vlei on the Cape Flats, various dams at Benoni, Transvaal, five
stations in the Crocodile—Jukskei System and the Apies River, Pretoria.

Thermocyclops emini (Mrazek)

Originally found in East Africa (Mrazek, 1896: 4; Sars, 1909: 50) it
has since been found in the Sudan and in Egypt (Gurney, 1911: 32) and in the
Transvaal and Portuguese East Africa (Hutchinson et al., 1932). The present
samples were both from the Vaal Barrage.

Lovenula barnardi Sars
Originally described from Ovamboland, South West Africa (Sars, 1927:
92) this species was present in a number of the samples collected along the
Vaal River. |
Lovenula falcifera (Lovén)

First described from Natal and later found near Cape Town (Sars, 1927:
86) and in a number of places in the Transvaal (Hutchinson e¢-al., 1932). This
species was found only in the Cape Province in the present collections: in
temporary waters on the Cape Flats, Berg River localities and the Riet Vlei,
Milnerton.

Paradiaptomus lamellatus Sars

The type locality is Knysna, Cape Province (Sars, 1895: 46) and this
species is recorded by Sars (1927: 94) also from near Cape Town. Brady (1907:
183) records it from Natal and Rithe (1914: 27) from the Cape. The species was
found in three of the present samples, all from the Cape.

ee ee

o89

SOME SOUTH AFRICAN FRESH WATER COPEPODA

‘sSa] WIJ pue sturwes umnpnovydaser SuIMoyYs JUsUISaS jeques jo jUSUISaS 1DvI0Y}
ysey “Qg “Y ‘s81q ‘Sa_ yIINo; Jo podopus jo yuouiSes jeurus} “6 ‘E ‘s81q ‘ssa_ YIINOF jo eppeurey Ssunrun ‘g *% ‘SSI “TWIeI [BOINy
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520 ANNALS OF THE SOUTH AFRICAN MUSEUM

Metadiaptomus transvaalensis Methuen
Metadiaptomus transvaalensis Methuen 1910: 160, pl. 16, figs. 46a—c, pls. 17-18.
Diaptomus meridianus Douwe 1912: 24.
Paradiaptomus transvaalensis (Methuen) Hutchinson et al. 1932.
Diaptomus rigidus Sars 1927: 101, pl. 8, figs. 13-18.

This is the type species of the genus and was first described (Methuen,
1910) from Lake Chrissie and other parts in the eastern Transvaal. Hutchinson
et al. (1932) also record it from this area and other parts of the Transvaal. Van
Douwe’s 1912 specimens from Great Nama Land are probably the same species
and the present samples were mostly from the Vaal River, and also from one
station in the western Cape Province at Piquetberg on the Berg River. Only
two battered females were present in the last sample.

Metadiaptomus capensis (Sars)

Diaptomus capensis Sars 1907, Riihe 1914: 26, fig. 8a, b. Sars 1927: 98, pl. 8, figs. 1-12.
Paradiaptomus capensis (Sars) Hutchinson e¢ al. 1932.

This species as far as I know has only been recorded from the Cape
Province, it was present in three of the samples, the Zeekoe Vlei and a tem-
porary pool on the Cape Flats and Sirkel’s Vlei near Cape Point.

Metadiaptomus colonialis (Douwe)

Diaptomus colonialis Douwe, 1914.
Paradiaptomus colonialis (Douwe) Hutchinson et al. 1932.

The specimens described by Van Douwe were from South West Africa
and those collected by Hutchinson et al., from various parts of the Transvaal.
Eight of the present samples were from inside the Transvaal and three from
its southern border.

Metadiaptomus purcellr (Sars)

Diaptomus purcelli Sars 1907: 12, pl. 2, figs. 3-10. Sars 1927: 103, pl. 9, figs. 1-8.
Paradiaptomus purcelli (Sars) Hutchinson ¢é al. 1932.

All records of this species seem to be from the south-western bape Province
and it is characteristic of temporary pools on the Cape Flats.

Tropodiaptomus spectabilis (Kiefer)

Diaptomus spectabilis Kiefer 1929b: 311, figs. 5, 6. Hutchinson e¢ al. 1932.
Tropodiaptomus spectabilis (Kiefer) Relies 1932.

This is a Transvaal species and was collected from two Trae localities.

Eudiaptomus sp.

Three females and one male of an undetermined species of Eudiaptomus
were present in a sample from the Hartebeespoort Dam.

SUMMARY

Eleven species of Cyclopoida and eight species of Diaptomidae are recorded
from inland waters of the Cape Province and Transvaal. Figures of Thermo-
cyclops oblongatus (Sars) are given and it is shown that TJ. znfrequens Kiefer is
a synonym of this species.

i
A

SOME SOUTH AFRICAN FRESH WATER COPEPODA 521

ACKNOWLEDGEMENTS

We are indebted to Dr. A. D. Harrison and the National Institute for
Water Research of the South African Council for Scientific and Industrial
Research for the opportunity of examining this material. We are grateful to
Dr. J. R. Grindley for letting us examine specimens described by G. O. Sars
in the collections of the South African Museum.

The Trustees of the South African Museum acknowledge gratefully the
receipt of a grant from the Council for Scientific and Industrial Research
towards the cost of publication.

REFERENCES

Brapy, G. S. 1904. On Entomostraca collected in Natal by Mr. James Gibson. Proc. zool. Soc.
Lond. 1904 (2): 121-128.

Brapy, G. S. 1907. On Entomostraca collected in Natal by Mr. James Gibson. (Part IT.)

_ Ann. Natal Mus. 1: 173-186.

Brapy, G. S. 1910. On some species of Cyclops and other Entomostraca collected by Dr. J. M.
Dalziel in northern Nigeria. Ann. trop. Med. Parasit. 4: 239-250.

DouweE, C. VAN. 1912. Neue Stisswasser-Copepoden aus Siidafrika. Jn Schultze, L. Zoologische und
anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stidafrika 1903-1905.
5: 21-32. Jena: Fischer. (Denkschr. med.-naturw. Ges. Jena 17: 21-32.)

Douwe, C. VAN. 1914. Copepoda. Jn Michaelsen, W., ed. Beitrdége zur Kenntnis der Land und
Stisswasserfauna Deutsch-Stidwestafrikas. 1: 93-103. Hamburg: Friederichsen.

Gurney, R. 1911. On some freshwater Entomostraca from Egypt and the Soudan. Ann. Mag. nat.
Hist. (8) 7: 25-33.

Hutcuinson, G. E., Pickrorp, G. E. & Scuuurman, J. F. M. 1932. A contribution to the
hydrobiology of pans and other inland waters of South Africa. Arch. Hydrobiol. 24: 1-154.

Kierer, F. 1928a. Beitrage zur Copepodenkunde. (VIII.) Zool. Anz. 76: 5-18.

Krerer, F. 1928b. Beitrage zur Kenntnis der freilebenden Copepoden Marokkos. Bull. Soc.
Sci. nat. Maroc 8; 87-106.

Krerer, F. 1929a. Crustacea Copepoda. II. Cyclopoida Gnathostoma. Tierreich 53: i-xvi, 1-102.

KieFrer, F. 1929b. Neue und wenig bekannte Siisswasser-Copepoden aus Siidafrika. Zool. Anz.
80: 309-316.

KreFer, F. 1932. Versuch eines Systems der Diaptomiden (Copepoda Calanoida). Zool. 7b.
(Syst.) 63: 451-520.

MeruHueEn, P. A. 1910. On a collection of freshwater Crustacea from the Transvaal. Proc. zool.
Soc. Lond. 1910: 148-166.

MrAzexk, A. 1896. Die Copepoden Ost-Afrikas. In Deutsch-Ost-Afrika 4 fig. 1 (Mobius, K., ed.
Die Thierwelt-Ost-Afrikas . . . Wirbellose Thiere) (11): 1-11. Berlin: Reimer.

Rtue, F. E. 1914. Die Siisswassercrustaceen der Deutschen Siidpolar Expedition 1901-1903
mit Ausschluss der Ostracoden. Dtsch. StidpolExped. 16 (Zool. 8): 4-66.

Sars, G. O. 1895. On some South African Entomostraca raised from dried mud. Skr. Vidensk-
Selsk., Christ. 8: 1-56.

Sars, G. O. 1907. On two new species of the genus Diaptomus from South Africa. Arch. Math.
Naturv. 28 (8): 1-17.

Saps,G. O. 1909. Zoological results of the third Tanganyika expedition, conducted by Dr.
W. A. Cunnington, F.Z.S., 1904-1905. Report on the Copepoda. Pros. zool. Soc. Lond.
1909: 31-77.

Sars, G. O. 1927. The freshwater Entomostraca of the Cape Province (Union of South Africa).
Part III. Copepoda. Ann. S. Afr. Mus. 25: 85-149.

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INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TaBLeE oF CONTENTS and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 4? in. = 7 in. (7# in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

Situ, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmiTH, C. D. 1954. South African Plonias. In Brown, X. Y. Marine faunas. and ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).
Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:

Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.

When another species has been called by the same name:

[non] Plonia capensis: Jones, 1957: 27 ( = natalensis West).

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MUS. COMP. ZooL.
LIBRARY

MAY 26 1967

HARVARD
THE FISHES OF TRISTAN DA QCUNEAN,
GOUGH ISLAND
AND THE VEMA SEAMOUNT

M. J. PENRITH

March 1967 Maart
Volume 48 Band

Part 22 &Deel

ANNALS OF THE SOUTH AFRICAN MUSEUM

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THE FISHES OF TRISTAN DA CUNHA, GOUGH ISLAND

AND THE VEMA SEAMOUNT
MUS. COMP. ZOOL.

By LIBRARY
M. J. PENRITH* a
: MAY 26 1967
South African Museum, Cape Town
HARVARD
(With 2 text-figures and 1 plate) UNIVERSITY
CONTENTS
PAGE
Introduction : , ‘ ». 52g
Material : : : : » 525
Systematic account . : ») {926
Zoogeography . ; : an, ALE
Summary . , : ‘ = SAS
Acknowledgement : : . 546
peieremees, 6.10.13) sy A. eb
INTRODUCTION

In the south-east Atlantic Ocean there are three known areas of very
shallow water (not counting the African coast). Two of these are around the
shores of islands, the Tristan da Cunha group and Gough Island, while the
third is the Vema Seamount.

Tristan da Cunha

The island of Tristan da Cunha, together with the two smaller islands of
Inaccessible and Nightingale, lies almost midway between South Africa and
South America (37° 05’ S., 17° 40’ W.). It is permanently inhabited and has
been the subject of several scientific expeditions and reports.

Capt. Dugal Carmichael, F.L.S. (army surgeon to the garrison in the
island, 1816-17) was the first to publish a report on the flora and fauna of
Tristan. In this paper (1818) he described four species of fishes from the island
and listed five others as being common to both Tristan and the Cape. Car-
michael’s work was referred to by Regan (1913a) but no further fishes were
added to the nine already known from the island until Barnard (1923, 1925)
listed two more species which he considered to be conspecific with South
African species. Norman (1935a) described a further species believed by him
to be endemic to the island group. After the Norwegian expedition of 1937-8,
Sivertsen (1945) could add a further seven species to the fish fauna of the island,
two of them being new species and believed by him to be endemic to Tristan
da Cunha. Finally, Rowan & Rowan (1955) added three more records of
oceanic fishes from the island.

* Seconded from the Council for Scientific and Industrial Research Oceanographic Unit,
University of Cape Town.

523
Ann. S. Afr. Mus. 48 (22), 1967: 523-548, 2 figs., 1 pl.

524 ANNALS OF THE SOUTH AFRICAN MUSEUM

At present the known fish fauna of Tristan consists of 19 species which
Sivertsen (1945) divided into an oceanic component and a coastal component.
(In the list below, the first reference to the species occurring at Tristan is given
in parenthesis after the author of the species.) The islanders’ common name for
the species is also given where known.

(a) Oceanic species
Prionace glauca (Linn.) (Sivertsen, 1945) . . . . Blue shark
Maurolicus muellert (Gmelin) (Barnard, 1925) . . —
Myctophum humboldti (Risso) (Sivertsen, 1945) . .
Scomberesox saurus (Walbaum) (Sivertsen, 1945) . . —
Exocoetus exiliens Bloch (Carmichael, 1818) . . . —
Alepisaurus ferox Lowe (Rowan & Rowan, 1955) —
Notopogon lillie: Regan (Sivertsen, 1945) . . . . —

Seriola lalandi Valenc. (Rowan & Rowan, 1955) . Yellowtail
Polyprion americanus (Bloch) (Rowan & Rowan, 1955) Steambrass
Thyrsites atun (Euphrasen) (Carmichael, 1818) . . Snoek

(b) Coastal species
Artosoma australis (Barnard) (Barnard, 1923). . . _
1Gaidropsarus insularum Sivertsen (Sivertsen, 1945). .. =
Acantholatris monodactylus (Carmichael) (Carmichael,

1818) i. Ne Gudek elan 60 2alehtt en
Bovichthys diacanthus (Carmichael) (Carmichael, 1818) Klipfish
*Decapterus longimanus Norman (Norman, 1935a) . . Mackerel
Labrichthys ornatus (Carmichael) (Carmichael, 1818) Concha or Conger
3Helicolenus tristanensis Sivertsen (Sivertsen, 1945) . Soldier

Sebastichthys capensis (Gmelin) (Carmichael, 1818) . Soldier
Sertolella antarctica (Carmichael) (Carmichael, 1818) Bluefish
Seriolella christophersent Sivertsen (Sivertsen, 1945) . Stumpnose

The identity of the fish recorded by Carmichael as the ‘Roman fish’
Sparus , 1s not known. The South African ‘Roman’ is a Sparid (Chryso-
blephus laticeps) but has not been recorded from Tristan.

Gough Island

Gough Island lies slightly farther south than Tristan (40° 20’ S., 9° 55’ W.),
and is not permanently inhabited, although the South African Government
has maintained a weather station on the island for several years.

Although several expeditions have called at the island, it has not been
subject to the same degree of collecting as has Tristan. Only one expedition,
the Gough Island Scientific Survey 1955-6, has spent much time on the island,
but they have not published any data on the fishes obtained.

1 As G. capensis (Kaup) by Barnard (1925).
* Probably the fish recorded by Carmichael as Scomber trachurus.
® Recorded by Carmichael as Sebastichthys maculata.

Fy ae

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 525

Accounts of fish collected by the Scotza, which called on the way back
from the Antarctic (Regan, 1913)) and by the R.V. William Scoresby (Norman,
19350), have been published.

The fishes recorded are:

1Helicolenus tristanensis Sivertsen (Regan, 19350)
Sebastichthys capensis (Gmelin) (Regan, 1913))

Bovichthys diacanthus (Carmichael) (Regan, 19136)
Caesioperca coatst Regan (Regan, 1913))

Acantholatris monodactylus (Carmichael) (Norman, 19350)

Holdgate (1958), in a popular account of the Gough Island Scientific Survey,
listed the following species of fish under their Tristan common names: five-
fingers, soldiers, klipfish, congers.

Vema Seamount

This exceptionally shallow seamount was discovered in 1957 by the
R.V. Vema, and traverses were made then as well as in 1959 by the Vema and
in 1963 by the R.V. Robert D. Conrad. In November 1964 a detailed survey of
the peak was undertaken by the South African National Committee for
Oceanographic Research, using the diamond prospecting tug, Emerson K
(Simpson & Heydorn, 1965). ‘The peak was found to have a position 31° 38’ S.,
8° 20’ E. and a least depth of only 14 fathoms, with a well-defined plateau
about five square miles in extent and of approximately 35 fathoms depth.
Subsequent to the visit by the Emerson K there have been many visits by com-
mercial fishing vessels from Cape Town as well as a short visit by the R.V.
Africana II in April 1965.

MATERIAL

The material used for the present paper was obtained from several sources.
The South African Museum has been slowly building up a collection of Tristan
fish for many years, mainly obtained as donations from Mr. Keytel (1907),
the rock-lobster survey of 1949, and small but valuable donations from Tristan
and Gough Islands by the Division of Sea Fisheries and Capt. M. T. Scott,
master of the rock-lobster fishing vessel, Tristanza.

Specimens of fishes from the Vema Seamount were obtained from the
Division of Sea Fisheries (those from the Emerson K collection, and commercial
fishing vessels, and a collection made from R.V. Africana IT by Mr. A. C.
Paterson), as well as directly from the fishing companies.

These collections have resulted in several further species being added to
the fish fauna of Tristan da Cunha and Gough Island as well as adding to our
knowledge of the species already recorded from the islands, and in addition
have enabled their fauna to be compared with that from the Vema Seamount.

1 Recorded as Sebastes maculatus C. & V.

526 ANNALS OF THE SOUTH AFRICAN MUSEUM

SYSTEMATIC ACCOUNT

Exocoetidae
Cypsilurus lineatus (Cuvier & Valenciennes)

Cypsilurus lineatus (Cuvier & Valenciennes) Bruun, 1935: 47.

Only one specimen, of 382 mm standard length, was examined. The fish
was found in a drum of unsorted material from Tristan in the Department of
Ichthyology, Rhodes University.

The body proportions as thousandths of standard length are as follows:
pectoral fin 675, pelvic fin 309, dorsal height 111, depth 157, head 199, pre-
dorsal distance 728, preventral distance 560, preanal distance 754, snout 50,
eye 65. Fin counts are dorsal 12, anal 10 and pectoral 15.

This is an extremely large specimen of the species.

Gadidae

Gaidropsarus Rafinesque

Kaup (1858) described a small gadid as Motella capensis but gave only a
brief description and no locality, although on the basis of the trivial name it is
believed to be South Africa.

Small rocklings were described from St. Paul Island as M. capensis by
Kner (1868) and Sauvage (1879), while Barnard (1925) published a description
of material in the South African Museum which he also assigned to Kaup’s
species. |

Sivertsen described specimens from Tristan and showed that while they
were very similar to those described from St. Paul Island, they showed marked
differences from the South African specimens as described by Barnard. Since
South Africa was believed to be the type locality of capensis, Sivertsen proposed
a new name, G. znsularum for the Tristan and St. Paul Island specimens.

On re-examining the material used by Barnard, I find that only one of
the two fish he examined exactly fits the description he published, and that the
two fish he examined appear to belong to two distinct species. This view is
shared by Mr. A. Wheeler of the British Museum (Natural History), who has
also examined the South African Museum material, and Professor J. L. B.
Smith, who informs me that he has reached a similar opinion from examination
of material in his collection (Table 1).

TABLE I

Comparison of Gaidropsarus

Table 1a gives body proportions of two species of Gaidropsarus in millimetres and as
thousandths of standard length in parenthesis.

Table 1b compares body proportions of Gaidropsarus from various sources.

12719 G. insularum from False Bay
12528 G. capensis. Port Elizabeth
7858 G. capensis. East London
A G. insularum from Tristan da Cunha (from Sivertsen, 1945)
G.

B capensis from South Africa (Barnard, 1925)

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 527

TABLE 1a
12719 12528 7858
Standard length .. 153 gi 156
Wepth*.. : : ~ oo naiaeie. a5 (r65)eee7 73)
Head ; t ‘ are '20-ye (rQO)e 20-5 (206) 5 a8), W(212)
Pe 46 90) 398 (42)e.) 573. (34)
Snout ; : : : 8 (52) G*S. (75) 9:3 (60)
Naferorbital . .° . 6™ (39) Ar (4Qp 4°5 (29)
Bae lie le SF 5S (SeieuenO> (1FO)es) E7-2.( bro)
mee ott dorsal. . 19 (85) 12 (rg2)*‘ 19 (122)
me cians ts). 77 (500) 44 (485)y -76-5 (490)
TABLE 1b

A 12719 B 12528 7858

Depth/length : ; é . 64-7 62 6 6 6
Head/length ee is cae 54 4s 44 43
ieee... : é . 63-7 64 5 5¢ 64
Eye/interorbital . ; ; . I -14 iz +1 +1 14
and Pelvic ray/Origin ray to vent 2 3 3 3 3
Caudal peduncle/head : . 2%-a24 2k = 34 34
Rays, first dorsal ; : - 47-49 +45 55 65 56
Rays, second dorsal _ . : wy Gal 64 62 45 44. 43
Raywiwamal fine... «.- « «©, 52754 50 40 37 39

One specimen from Kalk Bay in the Cape Peninsula as well as some
juveniles from Table Bay and Lambert’s Bay fitted the description of the Tristan
fishes as given by Sivertsen and the St. Paul Island fishes as described by
Kner and Sauvage, while the specimens from East London and Port Elizabeth
are Clearly different, and are assumed to be G. capensis.

Gaidropsarus insularum Sivertsen

Motella capensis (non Kaup) Kner, 1868: 279. Sauvage, 1879: 42.
Gaidropsarus capensis (non Kaup) Barnard, 1925: 323 (partim). Blanc, 1961: 145. Blanc & Paulian,

1957: 327-
Gaidropsarus insularum Sivertsen, 1945: 8.

Recorded from Tristan da Cunha, St. Paul Island, the Cape Peninsula
and west coast of South Africa.

Body proportions and fin counts given in Table 1.

Gaidropsarus capensis (Kaup)

Motella capensis Kaup, 1858: go.
Gaidropsarus capensis (Kaup) Barnard, 1925: 323 (partim).

Depth 6 in standard length, head about 44. Eye equal to interorbital
about 6 in head. Second pelvic ray ? distance origin pelvic to vent. Caudal
peduncle 3} in head. Fin counts: Free dorsal rays 55-65, Dorsal rays 42-45,
Anal rays 36—40.

Description based on two specimens, one of 91 mm standard length

528 ANNALS OF THE SOUTH AFRICAN MUSEUM

deposited in the South African Museum (Reg. No. SAM 12528), collected in
Algoa Bay at a depth of 25 fathoms by the Pieter Faure, and a second of 156 mm
standard length in the collection of the Department of Ichthyology, Rhodes
University, Grahamstown (Reg. No. 7858), collected at East London.

Body proportions and fin counts are given in Table 1.

G. capensis differs from insularum in having a relatively longer first dorsal
base (12—13°5% of body length as opposed to about 8-5°%), a longer head,
larger eye, slimmer caudal peduncle and rather marked differences in fin
counts.

Berycidae
Beryx decadactylus Cuvier

Beryx decadactylus Cuvier. Fowler, 1936: 542.

One specimen obtained from Captain Scott, caught in “Tristan waters’.
Tears in the mouth suggest that the fish was caught with hook and line but
depth of capture is unknown.

The fish is 362 mm standard length and has the following body proportions
(expressed as thousandths of standard length): head 359, depth 472, eye 152,
maxilla 185, base of anal fin 348, base of dorsal fin 290. Fin counts are dorsal
IV 19, anal IV 27, pectoral 15, pelvic I 10.

B. decadactylus has not been recorded from South Africa although the
related B. splendens is trawled quite regularly. In the course of this work an
unrecorded specimen of B. decadactylus was found in the collection of the
South African Museum, trawled off Table Bay in 1948. It was 345 mm in
standard length and had the following body proportions (in thousandths of
Ls): head 322, depth 446, eye 125, maxilla 174, base of anal fin 359, base of
dorsal fin 293. Fin counts are dorsal IV 20, anal IV 27, pectoral 15, pelvic I 10.

Macrorhamphosidae
Notopogon lillies Regan

Notopogon lillies Regan, 1914: 14. Mohr, 1937: 50.

Sivertsen (1945) records two specimens of Notopogon which were found
washed up dead on Tristan. He assigned them to WV. Jilliei, a species known from
South Africa, Australia and New Zealand. It should be noted, however, that
his published figure (fig. 5) resembles V. macrosolen Barnard rather than N.
lilliei, especially as neither the figure nor the text makes any reference to the
dorsal brush which is a conspicuous feature of WV. lilliei but is absent in WV.
macrosolen.

Chilodactylidae

Sivertsen (1945) recorded only one species of fish belonging to the family
Chilodactylidae, Acantholatris monodactylus (Carmichael) ; called by the islanders
the ‘Fivefinger’.

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 529

He claimed that the islanders spoke of a second fish very like the fivefinger
which they were said to call the ‘yellowtail’. Rowan & Rowan (1955) have
shown that the fish called yellowtail is Sertola lalandi. No additional species of
chilodactylids have been received from Tristan but a species which appears to
be new has been obtained from Vema Seamount.

An additional species which appears to be undescribed has been obtained
from Gough Island, but is not described here as only one specimen was obtained.
It is known to fishermen as the ‘funny fish’.

Acantholatris monodactylus (Carmichael)
(Plate XXIs)

Chaetodon monodactylus Carmichael, 1818: 500.
Cheilodactylus carmichaelis Cuvier & Valenciennes, 1830: 360. Kner, 1868: go.
Chilodactylus monodactylus (Carmichael) Sauvage, 1879: 23; Regan, 1913a: 466. Norman, 1935):

57:
Acantholatris monodactylus (Carmichael) Gill, 1863: 119. Sivertsen, 1945: 10.

This species is known from Tristan and Gough Islands and has also been
found to be common at the Vema Seamount. Gunther (1860) recorded this
species from Chile but Norman (19354) has shown this to be a misidentification,
the Chilean species being the related A. gayi (Kner). In Table 2 below are given
body proportions and counts of several examples of the species from Tristan
da Cunha, Gough Island and the Vema Seamount.

TABLE 2

Body proportions and fin counts of Acantholatris monodactylus in thousandths of standard
length.

Origin of the specimen is:

Cat. No. SAM 10500 Tristan da Cunha
10501 ‘Tristan da Cunha
11764 No history, but register says ‘Very likely Tristan’
21284 ‘Tristan (perhaps!!)
22890 Gough
24288 Vema Seamount
24291 Vema Seamount

Cat. No. SAM 11764 11784 11764 10500 10501 10501 21284 10501 24291 24288 22890

ERG ats ANT 50. eR iieoe.) | BiG) 2000 9192-9). 936" 91955) 1/.. 851 1) 92). 3929... 1336
Depth - 405 385 405 389 379 391 396 391 370 381 405
Bee i, PW GA. 78 82 75 75 76 67 62 62 59 56
Interorbital. . . 69 78 76 75 84 92 93 88 74. 79 88
Base of dorsal. . 656 620 634 657 625 630 638 636 636 £663 _~ 605
Paid 8s, TO 177 - 172 154 160 153 147 . 148. 153 ~— 169

Length of pectoral 351 B40) 820. 9349 Bas6y 3959 349 G00 325. 909 . 320
Snout to dorsal

Peer 6 351 334 329 . 315 330. 344 379. 360. 343... 350 . 334
Snout to anal

Sse 879 9700 715, \ 680... 6o3,).705 » 687.679. 674. 665 736
Snout to pectora

Sues - 313 308. 323 303. 334. 332°. 942 340 332 3932 . 356
Worsal spines . . 17 16 ier 17 16 07 17 17 17 Li) 17
Dorsal rays,... . 25 25 25 25 26 25 26 25 25 25 2D
Analspines. . . = 3 3 3 3 3 3 3 3 3 3 3
Pai TAYSE 3) 2 \ , TZ II 12 II II II II II II II 10

Gillrakers . . .7+14 8+15 8+15 8+14 8415 7+15 7+15 7HI15

530 ANNALS OF THE SOUTH AFRICAN MUSEUM

One specimen (SAM 22890) obtained by Captain Scott at Gough Island
has been assigned to A. monodactylus although the fin counts are lower than
normal for the species. The body proportions fit within the range for A. mono-
dactylus however and coloration in formalin is identical.

Acantholatris vemae sp.n.

(Plate XXIa, Table 3)

MATERIAL

Two specimens from the Vema Seamount collected by the Emerson K,
November 1964, donated by the Division of Sea Fisheries, Sea Point (SAM
24.290, 24292).

Five specimens (all gutted) from the Vema Seamount collected by com-
mercial fishing vessels and donated by Messrs. Friedman & Rabinowitz (Pty.)
Ltd. (SAM 24280, 24293, 24296-8).

Two specimens from the Vema Seamount collected by Mr. A. C. Paterson,
April 1965, and donated by the Division of Sea Fisheries (SAM 24525, 24527).

Depth of body about 3, length of head about 3? in standard length. Eye
diameter 5 in length of head, 2 in snout and 14 in interorbital width.

Dorsal fin XVII 24-26, the eighth spine longest, about equal to snout,
4 in body depth. Last spine 24 in first ray. Base of dorsal a little over 14 in
body length, distance snout to origin dorsal 34 in body, about equal to length
of pectoral.

Anal fin III 10-11, base short only + of dorsal base. Second spine very
stout, only a little longer than third, shorter than first ray.

Pectoral having 9 branched, 6 simple rays, first simple ray elongate,
+ body length, free portion 34 in length of ray.

Forehead scaled to level of posterior nostril, both nostrils of similar size,
round.

Scales fairly large, very thin, lateral line 55-58, transverse 7g, cheeks
scaled. Gill rakers, upper 6—7, lower 14-16, total 21-23. |

Colour: in formalin, dark blue-black dorsally; fading to silver below.

Edge of fins dark, caudal with a few dark bands.

In life said to be bright blue.

Detailed body proportions of the seven specimens are given in Table 3.

TABLE 3
Catalogue No.: 24289 24290 24292 24293 24206 24297 24208 24525 24527

Standard length . . 327 320 385 249 72 338 392 384 383
Head. 28) awe. 288 271 260 277 255 264 263 266 253
Depth ... 2... 345 347 343 539 352 329 349 309 BS)
VE Wie ie eet eee 58 57 52 56 54 53 51 50 55
Interorbital 37; 80 75 75 76 75 71 79 78 78
Base of dorsal . .. 624 660 673 650 683 663 666 690 682

Basé-ofanal oi)or2%. « 165 161 174 169 Dye 9s) 078 168 174 175

Se re oe ee be) ee eee el

E

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 531

TABLE 3 (continued)
Catalogue No.: 24289 24290 24292 24293 24206 24297 24298 24525 24527

Length of pectoral . 327 328 204. 321 328 34.0 319 354. 329
Snout to dorsal origin 306 306 288 297 272 278 283 204 287
Snout to anal origin 686 627 607 650 650 642 654.

Snout to pectoral. . 303 278 299 273 253 255 268 263 264
Dorsalspines: . - - 17 17 1a 17 iy 17 17
Wormaliays) =< .- . 24 25 26 24 25 24. 25
Amalspines 9..: . . 3 3 3 3 3 3 3
PAGAL EAS he sw 5. « 10 10 i II II Hi II

Body proportions (in thousandths of standard length) of Acantholatris vemae sp.n. from the

Vema Seamount.
Dates of collection of the various specimens given in the text.
SAM 24527 the Holotype.

The specimen deposited in the South African Museum (Reg. No. SAM
24527) is designated as the type.

A. vemae is very close to A. monodactylus (both having 9 branched and 6
simple pectoral rays) which is also found on the Vema Seamount. It differs,
however, in colour and colour pattern, lacking the six dark vertical bars found
in monodactylus; also it has a markedly shorter head and slightly less deep body
when compared to monodactylus. As a result of the shorter head, the distances of
the snout to the origin of dorsal fin, the origin of anal fin and the origin of
pectoral fin are proportionately less. The slope of the forehead in vemae too
is steeper, and whereas it has the upper corner of the pectoral! fin angular it is
rounded in monodactylus. Finally it has much thinner lips than monodactylus.

It differs from A. gayi and A. aspersus in the pectoral not reaching the soft
anal.

Latris lineata (Bloch & Schneider)

Cichla lineata Bloch & Schneider 1801: 342.
Latris hecateia Richardson, 1839: 99. Richardson, 1842: 106. Gunther, 1860: 86. Kner, 1868: 95.
Sauvage, 1879: 17. Angot, 1951: 19. Blanc & Paulian, 1957: 331. Blanc, 1961: 152.

Two specimens, SAM 22623, 22891, both collected by Captain Scott at
Gough Island. The former is a skin with standard length of about 900 mm and
the latter a complete specimen of 904 mm.

The body proportions of the 904. mm specimen (as thousandths of L’s)
are as follows: head 324, depth 303, pectoral fin 192, snout 149, base of dorsal
fin 638, base of anal fin 207.

Fin counts are dorsal XVIII 35, anal III 26, pectoral 9 + 9.

Counts of the skin are dorsal XVIII 36, anal III 26, pectoral 9 + 9,
gill rakers 8 + 16.

L. lineata was originally described from Australia and New Zealand, but
has been recorded from St. Paul Island by Kner (1868), Sauvage (1879) and
Angot (1951). This is the only species that is without doubt common to the
Atlantic islands and Australasia.

532 ANNALS OF THE SOUTH AFRICAN MUSEUM

Serranidae
Polyprion americanus (Bloch & Schneider)

Amphiprion americanus Bloch & Schneider, 1801: 205.
Polyprion americanus (Bloch & Schneider) Barnard, 1925: 488. Smith, 1949a: 199. Blanc, 1961:

I .

Pubes cernum Gunther, 1859: 169. Angot, 1951: 15.

As noted above, Sivertsen listed two species of fish known to the Tristan
islanders by common names that he was unable to identify due to lack of
specimens. The fish called by the islanders the steambrass was said to be a
very large fish, nearly as large as a shark (+ 2 metres) ‘but much broader and
with soft rays in the fin’. Rowan & Rowan (1955) record an 8-foot long speci-
men of Polyprion americanus from Tristan and state that it is the species the
islanders refer to as the steambrass.

Specimens were received from the Vema Seamount. Two examples from
the Seamount of 465 and 602 mm standard length were found to have the
length 2#— 83 times depth and 22— 2% of head length. In both the pectoral fin
was slightly greater than half the head, while the longest dorsal spine went
24— 2% into head. The species occurring at Vema is thus clearly P. americanus
rather than the related P. prognathus.

Ephinephelus aeneus (Geoffroy St. Hilaire)

Serranus aeneus Geoffroy St. Hilaire, 1809: 317. Fowler, 1936: 756.
Epinephelus aeneus (Geoffroy St. Hilaire) Norman, 1935): 9.

One specimen (SAM 24299) presented by Messrs. Friedman & Rabino-
witz, Ltd., one of several landed by commercial fishing vessels from the Vema
Seamount.

Body proportions as thousandths of the standard length of 465 mm
are as follows: head 402, depth 331, length pectoral fin 219, height longest
(fifth) dorsal spine 123, eye 62, interorbital 64, snout 114, pelvis 172.

The cheeks are scaled with cycloid scales, as is body above lateral line;
scales ctenoid below lateral line. Caudal slightly rounded. Bottom opercular
spine posterior to top spine. Pre-operculum serrated, angle produced with
very long serrations, operculum acutely pointed.

Mandible with two rows of teeth.

Colour in formalin: Dark brown above shading to lighter below. Sparse
scattered very dark brown spots. Unpaired fins dark edged, paired fins light
grey-brown. Three dark stripes on face; one from posterior of eye to lowest
opercular spine, one from below eye to edge of operculum opposite base of
pectoral fin and the third above maxilla to pre-operculum below the angle.

Carangidae
Seriola lalandi Cuvier & Valenciennes
Seriola lalandi Cuvier & Valenciennes, 1833: 208. Barnard, 1925: 555. Smith, 1949@: 221.

Rowan & Rowan, 1955: 129.
Seriola pappei (Castelnau) Smith, 1959: 256.

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 533

One specimen was donated by the Division of Sea Fisheries, caught at
the Vema Seamount. It appears to be a very common fish there, as one com-
mercial fishing vessel returned with over a thousand large specimens.

It has also been recorded from Tristan da Cunha (Rowan & Rowan, 1955).

Decapterus longimanus Norman
Decapterus longimanus Norman, 19354: 255.

Two specimens from the Vema Seamount (SAM 24294) presented by
the Division of Sea Fisheries. The species was formerly considered endemic
to Tristan da Cunha.

The two specimens are the smallest yet recorded being 215 and 225 mm
in standard length. Body proportions of the 215 mm specimen in thousandths
of standard length are: head 288, depth 204, pectoral fin 260, base of soft
dorsal 400, base of anal 335, gillrakers 44, scutes 42.

Measurements could not be made on the other specimen as it was rather
damaged, having been obtained from the gut of a larger fish.

Emmelichthyidae

Plagiogeneion rubiginosus (Hutton)

Therapon rubiginosus (Hutton) 1876.
Plagiogeneion rubiginosus (Hutton) McCulloch, 1914. Barnard, 1927. Smith, 19494.

A single specimen was donated by Messrs. Friedman & Rabinowitz
(Pty.) Ltd. The fish was about 330 mm in standard length and was caught by
handline on the Vema Seamount. Unfortunately the fish had been cleaned
and somewhat distorted in freezing so that accurate measurements were not
possible. It could readily be distinguished from the related P. macrolepis by its
lateral line count of 68 as opposed to about 50 in macrolepis.

P. rubiginosus is recorded from Australasia and St. Paul Island, is not
uncommon off the east coast of South Africa (Smith, 1949) and a single
specimen has been recorded off Table Bay (Barnard, 1927).

Pentacerotidae

Pentaceros richardsoni Smith

PenNceros richardson A. Smith, 1844 (Plate XXI and 2 pages unnumbered text). Follett &
Dempster, 1963: 315. Smith, 1964: 572.

Pentaceros knert Steindachner, 1866: 208.

Pseudopentaceros richardsoni (Smith) Barnard, 1927: 621. Smith, 1949a: 242.

Griffinetia nelsonensis, Whitley & Phillips, 1939: 233.

Only a partial bibliography is given here; a more complete one is given
by Follett & Dempster (1963). It should be pointed out, however, that in

Follett & Dempster the references to Andrew Smith’s original description are
incorrect. The plates each with a page or more of unnumbered text were

534 ANNALS OF THE SOUTH AFRICAN MUSEUM

published separately between 1838 and 1847 but were not dated while only
the index and title page were dated and published in 1849. (See Waterhouse,
1880, and Barnard, 1956.)

Pentaceros richardsont was for long thought to be an extremely rare species of
fish, being known from the type, an adult caught off Cape Point, South Africa,
on a deep handline, and a few juveniles from Australia and New Zealand.
Recently, however, many medium-sized fish have been obtained from the
North Pacific (Follett & Dempster, 1963, give a complete set of published
records) while Smith (1964) has described in detail a second adult fish from
the Cape of Good Hope, and pointed out that the fish described from Cape
Horn by Steindachner as P. kneri is clearly a juvenile of P. richardsoni.

A juvenile of P. richardson obtained at Tristan was presented to the
Museum by Captain Scott. Unfortunately there are no data as to its method or
depth of capture. Measurements and counts of this juvenile are given in Table 4,
and compared with published measurements and with those of two specimens
in the South African Museum; a large adult specimen trawled off Cape Town
in 300 fathoms during March 1961 (SAM 23076) and a slightly smaller speci-
men trawled off Cape Columbine in April 1965 (SAM 24541) both presented
to the Museum by Messrs. Irvin & Johnson, Ltd.

TABLE 4
Standard length. Sia use. Pe UO be 2407 254? 803 4514 358°
Head ‘ : . : : : 5 25 325 319 338 321 319
Depths 7: : é ; , ; PAIS 433 422 425 418 422
Eye 1 AU WEN aote * od 87 79 83 100 gI 76
Interorbital distance ye 2) 1), eet a LS 117 118 135 109 106
Least depth caudal peduncle . . 93 100 102 112 106 109
Dorsal spines . : : , : ee ESTE, 14 14 14 14 14
Dorsal rays. : ; : : . 89 9 9 10 9 10
Amalsspines 1) tae 2 ear oe aan 4 4 4 4 4 4
Anal rays Wan Pre gee eT. a. tS 7 7 7 8 8 7
Pectovally fhe vcigits sc deenk aie tenes 17-18 17 5] 18 rz 18
Gillrakers total , : ; ; 4 OR PR Eee = == —
Lateral line scales . : : ‘ 68-69 71 76 — 86 a=

Body proportions (in thousandths of standard length) and counts of Pentacerosrichardsoni
from various localities.

* Japan (Abe, 1957).

2 North Pacific (Welander et al., 1957).

3 Tristan da Cunha (Reg. No. SAM 21791).

4 Cape of Good Hope (Reg. No. SAM 23076).
5 Cape of Good Hope (Reg. No. SAM 24541).

P. richardsoni is probably found throughout the world in the warm temper-
ate waters, but the published records of its occurrence are confined to a few
scattered localities. At most of these places it has been found more than once.
Both Follett & Dempster (1963) and Smith (1964) suggested reasons for this,
but it seems likely that the known distribution of the species bears no relation

a we!

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 535

to the true distribution but is due mainly to the widely scattered areas of
intensive fishing and the introduction of new fishing techniques in various areas.

Thunnidae
Thunnus spp.

Several species of the genus Thunnus have been landed by commercial
fishing vessels from the Vema Seamount. Unfortunately all specimens have
been brought ashore gutted and without heads or fins. Many small tuna
(about 50—75 lb. whole weight) had bright yellow peduncle keels which is
characteristic of the southern bluefin (T. maccoyi Castelnau) (known from South
Africa and Australia), while others were stated by the fishermen to be yellow-
fins (J. albacares (Bonaterre) ) and bigeyes (7. obesus (Lowe) ). Mr. A. Heydorn
(Division of Sea Fisheries) (fers. comm.) informs me that he saw a yellowfin
while diving on the Vema Seamount.

Until whole examples are obtained nothing definite can be stated con-
cerning the species of tuna congregating about the Seamount.

Stromateidae
Palinurichthys antarcticus (Carmichael)

Perca antarctica Carmichael, 1818: 501.

Seriollella antarctica (Carmichael) Regan, 1913a: 467.

Seriolella antarctica (Carmichael) Sivertsen, 1945: 21.

Palinurichthys porosus (Carmichael) Barnard, 1948: 395. Smith, 19494: 304.
Mupus perciformis (Mitchill, 1818: 244). Smith, 1949): 843.

The Tristan bluefish was first described and figured by Carmichael (1818).
It was not discussed by Regan in his review of the Stromateidae (1902), but
in a later paper Regan (1913) reviewed the fishes described by Carmichael and
suggested, but without giving reasons, that antarctica was ‘rather closely related
to Sauvage’s Seriollella velaini from the Island of St. Paul’.

Sivertsen (1945) placed antarctica in the genus Seriolella, but again no
reasons for doing so were given; nor was the generic definition modified.
According to Regan (1902) the genus Seriolella had inter alia the following
characters: a fin formula of VIII 27-40; anal III 19-24, and a lateral line
concurrent with the dorsal profile. The fin formula of antarctica on the other
hand is dorsal IX 18-20, anal III 14-15, and the lateral line becomes straight
before the caudal peduncle.

The correct genus for antarctica would thus appear to be Palinurichthys
Bleeker 1859.

A specimen of P. antarcticus was donated to the Museum by the Director,
Division of Sea Fisheries, after it had died while on display in the Sea Point
Aquarium. The fish was unquestionably a Tristan bluefish although actually
obtained from Gough Island. When compared with other Palinurichthys material
in the Museum collection it showed a marked resemblance to two similar sized
fish in the collection recorded by Barnard (1948) as P. porosus (Richardson).

536 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 5
24532 19573 19557 23311 } :

Standard length . . . 884 436 453 432 35 34
Head we eee 339 351 356 358 370 380
Depths. WV des SAT 344 335 358 400 380
Length anal fin base. . IgI 222 212 226 230 235
Diameter eveke ts. f-.,us 75 66 75 2 103 103
Length pectoral fin’. . — 266 274 306 260 265
Length pelvic fin hee es 152 160 166 158 245 235
Snout Te LER ve 105 88 95 102 86 88
Wiascillciaals wih menciiiot cc 153 138 150 150 145 147
Pectoral.rays’? 1, te oy. 21 QI 21 20 —_ —

Dorsalispines’)) 3) OP or Bar 8 Se
Donsalmaysys rue) eflioms, 19 19 18 20 19 19
ATV SITES Hk lees: 3 8 3 3 3 +s
Anal rays ope Meat ce 14. 15 14 14 14 14

Body proportions in thousandths of standard length and counts of P. antarcticus from the
Cape and Gough Island and P. porosa from the Kermadac Islands.

SAM 19573 NW. of Cape Town
19557 NW. of Cape Town
23311 Gough Island
24532 Vema Seamount

1 Kermadec Islands. Collection of the British Museum (Natural History).

Body proportions and counts are given (Table 5) for the Gough Island
fish, the two Cape fish and a larger specimen caught at the Vema Seamount
by Mr. A. C. Paterson and presented by the Division of Sea Fisheries. As far
as proportions and fin counts are concerned it is clear that there are no signifi-
cant differences between the fish that could be construed as being of a specific
nature. |

In all four examples the pectoral fin is long and falcate, reaching the anal
origin. The preoperculum has a curved scaled patch and the operculum is
completely scaled. There is also a small patch of scales on the head above the
preoperculum, as shown in Barnard (1948, fig. 14). The mouth is large, reaching
the posterior edge of the pupil. The origin of the dorsal fin is posterior to the
pectoral fin base.

P. porosus is known from the coast of Australia and the Kermadec Islands.
Seven juveniles from the latter locality, kindly lent by the British Museum
(Natural History), were compared with P. antarcticus. Several differences in
body proportions were apparent (Table 5), as well as the shape of the pectoral
fin. Regan (1902), however, mentioned that the stromatid fishes changed
considerably in body shape during growth, especially in that the pelvic fin
becomes shorter and the pectoral fin becomes falcate in shape.

Although there are marked differences in the size of the fishes examined
(over 430 mm as opposed to about 35 mm in total length), the similarity in
pattern of the scales of the head suggest that P. porosus is at least very closely
related to P. antarcticus.

In recent years large numbers of stromatid fishes assigned to the species

\

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 537

P. porosus have been reported from Australian waters (Cowper, 1960) but no
redescription or figure of a specimen larger than the type of 140 mm has been
published.

Two stromatids have been recorded from St. Paul Island, Seriolella
velaint which has a fin formula of dorsal VIII 27 and anal III 20 (Sauvage,
1879), and P. porosus, which was recorded and briefly described by Angot
(1951). His brief description could apply equally to porosus or antarcticus, but
the sketch shows a fish with a long falcate pectoral fin characteristic of antarcticus
rather than the short rounded fin as shown in Richardson’s figures of the type
of porosus.

In the North Atlantic there is another related species, P. perciformis
(Mitchill) which Smith (1949)) recorded from the west coast of South Africa.

This species appears to be generally smaller than antarcticus; it is said to be
dark green in colour (Jordan & Evermann, 1896) whereas antarcticus is blue;
the dorsal spines after the fourth are subequal (Regan, 1902) whereas in
antarcticus and porosus they decrease in size and, finally, the pectoral fin in
perciformis (as figured by Merriman, 1945) is markedly shorter and less falcate
than in antarcticus, even at the largest recorded size (284 mm L’s).

It would appear therefore that there are two closely related species of
Palinurichthys in the Atlantic Ocean, a northern species P. perciformis (Mitchill,
1818) and a southern species P. antarcticus (Carmichael, 1818). P. antarcticus has
been found in South Africa, at Tristan and Gough Islands, the Vema Sea-
mount and also appears to be present at St. Paul Island in the southern Indian
Ocean. If adult specimens of porosus and antarcticus are compared and found to
be conspecific P. porosus (Richardson, 1845) must be relegated to the synonomy
of antarcticus.

Mupus imperialis Gocco

Mupus imperialis Cocco 1833: 20. Smith, 1949): 843.
Mupus ovalis Smith, 19494: 303.
Lirius ovalis Barnard, 1948: 392.

This species seems very common on the Seamount and grows to a large
size, probably at least to 1 metre in length. In view of its rounded head it is
called ‘stumpnose’ or ‘biskop’ by South African fishermen visiting Vema. Two
specimens have been examined. In thousandths of standard lengths of 590 mm
and 762 mm the present specimens have the following respective body pro-
portions: head 289-297, depth at pelvic origin 334-343, max. depth 393-370,
snout 76—79, eye 68-66, pectoral fin 263-244, base of dorsal fin 728. Counts
were dorsal VI-VII 28, —28, anal III 20-21.

M. imperialis has been recorded from wide areas of the Atlantic and the
Mediterranean and appears to grow to a large size. Sivertsen (1945) described
as new a stromatid fish collected at Tristan which he named Seriolella christo-
phersent. From the description I can find no significant characters whereby this
species differs from M. imperialis. Both have similar body proportions, especially

538 ANNALS OF THE SOUTH AFRICAN MUSEUM

the large eye and rounded head, rather small mouth and a body which is
deepest posterior to the pelvic fin origin. The counts for the unique specimen of
christophersent are all within the range for M. imperialis.

Coryphaenidae
Coryphaena hippurus Linnaeus

Coryphaena hippurus Linnaeus, 1758: 261. Fowler, 1936: 649. Gibbs & Collette, 1959: 117.

Two small specimens of this species were collected at the Vema Seamount
by Mr. Paterson of the Division of Sea Fisheries. Both specimens (SAM 24531)
are immature, being only 352 and 375 mm in standard length, but are clearly
hippurus rather than the related equiselis, having total gillraker counts of 9-10
and 62-63 dorsal rays.

Scorpaenidae
Scorpaena scrofa Linnaeus

Scorpaena scrofa Linnaeus, 1758: 266. Smith, 1957: 51
Scorpaena natalensis Regan, 1906: 5. Barnard, 1927: 902. Smith, 19494: 371.

One specimen from a commercial fishing vessel at the Verma Seamount.
Body proportions as thousandths of standard length (313 mm) are as follows:
head 418, depth 313, eye 80, interorbital 73, snout 130, dorsal base 534, anal
base 125, pectoral 265.

Congridae
Ariosoma australis (Barnard)
Congermuraena australis Barnard, 1923: 442. Barnard, 1925: 190. Sivertsen, 1945: 4.

Arissoma balearica (non de la Roche) Smith, 19494: 393 partim.
Congermuraena ?habenata (Richardson, 1845: 109). Sivertsen, 1945: 4.

Barnard (1923) described two species of congrid eels as Congermuraena
australis and C’. albescens, the former from shallow water at the Cape of Good
Hope and Tristan da Cunha and the latter from deep water at the Cape.
Sivertsen (1945) obtained one specimen at Tristan, and, although he assigned
it to C.. australis, he stated that there were minor differences between his specimen
and Barnard’s description, and suggested that the Tristan species may be
C’. habanata Richardson, known from New Zealand and St. Paul Island.

Smith (1949a) reduced both australis and albescens to the synonymy of the
Mediterranean species Ariosoma balearica (de la Roche) but gave no reasons
for doing so.

A careful examination of A. australis from shallow water at the Cape and
Tristan and A. albescens (from deep water off the Cape) has shown the two eels
to have clear differences. Body proportions are given in Table 6. There is little
difference between the fish in so far as the body proportions are concerned
except that the eye in albescens is proportionally smaller and the interorbital
width greater than in australis from the Cape and with the Tristan fish having
an intermediate eye size but a narrow interorbital width.

es ee ee! ee ee ee rl ee Eee

}
Ht
b,

it eS ee *

ae

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 539

In general appearance, however, the fish from Tristan are very similar to
australis, which differs from albescens in the following characters: in australis the
rays in the vertical fins are clearly visible, the fin has a dark edge, the lateral
line has one tiny pore to each myotome, the upper dentition has a rather

Lb

Fig. 1
Upper tooth patterns of a, Ariosoma australis syntype;
b, Ariosoma albescens holotype. (Diagrammatic and
not to scale.)

elongate, slender pattern (fig. 1a), the body is well formed and firm, and I can
find no trace of the epidermal processes stated by Castle (1960) to occur on the
body of A. balearica. In albescens the vertical fins are enclosed in a thick gelatinous
envelope, the fins are pale edged, the lateral line is a distinct ridge and has a

540 ANNALS OF THE SOUTH AFRICAN MUSEUM

large lower pore and a small upper one to each myotome, the pattern of the
upper dentition is shorter and heavier (fig. 1b), there are distinct epidermal
processes present and, finally, the body is soft and flabby. It is unlikely that the
differences are due to changes taking place during growth, as a specimen of
australis at 280 mm was found to be fully mature and to have ripe ovaries
containing large eggs.

Smith’s (19492) combining of australis and albescens is therefore considered
to be incorrect.

A. balearica is known from the tropical Atlantic and the Mediterranean
(Fowler, 1936) and is very similar to A. australis. It, too, has a very indistinct
lateral line which will separate it from A. albescens. There is also very little
difference between australis and A. habenata from New Zealand and St. Paul
Island as pointed out by Sivertsen (1945) and between the New Zealand deep-
water species Pseudoxenomystax hirsutus Castle (1960) and the South African
deepwater species A. albescens. Castle divides the two species by the position of
the origin of the dorsal fin, stating that it lies over the gill slit in hirsutus and at
a level half-way along the pectoral in Smith’s specimens of ‘balerica’. In the
" type of albescens (designated in the museum catalogue as SAM 12775) the
origin of the dorsal is above the pectoral origin.

TABLE 6

Catalogue No. SAM: 12781 12781 12780 12781 18097 12782 24551 12775 18097 18097

Standard length. ... 206° 2925 238 274 261 186 960s pieeee
Depth - os ee se 590167 6 BU 85547 5G | Oa
Head 2.0: . se). 660 160 J t6Oirspoee 760. 161 | yheiee
Head‘to vent «©. . «398 9) 301 #)406))) 392 69403 976° “AOR eigen
Vent to tail’)... . (606° «| “G089 V5g2. "G07 597° «635 «= 5 eee
Evel Gore Wer aaa. shag 38 38 40 32 34 33 25 26 au
Imtcronoitala: amt a aul 16 13 16 II 16 18 37 26 26
Snout to oyigin dorsal. .. 180 182 \ 389 (972 “(162.0 178 460 eee
Snout to origin anal 2147 Ope 420° 405, 403 — 481 — —

teloiie cullslviv ae a be wal 18 107) 18 18 21 19 25 — —

Body proportions (in thousandths of standard length) of Ariosoma australis from South
Africa and Tristan da Cunha and A. albescens from South Africa.

SAM 12780 A. australis paratype, collected Kalk Bay
12781 A. australis, Kalk Bay
18097 A. australis, Kalk Bay
12782 A. australis, Tristan
24551 A. australis, Tristan
12775 A. albescens, holotype, deep water off Cape Town
23189 A. albescens, deep water off Cape Town

The systematics of this group of eels is in a very confused state, there being
little agreement even at the generic level, and until such time as a careful
revision of the family is made on a world-wide basis it will be impossible to
decide whether A. habenata, A. australis from the Cape, A. australis from Tristan
and several other species are in fact distinct species or are only geographical |
variants. It is considered advisable therefore that the existing species be retained

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 54.1

in the meantime. It would be surprising, however, if the Tristan and St. Paul
Island eels are specifically different.

Lagocephalidae
Sphaeroides cutaneus (Gunther)

Tetrodon cutaneus Gunther, 1870: 287. Barnard, 1927: 971.
Sphaeroides dubius von Bonde, 1923: 40.
Sphaeroides cutaneus (Gunther) Smith, 19494: 417.

This large species of puffer fish is found throughout the warm, temperate
and tropical seas. It is known from the Cape and St. Helena Island.

Three specimens from Vema were collected and presented by Mr. Paterson
(SAM 24524). All three were a dirty grey-green dorsally with a white belly.
Dorsal, anal and pectoral fins were pale yellow. The pectoral fin has a charac-
teristic shape, being broad, rounded on the lower distal corner, and pointed
at the upper. The species is capable of very extensive inflation and all three
examples examined were much distorted due to this.

ZOOGEOGRAPHY

The relative geographical positions of the islands of Tristan da Cunha,
Gough, St. Paul, Marion and Crozet, the Vema Seamount and South Africa
are shown in figure 2. Tristan, Gough and St. Paul islands lie on very similar
latitudes, whereas the Vema Seamount is roughly as far north of Tristan as the
Marion and Crozet groups are south. The Vema Seamount, however, has
several fish in common with Tristan whereas the Marion and Crozet groups
have no fish in common with Tristan or St. Paul. This, as was suggested by
Sivertsen (1945), is almost certainly due to hydrographic conditions, mainly
water temperatures.

e Mauritius
* Reunion

Vem e

Port Elizabeth

Amsterdam

4 Tristan
t. Paul»

+ Gough

E Prince s.. Crozet
ae .
Marion e award

Fig, 2
Shallow-water areas in the southern Atlantic and Indian Oceans.

542 ANNALS OF THE SOUTH AFRICAN MUSEUM

There are two areas of sudden temperature change in the Southern Ocean,
the Sub-Tropical Convergence, where the surface temperature rapidly changes
from about 13°C to about 9°C and a more southerly line, the Antarctic Con-
vergence, where the surface temperature again drops, from 6°C to about 2°C.
The Antarctic Convergence is quite narrow but the Sub-Tropical Convergence
is much broader and Sverdrup ¢é¢ al. (1942) refer to it as a Region of Conver-
gence. The Sub-Tropical Convergence fluctuates in latitude, at times the whole
zone is 5-6° South of the Island and at other times Deacon (1937) suggests
that it may be north of Tristan. Temperatures taken at Gough Island at a time
when he suggested this do not support it, however.

Gough Island is approximately 3° 20’ south of Tristan, thus it may lie
north or south of the Convergence according to the conditions prevailing in
the ocean. Both, however, probably normally le north of or within the Con-
vergence zone, while Vema always lies well to the north and the Marion and
Crozet group well to the south.

There is little published data available giving water temperatures at
Tristan and even less for Vema or Gough. Christophersen & Schon (1942)
published detailed sea surface temperature records taken during the Nor-
wegian expedition to Tristan. The main features were:

Temperatures in °C

Month Average Lowest Highest
December 1937 are 15°7 15:0 16-4
January ogo). | tao es ae Oey 14°8 17°9
Hebruary 1Q¢on. 07... 1776 19°8
March 1938 Pe LE PETG 18-0 20°0

The above are all surface temperatures taken from the shore. Two stations
worked by R.V. Discovery gave surface temperatures of 14°59°C (Station 4) in
January and 12-95°C (Station 397) in May. Station 4 was taken close inshore
where the bottom was only 37 metres. A subsurface sample at 35 m gave a
temperature of 13°64°C.

Gough Island too appears to have a very deep thermocline, stations worked
by the R.V. William Scoresby in June 1927 showed the following:

Temperatures in °C’ at Sample Depth

Station Date om 100 m 150 m
W/s 122 7-8/6/27 11°59 11°69 9°89
125 9/6/27 11°63 11°49 10°99
126 10/6/27 11°79 11°70 9°69
130 12/6/27 12°09 12°09 10°71

In April 1965, the supply ship R.S.A. recorded surface temperatures at
Gough Island ranging from 11°8°C to 137°C over two days (Crawford, pers.
comm.).

The main fishing areas at Tristan and Gough are in shallow (under 50
metres) water. At this depth it would seem that there is normally but little
difference between surface and ambient temperature of the bottom living
species.

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 543

The surface temperatures at Vema are significantly higher, the Emerson K
found surface temperatures of between 18-3°C and 18-8°C in November 1964
(Simpson & Heydorn, 1965) and in April 1965 the R.V. Africana IT obtained
the following temperatures on and about the Seamount:

Sounding Sampling depth Temperature

Station (metres ) (metres ) "6
A.3617 go O 20°44
50 20°13
80 18°50
A.3620 goo O 20°37
50 20°35
100 16°15
A.3624 480 fo) 20°40
50 20°38
100 16-63
A.3625 73 fo) 20:36
59 20°33
A.3626 goo O 20°35
50 20:28
100 15°51!

There is a more rapid decrease in water temperature in the first hundred
metres at Vema than at the two southern islands, but the ambient temperature
of the bottom living species (main fishing areas are 70-80 metres deep) is still
several degrees higher than at Tristan or Gough.

It is probable that this bottom temperature is close to the maximum for
the fish at Vema which are more normally associated with cooler water
(Acantholatris spp. and Palinurichthys antarctica) and yet still suitable for sub-
tropical species (Sertola lalandiit and Epinephalus aeneus).

The surface water temperature at Vema throughout the year, and at
Tristan occasionally in the summer is suitable for tropical and subtropical
surface pelagic species (Coryphaena hippurus, Sphaeroides cutaneus and Exocoetidae) .
Cool water surface species, however, such as Thyrsites atun, common at Tristan,
appear to avoid the warm water and have not been found at Vema.

TABLE 7
South
Tristan Gough Vema Africa St. Paul
(a) Coast fishes

Gaidropsarus insularum. . + 4 qe
Acantholatris monodactylus — +- ae
Acantholatris vemae .
NG + aS
Bovichthys diacanthus
Decapterus longimanus . .
Labrichthys ornatus . . .
Helicolenus tristanensis .
Sebastichthys capensis
Scorpaena scrofa
Epinephalus aeneus 4
Caesioperca coatsi. . . . +
Ariosoma australis. . 2°. + +

tS ee
fe
+ Ser
-+-

++

544 ANNALS OF THE SOUTH AFRICAN MUSEUM

South
Tristan Gough Vema Africa St. Paul
(6b) Oceanic fishes

Prionace glauca + +
Alepisaurus ferox . +

Maurolicus muelleri . a +
Myctophum humboldti . -- +
Scomberesox saurus + -.

Exocoetus exsiliens . -|-

Cypstlurus lineatus ae a5

Beryx decadactylus . + 4.

Sertola lalandi . . o+ a +
Plagiogeneton rubiginosus . + +

Pentaceros richardsoni ao +

Polyprion americanus + + +

Notopogon ?macrosolen . 4. =

Thunnus spp. - +

Thyrsites atun . =5 a8 +.
Mupus imperialis . + + ae
Palinurichthys antarcticus . + + + =e "y
Coryphaena hippurus . . . a aa
Sphaeroides cutaneus . . . > —

Distribution of fishes, known from the Islands of Tristan da Cunha and Gough and the
Vema Seamount.

Sivertsen felt that there was a large number of fish endemic to Tristan
(33% of the known fish) and a greater similarity with St. Paul Island than
South Africa.

Table 7, which lists all the fish known from Tristan, Vema and Gough, is
divided into oceanic and coastal species in much the same manner as arranged
by Sivertsen. It is never easy, however, to distinguish between oceanic and
coastal species and some, considered to be coastal by Sivertsen, are here listed
as oceanic. Thirteen species are considered coastal, of these eight are found at
Tristan, seven at Gough and six at Vema. Three of these species also occur
definitely at St. Paul Island and four in South Africa. None of the species
definitely occur at all five localities and only one at Tristan, Gough and Vema.

It is not agreed with Sivertsen that Tristan has a large proportion of
endemic fish, in fact it is a surprising conclusion that this isolated island has
no endemic fish species. Vema, however, appears to have one, as does Gough.
It is also surprising that such an apparently successful species as A.
monodactylus has reached Vema but not South Africa.

Almost all the oceanic species listed in Table 7 are known from Tristan,
only the tropical species being absent. All species recorded from Tristan are
also known from South Africa, a few from Vema and almost none from Gough.
This pattern is almost certainly due more to the varying intensities to which
the various areas have been fished than to any real differences.

Some of the species recorded from Tristan are deep-living bathypelagic
species that have either been caught on the surface at night or have come into
shallow water due to the abrupt elevation of the ocean floor. Such species are
Maurolicus muelleri, Myctophum humboldti, Alepisaurus ferox, and Pentaceros rich-

THE FISHES OF TRISTAN, GOUGH AND THE VEMA SEAMOUNT 545

ardsoni. All are species which have been found in widely scattered parts of the
world.

Other fish again are surface-living pelagic forms not normally found close
to land but presumably attracted by the rich food available at the edges of
the oceanic islands and in the shallow water over the Vema Seamount. Fish of
this type recorded are Coryphaena hippurus and Thunnus spp. from Vema, and
Scomberesox saurus, Prionace glauca and Exocoetus exsiliens from Tristan.

Another group of pelagic fishes are also wide ranging but normally congre-
gate close to land. Seriola lalandi, Thyrsites atun and Sphaeroides cutaneus are of this
type. The first has been taken at both Vema and Tristan, the second only at
Tristan and the third only at Vema. All three, however, are known from South
Africa and the east coast of America. Palinurichthys antarcticus is possibly a
similar fish but has a deep pelagic habitat rather than occurring close to the
surface. It appears to have a very wide distribution, being known from all three
areas discussed, South Africa, probably St. Paul Island and possibly Australia
and New Zealand.

Some of the species which have a wide geographic distribution are small,
weakly swimming, species, e.g. Labrichthys ornatus which is known from Tristan
and St. Paul islands, and probably also Gough (Holdgate 1948) and Vema
(Heydorn, fers. comm., underwater sighting). Other widely distributed, but
weakly swimming species are Ariosoma australis which occurs in South Africa
and Tristan, and possibly St. Paul as well and Gazdropsarus insularum known
from South Africa, Tristan and St. Paul. Both the latter two species have
pelagic larval stages.

Rowan & Rowan (1955) mention large fish seen jumping in the vicinity
of Tristan. At times, in summer, the surface water temperature at Tristan
reaches a temperature quite suitable for Tuna, Blue and White Marlin and
Broadbill Swordfish. Jsurus glaucus has been recorded from St. Paul (Blanc,
1961).

It is interesting that no elasmobranchs other than Prionace glauca have been
recorded from the areas discussed, although ‘spiny dog fish’ Squalus fernandinus,
are common at St. Paul (Blanc & Paulian, 1957).

None of the gadids, thought by Rowan & Rowan (1955) to belong to at
least two species, have been obtained. They thought that one belonged to the
genus Haloporphyrus but as they obtained only damaged specimens washed
ashore by storms, identification was not possible.

SUMMARY

The fishes known from the newly discovered Vema Seamount are described
and compared with those found off Tristan da Cunha and Gough Island. A
new species of chilodactylid, Acantholatris vemae, is described from Vema. The
genus Gaidropsarus from Tristan and South Africa is discussed and the South
African material shown to belong to two species, G. insularum, also known from

546 ANNALS OF THE SOUTH AFRICAN MUSEUM

Tristan and St. Paul Island, and G. capensis from the south-east coast of South
Africa.

An attempt is made to relate the fish fauna to what is known of the hydro-
graphic conditions around the islands.

ACKNOWLEDGEMENTS

For the material on which this paper is based I am indebted to the Director
and staff, Division of Sea Fisheries, Sea Point (especially Messrs. Heydorn,
Nepgen and Paterson), Capt. M. T. Scott, Messrs. Friedman & Rabinowitz
(Pty.) Ltd., and Messrs. Irvin & Johnson (Pty.) Ltd., all of Cape Town.
Material was lent by the Trustees of the British Museum (Natural History)
and Professor J. L. B. Smith of Rhodes University, Grahamstown. |

Unpublished data on water temperatures were supplied by Mr. A.
Crawford, Mr. G. Stander and Mr. M. Orren, while Mr. A. Wheeler of the
British Museum helped with much useful information.

Finally I am grateful to my wife, Dr. Mary-Lou Penrith, for checking the
manuscript, and to the Librarian, Council for Scientific and Industrial Research,
who obtained photocopies of several papers not available to me.

The Trustees of the South African Museum are grateful to the Council for
Scientific and Industrial Research for a grant to cover the cost of publishing
this paper.

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Ann. S. Afr. Mus., Vol. XLVIII Plate XXI

A. Acantholatris vemae sp.n. Type of 383 mm. standard length.

B. Acantholatris monodactylus. Specimen of similar size from Vema seamount.

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TaBLeE oF CoNTENTs and Summary. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 4? in. = 7 in. (74 in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

Situ, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945-

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmitH, C. D. 1954. South African Plonias. In Brown, X. Y. Marine faunas. and ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 ( = natalensis West).

INDEX TO GENERA AND SUBGENERA

(SYNONYMS IN ITALICS)

A

Acanthocyclops, 515, 517

Acanthodes, 405

Acantholatris, 524, 525, 528, 543, 545

Aceratherium, 302, 364

Achiropsetta, 181, 184, 186

Acontiostoma, 206

Acryptolaria, 6, 7, 18

Adenota, 296, 297

Admetophoneus, 235

Aelurognathus, 413

Aepyceros, 294, 296, 298, 301

Aequorea, 434, 461, 484

Aethomys, 300

Afrochoerus, 297

Afrocyclops, 516

Afropisidium, 81

Agama, 420

Agastra, 4.72

Aglaophenia, 494, 495

Agyneta, 71

Alcelaphus, 293, 294, 295, 297, 298, 300, 301

Alepisaurus, 524, 544

Aleterix, 299

?Alkmaria, 117

Allorchestes, 208

Alopecognathus, 124, 125, 126, 136, 143,
150, 155, 158, 162

Alopias, 280

Amage, 98, 99, 101, 102, 105, 112, 113

Ampharete, 98, 99, 101, 102, 105, 112, 114

Amphibolurus, 420

Amphicteis, 98, 99, 100, 101, 102, 105, 112,
113

Amphicteis, 113, 114

Amphiprion, 532

Amphisamytha, 101, 114

Amphisbetia, 6, 25, 491, 494, 495

Amphithoe, 208

Amphitrite, 113

Amythas, 98, 112, 117

Amythasides, 118

Anancus, 292, 293, 295, 296, 297, 315, 319

Anchitherium, 324

Aneugomphius, 414, 415

Anniella, 420

Anobothrus, 101, 102, 115

Antennella, 19, 492, 494, 495

Anteosaurus, 4.12

Antidorcas, 294, 298, 300, 301

Aonyx, 297, 299, 362

Aora, 208

Aphaneromma, 408

Archidiskodon, 280, 295, 296, 297, 298, 301,
315, 319

Arctocephalus, 280

Argobuccinum, 459

Ariosoma, 524, 538, 543, 545

Aryandes, 112, 117
Asabellides, 101, 112, 115

Asthenargus, 71

Atelodus, 293, 294
Auchenoplax, 99, 112, 118
Australopithecus, 295, 300
Auxis, 179

Avenantia, 235

Bathyergus, 301
Bathygobius, 190
Beatragus, 297, 363
Beryx, 528, 544
Bicorona, 433, 440, 484
Bidenichthys, 214
Bienotherium, 417
Bigois, 72

Bimeria, 434, 449
Bolosaurus, 410

Bos, 293
Bothrodendron, 268
Bougainvillea, 434, 449
Bovichthys, 524, 525, 543
Branchiosabella, 114
Brucella, 18

Bubalus, 293
Bularchus, 297

Bullia, 461
Bunolistriodon, 302

C

Caesioperca, 525, 543
Calamphora, 39

Calicella, 477

Calotes, 420

Campanularia, 434, 471, 480, 484
Campanularia, 477, 4.78, 481, 483
Campanulina, 463

Campomma, 463

Cangoderces, 62, 63, 65, 73
Canis, 280, 293, 296, 297, 299, 300, 301
Caprella, 209

Capreolus, 294

Captorhinus, 409

Caracal, 2906

INDEX TO GENERA AND SUBGENERA

Carcharias, 280

Carcharinus, 268

Caulerpa, 211, 214

Cephalophus, 293, 301, 302, 362

Ceramius, 219

Ceratella, 44.4, 446

Ceratotherium, 279, 280, 295, 296, 297, 300,
301

Cerceris, 219

Cercopithecoides, 299, 300

Chaetodon, 529

Chalicotherium, 319

Chara, 92

Cheilodactylus, 529

Cheiracanthus, 405

Chilifer, 57

Chilodactylus, 529

Chlamydosaurus, 420

Chrysemys, 419

Chrysoblephus, 524

Chrysotricha, 300

Chthoniella, 74

Cichla, 531

Cistecephalus, 402, 414

Clava, 434, 452

Clibanarius, 459

Climatius, 405

Clinus, 215

Clytia, 434, 477, 484

Clytia, 472, 476

Cnidonema, 444

Cnidoscyphus, 52, 54

Condylus, 404

Congermuraena, 538

Connochaetes, 298, 300, 301, 302

Cordylus, 419

_ Coryne, 444.

Coryphaena, 538, 543, 544, 545

Coryphopterus, 192

Crassatella, 453

Crateritheca, 6, 26

Creodonta, 302

Crocidura, 298

Crocodilus, 294, 295

Crocuta, 295, 296, 297, 299, 300, 301

Crossarchus, 299

Crossostoma, 113

Cryptolaria, 18

Cryptolaria, 7, 9

Cryptomys, 299, 300

Ctenogobius, 189

Cyclops, 516

Cycloptychius, 272

Cyclostigma, 268

Cynariognathus, 124, 144, 151, 155, 158, 164

Cynariops, 413

Cynictis, 300

Cynognathus, 400

Cypsilurus, 526, 544.

D

Damalavus, 293, 302, 362
Damaliscus, 280, 296, 297, 298, 300, 302
Daptocephalus, 412
Dasymys, 299

Dasyurus, 420

Decapterus, 524, 533, 543
Decennatherium, 363
Dehitella, 445
Deinotherium, 294, 295, 296, 297, 302, 315
Dendromus, 300
Desmodillus, 299

Deto, 196

Deuterosaurus, 233
Diadectes, 400, 409
Diademodon, 400, 415
Diaptomus, 520
Diarthrognathus, 416, 417
Dicerorhinus, 293, 362
Diceros, 295, 296, 297, 298, 300, 301
Dicoryne, 434, 451
Dictyocladium, 6, 28
Dicyclocoryne, 443
Dicynodon, 400, 412
Didelphis, 400, 420, 423
Didymophyllum, 268
Dimetrodon, 400, 411
Dinopithecus, 294, 363
Diogenes, 459

Diphasia, 6, 28
Doliosauriscus, 235
Dorcatherium, 302
Dynamena, 6, 29, 491, 494
Dynamena, 26, 27
Dynamenella, 196, 197

E

Eccasaurus, 128

Echidna, 400, 420

?Eclysippe, 118

Edops, 407

Elephantulus, 298, 299, 300

Elephas, 292, 293, 294, 315, 319

Eleutheria, 444.

Emys, 419

Endothiodon, 126

Eogyrinus, 407

Epinephelus, 532, 543

Equus, 279, 280, 286, 293, 294, 295, 296,
297, 298, 299, 300, 301, 316, 322, 323,
328, 331, 334, 343, 344, 346, 379, 381,
388

Equus, 389

Eremias, 419

Euchambersia, 4.14.

Eucheilota, 463

Eucyclops, 515, 516

INDEX TO GENERA AND SUBGENERA

Eucyclops, 517
Eudendrium, 427, 434, 454
Eudendrium, 451
Eudiaptomus, 516, 520
Eupagurus, 459
Eurychora, 57
Eurygnathohippus, 298, 349, 350, 389
Eurystheus, 208

Eusamytha, 118
Eusthenoptheron, 406
Euthecodon, 294
Euthynnus, 179

Exocoetus, 524, 544, 545
Exosphaeroma, 197

F

Felis, 420, 423
Felix, 299
Filellum, 6, 10
Flabellipecten, 287
Fusus, 459

G

Gaidropsarus, 525, 526, 543, 545

Gazella, 293, 294, 296, 297, 298, 300, 301,
302, 362

Georychus, 301

Giraffa, 293, 294, 295, 296, 297, 298, 301, 315

Glanosuchus, 122, 413

Glossogobius, 268

Glyphanostomum, 112, 118

Gnatho, 224

Gobius, 190

Gomphotherium, 297

Gorgon, 297

Gracilaria, 444

Grammomys, 299, 300

Griffinetta, 533

Grubianella, 112, 113

Gynutoclinus, 211, 215

Gypsorhychus, 300

H

Haemilla, 58, 73, 74
Halecium, 434, 461, 464, 489, 494
Haltacris, 203

Halicore, 423

Halonia, 268
Haloporphyrus, 545
Halopteris, 493, 495
Harpethrix, 74
Hastimima, 268

Hebella, 6, 10, 55
Helicolenus, 524, 525, 543
Helladotherium, 314
Hemitragus, 294
Herpestes, 299, 301
Heterobranchus, 116

Hincksella, 6, 22

Hinia, 461

Hipparion, 273, 293, 294, 302, 316, 320, 321

Hippopotamus, 280, 293, 294, 295, 296, 297,
298, 301, 317, 319, 362, 363

Hipposaurus, 237

Hippotigris, 342, 389

Hippotragus, 294, 296, 297, 298, 302

Homo, 296, 297, 301, 319, 423

Homoioceras, 279, 280, 296, 297, 298, 301

Homotherium, 294, 295

Hyaena, 280, 293, 295, 299, 300, 301, 302,
362

Hyale, 207

Hydractinia, 434, 457

Hydractinia, 461

Hydrocorella, 434, 458

Hydrodendron, 490, 495

Hypania, 101, 112, 113

Hypaniola, 114

Hypsohipparion, 289, 332, 333, 334 337, 345:
346, 369, 377, 379, 387, 389, 392, 393

Hystrix, 293, 296, 299, 300, 301

I

ais, 196, 199
Ictidosuchops, 414

Irana, 103, 106, 107
Ischnacanthus, 405
Ischyrocerus, 208

Isocladus, 196, 197, 200
Isolda, 98, 99, 100, 103, 106
Isurus, 280, 545

J
Jaeropsis, 196, 200, 209
Jasus, 489
Jonkeria, 412

K

Kannemeyeria, 412
Katsuwonus, 179
Keratocephalus, 252
Kingoria, 412

Knorria, 268

Kobus, 294, 296, 298, 362
Kollasia, 408

L

Labidosaurus, 409
Labrichthys, 524, 543, 545
Lacerta, 419

Lafoea, 6, 9, 13, 18, 491, 494
Lamna, 280

Laomedea, 4.78, 480

Larifuga, 74’

Latris, 531, 543

INDEX TO GENERA AND SUBGENERA

Leggada, 299, 300

Lepas, 480

Lephthyphantes, 70, 73, 74

Lepidodendron, 268

Lepidopsetta, 181, 186, 187

Lepidostrobus, 268

Leptailurus, 299, 301

Leptocyclops, 517

Leptotrachelus, 413

Lepus, 299, 301, 423

Leuckartiara, 434, 449, 461

Libyhipparion, 287, 340, 341, 342, 348, 350,
389

Libypithecus, 294

Libytherium, 280, 293, 294, 295, 296, 297,
298, 301, 315, 340

Limnosceles, 409

Linyphia, 71

Lirius, 537

Lovenella, 434, 464

Lovenula, 516, 518

Loxodonta, 298, 300, 301, 315

Loxosceles, 61, 73, 74

Lunatoceras, 302

Lutra, 294

Lycaenodon, 413

Lycaon, 301

Lynx, 301

Lyriocephalus, 420

Lyrocephalus, 408

Lysippe, 105, 112, 114

Lysippides, 101, 112, 114

Lystrosaurus, 412

Lytocarpus, 10, 472, 494, 495

M

Mabuia, 419

Macaca, 293, 362
Machairodus, 300
Macrocyclops, 515, 516
Makaira, 167

Makapania, 300, 301
Malacothrix, 299, 300
Mancopsetta, 181

Manis, 301

Mastodon, 286, 293, 294, 302, 315, 319, 364
Mastomys, 299, 300
Maurolicus, 524, 544
Medusa, 478

Megalichthyes, 407
Megalops, 268
Megalotragus, 298
Megantereon, 299, 300, 301
Melinna, 99, 100, 101, 102, 103, 106
Melinnampharete, 112, 117
Melinnexis, 103, 107
Melinnides, 107

Melinnoides, 99, 118 |
Melinnopsides, 106, 107

Melinnopsis, 100, 106, 107

Melinnopsis, 107

Melita, 206, 209

Mellivora, 301

Menelikia, 294

Mentzichthys, 267, 270

Merona, 434, 452

mat enc 331, 338, 359, 360, 384, 386,
3

Merycopotamus, 286, 294, 314

Mesacanthus, 405

Mesochoerus, 295, 297, 298, 301

Mesocyclops, 515, 517

Mesocyclops, 517

Metadiaptomus, 516, 520

Metaschizotherium, 295, 296, 297, 300, 319

Metridiochoerus, 294, 295, 297

Microcyclops, 515, 517

Microsamytha, 101, 112, 117

Mnemeiosaurus, 235

Monishia, 190

Monocoryne, 433, 435, 484

Monostaechas, 19, 492

Morganucodon, 418

Moschosaurus, 265

Motella, 526, 527

Moyanus, 106

Mugga, 102, 112, 118

Munna, 196, 203

Mupus, 537

Mupus, 535; 544

Mus, 420

Mustela, 420

Myctophum, 524, 544

Myorycteropus, 302

Myosorex, 298, 299, 300

Myotis, 299

Myriothela, 433, 437, 484

Mystromys, 299, 300

N

Nannippus, 331

Nassa, 459, 461

Nemertesia, 19

Neohipparion, 330, 337

Neosabellides, 100, 105, 112, 117
Nesotragus, 362

Nitella, 92

Notochoerus, 294, 295, 296, 297, 298, 301
Notohipparion, 292, 330, 333, 347, 348, 389
Notopogon, 524, 528, 544

Notosollasia, 415

Nyanzachoerus, 295

O

Obelia, 434, 483, 491, 494
Oeorpata, 106, 107
Okapia, 296, 297

INDEX TO GENERA AND SUBGENERA

Oligokyphus, 417
Omochoerus, 293, 294, 363
Ophioceps, 420

Ophiodes, 491

Ophiodes, 4.90

Ophiodissa, 490, 491
Orchestia, 207

Oreotragus, 280, 301
Orthopyxis, 472, 476
Orthostonyx, 297, 298
Orycteropus, 293, 296, 302
Oryx, 293, 294, 296, 297, 301, 363
Osteolepis, 4.06

Otomys, 301

Oudenodon, 400

lig
Pabits, 98, 112, 118
Pagurus, 459
Paiwa, 114

Palaeogyrinus, 407

Palaeoloxodon, 280, 298, 315, 316, 317

Palaeotomys, 299, 300
Palaeotragus, 293, 302, 362
Palinurichthys, 535, 543, 544, 545
Panthera, 296, 297, 299, 300, 301
Papio, 299, 300
Paracyclops, 515, 517
Paradiaptomus, 516, 518
Paradiaptomus, 520
Parahippus, 331

Paramage, 100, 112, 116
Paramphicteis, 113
Paramoera, 206
Paranthropus, 297, 299
Paranthura, 197

Parapapio, 299, 300
Parascyphus, 491, 494, 495
Parawaldeckia, 205
Parawrightia, 452
Parhypania, 101, 112, 113
Paridotea, 196, 198, 209
Parotomys, 301

Pasythea, 31

Pavoclinus, 211, 212, 217
Pedetes, 296, 299

Pelea, 296, 302

Pelomys, 300

Pelorovis, 297

Pentaceros, 533, 544

Perca, 535

Perigonimus, 449

Petraites, 214
Phacochoerus, 294, 297, 298
Phanotea, 71, 73, 74
Phenacotragus, 297, 300, 301
Phialella, 464

Phialidium, 4.77, 478

Philanthus, 227

Pholidosteus, 406

Phrynocephalus, 420

Phthinosuchus, 235

Phylactotheca, 490

Phylica, 205

Phyllamphicteis, 100, 112, 114

Phyllocomus, 99, 100, 101, 102, 103, 112, 115

Physignatus, 420

Phyxelia, 73

Pisidium, 77

Plagiogeneion, 533, 544

Platystega, 408

Pliohippus, 322, 331, 359, 361

Plumularia, 493, 495

Podocoryne, 434, 461

Polyprion, 524, 532, 544

Pontogeneia, 206

Porcellio, 196, 205

Porrhomma, 71

Potamochoeroides, 300, 301

Potamochoerops, 299

Potamochoerus, 296, 297

Prionace, 524, 544, 545

Pristerodon, 399, 401, 402, 412, 413, 424

Pristerognathoides, 124, 125, 138, 144, 152,
155, 160

Pristerognathus, 122, 123, 124, 125, 126, 138,
145, 158, 160

Pristis, 268

Proamblysomus, 298, 299

Procavia, 299, 300

Procolophon, 400, 410

Pronolagus, 300

Propalaeochoerus, 302

Protopterus, 294.

Psammogobius, 192

Pseudopentaceros, 533

Pseudosabellides, 115

Pseudoxenomystax, 540

Psilophyton, 268

Pterampharete, 100, 101, 116

Pterolysippe, 114.

Ptomalestes, 124, 14.5, 153

Pultiphagonides, 297

Pycnotheca, 494, 495

R

Raja, 497

Raphicerus, 300, 301

Redunca, 280, 293, 294, 296, 301
Rhabdomys, 299, 300
Rhadinichthys, 272

Rhinoceros, 314

Rhinolophus, 299

Rhinoptera, 280

Rhizorhagium, 434, 452
?Rytocephalus, 113

INDEX TO GENERA AND SUBGENERA

S Stegolophodon, 277, 280, 315, 319
Stenothoe, 206
?Sabella, 116 Stereotheca, 26, 27
Sabellides, 100, 101, 102, 103, 112, 116 Sternothaerus, 294
Sabellides, 106, 115, 116, 117 Stigmaria, 268
Salacia, 6, 31, 55, 492, 495 Stipidium, 359
Samotherium, 293, 302, 362 Strepsiceros, 294, 296, 297, 298, 301, 302
Samytha, 112, 116, 117 Struthio, 280, 293
Samytha, 118 Struthiocephalus, 251, 412
Samythella, 112, 118 ; Stylochoerus, 298
Samythopsis, 112, 116 Stylohipparion, 277, 286, 292, 293, 294, 295,
Sarda, 179 296, 297, 298, 299, 300, 315, 316, 333,
Sarsia, 433, 444 334, 336, 337, 338, 340, 341, 342, 343,
Scalopocynodon, 415 346, 347, 348, 350, 361, 387, 389, 390,
Scandia, 6, 15 393
Schistocomus, 100, 101, 115 Styloniscus, 196, 203
Scomber, 524 | Suncus, 298, 299, 300
Scomberesox, 524, 544, 545 Suricata, 299
Scomberomorus, 179 Sylvicapra, 298
Scorpaena, 538, 543 Symplectoscyphus, 7, 51
Scotussa, 73 Syncerus, 294, 296, 298, 300, 301
Scylacosaurus, 400, 413 Syncoryne, 443
Scymnognathus, 413 Synodontis, 294
Scymnosaurus, 123, 124, 125, 126, 139, 146, Synthecium, 6, 24
153, 158, 160, 164 Synthecium, 22
Sebastes, 525 Syodon, 235
Sebastichthys, 524, 525, 543
Selaginites, 268 Fis
Serengeticeros, 296, 297
Seriola, 524, 529, 532, 543, 544, 545 Tapinocephalus, 121, 237, 413
Seriolella, 524, 537 Tapinochoerus, 297, 298, 300, 301
Seriolella, 535 Tarsius, 423
Serranus, 532 Tatera, 299, 300
Sertularella, 6, 7> 345 55, 492, 494 Taurotragus, 294, 2096, 297; 298, 300, 301,
Sertularella, 22, 51 317
Sertularia, 7, 49, 492, 494. Teleoceras, 302
Sertularia, 25, 26, 29, 37, 477, 478 Terebellides, 103
Seymoria, 408 Testudo, 280
Simopithecus, 296, 297, 300, 301, 321 Tetrodon, 541
Singularia, 268 Teutana, 68, 73, 74
Sivachoerus, 294 Thallomys, 299
Sivatherium, 279, 280, 294, 295, 298, 301 Thaumantias, 480
Smeringopus, 67, 73 Thaumantias, 478
Socarnoides, 205 Thecocarpus, 10
Solanderia, 434, 444 Therailurus, 299, 300
Sosane, 101, 112, 115 Therapon, 533
Sosanopsis, 101, 102, 112, 115 Theridion, 69, 73, 74
Sparus, 524 Theriodesmus, 126
Speleoderces, 62, 73, 74 Theriognathus, 415
Speleomontia, 74 Therioides, 124, 142, 148, 155
Speleosiro, 74 Thermocyclops, 515, 517, 518, 520
Spermophora, 65, 73, 74 Thrinaxodon, 415
Sphaeroides, 541, 543, 544, 545 Thryonomys, 296
Sphenodon, 399, 403, 404, 419 Thuiaria, 25
Spirophyton, 268 Thunnus, 179, 535, 544, 545
Squalus, 545 Thyroscyphus, 7, 52, 474, 476
Staurocladia, 433, 444 Thyroscyphus, 39, 491
Steatomys, 299, 300 Thyrsites, 524, 543, 544, 545

Stegodon, 280, 295, 315, 319 Titanophoneus, 235

INDEX TO GENERA AND SUBGENERA

Tragelaphus, 294, 296, 297, 300, 301, 302,

317, 363
Tragocerus, 293, 294, 302
Trichoniscus, 203
Trichosurus, 4.20
Triconodon, 418
Tridentata, 50
Trigonephrus, 280
Trionyx, 294
Trioracodon, 418
Trirachodon, 415
Trochosaurus, 4.13
Troglohyphantes, 71
Trogonophis, 420
Tropiocolotes, 402, 404, 419
Tropocyclops, 515, 517
Tropodiaptomus, 516, 520
Tubularia, 427, 433, 434, 441
Tupinambis, 419
Turbo, 459
Turicius, 293

U

Ulemosaurus, 235, 236

V
Varanus, 419
Verticillina, 477
Vulpes, 299

xX
Xenohystrix, 300
Xiphias, 178

Z

Zinjanthropus, 319

Zinnosaurus, 121, 122, 125, 128, 130, 143,
148, 161, 165

Zonurus, 4.04

Zygophylax, 6, 15, 55

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