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ANNALS OF THE
SOUTH AFRICAN MUSEUM
VOLUME XLVI
With the Compliments of
The Director
IMPERIAL WaR MUSEUM,
LAMBETH RoOaD,
LONDON, S.E.1.
ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XLVI
PRINTED FOR THE
TRUSTEES OF THE SOUTH AFRICAN MUSEUM
1961-1963
@ PRINTED IN SOUTH AFRICA BY
THE RUSTIGA PRESS, PTY., LTD., WYNBERG, CAPE
Fis! OF CON TEN ES
BARNARD, K. H.
Deep sea Mollusca from west of Cape Point, South Africa (published April 1963)
Boonstra, L. D.
The dentition of the titanosuchian dinocephalians (published January 1962)
Crompton, A. W.
On the dentition and tooth replacement in two bauriamorph reptiles (published
February 1962)
Crompton, A. W.
Tooth replacement in the cynodont Thrinaxodon (published September 1963)
Day, J. HH.
Polychaete fauna of South Africa: part 7. Species from depths between 1,000 and
3,330 metres west of Cape Town (published March 1963)
FREEDMAN, L.
New cercopithecoid fossils, including a new species, from Taung, Cape Province,
South Africa (published April 1961)
GRINDLEY, J. R.
The Pseudodiaptomidae (Copepoda: Calanoida) of Southern African waters,
including a new species, Pseudodiaptomus chartert (published March, 1963)
Haropine, J. P.
Some South African Cladocera collected by Dr. A. D. Harrison (published April
1961)
Eiesse, A. J.
Supplementary contributions to the revision of the Bombyliidae (Diptera) of
Southern Africa: the genus Systropus (published March 1963)
Mitiar, R. H.
Further descriptions of South African ascidians (published January 1962)
Mitxiarp, N. A. H.
The Hydrozoa of the south and west coasts of South Africa. Part I. The Plumulariidae
(published June 1962)
scorr, K. M. F.
Some new caddis flies (Trichoptera) from the western Cape Province—III (pub-
lished April 1961)
Sseorr, K. M. F.
Some Ecnominae from the Transvaal and South West Africa (Trichoptera:
Psychomyidae) (published June 1963)
Secon, i. M.-F.
Some new caddis flies (Trichoptera) from the western Cape Province—IV. Some
Hydroptiliidae (published June 1963)
SmiTH, J. L. B.
Nomenclatorial change for a long-known South African fish (published February
1962)
SOUTHWARD, E. C.
Pogonophora from South Africa (published April 1961)
Stock, J. H.
South African deep sea Pycnogonida, with descriptions of five new species (published
March 1963)
TREWAvAS, E.
A new cichlid fish from the Limpopo Basin (published June 1961)
Unmann, E.
Dactylispa capicola (Péringuey) und Verwandte. (203. Beitrag zur Kenninis der
Hispinae (Coleoptera, Chrysomelidae) ) (published February 1962)
WINTERBOTTOM, J. M.
The South African sukspecies of the buffy pipit, Anthus vcelensis Shelley (published
March 1963)
469
ha
Pes
‘
ERR CIA a 9 Rh CY arma Fg alc
x : Mies Ny Pa PALO
i esting
AOTC RET nek ie
nee
ris Las wae
BALD S92
*
NEW SUB-FAMILY PROPOSED IN THIS VOLUME
Halopterinae Millard, 1962 (Plumulariidae) 268, 272
NEW GENERIC NAMES PROPOSED IN THIS VOLUME
Chetia Trewavas, 1961 (Cichlidae), 53
Corhiza Millard, 1962 (Plumulariidae), 272, 275
Pantopipetta Stock, 1963 (Colossendeidae), 334
INDEX TO GENERA AND SUBGENERA
(SYNONYMS IN ITALICS)
A
Abra, 413, 418, 450
Abyssochrysos, 410, 414, 417, 439
Acladia, 268, 286, 289
Acropterus, 35, 44
Admete, 4.00, 415, 429
Aglaophamus, 354, 362
Aglaophamus, 362
Aglaophenia, 273, 274, 303
Aglaophenia, 269, 312, 313, 315
Agnesia, 174, 218
Agnosaurus, 101
Alloeocarpa, 182, 218
Alona, 35, 44
Amaroucium, 119, 120, 122, 124, 125, 130
Amphitrite, 354, 368
Anabolia, 18
Anoplodactylus, 321, 340
Antenella, 269, 270, 273, 274, 281
Antennopsis, 271, 318
Antennopsis, 275, 281
Antennularia, 272
Antennularia, 299
Anthus, 341
Aphrodita, 354, 357
Aplidium, 119, 139, 217, 218
Archaeosuchus, 69
Arg yrobothrus, 476
Aricidea, 354, 364
Ascidia, 119, 172, 217, 218
Ascidia, 171
Ascorhynchus, 321, 323
Astyris, 434
Athripsodes, 15
Atlanta, 411, 415
Australopithecus, 1, 8, 14
B
Basilissa, 411, 414, 441
Bauria, 231, 232, 233, 241, 249
Bauroides, 232
Belonogaster, 397, 398
Boltenia, 119, 198, 218
Bosmina, 35, 43
Botrylloides, 177, 217
Botrylloides, 175
Botryllus, 119, 175, 218
Botryllus, 177
C
Cadulus, 412, 413, 446
Calliostoma, 411, 416, 442
Calliotropis, 410, 411, 414, 415, 416, 417, 440
Camptocercus, 35, 44
Cancellaria, 409, 417, 429
Captorhinus, 253
Cardiapoda, 411, 414
Cavolinia, 411, 413, 414, 415, 416, 417
Cercopithecoides, 4.
Ceriodaphnia, 35, 42
Cerithiella, 410, 414, 439
Charitodoron, 409, 413, 414, 415, 416, 417,
432
Chauvetia, 433
Chetia, 53
Chrysodomus, 433
Chydorus, 35, 45
Ciona, 171, 218
Cladocarpus, 273, 274, 304
Clathurella, 428
Clathurella, 426
Clavatula, 408, 413, 414, 415, 416, 417, 420
Clavelina, 139, 218
Cnemidocarpa, 119, 187, 218
Colella, 162
Colossendeis, 321, 326
Columbarium, 410, 413, 416, 438
Columella, 434
Conops, 395
Corella, 174, 218
Corhiza, 272, 273, 275, 291, 318
Cricodon, 516
Cryptodon, 450
Ctenicella, 203
Cuspidaria, 413, 415, 418, 450
Cuvieria, 411
Cynthia, 190, 193
Cynthiopsis, 195
Cystodites, 141, 14.7, 217
Cythara, 408, 414, 415, 416, 417, 422
D
Dactylispa, 223
Daphnella, 409, 413, 415, 427
Daphnia, 35, 36
Dentalium, 411, 412, 413, 414, 415, 416, 417,
418, 444
Deuterosaurus, 60
Dextrocarpa, 180, 218
Diacria, 411, 414, 417
Diadectes, 252
Diademodon, 233, 251, 252, 253, 480, 502,
515, 519
Diaphanosoma, 35
Diarthrognathus, 231, 251
Didemnum, 113, 119, 163
INDEX TO GENERA AND SUBGENERA
Dimetrodon, 232, 233
Diplosoma, 167, 171, 218
Distaplia, 149, 218
Distoma, 143, 147, 154, 157, 158, 160
Doridoxa, 411, 416, 4.43
E
Echinisca, 35, 44.
Ecnomus, 454
Epidiopatra, 363
Epizoanthus, 436
Ericiolacerta, 231, 232, 233, 234, 249, 480,
482
Eucoelium, 145
Eudistoma, 113, 119, 139, 154, 155, 217
Eudistoma, 162
Eugyra, 119, 206
Eunoe, 354, 359
Euphione, 354, 355
Euryalona, 35, 44
Eurycercus, 35, 44
Euthelepus, 370
F
Falsilunatia, 410, 415, 439
Fluxina, 441
G
Gamaster, 209
Gastropteron, 411, 416, 417
Gattya, 268, 270, 273, 281, 295
Goodsiria, 178
Guivillea, 408, 409, 415, 416, 417, 429
Gymnobela, 409, 413, 416, 428
Gynandrocarpa, 178, 218
Ja
Halicardia, 413, 450
Halicordia, 450
Halicornaria, 273, 274, 307
Halocynthia, 119, 201
Halopteris, 268, 269, 270, 273, 280, 281, 282,
295, 303
Haplochromis, 53, 55
Harmothoe, 354, 357
Helicopsyche, 23
Heptabrachia, 48
Hermonia, 354
Herse, 411, 417
Heterocalanus, 374
Heteroplon, 267, 268
Heteroplon, 285, 286
Heterotheca, 267, 268
Heterotrema, 128, 130
Hispa, 223, 227
Hyalinoecia, 354, 363
Hydroptila, 475
Ilyocryptus, 35, 43
4)
Jonkeria, 76
K
Kirchenpaueria, 273, 274, 292
Kirchenpaueria, 295
L
Labrus, 258
Lachesis, 433
Laetmonice, 354, 355
Lamellaria, 410, 415, 439
Lamellisabella, 47
Lechia, 412, 414, 446
Leanira, 354, 360
Leda, 412, 414, 416, 417, 448
Leptecho, 15, 23, 32
Leptoclinides, 167, 170, 217
Leptoclinum, 171
Leydigia, 35, 45
Lichnochromis, 54
Limopsis, 413, 414, 415, 416, 417, 418, 449
Lissoclinum, 119, 168
Lumbriclymene, 354, 366
Lumbrinereis, 354, 363
Lutjanus, 258
Lytocarpus, 273, 274, 312
M
Macellicephala, 354, 355
Macroclinum, 135
Macrothrix, 35, 43
Maldanella, 354, 355, 365
Malletia, 412, 416, 417, 418, 447
Mangilia, 409, 413, 415, 416, 427, 428
Melinodon, 232
Metandrocarpa, 119, 183
Metrocarpa, 177
Microcosmus, 196, 217, 218
Microgomphodon, 232
Microhelodon, 232
Mohnia, 433
Moina, 35, 43
Molgula, 119, 203, 211, 218
Moniliopsis, 408, 415, 417, 421
Monoculus, 43
Monospilus, 35, 45
Monostaechas, 269, 270, 272, 273, 278, 280,
2Q1
Monotheca, 301
N
INemertesia, 271,273,274, 202. 207
Neanthes, 354, 360
Neosabellides, 354, 367
Nephthys, 354, 362
Neptunea, 409, 415, 416, 417, 432
Nereis, 354, 360
INDEX TO GENERA AND SUBGENERA
Nothosaurus, 252
Notomastus, 354
Nucula, 412, 418, 446
Nux, 409, 414, 416, 432
Nymphon, 321, 322
O
Octopus, 412, 413, 414, 416, 417
Oligokyphus, 233, 518
Odcorys, 410, 414, 415, 416, 417, 439
Orthotrichia, 469, 470
Orthotrichiella, 474
Oswaldella, 273, 274, 2905
Oxyethira, 469, 476
IP
Pantopipetta, 321, 334
Papio, I
Paragattya, 270, 281, 318
Paraonis, 354, 363
Parapapio, 2
Parapapio, 1
Parascapanodon, 69
Pista, 354, 369
Pipetta, 336
Phalacrostemma, 354, 355, 367
Philbertia, 409, 415, 426
Physocephala, 395
Pleisiosaurus, 252
Pleurotoma, 423
Pleurotomella, 427, 429
Pleuroxus, 35, 45
Plumularia, 268, 269, 273, 274, 285, 286,
289, 293, 300
Plumularia, 285, 286, 292
Poecilochaetus, 354, 365
Polyandrocarpa, 180, 186, 217
Polycarpa, 186, 217
Polycarpa, 186
Polycitor, 145, 147, 155
Polycitor, 160, 162
Polycitorella, 119, 143
Polyclinum, 113, 135, 217
Polynices, 410, 413, 414, 416, 417, 439
Polysyncraton, 119, 165
Pomadasys, 257
Prionium, 16, 18, 30
Prionospio, 354, 365
Pristipoma, 258
Pronucula, 446
Pronucula, 446
Prosipho, 409, 415, 416, 417, 433
Psammaplidium, 124, 127
Pseudodiaptomus, 373
Pseudodistoma, 138, 217
Pseudodromia, 143
Pycnotheca, 273
Pyrene, 410, 415, 417
Pyura, 193, 218
S
Sabellides, 367, 368
Sarepta, 412, 418, 449
Scala, 410, 415, 439
Scalenodon, 233, 480, 502, 518
Scaloposaurus, 250, 251
Scapanodon, 63, 69
Schaphander, 411, 414, 415, 416, 417
Schapholeberis, 35, 40
Schizotrichia, 268, 269, 270, 273, 292
Schmackeria, 374, 381, 387
Scirpus, 472, 473, 474
Serranochromis, 53, 55, 56
Serranochromis, 54
Sertularia, 272
Sesamodon, 232
Setodes, 15, 27
Sigillina, 119, 152
Silicula, 448
Simocephalus, 35, 41
Stmosa, 41
Sipho, 433
Solariella, 411, 414, 442, 443
Solariella, 440
Spergo, 428
Spiophanes, 354, 365
Stomazoa, 139
Streblosoma, 354, 369
Strepsiceros, I
Struthiocephalus, 103
Styela, 119, 190, 217, 218
Surcula, 408, 413, 421, 428
Surcula, 423
Sycozoa, 147, 218
Symplegma, 217
Syndesmya, 450
Synoicum, 119, 131
Systropus, 393
ar
Terebellides, 354, 371
Terebra, 408, 417, 418
Terebratula, 413
Tethyum, 187, 190
Tetrazona, 119, 146
Thalassocyon, 410, 414, 415, 416, 439
Thecocarpus, 273, 274, 310, 312, 318
Thecocaulus, 268, 269, 270
Thecocaulus, 286
Thrinaxodon, 232, 239, 252, 254, 479
Thyasira, 413, 450 —
Tribolodon, 232, 252
Trididemnum, 170, 217
Trirachodon, 516
Trochus, 4.43
Trophon, 410, 413, 414, 415, 416, 417, 435
Turbonilla, 410, 413, 439
Turris, 418, 419
Turris, 420
INDEX TO GENERA AND SUBGENERA
Typhlomangelia, 408, 416, 422
Typhlosyrinx, 409, 414, 416, 417, 423 Watsoniella, 232
U Weismanella, 374
ill
Ulemosaurus, 60 Wyvillea, 429
V
Ventromma, 295 Yoldia, 448
foo e
ANNALS
OF THE
Pew lr AFRICAN MUSEUM
VOLUME XLVI
PART I
NEW CERCOPITHECOID FOSSILS, INCLUDING A NEW SPECIES,
FROM TAUNG, CAPE PROVINCE, SOUTH AFRICA
By
LEONARD FREEDMAN
Department of Anatomy, University of the Witwatersrand, Johannesburg
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[| CONTINUED ON. INSIDE BACK COVER
NEW CERCOPITHECOID FOSSILS, INCLUDING A NEW SPECIES,
FROM TAUNG, CAPE PROVINCE, SOUTH AFRICA
By
LEONARD FREEDMAN
Department of Anatomy, University of the Witwatersrand, Johannesburg
(With 2 figures in the text and plates I-X)
INTRODUCTION
In the collection of the Department of Anatomy, University of Cape Town,
Professor L. H. Wells recently found a number of blocks of breccia and partially
developed fossils from ‘Taung, Cape Province, South Africa. These had been
collected by Professor M. R. Drennan in 1925, a few months after the discovery
of the type skull of Australopithecus africanus, and are almost certainly derived
from the same cave deposit as the Australopithecus skull. Professor Wells arranged
for some of this material to be further developed by Mr. J. W. Kitching of the
Bernard Price Institute for Palaeontological Research, Johannesburg. The only
identifiable non-primate fossil recovered from this material by Professor Wells
is a juvenile lower jaw of a large antelope most nearly resembling the kudu
(Strepsiceros strepsiceros). Five cercopithecoid specimens from this collection
which have been transferred to the South African Museum, Cape Town, have
been handed on to me for description. These constitute a significant addition
to the cercopithecoid material available from Taung.
The new cercopithecoid specimens comprise a fairly complete skull,
with some upper teeth, of a female (S.A.M.11728); most of the right half of a
juvenile skull, with two molars (S.A.M.11729); the damaged lower part of a
calvaria and the left posterior part of a palate and maxilla, with one molar,
of an individual of unknown sex (S.A.M.11730) ; a bilaterally crushed but almost
complete immature female mandible with teeth (S.A.M.11731); three upper
incisors in a small anterior fragment of premaxilla (S.A.M.11732).
Two of these specimens (S.A.M.11728 and S.A.M.11730) appear to
represent a hitherto unrecorded species. A reconsideration of the material from
Taung previously studied (Freedman, 1957) has led to the conclusion that two
other specimens (T.10 and T.13, Transvaal Museum, Pretoria), may now be
assigned to this species.
SPECIMENS S.A.M. 11728 anp S.A.M. 11730
Papio wellsi sp. nov.
Parapapio antiquus (non Haughton) Freedman, 1957 (partim: T.10). p. 174.
Papio izodi (non Gear) Freedman, 1957 (partim: T.i3). p. 180, pl. 27, fig 56.
Holotype. A fairly complete female skull (S.A.M.11728), with only the v
right P§, P4, M? and M3 present.
I
Ann. S. Afr. Mus. 46 (1), 1-14, pls. 1-10, 2 figs.
BTN UN? 06T
2 ANNALS OF THE SOUTH AFRICAN MUSEUM
Repository of type. South African Museum, Cape Town, South Africa.
Type site. Limestone Deposits, ‘Taung, Cape Province, South Africa.
Referred Material. ‘The following specimens from Taung are referred to
P. wells:
Specimen T.13: The left half of a skull with M1-M®, and also the right
M® in a small piece of maxilla, probably of a female.
Specimen T.10: A damaged calvaria and posterior part of the muzzle,
with the left P*-M? and right M?, of a juvenile—probably male.
Specimen S.A.M.11730: The badly damaged base of a calvaria and left
maxilla with a molar (probably M?) of unknown sex but most likely a male.
Diagnosis. P. wellst is a small species of Papio which in norma lateralis
shows a deep concavity between glabella and rhinion. The muzzle is
characterized by fairly well-developed maxillary ridges and large maxillary
fossae of variable depth. The calvaria is considerably flattened and there
appears to be appreciable sexual dimorphism in skull size. The teeth are of the
Papio-Parapapio type; all female teeth are smaller than their male equivalents.
M3 shows some reduction of the disto-buccal cusp (metacone).
Description. For purposes of description, skulls have been oriented so that
the occlusal plane is horizontal.
The type specimen of P. wells: consists of a fairly complete female skull of
a small baboon-like species. ‘The skull lacks the anterior tip of the muzzle, the
left orbit and both zygomatic arches. Also, the muzzle is bent at an angle to
the calvaria. Much bone has been lost from the calvaria—particularly the
posterior, left parietal and basal portions—but a good endocranial cast has
been exposed in most of these areas. From the almost unworn state of M?,
it would appear as if this tooth had only just come into functional position
and the individual had thus only just become fully adult. ‘The other good female
specimen, T.13, consists of most of the left half of the skull but lacks the anterior
tip of the muzzle, most of the nasal bones and the top of the calvaria.
Viewed in profile (Plate I) the muzzle of S.A.M.11728 is seen to be
elongated and of about the same length as the calvaria. The region between
glabella and rhinion forms a deep concavity and the reduction in height is
particularly rapid in the inter-orbital region. The portion of the nasal profile
just posterior to the nasal aperture is almost horizontal and slopes inferiorly
only very slightly. Specimen T.13 (Plate II) has lost most of both nasal bones,
but it seems clear that the profile shape was very similar to that of the type
specimen.
The muzzle dorsum of the type specimen is short antero-posteriorly and
slopes down steeply from either side of the rounded nasal bones to the maxillary
ridges. From these ridges, which are not particularly well developed in this
specimen, the maxillae slope down almost vertically to the alveolar margin,
forming large but only slightly excavated bilateral maxillary fossae. In T.13
NEW CERCOPITHECOID FOSSILS INCLUDING A NEW SPECIES 3
the shape of the muzzle dorsum is similar to that of the type but, on the sides
of the muzzle, there are very large, deeply excavated maxillary fossae.
Both $.A.M.11728 and T.13 present fairly large orbits. The supra-orbital
tori are only slightly developed in the type specimen but well developed in
T.13. In $.A.M.11728, the palate is relatively long and narrow, with fairly
deep sides and a flat roof (Plate III) ; the posterior palatine foramina are large.
The palate in T.13 is damaged but its shape appears similar to that of the type
(Plate IV).
The calvaria of the type specimen is considerably flattened (Plate I).
The supra-orbital tori are slightly raised anteriorly and there is a slight
ophryonic groove running across the calvaria posterior to them. The post-glenoid
process of this specimen is broad mesio-laterally. Anterior to it there is a
prominent groove which is deep laterally but becomes shallower medially
(Plate III). Anteriorly this groove is bounded by the posterior edge of the
inferiorly projecting root of the zygomatic process of the temporal bone. The
post-glenoid process, the root of the zygomatic process of the temporal bone
and the glenoid fossa are all situated relatively high above the level of the
occlusal plane (Plate 1). In T.13, the calvaria also appears to have been
flattened (Plate I1) but the glenoid fossa region is not present for comparison
with that of $S.A.M.11728.
The nasal bones of the probable male, T.10, are considerably damaged
but its profile contour (Plate V) must have been of the Papio type (see
‘Discussion’ below) and similar to that of the type specimen. In spite of
considerable damage to this general region, the muzzle dorsum would appear
to have been flattened and have had almost vertical lateral surfaces. On the
left side, the vertical surface of the maxilla appears to have been pushed in,
but from the small posterior part of the right side preserved, it seems clear
that an excavated maxillary fossa was present. ‘The calvaria is flattened supero-
inferiorly and of very similar shape to that of the type specimen. Although
immature, T.10 is considerably larger than the females $.A.M.11728 and T.13.
The other probable immature male, S.A.M.11730, has only a small portion of
the palate and left maxilla present (Plate VI) but the very deep maxillary
excavation and the tooth size (see ‘Discussion’ below) suggest that it belongs
in this species.
In the females $.A.M.11728 and T.13 the dental arcade is horseshoe-
shaped but elongated antero-posteriorly (Plates III and IV); the male shape
cannot be determined. The male P* of T.10 (Plate VII) is very well preserved
and almost unworn. It is a typical bicuspid cercopithecoid premolar and has a
faint disto-lingual vertical groove. The two female premolars of the type
specimen are considerably worn and damaged and little can be seen of their
structure. The male and female molars are all typical bilophodont teeth of the
Papio-Parapapio type and several have grooves marking off mesio-lingual
cuspules (Plates III, IV and VII). The disto-buccal cusps of M? on both
female specimens show signs of reduction. (No male M? is known). On size
4. ANNALS OF THE SOUTH AFRICAN MUSEUM
(table 3) the molars of the female, T.13, are considerably larger than those
of the other female, S.A.M.11728. Nevertheless, it is apparent that the female
teeth of P. wellsi are considerably smaller than those of the males (table 2), the
difference being of the order usually found in the genus Papio.
The question of possible mandibular fragments and teeth of this species
is dealt with in the ‘Discussion’ below.
Measurements. The measurements of the female skull and the male and
female upper teeth of this species are given in tables 1, 2 and 3 respectively.
Discussion. In previous studies two baboon-like species, Parapapio antiquus
(Haughton) and Papio izodi Gear, and possibly a third, Parapapio jonesi Broom,
have been recorded from Taung, together with a fourth species of considerably
different character, Cercopithecoides williamsi Mollet. The material assigned to
P. wellsi must therefore be compared with the material which has been referred
to the first three of these species (Freedman, 1957).
The two known male individuals of P. wells: are immature as well as
damaged and fragmentary. They appear to agree with the females in general
| . Be ms. j { % 5S ems. J
FiGeas Ries
FicureE 1. Pantograph tracings of the muzzle profile Figure 2. Pantograph tracings of the
shapes of females of (from above downwards): muzzle coronal section shapes (taken
Parapapio antiquus (Tvl.639), Papio izodi (AD.g92), midway between glabella and rhinion)
Papio wellsi (S.A.M.11728) and Parapapio jonesi of females of (from above downwards) :
(S/ES-565). (Xa. Papio izodi (AD.992), Parapapio jonest
(STS.568), Parapapio antiquus (Tvl.639)
and Papio wellsi (S.A.M.11728). X I.
NEW CERCOPITHECOID FOSSILS INCLUDING A NEW SPECIES 5
morphology but (as described above) they differ in size to the degree usual in
Papio. Insufficient is thus known of them—and also of the males of some of the
species referred to— to make comparisons between the males of value at present.
The comparisons made below between P. wells: and other known cercopithecoid
species will thus depend mainly on comparisons between females.
Both Parapapio antiquus and Papio 1zodi are larger than P. wells in most skull
and some dental measurements (tables 1 and 3), P. zzodi being the largest.
In morphology P. wellst shows important resemblances to both P. antiquus
and P. izodi. Thus, in profile shape of the muzzle P. wells: is clearly of the
Papio type, with a steep drop in the inter-orbital region similar to that seen in
P. izodi (fig. 1). On the other hand, the muzzle shape of the new species in
coronal section (fig. 2) is very similar to that of P. antiquus. P. wellsi and
P. antiquus both have definite maxillary ridges and maxillary fossae, whereas
neither is present in P. 2zodi. The new species differs from both P. izod: and
P. antiquus in its generally smaller skull and dental size, its flatter calvaria and
the shape and situation of the structures in the glenoid fossa region as described
above. On muzzle profile shape and sexual dimorphism the group of specimens
referred to P. wellsi (S.A.M.11728, T.13, T.10 and S.A.M.11730) must clearly
be placed in the genus Papio. However, to suggest including them in the species
P. 1zodi would be to increase the range of variation, particularly morphological,
beyond reasonable limits.
P. wellsi is similar to Parapapio jonesi in skull size (table 1) and in muzzle
coronal section shape (fig. 2). However, it differs considerably from that species
in muzzle profile shape (fig. 1) and also in the shape and relative height of the
calvaria. In the teeth (table 3), the morphology of the two species is similar but
on size, particularly in the length dimensions, the P. wells1 molar measurements
are considerably larger.
Before this new species was established, T.13 was referred to P. tzodt
(Freedman, 1957). The presence, in a single specimen, of very large, deep
maxillary fossae and a slightly different muzzle dorsum shape, were not felt to
be sufficient grounds for the creation of a new species. Despite the size and
morphological differences from the type specimen, described above, it now
appears that this specimen is a female of the new species, P. wellsi. The main
difference between T’.13 and the type specimen of P. wellsi lies in the size and
depth of the maxillary fossae. In a large number of skulls of Papzo ursinus and
Papio cynocephalus which were examined for this feature, maxillary fossae were
found to be of very variable size and depth.
In that same study (Freedman, 1957) it was also suggested that the muzzle
shape of T.10, in profile, was probably that of a Parapapio. The profile shape
appeared to be of the Papio type but the inter-orbital region was considerably
damaged and the individual from which it came, immature. On the shape of
the muzzle in coronal section this specimen agreed with the Parapapio described
from Taung (P. antiquus) and disagreed with P. 1zodi, the only Papio then known
from Taung. It was thus decided that the shape in an undamaged adult of this
6 ANNALS OF THE SOUTH AFRICAN MUSEUM
type might well be similar to that of a Parapapio and the specimen was referred
to P. antiquus. The new species now being described is a Papio but has a muzzle
which, in coronal section, is similar to that of Parapapio antiquus. The apparent
contradiction of profile and coronal section muzzle shapes is now resolved and
T.10 can, with reasonable certainty, be referred to P. wellsi. On the size of
the skull and teeth the specimen seems clearly to be from a male.
TABLE I. Comparison of the skull measurements (in mm.) of Papio wellsi
females with those of similar female cercopithecoid specimens.*
Para- Papio
Papio papio | Parapapio Papio angust-
wellsa jonest antiquus ~zodt ceps
S.A.M. STS vn
11728] T.13 | 565 | 639 | T.17 |AD.992/AD.946/K.A.194
General
Greatest length | (134) | (130) | (135) | (143) (138 ) 139
Basal length (QI) 99
Calvaria
Height
Basion-bregma 54 58 60 (64)
Basion-glabella 64 70 72
Breadth
Greatest
temporal | (65) 68 69 72 67
Length
Inion-glabella (82) (88) | (go) (go)
Inion-basion (50) 47 53 48
Foramen magnum
Length (18)
Muzzle
Height
Anterior to P? 16 22 20 17 a0
Breadth
Anterior to M?| 43 A5 46 46 42 47 45
Anterior to P? | 29.5 35 36
Dorsal to M? 2 30 35 a1
Length
Muzzle (67) (65) 78 78
Palate (58) 51 63) (62)
Orbit
Inter-orbital (9) 10 8
Nasal
Breadth 15.5 ig 15.4
* The comparative measurements in this table, and also those in the
succeeding tables, are all from Freedman (1957). Professor L. H. Wells has
pointed out to me that the figure for the muzzle length of AD.946 was
erroneously given in that study as 88 mm. instead of 78 mm.
NEW CERCOPITHECOID FOSSILS INCLUDING A NEW SPECIES Vi
The new specimen, S.A.M.11730 is only a small maxillary fragment
(Plate VI). However, the deeply excavated maxillary fossa is strongly suggestive
of P. wellsi and, when the single M? is compared to its equivalents in tables 2
and 3, it is again seen to fit well as a male in that species.
Because of the resemblances and the small number of specimens of the
four baboon-like species from Taung, and particularly with the overlap of
characters between P. wellsi, and the previously described species Parapapio
TABLE 2. Comparison of the dimensions (in mm.) of the upper teeth of Papio
wellst males with those of similar male cercopithecoid specimens.
(no males known)
Papio angusticeps
Range*
* The ranges given in this and the succeeding tables are based on very
few (mostly 3-6) specimens.
jonesi, Parapapio antiquus and Papio izodt, it is difficult to identify positively a
number of specimens. For example, C.T.5360 and T.22, previously referred to
P. antiquus (Freedman, 1957), consist of upper teeth only and could, on size,
equally well belong in P. wellsi. Similarly, T.16 was previously described as a
female P. antiquus and could now also be referred to P. wellsi.
The possibility of even C.T.5356, previously considered a male of
P. antiquus, being a badly distorted male of P. wells cannot now be ruled out
completely. The distal reduction of M? of this specimen is mainly on the buccal
side and very similar to that seen in the female P. wellsi specimens. If this
specimen were referred to P. wellsi, this would resolve the apparently anomalous
situation of a female P. antiquus having an M2? very considerably reduced
8 ANNALS OF THE SOUTH AFRICAN MUSEUM
distally, and, the male of the same species having the distal portion of that
tooth only slightly reduced. However, it does not appear as if the specimen had
a Papio-like muzzle, although this region is badly distorted and damaged,
possibly sufficiently so to be misleading.
Further, the diagnostic features of the above mentioned four species lie
mainly in the skull and few specimens are known which include diagnosable
skull fragments associated with mandibles with teeth. In particular, no good
P. wellsi skull is associated with lower teeth. Hence, with the description of the
new species P. wellsi, the mandibles referred to P. antiquus (T.12 and T.23—
males; T.18, T.21 and T.27—females; A.D.944— ? sex) could all now equally
well be referred to P. wellsi. AD.943, a male specimen formerly referred to
P. antiquus, on tooth size and probable muzzle shape, the latter as deduced from
the case of the internal surface, should probably now be referred to P. wellsi. It is
therefore now apparent that the status of a number of specimens from Taung,
including almost all of the known mandibles, is best left in abeyance until
more material, especially associated skulls and mandibles, is discovered.
The presence at Taung of two similar species of Papio might at first glance
appear improbable, but it should be recalled that the limestone deposit at this
site covers a considerable period of time and the two species might not
necessarily have been in the area at the same time. The material described in
this paper almost certainly comes from the Australopithecus africanus type site,
whereas at least some of the material previously described from Taung came
from other parts of the deposit and may thus possibly be of different age.
Further, a fairly large number of different fossil cercopithecoids (fifteen species
of five genera), dating from roughly the same time period as that covered by
the Taung deposit, have been described from Southern Africa. From one of
these sites (Sterkfontein), three species of one genus (Parapapio jonesi, P. broomi
and P. white) have been recorded.
With regard to the other species of Papio described from Southern Africa,
P. wellsi shows resemblance to Papio angusticeps (known from Kromdraai and
Cooper’s Quarry, near Krugersdorp, Transvaal, South Africa) in the depth
and extent of the maxillary fossae. However, it is a smaller species, has less
well-developed maxillary ridges and a flatter calvaria. The teeth of the two
species are similar in size and morphology. The new species is very considerably
smaller than Papio robinson. (known mainly from Swartkrans, Transvaal,
South Africa) and Papio ursinus (the extant South African baboon), and also
differs from these two species in a number of morphological characters.
Of all the cercopithecoid species known from Southern Africa, P. wells:
appears to have its closest affinities with P. angusticeps. These two Papio species
are of generally similar morphology and are both characterized by large and
usually deep maxillary fossae. P. angusticeps is somewhat larger than P. wells
but it occurs at sites considered to be of more recent date than Taung.
However, P. wells: shows differences even to P. angusticeps, e.g. in its flattened
calvaria, which would appear to rule it out as a direct antecedent of that species.
NEW CERCOPITHECOID FOSSILS INCLUDING A NEW SPECIES
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IO ANNALS OF THE SOUTH AFRICAN MUSEUM
SPECIMEN S.A.M.11729
Description. This specimen (Plate VIII) consists of the right half of the skull
of a juvenile baboon of unknown sex. Most of the nasal and frontal bones are
missing and there are a number of fractures and slightly weathered areas on
the calvaria and muzzle. The base of the calvaria is considerably damaged
but the palate is well preserved. Of the dentition, only the right M? and the
buccal half of the right dm? are present.
The muzzle dorsum, on each side, is triangular in shape and slightly
concave. It slopes down rather steeply from the lateral edge of the rounded
nasal to the clearly marked maxillary ridge. From the maxillary ridge, the
maxilla slopes down even more steeply to the alveolar margin and a large,
slightly excavated, maxillary fossa is formed.
Features of note when this specimen is compared with Papio (P. ursinus
and P. cynocephalus) and Parapapio (M.3004, Parapapio sp.) specimens of
approximately the same dental age are:
(1) The size of the orbit is relatively large.
(2) The zygomatic arch is considerably better developed.
(3) The anterior part of the temporal line is rather well defined. (The
posterior part of this line is weathered away.)
(4) The mastoid region is well developed.
) The supramastoid crest leads to an occipital crest.
(6) The planum occipitale is more vertical than usual.
These differences may, in some of the above instances, be more apparent than
real as it is very difficult to get comparative specimens of precisely the same
age. Considerable time periods elapse between the eruption of successive
teeth in the cercopithecoids. However, morphologically the skull of this
specimen does give the impression of being robust and adult for its age.
The palate of S.A.M.11729 is fairly deep and smoothly concave from the
alveolar margin to the midline, which is raised into a crest anteriorly. The
posterior palatine foramen is situated in a deep recess. The fragment of dm?
appears similar to its equivalents in Papio and Parapapio; the single M? is well
preserved, almost unworn and of the Papio-Parapapio type.
Measurements. As the specimen is juvenile, no skull measurements were
taken. The dimensions of the single M! (in mm.) are: breadth (mesial) —8.5,
breadth (distal)—8.0 and length—g.5.
Discussion. On the shape in coronal section of the muzzle, S.A.M.11729
is similar tu Papio wells:, Parapapio antiquus and Parapapio jonesi. Reconstructions
of the frontal and nasal regions in plasticine to give the probable profile shape
did not give wholly satisfactory results, as the damage is so extensive that there
is a measure of variation in the shape which can be produced, depending
on how far forwards the supra-orbital tori and glabella are produced. However,
2B
NEW CERCOPITHECOID FOSSILS INCLUDING A NEW SPECIES Tok
the angle at which the remaining small portion of the nasal bones lies, and the
shape of the small piece of frontal lying at the supero-lateral corner of the orbit,
do seem to favour at least a shallow concave type of nasal profile.
The various notable features of $.A.M.11729, to which attention was
drawn above, cannot be compared fairly with the equivalents in the three
species under consideration. Firstly, usually only female skulls of those species
are available and secondly, the known specimens are often damaged in the
relevant areas. Further comparative material is necessary for the evaluation
of those features in $.A.M.11729. However, the general robustness of the
specimen may well be due to the specimen being a juvenile male and may
not have any taxonomic significance.
The dimensions of the single M! of S.A.M.11729 are similar to those in
the males and females of P. jones: (tables 2 and 3). When compared to their
equivalents in P. wells: and P. antiquus, the dimensions of this tooth are almost
all smaller than those of the females of both of those species (table 3), but,
of the two, they are appreciably closer to P. wellst. Compared with the male
equivalents in those two species (table 2), the tooth is considerably smaller
than those of P. wellsi (in which there is size sexual dimorphism of the molars)
but there is no known male M! of P. antiquus for comparison. However, as
there is little or no sexual dimorphism in the molars of any of the known
Parapapio species, the size difference should be about the same as for the female
of that species.
It is most difficult to come to any definite conclusion about the affinities
of S.A.M.11729. From the muzzle coronal shape, the specimen could belong
in P. wellsi, P. antiquus or P. jonest. ‘The reconstruction of the muzzle profile
shape is slightly more suggestive of a Papio but P. jonesi does show some concavity
in this region and the juvenile shape of Parapapio spp. generally is not well
known. The size of the single M! is most similar to the equivalent tooth in
P. jonest but, it is also not very much smaller than the equivalent tooth in the
P. wellsi female (T.13) and it should be remembered that only a few specimens
are involved in this comparison. On balance of evidence, it would appear that
S.A.M.11729 might be a juvenile female of P. wells: or, more likely, a juvenile
male or possibly female of P. jonesi. However, the provenance of P. jonesi at
Taung depends on a single specimen, consisting of a small piece of maxilla
and damaged teeth, and the possibility of the specimen belonging even in
P. antiquus cannot be ruled out completely.
SPECIMEN S.A.M.11731
Description. S.A.M.11731 is an almost complete mandible of an immature
female cercopithecoid. The specimen has been fractured slightly to the right
of the symphysis, and the right half has been pushed to the left and lies close
up against, and slightly above, the left half of the mandible (Plate IX).
I,—-M, are present on the left side. The premolars and molars are very
IS J, snnbiup
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ANNALS OF THE SOUTH AFRICAN MUSEUM
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12
NEW CERCOPITHECOID FOSSILS INCLUDING A NEW SPECIES 13
well preserved but the incisors and canines are cracked and damaged. On the
right, the incisors are also present but very badly damaged. The only other
tooth present on this side is M, and it is slightly damaged distally. M, has not
yet erupted on either side but the tooth can be seen, on both sides, lying in
its crypt. The tips of the cusps of that tooth are at about the level of the buccal
alveolar margin.
Anteriorly, the symphysis slopes down steeply and is flattened. A foramen
symphyseosum is present in the midline. The flattened anterior surface of the
symphysis is demarcated by a prominent oblique ridge on either side. The
body of the mandible is low and thin but shows some thickening posteriorly in
the region of M;. A large but very shallow mandibular fossa is present, on the
antero-inferior margin of which is the mental foramen. The ramus is about
as vertical as that of Papio ursinus but does not rise as high. The gonial angle is
inverted. The coronoid process lies slightly above the level of the condyle;
the sigmoid notch was probably not very deep. The teeth are all typical of
the female Papio and Parapapio and show no unusual features (Plates IX and X).
A small mesio-buccal cuspule is present on both M, teeth.
Measurements. As the specimen is immature, no measurements of the
mandible itself were taken. The dental dimensions and comparisons are given
in table 4.
Discussion. On molar size, which has often proved a most useful taxonomic
character in the Cercopithecoidea, $.A.M.11731 is smaller than the few known
female specimens previously referred to P. izodi and P. antiquus, the difference
being greater in the former case. In the incisors and premolars, the dimensions
of S.A.M.11731 and P. antiquus females are similar but there are no comparable
teeth of P. izodi. As compared to the female P. jones: teeth, the incisors of
S.A.M.11731 are larger but the premolars and molars are of about the same
size. No lower teeth or mandibles have so far been assigned to P. wellsz. It is
not possible to assess accurately what the adult dimensions of the mandible
of S.A.M.11731 would have been but, as far as can be judged, the size would
probably have been about the same as in P. jonesi females, slightly smaller
than in P. antiquus females (assuming T.18 and T.27 to have been correctly
assigned) and considerably smaller than in P. zzodi females.
As concluded above, the Taung mandibles are best left in abeyance
until more and better associated skulls and mandibles are recovered. However,
it would appear probable that, on size and the presence of a mandibular
fossa, the new immature, female, mandibular specimen, $.A.M.11731, might
well belong in P. wellst, but it could also reasonably be referred to P. jonest.
SPECIMEN S.A.M.11732
This specimen consists of a small portion of the left premaxilla with I?
and I? and an associated right I?, almost certainly of the same individual.
14. ANNALS OF THE SOUTH AFRICAN MUSEUM
The teeth are of the typical cercopithecoid pattern and show no unusual
features. Insufficient upper incisors are known from Taung for the affinities
of the specimen to be determined.
SUMMARY
1. Five new cercopithecoid fossils are described from Taung. ‘They almost
certainly come from the Australopithecus africanus type site.
2. Two of these specimens, plus two specimens previously described from
Taung, form the basis of a new species, Papio wellst.
3. Because of an overlap in size and morphological characters between
the four baboon-like species recorded from Taung, the identity of several
specimens, notably mandibles, cannot at present be determined.
ACKNOWLEDGEMENTS
My thanks are due to Professor L. H. Wells for handing me this material
for description and also for his most helpful criticisms of the manuscript.
Professor P. V. Tobias, Anatomy Department, University of the Witwatersrand,
Johannesburg, and Dr. J. T. Robinson, Transvaal Museum, Pretoria, also
kindly read through the manuscript and offered useful suggestions. Messrs.
A. R. Hughes and R. Klomfass, both of the Anatomy Department, University
of the Witwatersrand, Johannesburg, took the photographs which illustrate
this paper. This paper has been published partly with the aid of a grant from
the University Council Research Committee, University of the Witwatersrand.
REFERENCES
FREEDMAN, L. 1957. Ann. Transv. Mus., 23, 121—262.
Ann. S. Afr. Mus., Vol. XLVI Plate I
Papio wellsi, female (S.A.M.11728). Skull, lateral view.
Ann. S. Afr. Mus., Vol. XLVI Plate II
Papio wellsi, female (T.13). Skull, lateral view.
iy .'
,
q
Plate III
Ann. S. Afr. Mus., Vol. XLVI
]
Papio wellsi, female (S.A.M.11728). Skull, basal view
Plate IV
NMG
Ammen os) Adi. Mus... Vol; XacV:
lew.
asal \
b
3). Skull,
elit
Papio wellsi, female (°
fe
weds
=“
———
"MOIA [R19}P]
-Olojue UolUSseay =[[Nnyg *(o€ZL11
‘W'V'S) 9yew s[qeqoad ‘2sjjam o1vg
{Al ING
“MOIA [PIO] “TTNAS
(its
L) 2]ew
aTqeqoaid
“asjjam oidvg
A aN |
IATX “TPA “Sny “sty
Ann. S. Afr. Mus., Vol. XLVI
Papio wellsi, probable male (T.10). Skull, basal
view.
Ann. S. Afr. Mus., Vol. XLVI
Plate VII
Plate VIII
Juvenile cercopithecoid (S.A.M.11729). Skull, lateral view.
Anne oe Airs Mus... Vole SaaVI Plate IX
Immature female (S.A.M.11731). Mandible, lateral view.
Ammweoe Ati Mus, Vol: SV Plate X
Immature female (S.A.M.11731). Mandible, occlusal view.
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. Jn Michaelsen, W., ed. Beitrdge zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Hartman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8,1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuka, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.— All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name....
B. Form for bibliographic references to synonymic names.—The first reference following
any name in the synonymy should be tothe earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example :—
Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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XXVIII.
LOUD
TOK
INDEX
SOCK
POE
SOROX DEL,
SOO-CNe
SOUXVi
SOME
MX KVL
XXXVIII.
SOX
xi.
SLL.
XLI.
XL:
SIETVE
LV
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1914-1916
1Q17>1933
1917-1920
1921
1924-1925
1924-1926
1925-1927
1925-1928
1925-1926
1929-1938
1927-1928
1928
1929
1929-1952
1929-193!
L952 s1935
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BNINALS
OF THE
eOowWIm AFRICAN MUSEUM
VOLUME XLVI
PART II
SOME NEW CADDIS FLIES (TRICHOPTERA) FROM
THE WESTERN CAPE PROVINCE — III
By
Ke Mer. Scorr, Pa. D., F-R.E:S.
National Institute for Water Research
South African Council for Scientific and Industrial Research
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[ CONTINUED ON INSIDE BACK COVER
SOME NEW CADDIS FLIES (TRICHOPTERA) FROM
THE WESTERN CAPE PROVINCE — III
By
K. M. F. Scott, Pu.D., F.R.E.S.
National Institute for Water Research
South African Council for Scientific and Industrial Research
(With 8 figures in the text)
INTRODUCTION
This is the third paper in the present series, in which caddis flies from the
western Cape are being described. In it an account is given of the larval and
pupal stages of the three new species of Leptoceridae described in Part II
(Athripsodes prionii, A. bergensis and Leptecho helicotheca); the pupal skin of Athrip-
sodes tuckeri (Barnard) ? var. is also described, and all stages of a small species
of Setodes, S. barnardi, from the Great Berg River, which appears to be new.
The immature stages have been described in considerable detail with the object
of collecting sufficient information to build up keys for the identification of
larvae and pupae collected in field surveys.
The methods employed have been the same as in previous papers, and, as
before, holotypes will be deposited in the South African Museum, and paratypes
will be sent to the British Museum (Natural History). Terminology as before,
except that Ross’s term ‘stridulator’ is used instead of the more cumbersome
‘plaque d’appui’ (Barnard), or ‘process of the trochantin’ (Ulmer). A full descrip-
tion of the habitats of the species described will be found in Harrison and
Elsworth (1958) and Harrison (1958). (See addendum to the latter paper for
names used.) |
The author would like to express her grateful thanks to Mr. D. E. Kimmins
of the British Museum (Natural History) for much helpful advice, also to
Dr. A. D. Harrison of the National Institute for Water Research, and to
Professor J. H. Day of the Zoology Department, University of Cape Town, in
whose department the work was carried out.
The Trustees of the South African Museum gratefully acknowledge a grant
towards the cost of publishing this paper made by the S.A. Council for Scientific
and Industrial Research.
DESCRIPTION OF IMMATURE STAGES
Athripsodes prionii Scott
Fig. 1 A-R; Fig. 5 K-Q
Remarks: Larvae were collected from the Palmiet River at Elgin in
October and November, and from the Great Berg River near Assegaibos,
Ann. S. Afr. Mus. 46 (2), 15-33, 8 figs.
T9
SMT SUNTAN
institution YUN 7 Wet
16 ANNALS OF THE SOUTH AFRICAN MUSEUM
Driefontein, Groot Drakenstein and Bridgetown between April and October.
The only place at which they were common was at Groot Drakenstein in autumn
(April and May). The larvae are swimmers and live mainly among the marginal
vegetation, particularly the palmiet (Prionium serratum).
Larva (Fig. 1 A-M): Described from entire specimens. Case-bearing swim-
ming larvae; eruciform; length up to 8-0 mm. Head: hypognathous, oval,
clypeus somewhat asymmetrical with 2 pairs of lateral indentations and 10
bristles, a suture-like line present paralleling the epicranial arms; gular sclerite
pear-shaped, not fused with genae. Head golden brown with darker brown
patterning as indicated in figure 1 A-E. Anteclypeus membranous, pale. Eyes
large, black, set in a clear pale area, with 6 raised transparent cuticular plates
over each forming hyaline lenses. Antennae conspicuous, with bulbous proximal
segment and long distal segment terminating in a sensory bristle. Mouthparts:
labrum armed with 4 blade-shaped spines and 6 long setae, there are also 2
pairs of very short spines just below the central emargination. Mandibles unequal
in size, dark brown, heavily sclerotized; each mandible with 5 teeth, 2 small
brushes of stout bristles on the inner face of the larger left mandible. Maxillary
palps 3-segmented, 3rd segment bearing several small papillae; maxillary lobe
bordered with long blade-like spines, smaller spines and setae. Labium a low
cone, labial palps each tipped with a papilla. Thorax: pro- and mesonota
sclerotized, patterned with darker brown and cream as indicated in figure 1 B;
metanotum membranous except for 2 small oval patches bearing setae. The
patterning on both head and thoracic nota is variable, as is the ground colour,
which varies from dark brown, when the pattern scarcely shows, to pale
yellow, when it shows up very clearly. Each half of the pronotum is subdivided
by a groove. The mesonota are lightly sclerotized, with a pair of narrow dark
bars. The metathorax bears what appear to be 2 small gills on each side
(fig. 1 A). Legs: prothoracic legs short, stout; tibia and tarsus edged with
spines ; bristles, spines and hair-like setae fringe the femur. Stridulator stout, with
blunt apex. Mesothoracic legs armed with strong spines, interspersed with hair-
like setae along edge of femur. Metathoracic legs long, strong, with heavy double
fringe of long hairs used in swimming. All leg joints of one segment only except
trochanters which have two; femora of mid- and hind-legs also appear to have
two segments, though it is not clear whether this division is functional or merely
incipient. Coxopleurites, trochantin etc. of legs heavily sclerotized. Abdomen:
white; on segment 1 lateral humps covered with small scabrosities, and a pro-
minent dorsal hump bearing a patch of spicules; on segment 2 a small sclerite
on each side and 3 simple filiform gills (presegmental dorsal and ventrolateral
gills, and a postsegmental lateral gill) ; on segments 3 and 4, 2 gills (presegmental
dorsal and ventrolateral) ; no gills on segments 5-9. Lateral line extends from
segment 3 to 7; on segment 8 it is replaced by a row of small bifid tubercles.
Segment 9 is membranous but the tergite is bordered posteriorly with a row
of 6 strong black bristles. The anal prolegs are fused basally to form a 1oth
segment; each is short and bulbous with a lateral sclerotized patch and several
SOME NEW CADDIS FLIES FROM THE WESTERN CAPE PROVINCE I7
—————
M eS
=
—_—
=S
i
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i
|
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Zag
/
Fic. 1. Larva of Athripsodes prionii Scott.
A, lateral view of whole larva, with lateral tubercle enlarged. B, dorsal view of head with
pro- and mesonota. C, antenna. D, clypeus. E, head from behind showing gular sclerite.
F, right mandible. G, left mandible. H, labrum. J, posterior end of larva (right side).
K, anal claw. L, prothoracic leg. M, metathoracic leg. N, case of early instar of larva.
P, Q, cases of later instars. R, case of mature larva.
18 ANNALS OF THE SOUTH AFRICAN MUSEUM
stout black setae; anal claws have 4 barbs (1 large and 3 small); anus flanked
by patches of small spicules.
Case (fig. 1 N-R): Tubular, wider at anterior end, with hood. Small
larvae make their cases of sand grains, lining them with secretion; older larvae
utilize small bits of vegetable debris, charcoal, etc., with a few sand grains at
the posterior end; mature larvae make a very characteristic rounded case from
3 neatly shaped pieces of dead leaf (usually of Prionium serrata); one piece is
always ionger than the others and forms a hood at the anterior end of the case
and a projecting strip at the posterior end. The case is lined with secretion and
may be neatened round the front and hind openings by the addition of a few
small bits of leaf glued on with secretion; there is always a neat round hole at
the posterior end.
Pupa (fig. 5 K-Q): Labrum rounded, rising to a low point; setae compara-
tively short, some recurved; mandibles large, strong, inner edge finely serrated,
outer edge sinuous, ending in a strong condyle, one pair of lateral setae. Antennae
much longer than body, distal ends coiled. Fore-tarsi slightly, mid-tarsi heavily,
fringed. Dorsal plates present on abdominal segments as follows: on segment 1,
2 pairs of protuberances covered with small sclerotized points (these point in
opposite directions in each pair); on segments 3 and 4, 1 pair of hook-bearing
plates; on 5, 2 pairs; on 6, 1 pair. Segment g bears a pair of small lappets
tipped with forwardly directed setae. The plates on 3 and 4 each have 3 hooks,
the anterior plates on 5 and those on 6 each have 2 large hooks and 3-4 smaller
ones. There are minute scabrosities scattered on the dorsal surface of the
abdomen. Lateral lines extend from segment 3 to 8, where the two curl round
and meet ventrally. Anal appendages are long and stout, with rounded tips;
along the inner edges are small barbs and a few setae.
Athripsodes bergensis Scott
Fig. 2 A-L; fig. 5 A-J
Remarks: The larvae are common in the Great Berg River near Drie-
fontein and Groot Drakenstein in the late winter and spring (August to October).
They are crawlers with long, slender, rough-looking cases of brown sand-grains
to which pieces of stick or charcoal are often attached, and they are frequently
to be seen crawling actively about on top of the stones seeking food. They are
carnivorous and in captivity must be given insect larvae or other small animals
to eat to prevent cannibalism. It is possible that the sticks etc. attached to their
cases serve to discourage predatory fish, as in the case of Anabolia nervosa in
Britain. Pupae were found in October, clustered in numbers under marginal
stones; some were collected and bred out in the laboratory. The pupal cases
are not larval cases sealed off, but each is constructed from small white sand-
grains at the end of the old larval case, being first formed as an extension of it.
When completed however the pupal case is closed off at each end by a neat lid
SOME NEW CADDIS FLIES
FROM THE WESTERN CAPE PROVINCE
Serene rz iEsschpeiziue \\
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aigae uae
eH T
<qrage
CON)
pati
AL
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ie
Ani
Fic. 2. Larva of Athripsodes bergensis Scott.
A, lateral view of larva in case. B, dorsal view of head and thoracic nota. C, head from behind
showing gular sclerite, with eye further enlarged. D, dorsal view of head. E, right and left
mandibles (ventral view). F, prothoracic leg. G, mesothoracic leg. H, metathoracic leg.
J, lateral view of posterior end of larva (right side). K, anal claw. L, left maxilla and labium,
with labial palp further enlarged. (Brushes of hair omitted from left side of labium.)
20 ANNALS OF THE SOUTH AFRICAN MUSEUM
of mesh partly covered by sand-grains, so that the larval sclerites are retained in
the pupal case. Ross (1944, p. 210) states that in the entire family (Lepto-
ceridae) the pupal chamber has a slit and not a mesh in the closing cap at both
ends; while this no doubt applies to the American members of the group, it is
evidently not invariably the case in African species.
Reference to the tables and notes in Harrison and Elsworth (1958) and
Harrison (1958) will show that these larvae were apparently much more wide-
spread and common than is indicated here, having been recorded from stations
1 to g, often at all seasons of the year, and from most types of substratum. It was
found however on closer examination that the larvae collected included three
slightly different types, probably of different species, though that remains to be
proved. Only the records of larvae of the type actually bred out as A. bergensis
have been given here. It is hoped that the other types will be correlated later on.
Larva (fig. 2 A-—L): Described from entire specimens and from larval
sclerites in pupal cases. Length of larvae up to 8-0 mm. Head: hypognathous,
elongate oval, clypeus slightly asymmetrical with a single pair of lateral inden-
tations and 12 setae; gular sclerite long, narrow, widening anteriorly, not fused
with genae. Head capsule shining black except for conspicuous white patches
surrounding the eyes, along the posterior margin of the head and the anterior
border of the clypeus; anteclypeus, antennae and labrum pale yellow to brown
in colour. Antennae long, stout, with small rounded basal segment and terminal
sensory bristle. Eyes large and black, with 6 raised hyaline lenses as in A. prioni
(fig. 2 C). Sutures round clypeus and gular sclerite, and suture-line parallel to
epicranial arms, white and very conspicuous; maxillae and labium large, also
white and very conspicuous. Mouthparts: labrum with 2 pairs of long, curved,
blade-like marginal spines, also a smaller pair, centrally placed; a row of 6
setae across the upper surface and large brushes of setae on the lower. Mandibles
strong, prominent, the left mandible with 5 blunt apical teeth and a brush of
bristles on the inner face, the right mandible with 6 blunt apical teeth and no
brush. Maxillary palps 3-segmented, with 4 apical papillae; maxillary lobe with
a fringe of stout spines and setae along the inner margin. Labium a low cone
with silk gland opening at tip; labial palps each tipped with a small distal seg-
ment bearing a sensory papilla and two spines. Thorax: pro- and mesonota
sclerotized; pronotum brown to blackish, showing slight patterning and bearing
long setae anteriorly, the posterior border separated by a groove from the rest.
Mesonotum yellowish with a brown pattern as shown in figure 2 B. Meta-
notum membranous. Legs: prothoracic legs strong, fringed with spines and
setae; stridulator with rounded tip and two small setae; meso- and metathoracic
legs with long spines bordering tibia and tarsus, shorter spines along femur. All
joints of one segment except trochanters which have two; femora of mid- and
hind legs also show signs of division. Tarsal claws very long, specially on meta-
thoracic legs, each claw with a strong basal bristle. Abdomen : very long compared
with the head and thorax, tapering gradually, znd—4th segments widest. Large
dorsal and lateral humps on Ist segment, each of the latter bears a lightly
SOME NEW CADDIS FLIES FROM THE WESTERN CAPE PROVINCE 21
sclerotized oval setose plate. Anal sclerite present on gth segment, and just
posterior to it 4 long brown bristles. At the base of each anal appendage there
is a large sclerite with 4 strong bristles along its curved posterior edge. Anal
appendages fused to form 10th segment. There are small spicules and tracts of
hair lateral to the anus. Anal claws stout, each with a pair of auxiliary hooks,
and with 2 small sclerites at its base. Feebly developed lateral line present from
segment 3 to 7, followed by a row of chitinous points on 8. Gills absent.
Case (fig. 2 A): A long slender tube, length up to 11:0 mm., wider and
hooded at the anterior end, rough-looking, made of variously sized brownish
sand-grains, with a neat hole at the posterior end. A small stick or one or more
pieces of charcoal is frequently attached to the case, making it rather unwieldy.
The larva sometimes makes use of a hollow reed or anempty case, adding a hood
of sand grains.
Pupa (fig. 5 D-J): Labrum prominent, somewhat angular, with very long,
strong setae; mandibles small, slender, inner edge finely serrated, condyle small.
Antennae very long, distal ends coiled, fore- and mid-tarsi fringed. Protuberances
and dorsal plates present as in A. prionii, the presegmental plates usually with
4 teeth, occasionally 5 or 6; a pair of seta-bearing lappets on gth segment.
Strong lateral lines present from segment 3 to 8, where they meet ventrally.
Anal appendages long and slender, with bifid tips; near tip of each, 3 long stout
setae and several small processes.
Pupal case (fig. 5 A-C): Not an altered larval case, but a new cylindrical
case made of small white sand-grains; closed at each end with a grating made
of secretion, this is partly covered by two or three sand-grains. The larval
sclerites are thus retained within the pupal case. The case is affixed along one
side to the lower edge of a stone resting on clean sand near the margin of the
stream.
Athripsodes tuckeri Barnard ? var.
Fig. 5 R-X
Pupa (fig. 5 R-X): Described from the pupal skin of the g bred out in
January 1955. Labrum low, rounded, with slight anterior point; most setae are
missing, but those present are long and strong. Mandibles fairly short, with
broad base and tapering blade, inner edge serrulate, outer edge sinuous, ending
in a small condyle; one pair of lateral setae. Antennae long, distal ends coiled.
Mid-tarsi fringed. Dorsal plates present on abdominal segments as in A. prioni
and A. bergensis; 4-6 teeth on all presegmental plates. On gth segment 2 small
lappets tipped with setae; these lie on rounded lateral protuberances just anterior
to each of which, but on the ventral side of the body, there is a patch of small
scabrosities (fig. 5 X). Lateral lines extend from segment 3 to 8, where the two
curl round and meet ventrally. Anal appendages long and slender, each tip a
rounded spatulate lobe with three long setae arising from its base.
22 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 3. Larva of Leptecho helicotheca Scott. }
A, entire larva, mature. B, case of very young larva. C, case of mature larva, dorsal view.
D, case of same, ventral view. E, dorsal view of pro- and mesothoracic nota. F, dorsal view of
head. G, ventral view of head. H, left and right mandibles (dorsal view), with brushes on left
mandible further enlarged. J, lateral view of posterior end of larva (right side), with anal claw
further enlarged.
SOME NEW CADDIS FLIES FROM THE WESTERN CAPE PROVINCE 23
Leptecho helicotheca Scott
Fig. 3 A-J; fig. 4 A-P
Remarks: The larvae are fairly common in the Great Berg River near
Driefontein and Groot Drakenstein from late spring to early autumn (Novem-
ber to March); a few were also found in August. Adults were captured or bred
out from March to June. The larvae are crawlers with bright brown heads,
living in small snail-shaped cases made from sand-grains, the cases being
remarkably similar to those of species of Helicopsyche. Leptecho helicotheca larvae
were not found making use of the cases of other Leptocerids, though occasionally
other Leptocerid larvae were found inhabiting their discarded snail-shaped cases.
L. helicotheca larvae appear to feed chiefly on vegetable matter on and under
stones in comparatively deep pools and flats in parts of the river where the
current is slow and the water clear. In autumn a few are also found living on
the marginal vegetation. In captivity they can be reared successfully on alga-
covered stones in aquaria, though judging by the size and position of their eyes
it seems probable that they are actually omnivorous, including minute animals
in their diet as well as algae. When about to pupate the larvae move close
inshore; their pupal cases were found in autumn (March to May), clustered
under the edges of marginal stones. The pupal cases are initially built as an
extension to the larval case, but are straight, not coiled; thus, where they have
been left undisturbed, each pupal case still has the larval case situated at one
end; this facilitates recognition of the pupal cases, which are outwardly much
like those of several species of Athripsodes.
Larva (fig. 3 A, E-J; fig. 4 A-D): Described from entire specimens. Length
of fully grown larva approximately 3-0 mm., but the larvae are always curled
round and not easy to measure accurately. Head: hypognathous, almost oblong;
clypeus triangular, very wide anteriorly, short, with one pair of lateral indenta-
tions and 10 setae. Gular sclerite rounded anteriorly, tapering to a point behind,
not fused with genae. Head capsule bright chestnut brown, with a slightly paler
patch anterior to each eye; anterior margin of clypeus darker brown; sides of
genae thickened and pitted, anteclypeus pale, labrum brown. Antennae long
and stout, each with rounded basal segment and tipped with a sensory bristle.
Eyes large, black, situated fairly far forward, with hyaline lenses as described in
Athripsodes prionit and A. bergensis; there are however no suture-like lines paral-
leling the epicranial arms as in those species. Mouthparts: labrum with 2 pairs
of curved blade-like spines and 3 pairs of long straight setae. Mandibles strong,
the left one with 6 blunt apical teeth and two clumps of bristles on the inner
face; the right mandible somewhat smaller, with 5 teeth and no bristles.
Maxillary palp 4-segmented, with apical papillae; maxillary lobe fringed with
stout spines and bristles. Labium a slender cone with silk gland opening at tip,
labial palps stout, bearing papillae. Thorax : pronotum bright brown, sclerotized,
setose, with the posterior border separated by a fracture line from the rest;
24 ANNALS OF THE SOUTH AFRICAN MUSEUM
ai’
Ye
Fic. 4. Larva and pupa of Leptecho helicotheca Scott.
A, prothoracic leg of larva. B, portion of margin of femur further enlarged (long setae shown
cut short). C, D, meso- and metathoracic legs of larva. E, mandible of pupa, with portion of
serrated edge further enlarged. F, labrum of pupa (long setae shown cut short). G, dorsal
lobes from 1st abdominal segment. H, presegmental dorsal plate from 5th segment. J, post-
segmental plate from same. K, posterior end of pupal pelt, showing lappets and anal appendages.
L, ends of anal appendages further enlarged. M, pupal case. N, lid from anterior end of same.
P, posterior end of pupal case.
ee
SOME NEW CADDIS FLIES FROM THE WESTERN CAPE PROVINCE 25
mesonotum with a pair of thinly sclerotized plates, pale yellow and unpatterned,
each with 3 pairs of setae; metanotum membranous. Legs: prothoracic legs
strong, tarsus and tibia each with a spine and a fringe of small hairs; femur and
anterior portion of trochanter bordered with spines, long setae and short hairs
(fig. 4 A and B). Stridulator blunt, with an upturned rounded apex and 2 small
setae. Meso- and metathoracic legs with a few slender spines and long hair-like
setae; metathoracic legs longest. All joints of one segment except the trochanters
which have two, the femora of mid- and hind legs do however show a constric-
tion near the proximal end which may indicate a division. Tarsal claws long
and stout, each with a strong basal bristle. Abdomen light green, coiled round to
fit case, with large dorsal and lateral humps on 1st segment, on each of the latter
a pubescent, sclerotized plate. On the gth segment an anal sclerite bordered
posteriorly with bristles. Anal appendages fused basally to form roth segment,
a sub-triangular sclerotized plate at the base of each. Anal claws very small,
each with a pair of minute auxiliary hooks. Gills absent. Lateral line strongly
developed from 3rd to 7th segments, on 8th segment a long row of minute
sclerotized points.
Case (fig. 3 B—D): A neat snail-shaped case, 2:5 to 3:0 mm. in diameter,
made from variously sized sand-grains. The case is actually a tube, open at both
ends, and is always a dextral spiral; the larger opening is neatly finished with
smoothly rounded grains, forming a lip, next to which is an umbilicus. The
earliest stage collected was just a small twisted tube (fig. 3 B).
Pupa (fig. 4 E—P): Labrum low, rounded, studded with long setae; each
mandible with broad base and slender apex, inner margin minutely serrated,
condyle small, one pair of lateral bristles. Antennae much longer than body,
‘distal ends coiled; fore-tarsi slightly, mid-tarsi heavily, fringed. Dorsal plates
present on abdominal segments as in Athripsodes prionii; all presegmental plates
have 5-6 hooks. On 9th abdominal segment a pair of small lappets bearing long
setae. Lateral lines extend from segment 3 to 8, where they curl round and meet
ventrally. Anal appendages long and slender, with rounded recurved tips, set
with slender spine-like processes and three long setae.
Pupal case (fig. 4 M-—P): A straight cylindrical tube, roughly constructed of
white sand-grains and lined with secretion. Length 6-0 to 6:5 mm.; a cap at the
anterior end, edged with sand-grains and with a central membranous area
perforated by a meandrine slit. There is a slit in the posterior end of the case,
through which the larval sclerites can be ejected; this is similar to the slit in the
lid. They are not however always ejected, some at least often being retained
within the pupal case.
26
ANNALS OF THE SOUTH AFRICAN MUSEUM
E
===
hdd
SOME NEW CADDIS FLIES FROM THE WESTERN CAPE PROVINCE P47 |
DESCRIPTION
Genus SETODES Rambur
Rambur, 1842, Hist. nat. Ins. Névr.: 315.
Milne, 1934, Stud. N. Amer. Trich., 1: 18.
McLachlan, 1879, Rev. Syn. Eur. Trich.: 338 (partim).
McLachlan, 1884, 1st Add. Suppl.: 39-40 (group II).
Mosely, 1939, Brit. Caddis Flies: 178 (group II).
Kimmins, 1949, Entomologist, LX XXII, No. 1036: 201-4.
Setodes barnardi sp. n.
Fig. 6 A-J; fig. 7 A-J; fig. 8 A-L
A very small dusky gold species with narrow, pointed wings and annulate
antennae. The larvae are crawlers inhabiting neat tubular sand cases. I have
pleasure in naming this species in honour of Dr, K. H. Barnard, pioneer worker
on South African caddis flies.
Imago (dry, before putting into alcohol) (fig. 6 A—J): Head: dusky gold
with pale gold or white tufts of setae, sometimes interspersed with a few black
setae; antennae more or less strongly annulate black and white basally, about
twice wing length, basal segment as in figure 6 J. Eyes blackish, large. Maxillary
and labial palps fuscous. Thorax: dusky gold, with paler gold or white shoulder
tufts with a few black setae. Legs: tibial spurs 1, 2, 2; spur on fore-leg very small,
spurs on mid- and hind-legs unequal in size, the outer one on the mid-leg being
very short and on the hind leg over half the length of the inner. Fore-legs
brownish, mid-legs paler, hind-legs pale gold. Abdomen yellowish. Wings (fig.
6 A, B): length of ¢ fore-wing 4:0 mm., of 9 same. ¢ fore-wing long, narrow,
apex acute; membrane clear, slightly iridescent, somewhat thickened in the
pterostigmal area; pubescence plain dusky gold, a long fringe along the posterior
border; apical forks 1 and 5 present, stalk of fork 1 approximately equal to Ra;
media forks well beyond the anastomosis; thyridial cell long and very narrow,
anastomosis irregular, not clearly defined. g hind-wing narrower than the fore-
wing, acute, membrane clear, pubescence thin and pale, wing fringes very long
(fringe along posterior border nearly as wide as wing) ; only fork 5 present; base
of Rs obsolete. 2 wings similar, slightly wider.
<— Fic. 5. Pupae of Athripsodes bergensis Scott (A-J), A. prionit Scott (K-Q),
A. tuckeri (Barnard) ? var. (R—X).
A, pupal case of A. bergensis. B, lid of same. C, posterior end of case, with portion of grating
further enlarged. D, protuberances on 1st abdominal segment of pupa. E, anterior and posterior
dorsal plates of 5th segment. F, anal appendages. G, tips of anal appendages further enlarged
(setae of left side cut short). H, labrum (long setae shown cut to half actual length). J, mandible.
K, labrum of A. prionii. L, mandible of same. M, protuberances on 1st abdominal segment.
N, anal appendages. P, tip of anal appendage further enlarged. Q, anterior and posterior dorsal
plates of 5th segment. R, labrum of A. tuckeri ? var. (most setae missing). S, mandible of same.
T, anal appendages. U, tips of same further enlarged. V, anterior and posterior plates of 5th
segment. W, protuberances of 1st abdominal segment. X, scabrosity from patch on ventral
surface of abdomen.
28 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 6. Setodes barnardi sp. n., § and 9 imagos.
A, B, fore- and hind-wings of g. C, D, E, lateral, dorsal and ventral views of 3 genitalia.
F, G, H, lateral, ventral and dorsal views of 2 genitalia. J, base of J antenna.
SOME NEW CADDIS FLIES FROM THE WESTERN CAPE PROVINCE 29
Genitalia 3 (fig. 6 C—E): Ninth segment narrow dorsally, broad ventrally,
ventral margin (in lateral view) convex, slightly hairy, side-pieces slightly
produced, thickened marginally, set with long coarse setae; in ventral view the
posterior margin of the sternite is slightly excised and heavily sclerotized. Dorsal
apical margin produced as a narrow transverse band, sparsely setose, which
overlies the 10th segment. Apical margin of 1oth segment produced to form a
pair of long processes; these are joined basally, widely separated apically, with
a V-shaped emargination between them. Each process forms a thin, curved,
vertical plate, sinuous in lateral view and ending in a sharp beak-like apical
point. Beneath these the lower part of the roth tergite is divided to form a pair
of large, deep, transparent upper penis covers; these are sub-triangular in lateral
view, and in dorsal view they end in blunt points close to or overlying the
claspers. Each bears a few setae and a small dorsal process tipped with a seta.
¥
iS
arnt
a
cee
é.
ie
Fic. 7. Larva of Setodes barnardi sp. n.
A, larval case. B, head of larva from behind showing gular sclerite. C, dorsal view of head.
D, dorsal view of thoracic nota. E, left and right mandibles from above. F, maxillae and labium.
G, lateral view of posterior end of larva (right side). H, dorsal sclerite of gth segment. J, anal
claw.
30 ANNALS OF THE SOUTH AFRICAN MUSEUM
Aedeagus short, stout, downcurved, with a U-shaped sclerotized thickening.
Beneath it lie the paired, down-curved lower penis covers, ending in triangular
points. Claspers bent inwards, two-branched; the main part extends backwards,
then turns sharply inwards and upwards, ending in a sharp point; the branch
arises about a third of the way along it, and is upwardly directed, with a narrow
base and wider, triangular apex; it bears a number of strong setae.
Genitalia 2 (fig. 6 F-H): Eighth sternite flattened, ending in a wide shallow
excision; ninth sternite hollowed, with lateral margins of tergite forming flaps
on each side; ninth tergite transversely ridged, sparsely setose, fused to the
10th tergite which is rounded, with the central third backwardly produced and
emarginate; in lateral view it is beak-like. Below this are paired, hollowed out,
sclerotized plates, dorsally ridged, sharply triangular as seen from the side.
Lateral gonapophyses of 9th segment scoop-shaped, pubescent; between them
another plate, in lateral view sharply pointed, in ventral view a truncate
triangle.
3 holotype in alcohol, the genitalia mounted as a microscope preparation.
© paratype and g paratypes in alcohol.
This species is easily recognizable by the fact that fork 1 is lacking in the
posterior wing, only fork 5 being present, and by the tibial spurs, which are
I, 2, 2, instead of 0, 2, 2; the spur on the fore-tibia is small. The ¢ genitalia
also differ considerably from those of other African species of the genus,
showing most resemblance to S. baccata Kimmins; the 3 however lacks cerci and
shows greater complexity in the development of the roth tergite, the 9th sternite
is much broader and the main branch of the clasper differently shaped.
After much consideration it appeared best to include this species in the
genus Setodes, in spite of the fact that it differs from the generic diagnosis as
given by Kimmins (1949) in two respects, namely the absence of fork 1 in the
hind wing, and the presence of a small spur on the fore-tibia. Mr. Kimmins has
kindly informed me (personal communication) that his species S. baccata also
has a minute spur on the fore-tibia. In the other species of Setodes (restricted
sense) so far described, fork 1 is always present in the hind-wing, although it is
often quite small. In all other respects however S. barnardi appears to be a typical
Setodes, so has been included in that genus.
Locality: Larvae were collected from the Great Berg River near Drie-
fontein in October; adults were bred out in the laboratory from some of these
larvae as follows: October 1956, 1 3, 1 9; December 1956, 1 g, 1 9; January
1957, I g, 1 2. Most of the larvae were collected from the trailing marginal
palmiet (Prionium serratum), a few were found living under stones in the stickles.
Larva (fig. 7 B-J, fig. 8 A-C): Described from entire specimens. Length of
larvae up to 5:0 mm. Head: hypognathous, small, almost oblong, clypeus
asymmetrical with 2 to 3 pairs of lateral indentations and 12 setae; gular
sclerite rounded anteriorly, tapering to a point posteriorly, not fused with genae.
Head capsule and labrum light golden brown, set with small setae near the
margin of the clypeus. Antennae long, with small basal segment and tipped
a ea ee Mu
SOME NEW CADDIS FLIES FROM THE WESTERN CAPE PROVINCE 31
v,
eal
wadanne
OES az y
Be
ES Ky
Fic. 8. Larva and pupa of Setodes barnardi sp. n.
A, B, C, fore-, mid- and hind-legs of larva. D, pupal case. E, lid of same (inside view).
F, mandible of pupa. G, labrum (long setae cut short). H, pre- and postsegmental dorsal plates
of 5th abdominal segment. J, protuberances from 1st abdominal segment. K, anal appendages.
L, tips of anal appendages further enlarged.
32 ANNALS OF THE SOUTH AFRICAN MUSEUM
with a sensory bristle. Eyes large, dark, set well forward, with a pale patch just
behind each, and beneath hyaline lenses as in the other species described.
Suture-like line present paralleling the epicranial arms. Mouthparts : labrum with
two pairs of blade-like bristles anteriorly and 3 pairs of longer setae across the
top. Mandibles strong, short, heavily sclerotized, each with 5 blunt teeth, the
left mandible with a brush of strong bristles on the inner side. Maxillary palp
3-segmented, with apical papillae, junction with maxilla indistinct; maxillary
lobe fringed with setae and blade-like bristles. Labium rounded, with silk gland
opening at tip, labial palps stout, tipped with papillae. Thorax: pronotum light
brown, sclerotized, setose, with lateral fracture separating posterior edge;
mesonotum with a pair of thinly sclerotized yellowish-grey plates; metanotum
membranous, with an irregular band of setae across the ventral side. Legs:
small, slender, yellowish in colour, set with setae and bristles as in figure 8
A-C, but without special swimming hairs. Femora and trochanters of mid- and
hind-legs divided, also trochanters of fore-legs; tibiae and tarsi undivided;
claws long and strong, each with a basal bristle. Stridulator large, stout, blunt,
with a small seta. Abdomen: long, white, thick, without gills, lateral line very
poorly developed, chitinous points on segment 8 present but minute. On the
gth segment a semicircular dorsal sclerite bordered posteriorly with setae. Anal
appendages fused to form 1oth segment, anal claws minute, blunt, at the base
of each a large rectangular plate which is armed posteriorly with a frieze of
strong brownish-black spines set in 5-6 parallel rows. Anus flanked by rows of
small colourless setae.
Case (fig. 7 A): A neat, slender, curved tube about 5:0 mm. in length, of
small yellowish sand-grains, open at the ends and slightly hooded anteriorly.
Pupa (fig. 8 F-L): Labrum rounded, set with long bristles and smaller
setae; mandibles small, minutely serrulate along the distal third of the inner
border, condyles lateral; one pair of lateral bristles. Antennae much longer than
body, distal ends coiled; mid-tarsi broad and heavily fringed. Dorsal plates
present on abdominal segments as in Athripsodes species described; all pre-
segmental plates have 3 teeth. On goth abdominal segment a pair of setose
lappets; these are set on raised humps. Lateral lines extend from segment 3 to 8,
where they meet ventrally. Anal appendages slender with a few spines on the
proximal half; tips pointed and proximal to them 3 strong setae.
Pupal case (fig. 8 D, E): A cylindrical tube about 5:5 mm. long, neatly
made of yellowish sand-grains and pointed at both ends; a small opening is left
between the terminal sand-grains at each end, this is guarded by a membrane
whose inner edge forms stiffly projecting flaps. The result is rather like the
meandrine slit found in Leptecho helicotheca, but in Setodes barnardi the slit is not
visible from outside, being completely roofed over by sloping sand-grains with
only narrow cracks between them. In the diagram (fig. 8 E) the lid of the tube
is shown from the inside, and the band of secretion which fastened it to the case
can be clearly seen. The case is not completely lined by secretion, the secretion
merely being used to cement the sand-grains together.
3B
SOME NEW CADDIS FLIES FROM THE WESTERN CAPE PROVINCE 33
SUMMARY
A new species of caddis, Setodes barnardi sp. n. (Trichoptera: Leptoceridae),
is described from South Africa, together with its larval and pupal stages; also
the larval and pupal stages of Athripsodes prionii Scott, A. bergensis Scott, and
Leptecho helicotheca Scott, and the pupal skin of A. tuckeri Barnard ? var.
REFERENCES
BARNARD, K. H. 1934. South African caddis-flies (Trichoptera). Trans. roy. Soc. S. Afr. 21,
291-394.
BARNARD, K. H. 1940. Additional records, and descriptions of new species, of South African
alder-flies (Megaloptera), may-flies (Ephemeroptera), caddis-flies (Trichoptera), stone-
flies (Perlaria) and dragon-flies (Odonata). Ann. S. Afr. Mus. 32, 609-661.
BETTEN, C. 1934. The caddis flies or Trichoptera of New York State. Bull. N.Y. St. Mus. 292,
3-576.
Harrison, A. D., & Etswortn, J. F. 1958. Hydrobiological studies of the Great Berg River,
Western Cape Province. Part I. Trans. roy. Soc. S. Afr. 35, 125-226.
Harrison, A. D. 1958. ibid., Part II. Trans. roy. Soc. S. Afr. 35, 227-276.
Hicxin, N. E. 1943. Larvae of the British Trichoptera. 19. Setodes argentipunctella McLachlan
(Leptoceridae). Proc. R. ent. Soc. Lond. (A). 18, 109-111.
Hicxin, N. E. 1946. Larvae of the British Trichoptera. Trans. R. ent. Soc. Lond. 97, 187-212.
Hicxin, N. E. 1949. Pupae of the British Trichoptera. Trans. R. ent. Soc. Lond. 100, 275-289.
Hick, N. E. 1953. Larvae of the British Trichoptera. 38. Proc. R. ent. Soc. Lond. (A). 28, 111-113.
Kimmins, D. E. 1949. Some changes in the generic names in the family Leptoceridae (Order
Trichoptera). Entomologist 82, 201-204.
Kimwmins, D. E. 1956. New and little-known species of the Leptocerinae (Trichoptera) from the
African mainland (south of the Mediterranean region). Trans. R. ent. Soc. Lond. 108,
117-146.
Kimmins, D. E. 1957. New and little-known species of African Trichoptera. Bull. Brit. Mus.
(Nat. Hist.) Entomology. 6, 1-38.
Mose.ty, M. E. 1931. Some new Trichoptera from Africa and British Guiana. Trans. R. ent.
Soc. Lond. 79, 545-551.
Mosgty, M. E. 19394. The British caddis flies (Trichoptera). London: G. Routledge & Sons Ltd.
Mose ty, M. E. 1939). Trichoptera. (Mission scientifique de ’Omo, V.) Mém. Mus. Hist. nat.
Paris (N.S.) 9, 293-301.
NavaAs, L. 1931. Insectos del Museo de Paris. Brotéria, Zool. 27, 114-135.
Ross, H. H. 1944. The caddis flies, or Trichoptera, of Illinois. Bull. Ill. nat. Hist. Surv. 23, 1-326.
Scott, K. M. F. 1955. Some new caddis flies (Trichoptera) from the Western Cape Province
—I. Ann. S. Afr. Mus. 41, 367-380.
Scott, K. M. F. 1958. Ibid.—II. Ann. S. Afr. Mus. 44, 39-52.
Umer, G. 1955. Kocherfliegen (Trichopteren) von den Sunda-Inseln. Teil II. Arch. Hydrobiol.
(Plankt.) Suppl. 21 408-608.
wPRAS A
veo Pe ae i) |
ameriel).
?
shee
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
HartMAN, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8,1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
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Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XLVI
PART III
SOME SOUTH AFRICAN CLADOCERA COLLECTED
BY DR. A. D. HARRISON
By
J. P. Harpinc
British Museum (Natural History)
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-..1,, [CONTINUED ON INSIDE, BACK COVER
SOME SOUTH AFRICAN CLADOCERA COLLECTED
BY DR. A. D. HARRISON
By
J. P. Harpinc
British Museum (Natural History)
(With 14 figures in the text)
The material consisted of 61 tubes from the western Cape Province,
45 tubes from the Vaal River, and some of its tributaries, 58 tubes from the
Jukskei-Crocodile system, and 21 tubes from other parts of the Transvaal.
Dr. Harrison had already sorted the collections into species and made prelimi-
nary determinations, very considerably reducing the burden of identification.
Thirty-eight species of Cladocera were present, as listed below, those with an
asterisk seem. to be new to the South African fauna, but some of these may be
recorded in the literature under other names.
1. Diaphanosoma excisum 20. Bosmina longirostris
2. Daphnia barbata 21. Llyocryptus acutifrons*
3. Daphnia dolichocephala 22. Ilyocryptus sordidus
4. Daphnia magna 23. Macrothrix spinosa
5. Daphnia similis* 24. LEchinisca capensis
6. Daphnia longispina 25. EHurycercus lamellatus*
7. Daphnia obtusa 26. Camptocercus australis*
8. Daphnia pulex* 27. Acroperus harpae
g. Scapholeberis kingi 28. LEuryalona colletti
10. Scapholeberis aurita* 29. Alona affinis
11. Szmocephalus exspinosus 30. Alona bukobensis
12. Simocephalus serrulatus 31. Alona pulchella
13. Szmocephalus vetulus 32. Alona diaphana*
14. Ceriodaphnia reticulata 33. Leydigia microps
15. Certodaphnia quadrangula 34. Leydigia propinqua
16. Ceriodaphnia rigaudi 35. Pleuroxus aduncus*
17. Moina dubia 36. Chydorus globosus*
18. Moina belli 37. Chydorus sphaericus
19. Mona rectirostris* 38. Monospilus dispar*
None of the species is new to science, although the form of D. longispina
is one which seems to be typical for South Africa.
Diaphanosoma excisum Sars
This species was collected from the Vaal River in three places and also
from a vlei near Paarl. The specimens were not sufficiently well preserved for
the duplicature of the shell to be seen for certain, but the spinules on the edge
of the shell and the bristles near the post-dorsal corner leave no doubt of the
Ann. S. Afr. Mus. 46 (3), 1961, 35-46, 14 figs.
315)
MIT RSONIAN
meriTition JUN 7 1961
36 ANNALS OF THE SOUTH AFRICAN MUSEUM
identification. The species was collected from the Steenbras Reservoir by
Hutchinson ei al. (1932) and has been recorded from North, East and West
Africa as well. It is also known from Australia and the East Indies.
Daphnia barbata Weltner
This well-known African species was present in five tubes from the Vaal
Barrage, from the river below it, from the Kalkspruit near Balfour, and also
from the lower Berg River and the Cape Flats.
Daphnia dolichocephala Sars
Syn.: D. hodgsoni Sars
(Figs. 1-2)
Although there were only four specimens of this species from the Riet Vlei
off the Diep River, western Cape Province, and four specimens from the Vaal
Barrage, there was sufficient variation in the armature of the abreptor for it to
be possible to confirm Wagler’s (1936, p. 520) suggestion that D. hodgsoni and
D. doli:h cephala are one and the same species. The species resembles, in some
,
a 4
7 ’
Amel, rf
a Bo
4 % 5
4%
Fics. 1-2, Daphnia dolichocephala, Sars
1, head X72. 2, abreptor X 72.
SOME SOUTH AFRICAN CLADOCERA COLLECTED BY DR. A. D. HARRISON 37
ways, D. carinata and D. similis, but may be distinguished by the large number
of anal spines closely packed together and arising on the side of the abreptor,
a short distance from the dorsal edge (compare figs. 2 and 5). The species is
known from the Cape (Sars, 1916, p. 306) and from Kenya (Brehm, 1935,
P- 144). |
Daphnia magna Strauss
Specimens of this species were collected from a dam at the Benoni sewage
works in the Transvaal and from the De Hoop Vlei at Bredasdorp, western
Cape Province. The species is widely distributed throughout the Palearctic
and Ethiopian regions of the world, and in the northern and western parts of
North America.
Daphnia similis King
(Figs. 3-5)
This species was collected in small numbers in two places on the Jukskei
River. These specimens are very similar to those from Peru described by me
in 1955 (p- 330) as D. carinata, as I believed then that D. similis and D. carinata
e
oe ee ee eee ee
aS
Cee
Fics. 3-5, Daphnia similis King
3, head, lateral view; 4, head, dorsal view; 5, abreptor; all x 72.
38 ANNALS OF THE SOUTH AFRICAN MUSEUM
were synonyms. Brooks (1957, p. 32), however, gives reasons for believing that
they are distinct species. Certainly the shape of the antennule and rostrum of
specimens of D. carinata from Australia and Tasmania which I examined are
remarkably uniform and similar to Sars’s (1914) drawings, and in these respects
the South African specimens differ slightly, but I can find no other difference.
Brooks gives as an essential difference between D. carinata and D. similis the
different plane in which the dorsal ridge expands when it becomes larger.
This feature cannot be used with the present specimens since the dorsal ridge
is not expanded.
Daphnia longispina O. F. Miiller sensu lato
(Figs. 6-13)
About 40 females, some of them carrying ephippia, were collected from a
temporary vlei on the Wemmershoek Road, Paarl. These showed an extra-
ordinary range of shape of head (figs. 7-11, all taken from this sample and at
the same magnification). Figure 6 gives the general appearance of a whole
animal also from this sample at a smaller magnification. About 30 similar
specimens, mostly with tall helmets, were present in a sample from a flood-water
pool near the Berg River, Bridgetown, and other specimens were found in ones
Fics. 6-11, Daphnia longispina, O. F. Miiller, s. lato
6, female x25; 7-11, heads of females from the same sample as fig. 6 from a temporary
vlei near Paarl, x 45.
SOME SOUTH AFRICAN CLADOCERA COLLECTED BY DR. A. D. HARRISON 39
and twos at three other places on the Berg River and in a small lake near Paarl.
There is no doubt that these all belong to the same form of D. longispina as that
figured by Wagler (1936, p. 535, figs. 27-30), from the Rietfontein, Brak, and
Weltevreden Pans. Brooks (1957) has made a good case for treating D. longispina,
D. rosea, D. hyalina and D. galeata as separate species, and if more were known
about the present form it might be worth a new name. The very variable
nature of the shape of the head, however, makes it difficult at present to give
a satisfactory definition, and it seems best to treat it as a hyaline planktonic
form of D. longispina. The following features may be noted:
The body is colourless and hyaline with a long spine about equal in
length to the carapace projecting from the post-dorsal corner. The helmet,
if present, may be tall, but is usually evenly rounded. The dorsal outline
between the head and body is usually an even concave curve. This is
perhaps the most conspicuous feature of this form, and gives the Daphnia
a characteristic upright appearance. The eye is of moderate size and
usually at some distance from the ventral margin of the head, Spinules
are present on the posterior quarter of the dorsal margin and on the posterior
half of the ventral margin of the carapace. The ocellus may be small,
but is always present. The rostrum is pointed and the antennule is small
with sense hairs not reaching to the end of the rostrum. The shape of
the abreptor is shown in figure 12. The claw has three fine combs of the
typical longispina type (fig. 13).
Some juvenile specimens of a form of D. longispina were also found below
the Vaal Barrage.
Daphnia obtusa Kurz
Specimens of this species were collected from the Blaauwberg road bridge,
Milnerton, and from the Berg River at Hermon and Wellington. The specimens
from Milnerton had previously been sorted according to the length of the shell
spine into D. propingua and D. tenuispina, two species erected by Sars (1916,
p. 309) for obtusa-like Daphnias from the Cape. In this small sample there was
a whole range of forms from those without spines on the shell, which would be
called D. obtusa without question, to those with spines on the tail long enough
for D. tenuispina. These specimens, however, clearly all belong to the same species
and I consider them all to be D. obtusa; although there is a tendency for longer
spine forms to occur more frequently in South Africa than in Europe, I do not
think other names are justified for these forms. These specimens agree in every
respect with the description of D. obtusa given by Scourfield (1942) and Johnson
(1952, p. 446). The species is found in temperate waters in both hemispheres.
Daphnia pulex Leydig
Specimens which agree in every particular with Johnson’s (1952, p. 443)
description of the female and which run down to this species in Brooks’ key
40 ANNALS OF THE SOUTH AFRICAN MUSEUM
(1957, Pp. 30) were found at the Upper Laundry Dam at Benoni in the
Transvaal, and from the lower Berg River in the Cape at the farm Sanddrift.
This species is now known from all parts of the world except from the
Australian and Oriental regions; this is the first record from S. Africa.
Scapholeberts kingi Sars
This species was collected from six places on the Jukskei-Crocodile system
among marginal vegetation, and two femaies were also found in brown and
acid water in a vlei at Betty’s Bay. Sars (1916, p. 314) described specimens raised
from dried mud from Bergvliet.
5
el
13
yy
Fics. 12-13, Daphnia longispina O. F. Miller, s. lato
12, abreptor, X 150; 13, claw, x 600.
The species has been recorded from Senegal, Corfu, Germany, India,
China, Siam, the East Indies and Australia.
Scapholeberis aurita (Fischer)
Three females were collected from the marginal vegetation at the Vaal
Barrage, and one from the Riet Vlei off the Diep River.
The species seems to be new to South Africa. It is widely distributed in
Europe, Central Asia, North Africa, and North America, but is not a common
species anywhere.
SOME SOUTH AFRICAN CLADOCERA COLLECTED BY DR. A. D. HARRISON 41
Simocephalus exspinosus (Koch)
Syn.: S. australiensis Sars
A number of samples from the Transvaal including the sewage works at
Benoni and Pretoria, contained this species; it was also collected from the Salt
River and from the Riet Vlei in the western Cape Province.
This is a cosmopolitan species and I adhere to the opinion previously
expressed (Harding, 19574, p. 62) that there is no justification for regarding
specimens from the southern hemisphere as a separate species.
Simocephalus serrulatus (Koch)
Syn.: Szmosa capensis Sars
A few individuals were found in five samples from the Vaal, the Klein
Jukskei, and the lower Berg River; three of these samples also contained
S. vetulus.
This species is widely distributed in the temperate parts of the world, but
has not yet been recorded from Australia or New Zealand.
Simocephalus vetulus (O. F. Miller), sensu lato including S. vetuloides Sars
(Fig. 14, A-H)
This was the commonest species of Simocephalus in the collections, and was
present in 12 of the samples and in all the regions investigated. It is very difficult
: sae
NS
ro
SS
Fic. 14, Simocephalus vetulus (O. F. Miiller), s. lato.
Posterior margin of shell: A-H, South African specimens; I—P, specimens from England. x 30.
ie)
42 ANNALS OF THE SOUTH AFRICAN MUSEUM
to decide whether to recognize S. vetuloides as a distinct species from S. vetulus.
The only difference between the two given by Sars (1916, p. 313) is in the shape
of the posterior margin of the shell, S. vetulocdes having a protuberance which is
missing in §. vetulus. Most South African specimens have this protuberance as
can be seen from the camera lucida drawings of the posterior margins of the shells
of specimens from various South African localities (fig. 14, A-H). But the
specimens shown in figure 14, E, G and H, lack this protuberance. Figures I—-P
show a similar series of drawings of specimens from various English localities,
and while most of the specimens are without a protuberance, those in figures
I and O show one. The overlap seems to be too great to justify the separation
into two species, but there are samples in which there would be no difficulty in
allocating one or other name. Until breeding experiments have been carried
out, it will probably be impossible to decide this question.
Ceriodaphnia reticulata (Jurine)
Syn.: C. natalis Brady
C. reticulata var. minor Sars
Specimens of this species were present in samples from the lower Berg River
at Bridgetown, from among Potamogeton at a weir at Olifants Vlei, from the
marginal vegetation at the Hartebeestpoort Dam. Sars (1916, p. 316) dis-
tinguished a var. minor solely on the small size—‘reaching 0-9 mm.’, and this is
about the limit of size for the present specimens. No large specimens of C. reticulata
seem ever to have been collected from South Africa, but this hardly justifies
the use of a separate name. Lilljeborg (1900, p. 184) gives 0-8—1-44 mm. as the
range of length of the adult female of this species in Sweden, and refers also to a
population of small specimens whose sizes range from 0-5 to 0-8 mm. In a foot-
note on p. 188 he informs us that he was at first inclined to erect a new species
for this population and in a public lecture suggested the name C. minor; but
that later he considered this to be unjustified.
The species is widely distributed in the Palearctic, Nearctic, Neotropical
and Ethiopian regions.
Ceriodaphnia quadrangula (Miller)
Parthenogenetic females and a few males were present in brown and acid
water from a vlei at Betty’s Bay. The species has the same geographical distribu-
tion as C. reticulata.
Certodaphnia rigaudi Richard
Syn.: C. cornuta Sars
About one dozen specimens were collected on different occasions from the
Amato Dam, Benoni. The species is found in all the warmer regions of the world.
SOME SOUTH AFRICAN CLADOCERA COLLECTED BY DR. A. D. HARRISON 43
Moina dubia Guerne and Richard
Moina dubia was collected from the Zeekoe Vlei, Cape Flats; from the
Berg River at Wellington, and Piketberg; and from several places in the
Transvaal.
Sars (1916, p. 322) described the claw of the abreptor as being without
secondary denticles, but this is a very variable character in this species (see
Gauthier, 1955, p. 26), and the present specimens showed every gradation.
The species is cosmopolitan.
Moina belli Gurney
Five females were collected from the Aapies River at Onderstepoort.
Gurney (1905, p. 299) described the species from Kroonstad; Brehm (1938,
p- 212) described what is probably the same species under the name M. rutineri
from Kurdistan, and Stephanides (1948, p. 17) records the species from Corfu.
- Moina rectirostris Leydig
Syn.: Moina brachiata Sars not Jurine
This species was present in five tubes from the Transvaal and two tubes
from the Milnerton area of Cape Province. It is widely distributed in the
Palearctic, Nearctic, and Ethiopian regions. Sars (1916, p. 321) called this species
M. brachiata as he thought it was the species described by Jurine (1820, p. 131)
as Monoculus brachiatus; but as Gauthier (1955, p. 15) points out, Jurine described
two eggs in the ephippium of the female, which therefore could not have
belonged to this species.
Bosmina longirostris O. F. Miiller
Samples from the Vaal, Hennops and Berg Rivers contained this cosmo-
politan species, which is the Bosmina most often found in all parts of Africa.
Ilyocryptus acutifrons Sars
About 30 specimens were collected from the Sirkels Vlei near Cape Point.
This is a Palearctic and Nearctic species which seems not to have been recorded
from Africa before.
Tlyocryptus sordidus (Liéven)
This common species can probably be found in mud in any part of South
Africa, and was present in 12 of the samples. It is cosmopolitan.
Macrothrix spinosa King
This species was found in a number of places in the Transvaal and the
-Cape, eight samples in all. Sars records it from the Cape and it is known also
from Australia and South America.
44 ANNALS OF THE SOUTH AFRICAN MUSEUM
Echinisca capensis Sars
A few specimens were collected on three occasions from the Riet Vlei and
a flooded backwater of the Bushman’s River. It seems to be endemic to the
Cape Province.
Eurycercus lamellatus (O. F. Miller)
This large cladoceran was found in the vlei near Paarl, in the lower Berg
River at Hermon and at Southfield on the Cape Flats. It is common in the
Palearctic and Nearctic regions and is known from South America and North
and West Africa, but these are the first South African records.
Camptocercus australis Sars
A single female was collected from a vlei near Paarl. C. australis is known
from Australia, India, and Argentina. It differs from the C. rectirostris Schoedler
of Palearctic and Nearctic regions only in the absence of small teeth on the
post ventral corner of the shell. The latter species has been recorded from North
Africa.
Acroperus harpae (Baird)
Over 100 specimens were collected from the Great Usutu River, Transvaal.
This species appears to be cosmopolitan except for Australia. Many different
forms have been described, of which the form neglectus Lilljeborg best fits the
present specimens.
Euryalona colletti (Sars)
Only two females were collected, both from the Vaal. One at the Suikerbos
pumping station and one below the confluence with the Leeuwspruit. Previous
records are from Knysna and Zululand (see Harding, 19570).
Alona affinis (Leydig)
Specimens of this species were found on four occasions on various parts of
the Berg River and the Usutu River. It is a well-known cosmopolitan species.
Alona bukobensis Weltner
Considering that this species is known from most parts of Africa it is rather
surprising that only one adult female from Benoni and one juvenile from Paarl
can be identified as such. Outside Africa, Ekman (1905, p. 8) has described the
variety subantarctica from South Georgia.
Alona pulchella King
Specimens were found in various parts of the Vaal River and at Benoni.
The species, often recorded under the synonyms A. glabra or A. cambouei
4A
SOME SOUTH AFRICAN CLADOCERA COLLECTED BY DR. A. D. HARRISON 45
(see Harding, 1955; 1957a, p. 76), is known from all tropical and sub-
tropical regions of the world.
Alona diaphana (King)
This variable, but easily recognized, species was found in a number of
places on the Vaal River and the Klein Jukskei and at Benoni. It is known from
North, East and West Africa, and from South America and Australia, but does
not seem to have been recorded from South Africa before.
Leydigia microps Sars
Four females were collected from the Vaal River below its confluence with
the Klip River. This seems to be the first record of the species since Sars
(1916, p. 330) first described it from three places in the Cape Province.
Leydigia propinqua Sars
This was found in small numbers from a number of places on the Crocodile-
Jukskei system, in the Vaal, and in the Krom River at Stellenbosch. Originally
described from Sumatra, it has been recorded from South Africa (Sars, 1916,
p. 329) and East Africa (Jenkin, 1934, p. 285).
Pleuroxus aduncus ( Jurine)
This cosmopolitan species was the most common in the collections and
was present in large numbers in 22 of the tubes. These were all from the Vaal
and ‘Transvaal except for one sample with 6 specimens from the vlei at Betty’s
Bay. The specimens have from 1 to 3 teeth on the post ventral corner of the
shell, and also have a peg on the antennule as described by Jenkin (1934, p. 296)
for the var. makaliensis. This peg has been shown to be typical for P. aduncus
(Harding, 1955, p. 348) and is also present in the type specimens of P. inermis,
which is the only species of Plewroxus described for South Africa by Sars, and is
probably identical with P. aduncus.
Chydorus globosus (Baird)
Four females were found in the Klein Jukskei River. Although compara-
tively rare, this species is probably cosmopolitan.
Chydorus sphaericus (O. F. Miller)
This cosmopolitan species was found in nine of the samples and is clearly
present in all the regions investigated.
Monospilus dispar Sars
Three females were found in the vlei at Betty’s Bay in brown and acid water.
It is widely distributed in the Palearctic, Nearctic and Ethiopian regions, but
is not often found.
46 ANNALS OF THE SOUTH AFRICAN MUSEUM
REFERENCES
BrREHM, V. 1935. Crustacea. I. Cladocera und Euphyllopoda. Miss. sci. Omo. 2: 141-166.
BrEHM, V. 1938. Uber die Siisswasserfauna von Kurdistan. I. Zool. Anz. 121: 209-219.
Brooks, J. L. 1957. The systematics of North American Daphnia. Mem. Conn. Acad. Arts Sci. 13:
1-180.
EKMAN, S. 1905. Cladoceren und Copepoden aus antarktischen und subantarktischen Binnen-
gewassern. Wiss. Ergebn. schwed. Stidpolarexped. 5, 4: 1-140.
GauTHIER, H. 1955. Essai sur la variabilité, V’écologie, le détérminisme du sexe et la reproduction de
quelques Moina (Cladocéres) récoltées en Afrique et a Madagascar. Alger.
Gurney, R. 1905. On a small collection of freshwater Entomostraca from South Africa. Proc.
zool. Soc. Lond. 1904, 2: 298-301.
Harpinc, J. P. 1955. The Percy Sladen Trust Expedition to Lake Titicaca in 1937. Crustacea;
Cladocera. Trans. Linn. Soc. Lond. (Xool.) (3) &: 329-354.
Haroinc, J. P. 1957a. Crustacea: Cladocera. Res. sci. Explor. Hydrobiol. Lac Tanganyika 3, 6:
55-89.
Haropinc, J. P. 19575. The South African Cladoceran Euryalona colletti (Sars) and another
African species. Ann. S. Afr. Mus. 43: 245-247.
Hutcuinson, G. E., Pickrorp, G. E., and SchuuRMAN, J. F. M. 1932. A contribution to the
hydrobiology of pans and other inland waters of South Africa. Arch. Hydrobiol. 24: 1-154.
Jenkin, P. 1934. Reports on the Percy Sladen Expedition to some Rift Valley lakes in Kenya
in 1929. VI. Cladocera from the Rift Valley lakes in Kenya. Ann. Mag. nat. Hist. (10) 13:
281-308.
Jounson, D. S. 1952. The British species of the genus Daphnia (Crustacea, Cladocera). Proc.
zool. Soc. Lond. 122: 435-462.
Jurine, L. 1820. Histoire des Monocles qui se trouvent aux environs de Genéve. Geneve & Paris.
LittjEBorG, W. 1900. Cladocera Sueciae. Uppsala.
Sars, G. O. 1914. Daphnia carinata King, and its remarkable varieties. Arch. Math. Naturv. 34, 1:
I-14.
Sars, G. O. 1916. The fresh-water Entomostraca of Cape Province (Union of South Africa).
Part 1: Cladocera. Ann. S. Afr. Mus. 15: 303-351.
SCOURFIELD, D. J. 1942. The ‘Pulex’ forms of Daphnia and their separation into two distinct
series, represented by D. pulex (de Geer) and D. obtusa Kurz. Ann. Mag. nat. Hist. (11) 9:
202-219.
STEPHANIDES, TI’. 1948. A survey of the freshwater biology of Corfu and of certain other regions
of Greece. Prakt. Hellen. hydrobiol. Inst. 2, 2: i-viii, 1-263.
Wac ter, E. 1936. Die Systematik und geographische Verbreitung des Genus Daphnia O. F.
Miiller mit besonderer Berucksichtigung der stidafrikanischen Arten. Arch. Hydrobiol. go:
505-556.
References thus appear as follows: 7 :
AUGENER, H. 1913. Polychaeta. Jn Wirchaekent W., ed. Beitrége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Sedat chosen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
HartMan, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8,1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sct. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-500.
SYNONYMY. arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.—All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name....
B. Form for bibliographic references to synonymic names.—The first reference following
any name in the synonymy should be tothe earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example :-
Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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The Laboratory, Citadel Hill, Plymouth, U.K.
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[ CONTINUED ON INSIDE BACK COVER
POGONOPHORA FROM SOUTH AFRICA
By
Eve C. SouTHWARD
The Laboratory, Citadel Hill, Plymouth, U.K.
(With 3 figures in the text)
INTRODUCTION
The Pogonophora, a group of tubicolous marine animals related to the
Hemichordata, have aroused much interest in recent years. A curious feature of
the group is the total absence of an internal digestive system, a condition which
is unique among non-parasitic Metazoa. The Class name Pogonophora was
suggested in 1937 by Johansson who chose the name to describe the beard-like
appearance of the tentacles of the only species known to him then (Lamellisabella
zachst Uschakov). A new Phylum, Brachiata, was erected for the group by
Ivanov (19550), who had already (1951) assigned to the Pogonophora a peculiar
animal described much earlier by Caullery (1914, 1944) and described several
new genera and species himself (Ivanov, 1949, 1952).
Our knowledge of the distribution of these animals is still very incomplete.
They seem to be restricted to soft sediments in deep, or at least cold, water.
Ivanov’s specimens have come, so far, from the north-west Pacific and Arctic
Oceans. Others have been collected in the East Indies and in the east Pacific
off the coast of Central America (Caullery, 1944; Kirkegaard, 1956a & b;
Menzies, Ewing, Worzel & Clarke, 1959). The first species was recorded from
Europe in 1956 (Jagersten) and more have been found since (Southward &
Southward, 1958; Brattstr6m, 1959; Southward, 1959).
A collection of pogonophore tubes made by the R.S. Africana ITI, off the
west coast of South Africa, is of great interest as the first record from the south
Atlantic and the most southerly record so far.
The deep-trawling was done by courtesy of the Director, Division of
Fisheries, Cape Town. The tubes were sent to me by Dr. F. H. Talbot of
the South African Museum, to whom I am very grateful for the opportunity to
examine the specimens. The tubes seem to belong to three species, but only one
tube was occupied by an animal. This one animal, though incomplete, shows
enough of the important taxonomic features to allow it to be used as the type
specimen of a new species.
The Trustees of the South African Museum wish to acknowledge a grant
from the Council for Scientific and Industrial Research towards the purchase
of the deep-trawling equipment with which the specimens reported on were
collected.
Ann. S. Afr. Mus. 46 (4), 47-52, 3 figs.
47
104
48 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family POLYBRACHIIDAE Ivanov 1952
Heptabrachia talboti n. sp.
Material: One occupied tube (holotype) and 7 empty fragments; position:
33° 26’ S., 16° 33’ E.; depth: 2,268 m.; date of collection: 26.8.59; S.A.M.
No. A 19704.
Three empty fragments of tube; position: 33° 50’ S., 17° 21’ E.; depth:
1,097 m.; date of collection: 25.8.59; S.A.M. No. A 19702.
Two empty fragments of tube; position: 33° 50’ S., 16° 30’ E.; depth:
2,750-2,890 m.; date of collection: 27.8.59; S.A.M. No. A 193.
Description: ‘The tubes are brown and stiff anteriorly, white and soft posteriorly.
Their diameter is a little greater at the anterior end. The tube of the holotype
is 0-46 mm. in diameter at the anterior end and 0-41 mm. posteriorly, while
the variation among the empty tubes is from 0-34 to 0-56 mm. The longest
tube is not complete and is 13 cm. long. The anterior part of the tube is marked
with wide brown rings, separated by narrower yellow rings (fig. 1, A), but
towards the middle part of the tube the brown rings become narrower and less
distinct (fig. 1, B) and finally disappear. The posterior part of the tube is
whitish and its rather thick walls are encircled by slightly wrinkled marks about
I cm. apart. One short fragment of tube with stiff, transparent, yellow walls
may be the extreme anterior end of one of the tubes of this species, although
other species of Heptabrachia have the anterior end of the tube colourless, with
thin, limp walls (Ivanov, 1957).
The animal has lost the distal parts of the tentacles and much of the
posterior part of the body. Although parts of the specimen are crushed the
following details can be seen. At the anterior end about 15 tentacles were
attached to the cephalic lobe of the protosoma, or first segment (fig. 2, A, B),
and about half of these tentacles were lying along the mesosoma (an unusual
position which may have been produced during fixation). These backward-
pointing tentacles had to be removed to expose the mesosoma (fig. 2, A) and
this procedure slightly damaged the anterior part of the specimen, so that it is
difficult to decide how the bases of the tentacles were arranged. In this specimen
it is not possible to see any pinnules on the tentacles, but many pogonophores
have them and they may be found in other specimens.
The protosoma is separated from the mesosoma by a circular groove,
behind and over which the anterior mesosoma is drawn out into a collar-like
flap (fig. 2, A, C). On the tentacular side* the collar is divided by a deep groove
which extends back, with a flap on each side, as far as the bridle. This bridle
is a ridge of thickened cuticle which encircles the mesosoma except for a small
gap on the tentacular side (fig. 2, A, C). Behind the bridle the mesosoma is
smooth. Together the protosoma and mesosoma are 1°52 mm. long and the
* The tentacular side of the animal is ventral according to Ivanov (1955a), dorsal
according to Jagersten (1956).
POGONOPHORA FROM SOUTH AFRICA 49
greatest width of either is 0-32 mm. The third, last, and longest segment of the
body is the metasoma, which in this specimen is 12 mm. long and certainly
not complete. The first part (1-7 mm.) is very much contracted and twisted,
and the closely packed papillae on the tentacular side (fig. 2, A) are probably
normally arranged in two rows separated by a longitudinal groove, as they
are in many other pogonophores. The paired arrangement of the papillae on
this part of the body in other pogonophores has earned it the name of the
‘metameric’ region. In the present specimen each papilla is crowned with an
l-O MM
Fic. 1
Tube of Heptabrachia talboti n. sp. A, anterior. B, middle. C, posterior.
oval adhesive plate with one thickened border (fig. 2, A, E). Behind the meta-
meric region is a muscular region bearing a few scattered papillae, with adhesive
plates (in a mature specimen this is the region which contains the gonads),
extending for about 10 mm. and ending with two girdles of small toothed
platelets (fig. 2, D). The specimen is broken off at the second girdle and is also
damaged at the level of the first, so that the extent and shape of the girdles in
the complete animal are not known. Each girdle is made up of several irregular
rows of platelets (fig. 2, F, G), of the shape and form found in many other
pogonophores. The part of the metasoma behind the girdles (postannular
region) is missing; in other species it is often as long as the preannular
region.
50 ANNALS OF THE SOUTH AFRICAN MUSEUM
For a complete description it will be necessary to have a specimen with the
metasoma in good condition, in order to discover the true arrangement of the
papillae in all parts, and to clarify the arrangement of the girdles. Complete
tentacles are needed for measurement of their length and for investigation of
their pinnules, if present. |
However, with the material available, it is possible to place the species, with
reasonable certainty, in the genus Heptabrachia Ivanov 1952, and to conclude
that it is unlike any of the four species so far known. H. abyssicola, H. gracilis,
H. beringensis and H. subtilis were all described by Ivanov and he has summarized
their characteristics in a table (Ivanov, 1960, p. 214). H. gracilis is the closest in
1:O MM
A B
Fic. 2
Heptabrachia talboti n. sp. A, anterior end, tentaculate side, after removal of tentacles. B, anterior
end, before removal of tentacles. C, anterior end, non-tentaculate side. D, girdle region of
metasoma. E, adhesive plate. F, G, side and surface views of girdle platelet. Diagonal hatching
indicates damaged surfaces. Abbreviations: b. bridle, cl. cephalic lobe, g. 1 first girdle, g. 2
second girdle, gr. 1 groove between proto- and mesosoma, gr. 2 groove between meso- and
metasoma, mes. mesosoma, p. papilla, t. tentacle.
POGONOPHORA FROM SOUTH AFRICA 51
general appearance to H. talbotz, but, in addition to differences in size and the
number of tentacles, they differ in the size of the adhesive plates which are 15 to
20 pw long in H. gracilis and 30 to 40 pw long in H. talboti. Examination of more
specimens of H. talbott might disclose more differences, since all four of Ivanov’s
species have, on part of the preannular metasoma, an area or row of large
papillae set close together, whose arrangement and number varies with the
species. In H. talbott some large papillae are visible in the correct region but
their arrangement cannot be seen clearly. Another useful specific character
in this genus is the arrangement of the papillae on the postannular region,
which has been lost by the present specimen.
MM
O-l
A B
Fic. 3
A & B, two ends of tube type (a). C, tube type (0).
Empty TuBEs oF UNKNOWN GENERA AND SPECIES
Material: Three tubes; position: 34° 05’ S., 16° 58’ E.; depth: 2,690 m.;
date of collection: 9.12.59; S.A.M. No. A 319.
The tubes are of two types: (a) almost colourless, transparent, with one
end slightly wrinkled but not ringed and the other end marked with wide
greyish rings (fig. 3, A, B). There are two specimens of this type, one Io cm.
long by 0-195 mm. diameter, the other 2-5 cm. by 0-145 mm. (b) reddish
brown in colour, with well-marked brown rings, separated by narrow yellow
rings (fig. 3, C). The diameter is 0:12 mm. and the length is 2 cm.
The holotype and other specimens of Heptabrachia talboti, and the unidenti-
fied pogonophore tubes, are all in the collection of the South African Museum,
Cape Town.
52 ANNALS OF THE SOUTH AFRICAN MUSEUM
SUMMARY
The first member of the Pogonophora to be described from the south
Atlantic is Heptabrachia talboti n. sp., collected off the west coast of South Africa
in depths between 1,097 and 2,890 m. Empty tubes of two other pogonophores
are also described.
REFERENCES
BRATTSTROM, H. 1959. Pogonophora in the Hardangerfjord, Western Norway. Nature, Lond.,
183, 1458.
CauLuery, M. 1914. Sur les Siboglinidae, type nouveau d’invertébrés recueilli par l’expedition
du Siboga. C. R. Acad. Sci., Paris, 153, 2014-2016.
CaAuL_eRyY, M. 1944. Szboglinum Caullery 1914 type nouveau d’invertébrés, d’affinités a Breency
Siboga Exped., 25 bis, 26 pp.
Ivanov, A. V. 1949. A new representative of the class Pogonophora. Zool. Zh. S.S.S.R., 28,
78-84 (in Russian).
Ivanov, A. V. 1951. On the affiliation of the genus Szboglinum Caullery with the class Pogono-
phora. Dokl. obsch. Sobr. Ak. Nauk S.S.S.R., 76, 739-742 (in Russian).
Ivanov, A. V. 1952. New Pogonophora from far eastern seas. Zool. Zh. S.S.S.R., 31, 372-391
(in Russian). Translation in Syst. Zool., 3, 69-79.
Ivanov, A. V. 1955a. The main features of the organisation of the Pogonophora. Dokl. obsch.
Sobr. Ak. Nauk S.S.S.R., 100, 175-177 (in Russian). Translation in Syst. Zool., 4, 172-174.
Ivanov, A. V. 19555. On the assignment of the class Pogonophora to a separate phylum of the
Deuterostomia— Brachiata A. Ivanov, phyl. nov. Dokl. obsch. Sobr. Ak. Nauk S.S.S.R., 100,
595-596 (in Russian). Translation in Syst. Zool., 4, 177-178.
Ivanov, A. V. 1957. Neue Pogonophora aus dem nordwestlichen Teil des Stillen Ozeans.
Kool. Fb., Abt. 1, 85, 430-500.
Ivanov, A. V. 1960. Pogonophora. Fauna Rossii, 75, 1-271 (in Russian).
JAGERSTEN, G. 1956. Investigations on Siboglinum ekmani, n. sp., encountered in Skagerak, with
some general remarks on the group Pogonophora. ool. Bidr. Uppsala, 31, 211-252.
Jouansson, K. E. 1937. Uber Lamellisabella zachsi und ihre systematische Stellung. Zool. Anz.,
117, 23-26.
KiIRKEGAARD, J. B. 1956a. Pogonophora. Galathealinum bruuni n. gen. n. sp., a new representative
of the class. Galathea Rep., 2, 79-84.
KIRKEGAARD, J. B. 19565. Pogonophora. First records from the eastern Pacific. Galathea Rep.,
2, 183-186.
Menzigs, R. J., Ewinc, M., WorzeEL, J. L., & CLARKE, A. H. 1959. Ecology of the recent Mono-
placophora. Ozkos, 10, 168-182.
SOUTHWARD, E. C. 1959. Two new species of Pogonophora from the north-east Atlantic. 7. mar.
biol. Ass. U.K., 38, 439-444.
SouTHWARD, E. C., & SourHwarp, A. J. 1958. On some Pogonophora from the north-east
Atlantic, including two new species. 7. mar. biol. Ass. U.K., 37, 627-632.
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. Jn Michaelsen, W., ed. Beitrdge zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
HartTMAn, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sct. 8,1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-500.
SYNONYMY. arrangement according to Schenk, E. T. & McMaster, J. H.:
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pagination and illustrations (plates and figures).
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Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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ANNALS
OF CHE
SeOTivAPRICAN MUSEUM
VOLUME XLVI
PART V
A NEW CICHLID FISH IN THE LIMPOPO BASIN
By
ETHELWYNN TREWAVAS
British Museum (Natural History)
=
inane
AUG 1 8 1961
OT HSONIAN peroe
ISSUED JUNE 1961 PRICE 100
PRINTED FOR THE
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LIBRARY
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MSS. submitted for publication must be typewritten, double spaced with good
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conclusions. The position of the text-figures and tables must be clearly indicated.
ILLUSTRATIONS. Please must be kept to a minimum and made up to occupy
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REFERENCES. Harvard system—authors’ names and dates of publication given
in the body of the text; references arranged at the end of the paper in alphabetical
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SUM <eSan NORelG. ane: USEC:
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[CONTINUED ON INSIDE BACK COVER
A NEW CICHLID FISH IN THE LIMPOPO BASIN
By
ETHELWYNN 'TREWAVAS
British Museum (Natural History)
The fish to be described has been known as Serranochromis thumbergi
(Castelnau), but I recognized it as distinct some years ago when I handled a
specimen from the collection of the Transvaal Museum. I was unwilling to
describe it from a single specimen, but I have since received five more from
Mr. S. S. du Plessis which enable me to characterize the species. Although
related to Serranochromis it combines characters which suggest an independent
though neighbouring point of divergence from Haplochromis and I have given
it a new generic name.
ACKNOWLEDGEMENTS
I am grateful to Dr. V. Fitzsimons of the Transvaal Museum for the loan
of a specimen of this species and to Mr. S. S. du Plessis for supplying the five
which have been described as holotype and paratypes. I acknowledge too the
patience of interested fishery research and development officers who have
waited while I studied specimens of Haplochromis and related genera from the
Zambezi and neighbouring basins so that I could judge the systematic position
of this species on the broadest basis before naming it. I have seen types of most
of the species of this region and am satisfied that it can be identified with none
of them.
Chetia gen. n.
Type C. flaviventris sp. n.
From Cheti, a name given in Nyasaland to a yellow weaver-bird, in
reference to the yellow colour on the type species and by analogy with the
Afrikaans name kanariekurper.
Piscivorous cichlids resembling Haplochromis, but without any ventral
vertebral apophysis for the attachment of the air-bladder; differing from
Serranochromis in retaining a minor cusp on at least some of the teeth of the
jaws up to a standard length of 70 mm. and in having usually a lower number of
vertebrae (31-33) and of soft dorsal rays (11-12). As in both these genera, the
apophysis for the upper pharyngeal bones is formed from the parasphenoid
in the middle and the basioccipital at the sides. Some of the scales are finely
ctenoid in the young, but at the sizes examined most or all are cycloid, often
with a small group of fine granulations posteriorly. The caudal fin is truncate.
39
Ann. S. Afr. Mus. 46 (5), 1961, 53-56
54 ANNALS OF THE SOUTH AFRICAN MUSEUM
Lichnochromis, a monotypic Nyasan genus which also falls within most of
this definition, differs in its typical Nyasan colour-pattern of an oblique dark
band from nape to caudal, and in the emarginate and heavily scaled caudal
fin, as well as in the produced snout (more than half of the length of head)
and expanded, beak-like premaxillary bones.
Chetia flaviventris sp. n.
Serranochromis thumbergi (nec Castelnau), du Plessis & Groenewald, 1953, p. 41, 2 figs.
Description of six specimens 71-117 mm. in standard length. ‘The propor-
tions in square brackets are from du Plessis & Groenewald’s figure of a male,
evidently a larger specimen.
Proportions in hundredths of standard length:
Depth of body 29-6-33-3 [35-5]
Length of head 33-5—35-4 [31:3]
Length of caudal peduncle 15-17
Length of pectoral fin 21-7—24-7
Proportions in hundredths of length of head:
Length of snout 34-0—38-0
Diameter of eye 21-25~-26-0 [19:2], negatively allometric
Depth of preorbital 18-0—22°5 [23-1], positively allometric
Interorbital width 18-7—21-1
Length of lower jaw 43:°4—47°9
Length of premaxillary pedicels 30-0—33°8
Width of lower pharyngeal bone 25:0 (1 specimen)
Snout weakly and evenly decurved. Cleft of mouth at an angle of 20°—30°
with the horizontal; maxillary extending to below some part of anterior half
of eye. Five horizontal series of scales on the cheek. Gill-rakers on the anterior
arch 3 or 4 + 1 + 9 or 10. Teeth in 2 series with a few anterior teeth forming
a third series in upper jaw; in lower jaw in 2 series, the inner of up to 12
teeth; 44-50 teeth in outer series of upper jaw. All teeth conical, curved in the
specimens of more than 100 mm.S.L., part conical and part spear-shaped with
a single point in two of the others, some spear-shaped and some with a main
and a minor cusp in two specimens of 71 and 85 mm. s.l.
Lower pharyngeal narrow, with a short anterior blade and slender pointed
teeth with a minor cusp or shoulder.
Dorsal XV 11 (one) or XV 12 (five). Anal III 9 (three) or III 10 (three).
Caudal truncate, in larger specimens with rounded corners. Caudal peduncle
1*3 to 1-5 times as long as deep.
Scales 34 in a longitudinal series including the upper lateral line, 5-6
between origin of dorsal and lateral line, 5 or 6 between bases of pectoral and
pelvic fins.
Vertebrae 31-33, comprising 15 abdominal and 16-18 caudal.
A NEW CICHLID FISH IN THE LIMPOPO BASIN 5D
Colour pattern in preserved specimens: a shadowy mark between the eye
and maxillary (lachrymal mark); a dark opercular spot and a series of dark
blotches from this to the base of the caudal along the middle of the side, more
or less connected posteriorly to form a band; a less complete series on the upper
lateral line and a vague series from nape along base of spinous dorsal. [These
body-markings commonly appear in dead or dying specimens of Haplochromis
and its relatives and also in certain physiological or emotional states, and are
generally more marked in the young.] Soft dorsal and caudal with series of
small dark spots.
Life-colours (du Plessis & Groenewald, 1953): belly bright yellow (giving
it the vernacular name of kanariekurper); anal fin with bright red dots which
are more plentiful in males than in females.
Breeding: a mouth-brooder, the female carrying the eggs (du Plessis &
Groenewald, 1953).
Food: the adult dentition is piscivorous and du Plessis & Groenewald
report that ‘the fish is a predator, thriving on small fish, insects and snails’.
Since these authors identified this species with Serranochromis thumbergi, which
was once mistakenly reported as feeding on snails, this account of the kanarie-
kurper’s diet should not be accepted without confirmation on the Limpopo
populations.
Distribution: known only from the Limpopo basin, ‘mostly in the dry
western part of the Transvaal’, where ‘it can withstand the severe cold of the
highveld’ (du Plessis & Groenewald, 1953).
Size: my largest specimen is about 53” in over all length, and du Plessis
& Groenewald (1953) state that it does not grow large enough to be valued
for sport. Judging from the relative size of the eye and the depth of body their
figured male must be 7” or 8”.
Material described: B.M. (N.H.) 1957.11.16.1-3. holotype, g 106 + 25
mm.; paratypes, 73 + 18 & 93 + 22mm.; South African Museum: paratypes
q7i+17 (S.A.M. 23058) & 85-++21 mm. (S.A.M. 23059); all from Buffelspoort
Dam (district of Rustenburg, Transvaal) in the Sterkstroom, tributary of
Crocodile R., Limpopo system; presented by Mr. S. S. du Plessis. Transvaal
Museum no. 8676, 117 + 30 mm.
Distinguishing characters: this species has hitherto been misidentified as
Serranochromis thumbergi under which name two other distinct species have been
confused, §. thumbergi (Castelnau) and S$. robustus (Giinther). Both the latter
species are more clongate than C. flaviventris, the former having 37 vertebrae
of which 18-20 are abdominal, the latter 34-36 vertebrae, of which 16-18 are
abdominal. The extra length also shows in the scale-count, 39-40 in S. thumbergi
and 36-38 (39) in S. robustus, and in the-dorsal rays, XVII-XVIII 13-14 in
S. thumbergi, XV-XVI 14-16 in S. robustus. The smallest specimens of these
56 ANNALS OF THE SOUTH AFRICAN MUSEUM
species that I have seen, respectively 65 and 54 mm. in standard length, have
simple conical teeth as in the adult.
Neither S. thumbergi nor S. robustus is reliably reported from the Limpopo
basin.
The colours of Serranochromis thumbergi and S. robustus are described by
Messrs. G. Bell-Cross and M. A. E. Mortimer in a set of mimeographed notes
of which they have kindly sent me a copy. They refer to them respectively as
species B and C, but their identity is clear from the accompanying descriptions.
S. thumbergi is grey-green with no bright yellow colour ventrally. S. robustus
is much darker, but breeding specimens are described as having ‘a deep
chrome yellow flush on throat and belly’; Nyasa specimens seen alive by me
had not this yellow colour, but were rich green or blue, sometimes with black
horizontal bands.
REFERENCE
Du Puessis, S. S. & GROENEWALD, A. A. 1953. The kurper of Transvaal. Fauna G Flora, Transv.
prov. Adm., 3, 35-43-
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. Jn Michaelsen, W., ed. Beitrége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Harrman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8,1-58.
Harman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. arrangement according to Schenk, E. T. & McMaster, J. H.:
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Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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ANNALS
OF THE
SOU (eearnRiCAN MUSEUM
VOLUME XLVI
PART VI
THE DENTITION OF THE TITANOSUCHIAN
DINOCEPHALIANS
By
LIEUWE Dirk BoonstTRA
South African Museum, Cape Town
er cH
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[| CONTINUED ON INSIDE BACK COVER
PAT DEM TITION GF THE ‘TITANOSUGHIAN
DINOCEPHALIANS
By
LizuwE Dirk BoonstRra
South African Museum, Cape Town
[Accepted January, 1961]
(With 1 plate and 41 figures in the text)
CONTENTS
PAGE PAGE
INTRODUCTION . : E : . 57 DESCRIPTIVE (SPECIMENS ‘'TAXONOMIC-
GENERAL d : : : Puss: ALLY) 3 : : eer 5)
MATERIAL : : : : a 45O Titanosuchida <4, 69
DEscRIPTIVE (GENERAL). DE 'G@ Jonkeriidae . ; 76
Incisors : ; . €o0 Comparison wITH OTHER DINOCEPHALIA 97
Canines : . 63 COMPARISON WITH CONTEMPORARY
Postcanines . , 3 ; : 63 ‘THERAPSIDS . : : : i ALOS
ToorH REPLACEMENT ; ; . 65 OricIn oF DINOCEPHALIAN DENTITION . 104
Incisors. , ‘ . 66 ARTICULATION OF THE LOWER JAW 4108
Canines : : : . 67 ACKNOWLEDGEMENTS . : BA tits
Postcanines . - 68 REFERENCES - : : Hag i
INTRODUCTION
In the South African Dinocephalia the teeth are but poorly known. The
inadequate accounts hitherto given by the authors, who have studied this
sub-order, are chiefly due to the nature of the preservation of the studied material
and the rather rough methods of preparation hitherto employed.
Many specimens are recovered only when already naturally weathered out
of the entombing rock and lying exposed on the eroded surface of the rocks of
the Yapinocephalus zone as broken-up pieces weathered to various degrees.
In the cranial material thus collected the crowns of the teeth are seldom well
preserved, but exceptionally such material has yielded isolated well-preserved
crowns. Such is the case in specimens which when petrified had full sets of
teeth, but it is evident that a large number of skulls had on death lain exposed
before entombment and during this time many of the teeth had often either
fallen out of their sockets or had their crowns damaged or wholly broken off.
This is evident from the fact that in many cases we find empty alveoli filled by
matrix and by the fact that loose-lying teeth occur lying near parts of the skull
or even unassociated with any other skeletal material, the harder dental
substance being preserved and the less hard bone being lost. During thirty-two
years of collecting in the Tapinocephalus zone I have found that the whole
aif
Ann. S. Afr. Mus. 46 (6), 1962, 57-112, 1 pl., 41 figs.
+} 2% AGS
58 ANNALS OF THE SOUTH AFRICAN MUSEUM
teeth—crown plus root—fall out of the alveoli before petrifaction more
frequently in the Tapinocephalia than in the other three infra-orders; and in
the Titanosuchia the crowns are broken off at the level of the alveolar border
either before petrifaction or during the later weathering more frequently than
in the other infra-orders. I believe this to be due to the difference in implanta-
tion and the mode of replacement of the teeth in these infra-orders.
When crowns are preserved in situ the intractable matrix makes good
preparation difficult with the older hammer-and-chisel technique employed.
Improved newer mechanical methods—vibro-needles and sectioning with
rotating diamond-studded saws—and chemical solution now give better results
in suitable specimens. If available, radiography would be a further improvement.
GENERAL
The dentition of the sub-order Dinocephalia (Anteosauria, ‘Titanosuchia,
Tapinocephalia and Styracocephalia) can be readily distinguished from those
obtaining in all the other contemporary sub-orders of the order Therapsida by
the following distinctive character:
In occlusion the upper and lower incisors intermesh, whereas in all the
other contemporary therapsids with incisors (thus excluding the anomodonts)
the upper incisors pass labially of the lower incisors. This distinctive intermeshing
is developed further in the Titanosuchia, where the lower canine passes in front
of the upper canine to intermesh between the last upper incisor and the upper
canine (fig. 1). This process is carried still further in the Tapinocephalia and
the Styracocephalia where the whole battery of teeth intermesh.
VAN ANAY
Fic. 1. A diagram to illustrate the intermeshing of the incisors and canines in the Titanosuchia. |
Abbreviations used in this and subsequent figures:
1-5 = functional incisors. 1’—5’ = first replacing incisors. 1”—5” = second replacing incisors. 1°-5° = pre-
decessors to the functional incisors. An = angular. Ar = articular. C = canine. C! = replacing canine.
Cor = coronoid. D = dentary. F = lingual flange of the dentary. F. Pt. = fossa for the quadrate ramus of
the pterygoid. G = groove labial to the flange of the dentary. H = heel of the tooth. I.C. = inner condyle.
L.C. = lower canine. M = maxilla. m = maxillary tooth. m1! = replacing maxillary tooth. N = nasal.
O.C. = outer condyle. Pa = prearticular. Pal = palatine. PC = postcanine. PC! = replacing postcanine.
PCla = alveolus for replacing postcanine. Pm = premaxilla. Proc = processus musculus pterygoidus pos-
terius. Pt = pterygoid. Q = quadrate. Q.F. = foramen quadrati. QRPt = quadrate ramus of the ptery-
goid. R = root of the tooth. Sa = surangular. Sm = septomaxilla. Sp = splenial. St = stapedial recess on
the quadrate. T = talon of the tooth. U.C.. = upper canine. V = vomer.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 59
This intermeshing of the teeth is, I believe, a character of sufficient impor-
tance, when taken in conjunction with other evidence available from the rest
of the dinocephalian skeleton, to validate the old conception of grouping the
four infra-orders together in one sub-order—Dinocephalia, and to show that
the newer attempts in classification which are intended to refute this close
consanguinity are ill-advised and contrary to the facts.
Although the four infra-orders of the Dinocephalia are linked together by
this common character of intermeshing, there are certain dental features
(together with other skeletal characters) distinctive of each infra-order, but all
derivable from one common ancestral condition.
The dentition in the Titanosuchia can be readily distinguished from that
of the Tapinocephalia and Styracocephalia by the presence of large specialized
canines in both upper and lower jaws; from that in the Anteosauria, which
also have large specialized canines, by the fact that in the Titanosuchia the
lower canine is directed both outwards and forwards to intermesh between the
upper canine and the fifth upper incisor, whereas in the Anteosauria the lower
canine is directed upwards to lie inside the upper canine with its point housed
in a special recess in the maxilla.
The infra-order Titanosuchia has, on skeletal characters other than those
shown by the dentition, been subdivided into the three families Titanosuchidae,
Jonkeriidae and Dinartamidae.
The dental material at my disposal has not enabled me to recognize
differences in the dentition to substantiate this subdivision into families—a sub-
division which is undoubtedly valid for the former two families. The family
Dinartamidae founded on a single very poor specimen is of doubtful
validity.
MATERIAL
This report on the dentition of the Titanosuchia is based mostly on the
material in the collection of the South African Museum. This consists of 37
specimens, from isolated localities situated over the whole of the western
fossiliferous part of the Tapinocephalus zone, in which adequate evidence of the
dentition is preserved. Of these 37 specimens, 1 is from an unknown collector,
1 collected by Cloete, 1 by Cairncross, 1 by Hugo, 3 by Whaits, 3 by Haughton
and the other 27 by the author.
Although the majority of these specimens are poorly preserved, the total
of the determinable features gives us a fairly complete picture of the nature of
the dentition in the infra-order as a whole. The available facts are, however,
insufficient as reliable characters employable for taxonomic purposes within the
group. Such characters as have been used by previous authors for the establish-
ment of genera and species do not at present appear to be sufficiently distinctive
or constant and such classification must at the present stage be considered of
doubtful validity.
60 ANNALS OF THE SOUTH AFRICAN MUSEUM
DESCRIPTIVE
General
Before giving an account of the individual specimens, in an attempted
taxonomic order, the general features of the dentition of the infra-order as a
whole follows.
In the Titanosuchia a full and mature set of teeth gives the formula:
, 5 i peak)
la, Cy, PC ye rer.
Where previous authors have given different formulae for their specimens
and in the specimens at my disposal, which give a different count, these
differences should at present, without further confirmatory evidence, not be
attributed to any taxonomic distinction. That the different genera and species
Fic. 2. Jonkeria. The left second upper incisor S.A.M. E. x $.
A = anterior or inner view.
B = posterior or outer view.
C = lingual view.
D = labial view.
may have had different numbers of post-canines cannot, of course, be discounted ;
but in the light of our present knowledge such differences should rather be
considered to be due to imperfect observation, imperfect preservation, age of
the animal, or to the stage which the replacement of the teeth happens to have
reached in each individual jaw. Noted differences in the left and right sides of
the jaws of the same beast bear this out.
The Incisors
Both the upper and lower incisors have the structure which has been called
‘dinocephalian’ by various authors. What these authors really mean is that the
incisors have the general structure of the teeth of the Tapinocephalia (especially
the Russian Deuterosaurus and Ulemosaurus). Later in this paper it will be shown
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 61
that the titanosuchian incisor can be readily (figs. 2 and 3) distinguished from
the tapinocephalian teeth.
Essentially the distinctive structure of these teeth is the development in the
crown of a piercing point separated from a cutting and/or crushing surface.
Each tooth consists of a strong, long, curved, conical root; oval in section, and
a nearly equally long, curved, claw-like point or talon with a convex labial face
and a flattened lingual face with longitudinal striae and fairly flat sides. At the
Fic. 3. Jonkeria. Lateral view of the upper jaw teeth. S.A.M. E. x4. Partly restored.
junction of crown and root the crown has on its lingual or inner face an oblique
step, heel or cingulum. This heel forms an oblique ledge with its outer part
higher than its inner part in the upper incisors and its outer part lower and its
inner part higher in the lower incisors. In the unworn tooth the lingual edge
of this heel forms a fairly sharp ridge with coarse serrations; after some wear
the edge loses its serrations and the sharp edge becomes rounded; when greatly
worn the edge is lost and the whole ledge acquires a flat face. Concomitant with
this wear of the heel the talon is also worn away lingually to become more and
more slender and thereby increasing the size of the crushing flat face of
the heel.
62 ANNALS OF THE SOUTH AFRICAN MUSEUM
There are certain differences between the five upper inter se and between
the four lower incisors (fig. 4).
The first pair of upper incisors is more lightly built than all the others;
this lightness is mainly due to the fact that the teeth are compressed from side
to side, lying close together and, in occlusion, passing their talons in between
the pair of lower incisors. The fifth upper incisor has the rear face of the talon
modified to receive the lower canine with which it intermeshes. In the lower jaw
the fourth incisor is somewhat weaker than the anterior ones. Other differences
Fic. 4. Jonkeria. S.A.M. 9162. x 4.
A = anterior view. B = lateral view.
between the incisors can best be attributed to age or wear, but with more
material available some of these differences may prove to be of taxonomic
value.
The length of the incisor series varies in length: upper 73-106 mm. and
lower 68-94 mm., but the variation between the two sides of the jaw is such
that these measurements have a doubtful taxonomic value.
In occlusion the incisors of the two jaws intermesh (fig. 1) as follows:
The talons of the upper No. 1 pair lie in between the talons of the
lower No. 1 pair. |
Upper No. 2 lies between lower Nos. 1 and 2.
Upper No. 3 lies between lower Nos. 2 and 3.
Upper No. 4 lies between lower Nos. 3 and 4.
Upper No. 5 lies between lower No. 4 and the lower canine.
The outer half of the heel of upper No. 1, abuts against the inner half of
the heel of lower No. 1; the heel of upper No. 2 has its inner half abutting
THE DENTITION: OF THE TITANOSUCHIAN DINOCEPHALIANS 63
against the outer half of the heel.of lower No. 1 and its outer half against the
inner half of lower No. 2; similarly upper No. 3 abuts against the halves of
lower 2 and 3; upper No. 4 abuts against the halves of lower 3 and 4; upper
No. 5 abuts against the halves of lower 4. and the inner face of the lower canine.
The inner face of the lower canine shows in some specimens an oval worn face
where attrition by the upper No. 5 incisor takes place.
In both upper and lower jaws the incisors do not stand -yenacsllbe ¢ in the
jaws but are directed forwards so that in occlusion the tips of the talons protrude
anteriorly. |
The Canines
Each maxilla and dentale bears a single robust canine (figs. 3. and aye |
The upper canine has a large conical crown which curves moderately to .
strongly backwards and is moderately flattened from side to side to present a —
greater or lesser oval in cross section, its posterior face is rounded with no sharp
edge or serrations. The root is large and long and its presence causes the maxilla’
to bulge both externally as well as internally; internally the root opens on to
the inner face of the maxilla, where the large pulp cavity remains open for the
passage of nerve and nutrient vessels (fig. 31).
Where it leaves the jaw the canine diameters vary from 35 to 50 mm.
antero-posteriorly and from.21 to 36 mm. from side to side. The crown length
varies from 60 to 110 mm.
Between the last upper incisor and the canine there is a moderate diastema
+20 mm. which allows the lower canine to pass between these two teeth. In
occlusion the upper canine passes behind the lower canine and fits in a hollow
in the outer face of the dentale. In place of the lower canine lying lingually of
the upper canine in occlusion there are here situated the first 4-5 lower post-
canines.
The lower canine is smaller than the upper, shorter and less pointed and
curved, with its blunted point directed outwards to pass in between the upper
canine and the 5th upper incisor and to pass externally of the edge of the upper
jaw. Its intero-anterior face frequently shows an attritional face where the last
upper incisor wears against it. The outwardly directed lower canine has by
Broom been considered a distinctive character of Scapanodon, but it is a feature
common to all the known Titanosuchia, which in this character differ from all
other contemporary Therapsida. | | |
The Postcanines
The postcanines (fig. 5) form a long series in both jaws. The spacing is
close and, except where replacement is taking place, regular. They differ from
both the incisors and canines in that the teeth of the upper and lower jaws do
not intermesh. In occlusion the lower set lies lingually of the upper set, as is
normal, in the contemporary therapsids. The upper set is implanted close to
the outer edge of the maxilla, whereas the lower set lies close to the inner edge
64. ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 6. Parascapanodon. S.A.M. 12213 X $4.
A = right upper postcanine in labial and posterior view.
B = a postcanine in labial view.
C = the 11th right lower postcanine in labial and posterior
view.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 65
of the dentale, with on their outside, a groove for the reception of the upper set.
The upper postcanines are somewhat larger than the lower ones, but built on
essentially the same plan. In both jaws the crowns lie obliquely to the long axis
at an angle of about 40°, with the anterior edge directed inwards and the
posterior edge outwards. The first postcanine in both upper and lower jaws is
directed somewhat anteriorly and is usually the largest postcanine, the next four
to five are of the same size, and the rest gradually decrease in size backwards.
The postcanines in both jaws follow on their respective canines without a
diastema. As the lower canine passes anteriorly of the upper canine, and the
length of upper and lower series being approximately equal, it follows that the
last 4-5 upper postcanines have no lower antagonists. Furthermore in occlusion
the first 4-5 lower postcanines lie lingually of the upper canine.
Each postcanine (fig. 6) consists of a long root more or less oval in section,
with a spatulate crown roughly triangular in outline; the outer face is moderately
convex and the inner face flat to slightly concave. Both inner and outer faces
carry fairly coarse striae, which in unworn teeth extend to the edge which thus
becomes serrate.
In my specimens the length of the upper series varies from 169 to 188 mm.
and the lower set from 165 to 210 mm.
The full set of postcanines numbers 19-21 ; in those specimens with a shorter
set and a lesser number of teeth it would be reasonable to assume juvenility,
additional teeth being added posteriorly with increasing age.
TootH REPLACEMENT
In nearly every specimen replacement of teeth can be seen (figs. 7 and 8).
Evidence of this is, first, irregularity in the series, the presence of empty
alveoli, and, lastly, the presence of crowns in the process of eruption.
Replacement occurs in incisors, canines and postcanines, but due to the
great mass to be replaced replacement is less frequently seen in the large
canines.
There is evidence of at least two sets of canines and postcanines and of at
least three sets of incisors. Replacement may thus be a continuous process—
. there being no positive evidence of it ceasing.
In all the teeth the immediate replacing tooth erupts lingually of its
predecessor and is thus a younger member of the same tooth family. In the
Titanosuchia functional distichism thus no longer occurs.
Since in each tooth, generation follows generation independently, at any
given time the functional teeth are, except initially, never all of the same age
or generation. This is specially noticeable in the incisors, where in almost every
set of functional teeth very distinct stages of development and subsequent wear
can be seen. In many cases, from the stage of wear of the functional incisors,
the order of replacement can be deduced and this can then be verified by
noting the degree of development of their respective replacing teeth.
66 ANNALS OF THE SOUTH AFRICAN MUSEUM
Replacement of the Incisors
In the premaxilla (fig. 7) a cavity develops in the bone lingually of the
alveolus which houses each functional incisor. This cavity houses the germ from
which the replacing incisor develops. By the time the replacing crown reaches
a certain stage of development the inner or lingual wall of this cavity develops,
by resorption of the bony tissue, a perforation. This opening enlarges to form
a long oval fenestra and through this fenestra the crown of the replacing tooth
908209 OQ
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Fic. 7. Jonkeria boonstrat. S.A.M.9079 x t.
Ventral view of upper jaws.
emerges. As the replacing tooth continues its growth it moves labially into the
lingual wall of the alveolus of the functional tooth which is gradually reabsorbed,
until the new alveolus coalesces with the old alveolus. The old tooth has con-
comitantly suffered resorption and as it disappears the new tooth takes over its
alveolus. In the meantime the fenestra through which the replacing crown can
first be seen has closed up from above downwards so that the new tooth when
arriving in the old alveolus has its own lingual alveolar wall. The talon of the
young tooth, when just emerging, is flattened antero-posteriorly, roughly
triangular in outline with its inner face coarsely striate and since these striae
extend to the edges these are serrate. |
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 67
Noting the degree to which each replacing incisor crown has developed,
I have tried to determine if there is a regular order in the replacement. (This
can be controlled by noting the degree of wear in the crowns of the functional
incisors.) But as could be expected this was not possible.
In the dentary (fig. 8) the eruption of the replacing incisors appears to
follow a different course than noted in the upper incisors. This is due to the
fact that in the dentary there lies lingually to the erupting incisors a flange of
bone with a free upper edge, which thus forms a groove labially and at first
sight it appears as if the replacing incisors emerge from this groove.
Fic. 8. Jonkerta boonstrai. S.A.M. 9079. x $.
Three consecutive frontal sections through the
anterior part of the dentary to show the
replacement of the incisors and the canine.
When seen in section (fig. 9), however, it is clear that the replacing teeth
develop in cavities in the solid dentary labially of this groove. The mode of
development is thus similar to that of the upper incisors, viz. lingually of each
alveolus housing a functioning incisor a cavity is formed in which the tooth
germ develops and the further course of development parallels that already
described for the upper incisors.
Replacement of Canines
Little is known of the replacement of the canines (figs. 7 and 8). In the
upper jaw there is frequently a small hollow in the lingual wall of the maxilla
just above the inner alveolar edge of the functional canine. That this hollow
indicates the point of eruption of the replacing canine as do the fenestrae lying
68 ANNALS OF THE SOUTH AFRICAN MUSEUM
lingually of the incisors is proved by the one case (S.A.M. 11884) where on
both sides it houses the tip of the replacing canine. In two cases we have an
empty canine alveolus, but in-neither can a replacing tooth be determined with
certainty.
In the lower jaw a hollow lying lingually of the posterior end of the inner
free-edged flange of the dentary is frequently present and this in four cases
houses a small tip of the replacing canine.
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Fic. 9. Jonkeria. S.A.M. 9161.
A = dorsal view of mandible x }.
B = fracture through the jaw at the level of the first right incisor showing the dental flange
(F) with its free upper border and the groove (G) lying labially. x 4.
Replacement of the Postcanines
In both upper and lower jaws the emerging cusps of replacing teeth can
frequently be seen, each lying lingually of either a functional tooth of the same
family or of an alveolus from which its predecessor has disappeared (fig. 7).
The replacing crown-tips appear on the alveolar border without the develop-
ment of a fenestra in the lingual wall of the jaws as is the case in both incisors
and canines. Some, at least, of the replacing postcanines arise in separate alveoli
lying lingually of those of the functional teeth and these alveoli coalesce on the
loss of the functioning tooth. The erupting teeth appear as striate, serrate tips
triangular in outline.
Usually each series of postcanines has only a few replacing tips showing,
but in a few specimens a larger number—up to 9—have erupted at more or less
the same time. At the most 3 adjacent teeth have been seen erupting together,
but mostly the erupting teeth appear at irregular intervals.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 69
DESCRIPTIVE
(Specimens Taxonomically)
Titanosuchidae
Genera ARCHAEOSUCHUS and SCAPANODON
The dental material which constitutes the types of these genera is so
poorly preserved that no useful purpose will be served by attempting to add to
Broom’s descriptions.
Genus PARASCAPANODON
Parascapanodon avifontis
(Fig. 10)
S.A.M. 9127. Type. Voélfontein, P.A. Collected Boonstra 1929.
This specimen consists of parts of a skull including parts of the upper and
lower jaws together with good bones of the postcranial skeleton.
Apart from some roots of the upper incisors there is an incomplete right
mandibular ramus showing the roots of most of the teeth.
Of the incisors only the root of the fourth is present. The canine root is oval
in section (diams. 28 and 36 mm.) and the lingual wall of the alveolus shows
an opening for the replacing canine. The set of postcanines consisted of 21 teeth
of which the first is much larger than the rest. Nos. 8, 10, 12 and 15 are
replacing teeth lying lingual of alveoli from which the former set has fallen out.
The length of the postcanine series is 180 mm.
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Fic. 10. Parascapanodon avifontis. Type. S.A.M. 9127. x.
A = dorsal view of right mandible.
B = lateral view of right mandible.
70 ANNALS OF THE SOUTH AFRICAN MUSEUM
Parascapanodon sp.
S.A.M. 12213. Kroonplaas, B.W. Collected Boonstra 1959.
Under this number there are parts of at least three individuals found
together. Besides some skull pieces, a number of isolated loose-lying teeth and
a fibula, are preserved:
A (fig. 11). A fairly good anterior two-thirds of a mandibular arch in which
crowns of the teeth are preserved in a fair condition. The dental formula is
i4, cl, pel19g:
Incisors
On the left side the crowns are broken off just below the heel. Lingually
of No. 2 lies the erupted crown of its replacement. It lies in the groove just
Fic. 11. Parascapanodon sp. S.A.M. 12213 A. Xx}.
A = dorsal view of mandibles.
B = lateral view of the right mandible.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 71
anterior to the flange of the dentary and against the lingual alveolar wall of
the functioning tooth. Its inner face is striate. Lingually of No. 4 a pocket in
the alveolar wall contains the crown of its replacement.
In the right dentary the four incisors are well preserved—only the tips of
the talons being lost. ‘The heels of all four are very well preserved, and are very
little worn so that the lingual edge still forms a raised edge with a hollow
between this edge and the posterior face of the talon; and in No. 4 the posterior
surface of the talon still shows fairly coarse striae.
Lingual of Nos. 2 and 4 replacing crowns are present in their respective
pockets.
The fact that replacing crowns are already showing, while the functioning
crowns are still little worn, indicates that considerable time may elapse before
the actual replacement takes place.
Canines
The left canine is broken off at the base of the crown, which is here oval
in section with diameters 39 and 28 mm. Of the right canine the tip of the
crown is lost. It is a moderately sized tooth (basal diameters 38 x 28 mm.)
directed much outwards. On its antero-internal face there is an oblique groove
in which the 5th upper incisor fitted in occlusion.
Lingual to both canines there is a notch in the flange of the dentary and
a pocket in the internal alveolar wall for the emergence of the replacing canines.
The Postcanines
On the left dentary only a few crowns are preserved — the rest being broken
off below the spatulate crown. The presence of 19 postcanines can be deter-
mined occupying 197 mm.—No. 12 is represented by an empty alveolus and
there is a replacing crown lying postero-lingual to No. 17. No. 1 is large
(diams. 9 x 7 mm.), whereas No. 5 is only 7x6 mm. and the others approxi-
mately of this size.
In the right dentary the crowns of the postcanines are better preserved. The
original presence of 19 teeth occupying 188 mm. can be determined. No. 6
and No. 11 are represented by empty alveoli and lingual of alveolus 6 is a
replacing crown.
The crown of the 1st postcanine is widely spatulate, with a convex outer
and a flattened internal face; the edges are sharp and coarsely serrate. This,
as also the other postcanines, have their spatulate crowns lying obliquely to the
long axis of the skull, with the anterior edge lingually and the outer labially
directed. In lateral view they thus overlap where closely spaced.
The other postcanines are very similar to No. 1, except that their crowns
are smaller and the spatula more sharply triangular in outline. There is a
gradual decrease in size posteriorly.
The postcanines lie on the inner part of the upper dentary edge; external to the
tooth row there is a longitudinal groove for the reception of the upper postcanines.
ANNALS OF THE SOUTH AFRICAN MUSEUM
~I
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B (fig. 12). A mandibular arch in somewhat better condition than A.
In the left dentary the four incisors and the canine have the major part
of their crowns preserved; of the postcanines only the first five and last two are
preserved and they occupied 194 mm.
In the right dentary most of the crowns of the four incisors, canines and
the original 18 postcanines are preserved. The postcanine series measures
199 mm. and lingual of No. 7 there is a replacing crown and No. 13 is repre-
sented by an empty alveolus.
Fic. 12. Parascapanodon sp. S.A.M. 12213 B. Xf.
A = dorsal view of mandibles.
B = lateral view of right mandible.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 73
Incisors
The talons of the incisors are all lost except in No. 4 right. All the heels
are very well preserved. The heel of No. 2 left has been worn fairly flat, whereas
in all the others the lingual edge of the heel is still sharp; between this sharp
ridge and the posterior face of the talon the face of the heel is hollowed out. In
all, the inner corner of the heel is higher than the outer corner—the attritional
face thus lies obliquely to the frontal plane.
On the left side replacing crowns lie in pockets of the alveolar wall of
Nos. 2, 3, 4, and there is an empty pocket behind No. 1.
On the right side a strong replacing talon emerges obliquely from a pocket
behind No. 2; the pocket behind No. 4 contains a tip, and empty pockets lie
behind Nos. 1 and 3.
Canines
All except the tips of both canines is preserved. The canines are large teeth,
oval in cross-section. The diameters at the crown base are for the left 37 x 28 mm.
and for the right 43 x28 mm. Both canines are directed outwards. On the
antero-internal face of the left canine there is a hollowed-out attritional face
against which the upper 5th incisor worked.
The Postcanines
The spatulate crowns stand obliquely to the long axis of the skull; they are
triangular in outline with sharp edges and are coarsely serrate. The outer face
is convex and the inner fairly flat. The last five are appreciably smaller than
their predecessors in the series. The postcanines are implanted on the inner
upper edge of the dentary with a shallow groove externally for the reception of
the upper postcanines.
C (fig. 138). A weathered anterior part of the dentary shows the roots of all
four functional incisors, but all four have reached the age at which they are
to be shed. Lingually of each is a large empty alveolus from which the replacing
74 ANNALS OF THE SOUTH AFRICAN MUSEUM
tooth has been lost. In No. 3 the alveolus, which housed the replacing tooth,
has labially become confluent with the older alveolus housing the functional
older tooth of the family. In the other three the alveoli of the replacing teeth
are still separate from the older alveoli of the still functioning teeth.
The canine crown is broken off at its base where the diameters are
44x27 mm. The pulp cavity is large. The inner alveolar wall has a pocket for
the replacing canine.
Evidence of nine postcanines is preserved; Nos. 4 and 5 are represented by
empty alveoli and pockets for replacing teeth lie lingually of Nos. 1, 2 and 8.
Parascapanodon sp.
(Fig. 14)
S.A.M. 12219. Bosluiskraal, Laingsburg. Collected Boonstra 1959.
This specimen consists of a good left half of a snout and much of the post-
cranial skeleton.
The snout has its left half preserved in natural occlusion, but upper incisors
Nos. 4. and 5 and the upper canine have fallen out of their respective sockets.
The intermeshing of the incisors and canines is very clearly shown in this
specimen. The first upper incisor is a very slender tooth. The lower canine is
directed much outwards to pass outside of the maxillary edge.
A well-developed striate replacing crown lies in a pocket lingual of the
right upper No. 1 incisor and a less developed one behind the left No. 1 incisor.
Fic. 14. Parascapanodon sp. S.A.M. 12219. X f.
A = lateral view of snout. B = anterior view of the left half of snout as partly restored.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 75
Parascapanodon sp.
S.A.M. 1204. Letjiesbos, B.W. Collected Whaits 1908.
This specimen (fig. 15) consists of the mentum of the lower jaw in which
the incisors are fairly well preserved together with part of a canine crown and
three postcanine crowns.
Incisors
Most of the crowns of the eight incisors are present; the heel is imperfect
in one but present in the other seven where only the talons are partly lost, but
these can be restored with confidence as is shown in the figures.
Fic. 15. Parascapanodon sp. S.A.M. 1204. X t.
A = dorsal view of lower jaws. B = anterior view of lower jaws. C = lateral view of the
left mandible. Partly restored.
On the left side the four heels are excellently preserved in different stages
of wear. No. I is worn smooth and nearly flat, in No. 4 there is still an indication
of a lingual ridge, whereas in No. 3 and particularly in No. 2 the lingual ridge
is still sharp and the face of the heel somewhat hollowed out. If the degree of
wear is a criterion of age the order from oldest to youngest would be 1, 4, 3
and 2.
In the lingual wall of the alveolus of each functioning incisor there is a
pocket. In pockets Nos. 1, 3 and 4, talons of the replacing incisors are present,
76 ANNALS OF THE SOUTH AFRICAN MUSEUM
whereas that of No. 2 is empty. The replacing talons are in outline isosceles
triangles, with longitudinally striate faces. According to the degree of develop-
ment the order of eruption would be 1, 4, 3 and 2 agreeing with the degree of
wear of the functional incisors.
On the right side the heel of No. 4 is damaged, that of No. 3 is worn flat,
that of No. 1 fairly flat, and in No. 2 there is still a sharp lingual edge. The order
of wear is thus 3, 4, I, 2.
Lingually there are again four pockets in the alveolar wall. No. 2 is empty,
whereas the others house replacing talons. The replacing order is 3, 4, 1 and 2.
Lingual of the replacing teeth the free upper edge of the flange of the
dentary forms a high sharp edge.
Canines
The base of the left canine is preserved, with basal diameters of 33 and
25 mm. Lingual of the canine, at the edge of the dentary flange, a pocket
contains the replacing canine tip.
Postcanines
Only three spatulate postcanines have their crowns preserved, with a
replacing crown lying lingually of No. 1.
Jonkeriidae
Genus JONKERIA
Jonkeria cloetet
S.A.M. 731. Type Gamka River. Presented by Mr. Justice Cloete.
This type (fig. 16) consists of the anterior part of a left dentary, showing in
section the roots of 4 incisors, I canine and the first four postcanines. The
incisors were apparently fairly lightly built teeth. Lingually of each incisor the
Fic. 16. Fonkeria cloetet. Type S.A.M. 731. Xt.
A = dorsal view of left dentary.
B = lateral view of left dentary.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS fey
inner alveolar wall of the functioning incisors has been resorbed to form pockets
through which the replacing incisors would in time have appeared. According
to the development of these pockets the replacing order would have been 2, 4,
ace
The base of the canine is only slightly oval in outline with diameters of
31 and 25 mm.
Lingually the alveolar wall of the functioning canine also shows a pocket
and notch for the emergence of the replacing canine.
The roots of the four postcanines are large, with the first—with diameters
of 7 and 10 mm.—smaller than those lying posteriorly.
The flange of the dentary with its free upper edge is well shown.
Fic. 17. Fonkeria dubius. Type S.A.M. 2759. x.
A = dorsal view of right dentary.
B = lateral view of right dentary.
Jonkeria dubius
S.A.M. 2759. Type Platfontein of Abrahamskraal, P.A. Collected by Haughton
1913.
This type specimen (fig. 17) consists of the anterior part of a right dentary
in which only the roots of the teeth are shown in section. There are four roots
of functional incisors, No. 1 shows only the tip of a root lying in an alveolus
from which it is just about shed, and intero-lingually lies a well-developed
replacing incisor already occupying the functional alveolus. The other three
incisors, as well as the canine, have the inner alveolar walls strongly developed
as encasing sheaths. The flange of the dentary extends high up to almost meet
the inner face of the alveolar walls at a high level.
The canine has a large root, which in outline is a long pinched-in oval] with
diameters 47 and 27 mm.
78 ANNALS OF THE SOUTH AFRICAN MUSEUM
Of the postcanines the first 13 have their roots preserved. The first is the
largest (diameter 9 x 8 mm. whereas the others have on the average diameters
of 7x7 mm.).
The depression for the reception of the upper canine immediately behind
the lower canine forms a deep anteriorly directed groove.
On the inner face of the dentary, where the splenial has fallen away, there
are embedded an incisor and two postcanine crowns, possibly from the upper
jaw.
Jonkeria haughton
S.A.M, 4343. [ype Welgemoed, P.A. Collected Haughton 1916.
This type (fig. 18) consists of a fair skull without mandible, but with much
of the postcranial skeleton.
Inctisors
On the left the tip of the premaxilla has been lost and only sections of the
roots of the five incisors can be seen at depth. On the inner alveolar face there
are pockets for the replacements of Nos. 2 to 5 with crown tips of Nos. 3 and 4
lying in their pockets. The replacing order appears to be 3, 4, 5, 2, I.
In the right premaxilla the crowns of Nos. 4 and 5 are very well preserved,
whereas the other three are broken off at the alveolar border.
As preserved the incisors are directed very much anteriorly, but most of
this is due to the dorso-ventral crushing of the skull after death.
The two preserved crowns differ from those known in other forms. In
No. 4 the inner face of the talon runs obliquely from the heel to the tip without
the usual distinctive face on the heel itself, which in other forms is either flat
or hollowed out. In No. 5 this sloping inner face is also evident, but in this tooth
it is bipartite —the outer and greater part is for the reception of the lower canine.
Lingually of the right incisors there are four pockets in the alveolar wall
for the replacements 2-5; in Nos. 2 and 4 there lie the striate tips of the talons
of the replacing teeth. The replacing order appears to be 4, 2, 5, 3, I.
Canines
In the left maxilla there is an irregular empty alveolus from which the
canine has dropped out and lingually of this there is a shallow pocket for its
replacement. As this pocket contains no replacing crown it is reasonable to
assume that the functional canine has not been naturally shed but lost post
mortem, otherwise the animal would have been without a functioning canine for
an inconveniently long time.
The right canine has its crown very well preserved. As preserved, it is rather
short and greatly recurved and directed much outwards—this may be unnatural
and really due to the aforementioned dorso-ventral crushing the skull has
undergone. At its base the crown has diameters of 42 and 24 mm. and a crown
length of at least 60 mm. .
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 79
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Fic. 18. Fonkeria haughtoni. Type S.A.M. 4343. x4.
A = ventral view of the anterior half of the palate.
B = lateral view of the right side of the snout.
So ANNALS OF THE SOUTH AFRICAN MUSEUM
Postcanines
On the right side some of the crowns are preserved, whereas on the left
only the roots are to be seen in section. The backward slant of the preserved
crowns is again due to the crushing the skull has undergone.
On both maxillae there is evidence of 19 postcanines, occupying on the
right 187 mm. and on the left 188 mm.
On the right side no replacing teeth can be determined and No. 8 is
represented by an empty alveolus. On the left there are replacing teeth lingual
of Nos. 1 and 3 and lingual to the empty alveoli of Nos. 12 and 13 is another
replacement. Alveoli Nos. 16 and 17 contain two small tips and alveoli 10, 12
and 13 are empty.
Jonkeria boonstrat
S.A.M. 9079. Rietkuil, B.W. Collected Boonstra 1929.
This specimen (figs. 7 and 8), which has been referred to Janensch’s type,
consists of a weathered disarticulated skull, with good upper jaws and part of
the mandibular ramus.
Inctsors
In the right premaxilla the roots of five incisors are present, with pockets
in the inner alveolar walls of Nos. 1, 3 and 5, in which lie the crown tips of
replacements 1 and 3, whereas no crown tip has as yet been formed for No. 5.
the replacement order is 1, 9515,.%, ¢-
On the left side the roots of Nos. 1, 4 and 5 are definitely present, whereas
those of 2 and 3 are indefinite. Four pockets in the inner alveolar wall can be
seen, each housing a tip of the replacing tooth. The replacement order is 5, 2,
4, 1, 3. The lingual face of replacing No. 5, which is well developed, shows
coarse striae, and the edge, which in outline is elongated triangular, is coarsely
serrate with blunt rounded serrations.
Canines
The canine roots have the following diameters: left 40x39 mm., right
40 X30 mm. Lingually of each canine there is a pocket in the alveolar wall
indicating where the replacing canines are to appear.
Postcanines
Both maxillae show a good series of roots of functional postcanines and
crowns of the replacing set. On both sides the complete set consisted of 18 teeth,
with the first larger than the rest, which gradually decrease in size posteriorly.
On the right side the series measures 169 mm. and on the left 170 mm. in
length.
On the right side Nos. 9, 10 and 13 are represented by empty alveoli and
lingual to Nos. 1, 2, 5, 8 and 11 are pockets for replacements and lingual to
6, 9, 10, 13 and 15 there are tips of the replacing teeth showing.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS SI
On the left side No. 6 has an empty alveolus, linguo-anteriorly of No. 1
there is a fairly large pocket for a replacing tooth, and lingually of Nos. 1, 2,
3, 5, 6, 10 and 11 there are replacing crowns. The large pocket may be for a
tooth anterior to the one I am considering the first postcanine, which would
then be the second.
The erupted crowns of the replacing postcanines are triangular to spatulate
in outline with serrate edges composed of blunt rounded cusplets and the inner
face is finely striate.
The Lower Faw
Of the anterior part of the left dentary (fig. 8) I have cut three slabs in
frontal section to show the roots of the teeth in use and the stages of development
of the replacing tooth.
Section A shows the roots of the four functioning incisors, the canine and
the first postcanine. Lingually there are cavities behind Nos. 2 and 4 which
contain the beginnings of the cusps of the replacing incisors. Lingually of
Nos. 1 and 3 there are empty cavities as yet not showing any dentine of
replacing Nos. 1 and 3. Lingual of replacing No. 2 there is an empty cavity
presumably for the successor to replacement No. 2.
Lingually of the canine root is a pocket which contains the tip of the crown
of the replacing canine lying loosely in the pocket.
Section B still shows the functioning teeth roots; behind No. 2 a crescent of
dentine in a large cavity is that of replacement No. 2; replacement No. 1 is
represented by an empty cavity and lingual of replacement 2 the cavity for its
replacement is also still empty. Two empty cavities situated far lingually may
be for the second replacements of 3 and 4.
The replacing canine is here seen to occupy the antero-labial part of a
cavity.
Section C. Besides the roots of the functioning teeth this section shows that
labially of the root of the functioning No. 4 incisor there lies the still unresorbed
root remnant of an older fourth incisor. The two empty cavities are presumably
those of replacement No. 4 and the second replacement of No. 2.
Jonkeria vanderbylr
S.A.M. 11884. Skroefpaal, P.A. Collected by Boonstra and Rossouw in 1948.
This fairly well-preserved and complete skull, with the lower jaws preserved
in natural occlusion, has been referred to Broom’s type (figs. 19 and 20).
In separating the lower jaws from the upper, the crown of the right lower
canine has broken off and lies in its occlusional position, viz. between the upper
canine and the 5th upper incisor; similarly the crown of the upper right canine
is broken off near its base and lies in its occlusional position, viz. labial to the
first 4-5 lower postcanines.
82 ANNALS OF THE SOUTH AFRICAN MUSEUM
Incisors
Both premaxillaries carry five functional incisors. The No. 1 pair are
smaller teeth lying close together. On the left side the inner alveolar wall is
fenestrated behind Nos. 1, 3, 4 and 5, and in each pocket lies the crown of
a replacement tooth, triangular in outline and with its inner face striate. The
replacement order is 5, 1, 3, 4, 2.
In the dentaries four incisors are preserved on the left side and three on
the right side—No. 1 being lost. No replacements have been exposed.
Fic. 19. Fonkeria vanderbylt. S.A.M. 11884. x 4.
A = ventral view of the palate.
B = anterior view of the snout.
This specimen again very clearly shows the intermeshing of the upper and
lower incisors. The pair of slender upper Nos. 1 having their talons passing in
between the lower Nos. 1, with the outer halves of their heels meeting the inner
halves of the heels of the lower incisors; upper No. 2 passes between lower
Nos. 1 and 2, and No. 3 between 2 and 3; No. 4 between 3 and 4, and No. 5
between lower 4 and lower canine.
Canines
The upper canines are large, pointed and curve backwards. The length of
the crown is at least 90 mm. and the basal diameters are right 49 x 36 mm. and
left 45 X 34 mm.
In the pockets lingual of each canine lies the tip of the replacing canine.
The lower canines are smaller than the upper and are directed forwards
and outwards. The basal diameters are right 29 x 27 mm. and left 27 x 27mm..,
and the lengths 50 mm. and 45 mm.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 83
The lower canines intermesh between the upper fifth incisor and the upper
canine.
Postcanines
The upper postcanines are very poorly preserved —in fact, most have fallen
out after death and their alveoli are difficult to determine. It is clear, however,
that on both sides the first postcanine is much larger than those further back.
Fic. 20. Fonkeria vanderbyli. S.A.M. 11884. x 4.
A = dorsal view of mandibular arch.
B = lateral view of the snout.
In the dentaries the postcanines are fairly well preserved. On the right side
18 teeth occupy 180 mm.—Nos. g and 16 are represented by empty alveoli and
lingual of alveolus No. g lies a replacing crown. In the left dentary 19 teeth
occupy 180 mm. Nos. 18 and 19 are represented by empty alveoli. Lingual to
Nos. 6 and g lie replacing crowns and lingual to No. Io is an empty alveolus.
Jonkeria truculenta
S.A.M. 12030. Bosluiskraal, Laingsburg. Collected Boonstra 1956.
This specimen (figs. 5, 21, 22, 23, 24, 40 and 41) consists of a fairly good
though somewhat distorted skull, which I am referring to van Hoepen’s species.
84 ANNALS OF THE SOUTH AFRICAN MUSEUM
In both upper and lower jaws of the left side the dentition is particularly
well preserved and on the right side sectioning shows complementary features.
On the left side the dental formula is: i 2, c t, pc 73%.
Inctisors
In the left premaxilla (fig. 21) the root only of No. 1 is preserved, whereas
the other four incisors have most of the crowns present. The heels of Nos. 3
and 4 are worn flat, whereas in No. 2 the heel has
a fairly sharp ridge, bounding a hollow lying
between it and the talon, and in No. 5 the inner
face of the talon is bipartite with the outer part,
which receives the lower canine, the greater.
Lingual of each incisor there is a pocket in
the inner alveolar wall of the functioning incisors
of which Nos. 3 and 4 house replacing tips and
the others as yet empty. The replacing order is
T, 4) 3) 5. 2
In the right premaxilla I have cut two
frontal sections (fig. 24B):
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1. shows the roots of five incisors with No. 1 a
slender tooth. Five lingual pockets are
present, of which Nos. 2, 3 and 4 contain
the talons of the replacing incisors and the
other two are empty.
ears =
7 arse
2. again has the roots of the five functional
incisors; cavities 2 and 3 contain replace-
ment teeth and Nos. 1 and 4 are empty.
The replacing order is 2, 4, 3, 1 and 5.
~ In the left dentary (fig. 23) four incisors
+ are preserved; No. 4 has lost its crown; No. 3
F is well worn so that the occlusial surface of the
IG. 21. Jonkeria truculenta. ‘
SAN neo ee heel is smoothly flat and the talon so worn on
Ventral view of left half of snout. its lingual face that all that remains of the talon
is a weak cone; No. 2 is much less worn and
has a fairly strong talon; No. 1 just shows the tip of a talon which appears
to be piercing a plug of cancellous bone filling the alveolus.
Three frontal sections of the right dentary (fig. 244) show:
1. Just below the alveolar level four incisor roots; lingually of Nos. 2 and 4
there are pockets in whose anterior part lie sections across the talons
of the replacing teeth; lingually of No. 3 is an empty pocket. The flange
of the dentary lying lingually of these pockets is seen as a loose strip of
bone.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 85
2. Lying 9 mm. deeper this section shows replacement No. 2 lying in its pocket,
a cavity housed the replacement of No. 3 and cavity No. 4 is empty.
Lingually of the functional canine root lies a pocket housing the
replacing canine.
3. Here an empty cavity for replacement of No. 1 is seen and a replacement of
the third postcanine.
Bloc eo Fic. 22. Fonkeria truculenta. S.A.M. 12030. x t.
Inner face of left mandible.
Canines
The left upper canine is beautifully preserved. The crown length is
go mm. and at its base its diameters are 42 and 29 mm. It is thus a long,
moderately strong tooth and has only a moderate posterior curvature; its
posterior face is rounded with no indication of a cutting edge. The right upper
canine has diameters of 50 and 24 mm. A small pocket for the left replacing
canine lies linguo-posteriorly of the functional canine.
The left lower canine is fairly well preserved; its length is 41 mm. and its
basal diameters are 24 and 31 mm. It is thus quite a moderately sized tooth.
It is directed both forwards and outwards. Just behind its tip there is an oval
hole which is a result of resorption of the dentine. Its antero-dorsal face is
excavated to form a heel bearing some similarity to that typical of the incisors.
The fifth upper incisor fits with the postero-external part of its heel into this
heel of the lower canine. The mode of intermeshing is thus as is typical of the
incisors. Lingually of the canine base there is the usual notch in the flange of
the dentary for the replacing canine.
In the right dentary, sectioning shows a canine root with diameters
30 X 20 mm. and a pocket containing the tip of the replacing canine.
86
ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 24. Jonkeria truculenta.
S.A.M. 12030. x 4.
A = Three consecutive frontal
sections through the right
Fic. 23. Jonkeria truculenta. dentary.
S.A.M. 12030. xX f. B = Two frontal sections through
Dorsal view of left mandible. the right upper jaw.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 87
Postcanines
In the left upper jaw (fig. 21) a well-preserved series of postcanines follows
immediately on the canine without any diastema. ‘The length of the series is
179 mm. and there were 18 teeth. No. 1 is much larger (diameters 13 X 11 mm.)
than its immediate successors (diameters 10X8 mm.) and then the teeth
gradually decrease in size posteriorly (the last 5 x 4 mm.). No. 4 is represented
by an empty alveolus and No. 11 by the crown of a replacing tooth.
The bulbously spatulate crown is very well shown in a number of teeth.
The crowns lie obliquely to the long axis, with the posterior edge directed
outwards and the anterior edge inwards. The posterior edge is sharp and is
separated from the main bulbously swollen part of the crown by a shallow
longitudinal groove. The inner face of the crown is flattened.
In the left dentary (figs. 22, 23) a good set of crowns has been exposed
from the lingual side. The series of 18 teeth occupy a length of 179 mm. No. 8
is represented by an empty alveolus. No. 9 is represented by a nearly fully
erupted crown of a replacing tooth and so is No. 16. The functional lower post-
canines show on their lingual faces a moderately developed heel forming a
cingulum at the junction of crown and root; above this the inner face of the
crown is concave with longitudinal striae.
In the unworn replacing teeth those striae are more definite and extend
to the edges of the crown, which thus become serrate with rounded cusplets.
The first 4-5 lower postcanines lie lingually of the upper canine in occlusion
and the rest lingually of the upper postcanines with no indication of any inter-
meshing.
As the upper and lower series of postcanines occupy the same length of jaw,
follow immediately on their respective canines, and the lower canine lies in
front of the upper canine, it follows that the posterior upper postcanines (four
of them) have no lower antagonists. The first upper postcanine lies labially of
the 6th lower postcanine to constitute its antagonist.
Jonkeria ingens
S.A.M. 11573. Klein-Koedoeskop, B.W. Collected Boonstra 1940.
S.A.M. 11574. Klein-Koedoeskop, B.W. Collected Boonstra 1940.
S.A.M. 11575. Klein-Koedoeskop, B.W. Collected Boonstra 1940.
Twenty years ago I excavated three quite good skulls lying together in a
thin sandstone layer. In all three, few of the crowns of the teeth are preserved.
Of 11573 (fig. 25) I have cut a frontal section across the premaxillaries and
the other two have been serially cross-sectioned for the study of the internal
structure which is being published shortly. What has been seen of the teeth
confirms what has been described in the foregoing pages for other Jonkerta
species, with little of taxonomic importance.
88 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 25. Jonkeria ingens.
S.A.M. 11573. X ¥.
Frontal section through the premaxil-
laries to show the replacing incisors.
Fic. 26. Fonkeria ingens.
Fic. 27. Jonkeria sp. S.A.M. 11486. x t. S.A.M. 12248. x 4.
Lateral view of the right side of the snout. Dorsal view of right dentary.
Upper Incisors
In the right premaxilla the roots of the functional Nos. 2, 3, 4 and 5 are
present with No. 1 represented by an empty alveolus. Lingually of these lie a
well-developed replacement No. 1 and 3, the former in a separate alveolus and
the latter growing into the old alveolus; replacement No. 2 is just developed
and for replacements of Nos. 4 and 5 there are empty alveoli.
In the left premaxilla there is no trace of the functional No. 2, and No. 4
is being resorbed, Nos. 3 and 5 have well developed roots and so has the slender
No. 1. Replacement No. 1 is seen in its pocket and Nos. 2, 3, 4 in separate
cavities, and for No. 5 there is an empty alveolus.
2
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 89
S.A.M. 12248. Skoppelmaaikraal, Laingsburg. Collected Boonstra 1959.
This specimen (fig. 26) consists of the weathered disarticulated bones of
both jaws, of which the right dentary is almost complete and shows cross-
sections of the roots of the whole set of teeth.
Incisors
There are four functioning incisors and lingual to No. 2 there is an erupting
talon of a replacing tooth lying in a separate alveolus.
Canines
At its base the canine has the diameters 37 x 26 mm. and, lingual to it,
above the flange of the dentary, lies the pocket for its replacement.
Postcanines
There were 19 postcanines occupying 189 mm. of the jaw. The first post-
canine lies immediately behind the canine. No. 5 is represented by an empty
alveolus and lingual to the empty alveolus of No. 14 lies its replacing crown.
Jonkeria sp.
S.A.M. 11486. Mynhardtskraal, B.W. Collected Boonstra 1940.
This specimen (fig. 27) consists of a fair snout of which the left half has
been serially cross-sectioned and the right half prepared externally to show the
dentition.
Incisors
The crowns of the five right incisors are preserved in a fair state. The
postero-lateral face of No. 5 receives the antero-internal attritional face of the
lower canine, which clearly intermeshes between it and the upper canine.
In the right dentary the first three incisors are functional, whereas No. 4
has fallen out, but has a well-developed replacement crown occupying the
lingua] part of the alveolus. The tip of No. 2 replacement is also well erupted
and pockets for Nos. 1 and 3 are present. The replacement order is 4, 2, 3 and 1.
Canines
The upper canine is a large conical tooth only slightly recurved. The crown
is 82 mm. in length and the basal diameters 50 xX 22 mm. The root is very large
with a roomy pulp cavity, its length is at least 100 mm.
The lower canine is short (crown length 32 mm.) and weak with basal
diameters 26 x 21 mm. It is directed outwards to lie between the upper canine
and the upper fifth incisor against which it bears.
Postcanines
The upper postcanines follow on the canine without diastema. There is
evidence of 19 teeth occupying 183 mm. This specimen clearly shows that the
go ANNALS OF THE SOUTH AFRICAN MUSEUM
first postcanine differs from the rest; it is directed parallel to the direction of
the canine and it appears to be more pointed than spatulate. Its antero-posterior
diameter is 14 mm. as against 11 mm. in the second postcanine and 7 mm. in
the last postcanine. Nos. 14 and 18 are represented by empty alveoli with
presumably replacements lying lingually.
In the dentary there are also 19 postcanines occupying a distance of 176 mm.
The postcanines have their crowns situated obliquely to the sagittal plane.
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Fic. 28. Jonkeria sp. S.A.M. 5017. xt.
Dorsal view of mandibular arch.
This specimen again clearly shows that, since the postcanines follow
immediately behind both upper and lower canines and the lower canine passes
anterior to the upper one, the lower series cannot extend so far posteriorly as
the upper series even if they were of the same length. In fact the lower series is
7 mm. shorter and ends 50 mm. anterior to the level of the last upper post-
canine. The last 6 upper postcanines thus have no antagonists and the first 4-5
lower postcanines have the upper canine as antagonist.
Jonkeria sp.
S.A.M. 5017. Abrahamskraal, P.A. Collected Haughton 1917.
This specimen (fig. 28) consists of a poorly preserved skull lacking upper
jaws and two imperfect mandibular rami.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS OI
Incisors
On the right dentary there are roots of the four incisors and lingually of
Nos. 1 and 2 replacing talons are developed. The flange of the dentary extends
high up behind the replacing teeth as in Jonkeria dubius.
Canines
The base of the canine crown is seen in section and the diameters are
35 x 28 mm. Lingually there is a pocket in the alveolar wall and a notch in the
flange of the dentary for the emergence of the replacing canine.
Postcanines
On the right side there is evidence of a series of 17 postcanines occupying
173 mm. Nos. 7 and g are represented by empty alveoli lingual to which are
the crowns of their replacements. A further replacing crown lies lingual of No. 4.
On the left side the dentary is incomplete anteriorly to the 3rd postcanine.
Posteriorly there is evidence of 16 postcanines occupying 140 mm. Nos. 7, 9,
11 and 17 are represented by empty alveoli lingual to which lie erupting crowns
of their replacements.
Jonkeria sp.
S.A.M. 9161. Wakkerstroom of Wolwefontein, P.A. Collected Boonstra 1929.
The specimen (fig. 9) consists of parts of a broken-up skull, including
pieces of the upper jaw and much of the mandibular arch.
Incisors
In the right dentary four functional incisors are present, but both Nos. 1
and 3 are on the point of being shed with their respective replacements already
well developed. This is particularly so in No. 1, where a lucky fracture shows the
mode of replacement very well (fig. 98). The root of functional No. 1 is nearly
resorbed and in its alveolus lingually to the remains of the root lies the already
well-developed replacement. The pulp cavity of functional No. 1 is closed,
whereas that of its replacement is large and widely open. The talon of the
replacing tooth is well shown as well as the hollowed face of the heel with its
rounded lingual ridge. Lingual to the heel the inner alveolar wall is seen in
section and is thus a fairly thin sheet of bone; lingual to this there is a deep
groove and still further lingually the flange of the dentary with its free upper
edge is seen in section.
In the left dentary No. 1 incisor is represented by an empty alveolus in
whose lingual part there lies the talon of the replacing tooth.
Canines
On the left a section across the base of the canine crown is seen, which is
much laterally compressed with diameters 29 and 19 mm.; on the right there
is a broken crown directed much outwards with basal diameters 32 x 24 mm.
Q2 ANNALS OF THE SOUTH AFRICAN MUSEUM
Postcanines
On the right a series of 16 postcanines occupying 154 mm. is preserved.
No. 1 is much larger (diameters 11 x8 mm.) than its successors in the series.
Nos. 4, 11 and 13 are represented by erupting alveoli with replacing crowns
lingual to Nos. 4 and 13. No. 12 is the remains of the root of a functional tooth
with an erupting alveolus lingual to it.
Jonkerwa sp.
S.A.M. 9162. Wakkerstroom of Wolwefontein, P.A. Collected Boonstra 1929.
This specimen (figs. 4 and 29) consists of the anterior half of a skull in
which the bones of the jaws have become partially disarticulated and shifted
from their natural position. The figures given here are reconstructions.
qo00O0D ~.
Fic. 29. Jonkeria sp. S.A.M. 9162. x4.
Ventral view of the anterior part of the palate.
Incisors
In both premaxillaries the crowns of the incisors are considerably damaged;
there are five pairs with the first pair slender and situated close together. This
specimen shows the intermeshing of the upper incisors with the lower, as
already described, very well.
On both sides the five pockets for the replacing incisors have been
exposed.
In the right premaxilla the replacing crowns of Nos. 2 and 4 are visible
in their pockets and they show the elongate inner face of the talons with the
edges coarsely serrate. The replacing order is 2, 4, 5, 3, 1.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS
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93
Fic. 31. Jonkeria sp. S.A.M. 12024. X {.
A = dorsal view of the anterior end of the mandibular arch.
B = a fracture through the maxilla gives a longitudinal section through the left canine.
O4. ANNALS OF THE SOUTH AFRICAN MUSEUM
In the left premaxilla the replacing crowns are visible in pockets Nos. 1,
3, 4 and 5, and the replacing order is 5, 1, 3, 4 and 2.
On the right side the incisors in the dentary are in the natural occlusional
position with the upper incisors, and the intermeshing of the four lower with the
five upper incisors is very definite.
Canines
The lower canine in occlusion lies in front of the upper canine passing
outwards in between this and the upper fifth incisor. It is a moderately sized
tooth with its greater basal diameter 29 mm. On the right side there is a large
upper canine very little recurved; the length of the crown is at least 80 mm.
with basal diameters of 43 and 28 mm.
Postcanines
The crowns of the postcanines are only preserved in some of the more
anterior teeth. ‘They have the usual swollen spatulate crowns lying obliquely to
the long axis. On the right maxilla there is evidence of 17 teeth occupying
142 mm. Nos. g and 10 are represented by empty alveoli and lingual to Nos. 8,
9, 10, 12 and 13 replacing teeth are present.
Jonkeria sp.
S.A.M. 11980. Lammerkraal, P.A. Collected Haughton 1916.
This right dentary (fig. 30A) shows the incisor roots of Nos. 2, 3 and 4
and an empty alveolus for No. 1, lingual of which is a well-developed talon of
the replacing tooth; pockets contain the replacements of Nos. 2, 3 and 4.
The canine at its base measures 34 X 26 mm.
Nine closely packed postcanine roots are preserved with replacements lying
lingually of Nos. 1 and 2.
This may be a second specimen of Jonkeria cloetet.
Jonkeria sp.
S.A.M. 12021. Skoppelmaaikraal, Laingsburg. Collected Boonstra 1956.
This specimen (fig. 308) consists of a left premaxilla and a right dentary in
which the teeth are seen in cross-section.
Four functioning incisors, a canine and 19 postcanines are evident with a
replacing 18th postcanine.
Jonkerva sp.
S.A.M. 12024. Skoppelmaaikraal, Laingsburg. Collected Boonstra 1956.
The specimen (fig. 31) consists of weathered broken pieces of both upper
and lower jaws.
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THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 95
The interesting features shown are: in the left dentary incisors Nos. 1 and 2
are functional, with well-developed talons of their replacements lying lingually;
in Nos. 3 and 4 well developed replacing talons are visible lying loosely in the
alveoli. In the right dentary No. 1 alveolus is occupied by a very well-developed
replacing tooth, which is just coming into use. The inner edge of the heel is
unworn and is sharp with fine striae and the face of the heel is deeply hollowed
out.
In a piece of the left maxilla fracturing has revealed the root of the canine
in longitudinal section. The root is curved strongly posteriorly and has a roomy
pulp cavity, which opens on to the inner maxillary face just behind the strong
internal maxillary bulge.
Jonkeria sp.
S.A.M. 12151. Danskraal of Bloudraai, P.A. Collected Boonstra 1957.
This small dentary (fig. 32) is of interest in that it appears to be that of a
young reptile.
The canine is small with basal crown diameters 29 and 22 mm.
Notwithstanding its assumed youth there is already an empty postcanine
alveolus (No. 5), but No. 3 is a newly erupted crown.
No. 1 incisor has a pulp cavity widely open at its base, but lingually in the
same alveolus there is already a replacing talon with an undeveloped root.
One can thus with reason state that in youth the incisor replacement occurs
rapidly and probably often. In alveoli Nos. 2 and 4 lie talons just appearing
above the alveolar border, and in No. 3 there is a root lying loosely in the
alveolus. Lingual to No. 4 there is already a pocket for the next tooth.
Jonkeria sp.
S.A.M. E. This specimen bears a field number E, but both the collector and
the locality are unknown.
The specimen (figs. 2 and 3) consists of a left premaxilla and maxilla not
in articulation with four incisors broken away from the premaxilla and one
root still in the premaxilla.
Incisors
Of the incisors there is preserved: the root of No. 5; No. 4 has part of the
root and crown preserved and this is in position in the piece of premaxilla,
which also holds Nos. 3 and 2 in natural relation and the latter two consist
of good crowns with only the tips missing; No. 1 consists of a detached
crown.
All the incisors have the same general build although differing somewhat
in size and proportions.
96 ANNALS OF THE SOUTH AFRICAN MUSEUM
Measurements as reconstructed:
ji 2 3 4 5
Total height of crown and root ? 1@ fe) 110 ? ig
Height of crown 45 53 53 ? Ly
Maximum width of talon 14 15 18 ? ?
Maximum width of heel 13 16 17 ? ?
Ant.—post. length over heel 31 30 29 24 ?
Height of talon 26 32 30 ? ?
No. 1 is thus shorter and more slender than the others.
In the incisors the crown and root are roughly two curved cones of about
equal size with their bases applied to each other, so that in side view they
together have an outline resembling a sickle moon.
At the junction of crown and root there is on the inner face of the crown
a ledge, shelf or heel. The inner face of this shelf looks like a cingulum lying
obliquely with the outer part situated higher than the inner. This inner face
is convex and on the occlusal face presents a rounded edge lying behind the
half-moon-shaped depression lying between it and the inner face of the talon.
The roughly conical cusp lying below the heel is here referred to as the
talon. The inner face of the talon is flattened with a low median and lateral
ridges.
Canines
The canine is a large, somewhat recurved conical tooth, slightly laterally
compressed so that its section is oval. The length of the crown is 110 mm. and
at its base the diameters are 45 X 30 mm. From about half-way down the crown
its postero-external face forms a fairly sharp edge but without any trace of
serrations. It thus appears to be a remnant of a truly cutting edge.
Postcanines
The postcanines follow close on the canine without any real diastema. A
series of thirteen teeth is preserved of which the first twelve occupy a length of
118 mm. They decrease in size from front to back. The first postcanine has the
crown imperfectly preserved; it is directed anteriorly, whereas the others incline
posteriorly; it is the largest postcanine and its greater diameter at its base is
12 mm. as against the 8 mm. of No. 2.
The postcanines all have the same general build, viz. a quite long stem
oval in section, then a neck followed by a swollen spatulate crown triangular in
outline. The crowns lie obliquely to the long axis of the skull, with the anterior
edge lying lingually and the posterior edge labially. The outer face of the
spatulate crown is convex with longitudinal striae and sharp, somewhat everted
external as well as internal edges which are serrate when unworn. The inner
face is flattened to concave.
No. 6 has fallen out, but lingually there lies the erupted crown of its
3
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THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 07
replacing tooth with a flattish crown and serrated edges. No. 11 has its outer
face coarsely striate and the edge serrate.
COMPARISON OF THE TITANOSUCHIAN DENTITION WITH THAT OF
THE OTHER DINOCEPHALIAN INFRA-ORDERS
Earlier in this paper I have stressed a distinctive character common to all
the infra-orders of the Dinocephalia and mentioned certain striking features in
the dentition peculiar to each infra-order. A more detailed consideratior is,
however, called for.
ANTEOSAURIA
In the mature anteosaurian skull the dental formula is i 4, c 4, pc &.
The postcanine series thus has a small number of teeth.
Incisors
The incisors are long recurved conical teeth, the greatest length encoun-
tered being 100 mm. The housing of these greatly lengthened teeth is made
possible by a retreat of the premaxillary alveolar border. Although most
incisors studied have a simple conical form with an oval cross-section, there is
in the South African forms a faint indication in some teeth of a differentiation
of a talon and a slight heel, which is apparently more pronounced in some
Russian forms. A fact which would strengthen our argument for close consan-
guinity with the other dinocephalians.
The intermeshing of the incisors is similar to that of the titanosuchians,
but the fifth upper incisor is an antagonist of the lower canine by passing labially
to it in occlusion and not anteriorly and does thus not intermesh with it. The
replacement of the incisors in the Anteosauria has not been adequately studied,
but in a series of five frontal sections which I have cut in one specimen (fig. 33)
it is evident that the replacing incisor arises lingually of the functioning tooth
as it does in the Titanosuchia. The manner of eruption, however, shows certain
differences. The development of pockets in the lingual face of the alveolar wall
of the functioning incisors for the emergence of the replacing incisors, which is
so typical in the premaxillaries of the Titanosuchia, has not been found in the
Anteosauria.
In fig. 33 the five sections show:
Sections 1 and 2: The former section passes just below the alveolar border
and the latter just above. On the right side the roots of Nos. 2, 3 and 4 are those
of functioning incisors. On the left the functioning incisors Nos. 2, 3, 4 and 5
are seen and lingual of No. 5 lies a section cut through the tip of the lower canine.
Section 3: On both sides the young developing crowns of pair No. 1 appear
and on the left the remains of the root of an older member of the family of No. 2
incisor is seen as an as yet unresorbed remnant.
98 ANNALS OF THE SOUTH AFRICAN MUSEUM
Section 4: The developing right No. 5 incisor can now be seen.
Section 5: The replacing tooth of the as yet unerupted right No. 1 incisor
is already commencing its development, i.e. before the other older incisors
develop any replacements.
Canines
The upper canines are large slightly recurved pointed teeth with crowns
up to 120 mm. in height, flattened from side to side with a sharp posterior edge
in which no serrations have been seen. ‘The lower canines are not directed
outwards to pass in between the upper fifth incisor and the upper canine as they
Fic. 33. Anteosaurus abeli. S.A.M. 9123. XH.
Five consecutive frontal sections through the snout showing the
replacement of the incisors.
do in the Titanosuchia, but are directed upwards and have their tips received
in a hollow in the maxillary surface anterior to the base of the upper canine.
Practically nothing is known of the replacement of the canines, but in one
specimen an upper canine appears to have its crown about to be shed and lying
posterior to it is a pointed tooth which may be its replacement. If this
observation be correct then the replacement of the anteosaurian canine wou!d
be different from that of the Titanosuchia and be similar to that known in the
contemporary Therocephalia and Gorgonopsia.
Postcanines
In the Anteosauria the postcanines are inadequately known. They form a
short series with at most 8 teeth, irregularly spaced. The crowns are spatulate,
bluntly conical in outline and bulbously swollen from side to side.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 99
TAPINOCEPHALIA
The dentition of the South African Tapinocephalia is as yet inadequately
known (fig. 34). This is mainly due to the fact that in most of the jaws known
the teeth have been lost before entombment, and in many cases where they have
been petrified while still in the jaws they have on subsequent erosion dropped
out or have had the crowns broken off or otherwise damaged.
As the Tapinocephalia have in neither upper or lower jaw teeth specialized
as canines, the usual connotation cannot be employed. The teeth in the pre-
maxilla will be referred to as incisors and the others as maxillary and dentary
teeth. A dental formula on this basis for the Tapinocephalia would thus read:
Premaxilla 3-5, Maxilla 8-17
Dentary 14-19
Whether the differences in the count given here have much taxonomic value
is to be doubted, and in absence of the necessary facts I am more inclined to
suggest inadequate observation coupled with the probability that age is an
important factor. It is more than probable that the incisor count of 3-5 may,
as is the case in the Anteosauria, only signify a difference in age, with 5 incisors
the count for the mature animal.
All the teeth of the mature upper jaw intermesh with those of the mandible,
but this intermeshing becomes gradually less definite posteriorly.
All the more anteriorly situated teeth when mature have the same general
build, with the crown consisting of a piercing pointed cusp or talon and lingually
a cutting and/or crushing heel in general similar to those of the incisors of the
Titanosuchia. As will be evident from figure 34 the anterior or incisor teeth of
the Tapinocephalia differ, however, in detail from those of the Titanosuchia.
Some of the differences in the incisors are here enumerated:
(1) The roots are laterally compressed and the linguo-labial diameter greatly
increased thus giving a very flat oval cross-section (diameters 24-40 by
13-19 mm.);
(2) the crowns are similarly laterally compressed and the linguo-labial diameter
over the heel greatly increased (diameters 31-43 by 16-20 mm.) ;
(3) the talons are much shorter; in the Titanosuchia the crown is about the
same length as the root, whereas in the tapinocephalian incisors it 1s
about five-eighths ;
(4) the nature of the heel shows considerable differences; in the Tapino-
cephalia the occlusial face of the heel presents a fairly deep oval hollow
bounded labially by the talon, anteriorly and posteriorly by a sharp edge
with a slight spur lingually. The anterior (or inner) edge lies much higher
than the posterior (or outer) edge.
Further back in the series the tapinocephalian teeth gradually lose these
incisor characters and begin to look more like titanosuchian incisors, except
that the talons remain short. The labio-lingual diameter gradually decreases
100
ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 34. Tapinocephalus. S.A.M. K203. x $.
Four anterior teeth of the right upper jaw shown:
A = anterior (or inner) view.
B = posterior (or outer) view.
C = lingual view.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS IOI
in length, the sharp ridges on the heel become lower, the face of the heel loses
its concavity and even develops a median ridge. Still further posteriorly the
linguo-labial character becomes still less and the lingual edge of the heel begins
to look very titanosuchian-like, although much smaller in size.
Near the posterior end of the series the teeth lose the heel altogether and
the crown becomes spatulate and looks very like those of the postcanines of the
Titanosuchia (fig. 35).
We may summarize the above observations as follows:
In a series of functional teeth in the tapinocephalian jaw the posterior
teeth have spatulate crowns, which, as one proceeds anteriorly, step by step
Fic. 35. Struthiocephalus whaitsi. S.A.M. 3012. x4.
Six right upper teeth lying loose under a skull from which they have dropped.
A = posterior (or outer) view. B = lingual view.
develop a pointed talon and a heel, the heel commencing as a lingual thickening,
becomes, firstly, a cingulum, then a step; this step develops a bounding edge
with a face between it and the talon; the edge becomes sharper and the face
hollower and this hollow increases its linguo-labial diameter.
In one genus (Agnosaurus) evidence is preserved that a set of unerupted
small spatulate teeth is being replaced by a younger set, which has crowns
composed of a talon and heel. This replacement commences anteriorly and
proceeds posteriorly, but is not evident anterior to the 6th or 5th tooth in the
dentary.
T apinocephalus sp.
S.A.M. 12139. Rietfontein, P.A. Collected Boonstra 1957.
In a weathered skull of Tapinocephalus (fig. 36), where only roots and empty
alveoli are preserved, some features of the replacement of the teeth are shown.
102 ANNALS OF THE SOUTH AFRICAN MUSEUM
In the left premaxilla the roots of five functioning incisors are preserved.
Lingual to No. 5 there is a large replacing root lying in the same alveolus as the
functioning root which has been partly resorbed. In No. 4 a functioning root
lies loosely in its alveolus and lingual to it in a separate alveolus, a section across
the replacing root is present. Lingual of No. 3 there is an empty alveolus and
lingual of No. 2 there lies a root of its replacing incisor.
In the right maxilla replacing roots are preserved lying lingual to maxillary
teeth Nos. 2 and 1o. In the left maxilla the roots of the first two maxillary teeth
are seen in section; Nos. 3 and 4 are represented by a large confluent alveolus;
of No. 5 the small tip of a root is embedded in the bone. No. 6 is represented by
an empty alveolus. No. 7 is again a root-tip embedded in the bone. No. 8 is
represented by a small root-tip lying loosely in an alveolus confluent with the
Fic. 36. Tapinocephalus sp. S.A.M. 12139. XH. Fic. 38. Struthiocephalus sp. S.A.M. 12050. x4.
Ventral view of the anterior part of the palate A frontal section through the anterior end of the
showing tooth replacement. upper jaws to show the tooth replacement.
Fic. 37. Tapinocephalus sp. S.A.M. 12139. xX.
Three frontal sections through the left dentary showing tooth replacement.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 103
empty alveolus of No. 9. No. 10 is again a small root-tip lying loosely in the
alveolus. Nos. 11 and 12 are represented by empty alveoli.
I have cut three frontal sections through the anterior end of the left dentary
to determine the nature of the replacement (fig. 37).
Section 1. The 2nd functioning tooth shows a root being resorbed, with the
remnant of a still earlier root lying antero-internally. And lingual to both Nos. 1
and 2 lie sections through the talons of their replacing teeth. Lingually of
Nos. 10 and 12 is the tip of the talon of the replacement of No. 11.
Section 2 shows replacements of Nos. 1, 2, 3, 4, 6 and 8 and an empty cavity
lingual to No. 5. Further lingually, cavities for a second set of replacements are
seen for Nos. 2, 4, 5 and 6.
Section 3 shows replacements of Nos. 1, 2, 4 and 8 and a cavity for No. 3
first replacement and a cavity for No. 3 second replacement.
Struthiocephalus sp.
S.A.M. 12050. Modderfontein, Laingsburg. Collected Boonstra 1957.
I have cut some frontal sections through the snout of a good skull of
Struthiocephalus, but as most of the teeth have been lost before petrifaction the
evidence obtained of the replacement of the teeth is poor (fig. 38).
In the right premaxilla can be seen: a narrow root of No. 1 with lingually
to it the tip of the talon of its replacement.
No. 2 shows the partially resorbed root of the functioning tooth with lingual
to it a well-developed replacement.
Of No. 3 there is a large oval alveolus from which the functioning tooth
has dropped out, and in its lingual end lies the tip of the replacing talon.
No. 4 shows a well-developed, but as yet unerupted, crown with talon and
heel.
In the maxilla large irregular alveoli can indistinctly be seen with a section
through a replacing talon lying loosely in the first maxillary alveolus.
In the right half of the upper jaw there is evidence of twelve teeth of which
Nos. 1, 8, 9, 10 and 12 were functioning, No. 11 is represented by an empty
alveolus and Nos. 3, 4, 6 and 7 are represented by replacing teeth. On this
scanty evidence we may tentatively state that the replacement in the Tapino-
cephalia takes place lingually of the functioning teeth as it also does in the
Titanosuchia and the Anteosauria. The anterior (incisor) teeth do not emerge
through pockets in the alveolar walls of the functioning teeth as they do so
typically in the Titanosuchia. In this they agree with the condition, also ten-
tatively, seen in the Anteosauria.
COMPARISON WITH THE CONTEMPORARY T'HERAPSIDS
The relevant comparative facts are known in a number of the contemporary
Therocephalia and Gorgonopsia of the Tapinocephalus zone.
104 ANNALS OF THE SOUTH AFRICAN MUSEUM
In these older therapsids there are usually more (1-2) incisors, but the
number of postcanines is always much smaller than in the Dinocephalia.
Primitively there were two functional upper canines, but in most the upper
canines have been reduced to one only. All the teeth are simple, conical but
with serrations developed, usually on the posterior edge, in the incisors and
canines.
The dentition in these two sub-orders was thus carnivorous.
As in the Dinocephalia each tooth is replaced lingually by a member of
its own family and this replacement occurs more than once.
In the Dinocephalia there is a single alveolus for the upper canine, whereas
in these Therocephalia and Gorgonopsia there are two alveoli. Primitively
these two alveoli house functional canines at the same time, but in the other
forms the functional canine appears alternately in the two alveoli.
PELYCOSAURIA
In the older and more primitive Pelycosauria there may be as many as g
incisors and up to 45 maxillary teeth. In the later Sphenacodontia a pair of
teeth becomes functional canines.
All the teeth are simple, conical, but sometimes become serrate.
Each tooth is replaced lingually by a member of its own family.
Primitively the functional replacement of a tooth is by the replacing
member of an adjacent tooth—thus distichially, but in the more advanced
Sphenacodontia the functional replacement is by a member of its own family,
except in the case of the upper canines which are replaced alternately.
THE ORIGIN AND FURTHER DEVELOPMENT OF THE
DINOCEPHALIAN DENTITION
The story of the origin and further development of the dinocephalian
dentition starts from some such point as seen in the less specialized Pelycosauria.
Both upper and lower jaws have a large number of teeth —all being simple
conical structures, with the upper set, in occlusion, passing labially of the
lower set. The dentition is of an unspecialized ‘carnivorous’ nature.
Each functioning tooth is during life replaced a number of times by a
tooth, which arises lingually to it, but as each functioning tooth is lost its
function is not immediately taken over by the tooth arising lingually to it but
this function is assumed by the tooth arising in the adjacent alveolus (distichial
replacement).
This simple type of dentition is changed, when in the maxilla two of the
more anterior maxillary teeth become larger than the rest and are then called
‘canines’. Not much later a further change takes place, when each of the teeth,
not canines, is functionally as well as actually replaced by the tooth arising in
the jaw lingually to it. The pair of canines is replaced alternately, i.e. when
Pe
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 105
both are not functioning at the same time either the anterior or the posterior
one assumes the function of both. This is the stage reached by the Sphenaco-
dontia.
The next stage is one to be postulated for the ancestral therapsid or
pro-therapsid. Here the dentition would consist of about 7 upper incisors, a
pair of canines, which are the first two teeth on the maxilla, numerous post-
canines; one incisor less in the dentary, a single lower canine and a long row of
postcanines. All the teeth would be simple conical teeth, with probably serrated
edges in the incisors and canines.
From some such pro-therapsid stage the various primitive therapsid
sub-orders diverged. One of these branches represents the sub-order Dino-
cephalia. In the ancestral dinocephalian the dentition must have been as
follows: 5 upper and 4 lower incisors, a single upper and lower canine and
numerous postcanines. All the teeth were still simple conical structures with or
without serrated edges. Each functioning tooth was replaced by a tooth arising
lingually to it. In occlusion all the teeth of the upper jaw passed labially of the
teeth of the lower jaw and the dentition was definitely of a habit usually inter-
preted as being ‘carnivorous’. In the next stage, this carnivorous dentition with
upper teeth occluding labially of the lower teeth, changed to a carnivorous
dentition in which the incisor teeth commenced to intermesh with each other.
This development of intermeshing incisors was a step which definitely split
the Dinocephalia off from the other Therapsida and set them off on their own
separate course, and it is the possession of this distinctive character that we
consider to be a major justification of our considering the Dinocephalia to
constitute a distinct sub-order.
Although continuing to retain this common character the Dinocephalia
very soon started to develop divergent features. The first dichotomy appears to
have split off the infra-order Anteosauria from the rest; from the latter branch
a further dichotomy split off the Titanosuchia and still later the Tapinocephalia
and Styracocephalia developed as two divergent branches.
Anteosauria Titanosuchia Tapinocephalia Styracocephalia
ANTEOSAURIA
In the nature of its dentition the Anteosauria appear to have changed
least from the above postulated condition.
106 ANNALS OF THE SOUTH AFRICAN MUSEUM
The incisors have retained the simple conical structure, the canines are
retained and the postcanines are still simple teeth.
But certain specializations have taken place, viz. the original fairly small
incisors have become greatly elongated, and this increase in length is made
possible by a concomitant upward retreat of the premaxillary edge; the canines
have greatly increased in size and provision for the reception of the tip of the
lower canine is made by the development of a recess in the lower face of the
maxilla antero-internally of the base of the upper canine; the postcanines have
been reduced in number (maximum of 8) and the simple pointed crown has
become spatulate with a bulbous outer face.
These specializations have changed the original simple carnivores into
anteosaurs, with a highly efficient grab, pierce-and-tear dental instrument.
With a slinking habit (deduced from the structure of the limbs) such a dentition
would allow of a sudden pounce on its prey, and the tearing out of a lump of
flesh. The degeneration of the postcanines indicates that no shearing, cutting or
grinding took place prior to the deglutition of the lump of flesh helped by the
teeth on the palatine boss.
TITANOSUCHIA
In the Titanosuchia the development away from the ancestral condition
was much greater and this can be described as follows: The incisors have
completely lost their original simple pointed form and have become highly
specialized piercing, cutting and crushing teeth. The strong point, cusp or
talon situated labially is a highly efficient piercing instrument and it is held
very firmly in the jaw by a large strong root. The well-developed heel situated
lingually on the crown has a structure adapting it very well for cutting and
crushing. Although such incisors could execute a bite into flesh of no mean order
they can hardly be considered to be efficient piercing and cutting instruments
adapted for biting into flesh and the crushing abilities of the heel would serve
no reasonable purpose when applied to meat. For piercing, cutting and crushing
hard fibrous vegetable tissue these teeth would be highly efficient and well
adapted.
The large canines can reasonably be considered a relict of former car-
nivorous days, retained and even enlarged to be employed complementary to
the incisors for piercing and severing fibrous material. This is most probably the
reason why the lower canines have changed their direction to become teeth
intermeshing between the last upper incisor and the upper canine and thus
really forming part of the incisor-battery.
The postcanines have not been reduced in number and may even have been
increased and their serrate spatulate crowns forming a low row along the
margins of both jaws could very well have been of use in vegetable feeding.
TAPINOCEPHALIA
In the ‘Tapinocephalia the development away from the ancestral condition
has been greater than in both the Anteosauria and the Titanosuchia.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 107
In the former only the incisors intermesh and in the latter the canines are,
as a further step, included in this intermeshing, and now as a third step the
Tapinocephalia have the maxillary set of teeth intermeshing with the posterior
dentary teeth, with a concomitant loss of the specialized canines. The inclusion
of the maxillary teeth in this intermeshing commences anteriorly and then
gradually extends to the end of the tooth rows. This process includes the change
from spatulate teeth to teeth in which a talon and heel is developed. It has been
indicated how the change from spatulate teeth to talon-and-heel teeth has taken
place in the maxillary and posterior dentary teeth, but in no specimen have we
seen any indication that premaxillary and anterior dentary teeth (incisors) were
once spatulate and then became tooth-and-heel teeth. Though in some Russian
anteosaurs Orlov (1959) found that the conical incisors develop an attrition
face that may be the beginning of a heel.
The point now seems to be whether from a simple conical ancestral incisor
the Titanosuchia and the Tapinocephalia developed talon-and-heel incisors
independently or whether it was the Titanosuchia who took this step and handed
the condition on to the Tapinocephalia. If the latter alternative is accepted, then
it must be maintained that more or less at the same time, the large titano-
suchian canines also became talon-and-heel teeth fitting nicely into the tapino-
cephalian series. We have noticed the development of attritional faces in the
titanosuchian canines which can be interpreted as indicating such a develop-
ment of talon-and-heel teeth.
Certain differences in the structure of the incisors of respectively Titano-
suchia and Tapinocephalia have already been pointed out. Now, do these
differences indicate that talon-and-heel teeth have been developed on two
separate occasions independently of each other, or do the similarities rather
point to a common origin?
The tapinocephalian incisor can be readily derived from the titanosuchian
incisor. If in the titanosuchian incisor the length of the talon is reduced, the
labio-lingual diameter of the heel increased and the inner edge of the heel
raised to form a high and sharp cutting edge then we have a tapinocephalian
incisor.
If such a development took place it could have been caused by a change
in the nature of the vegetable food. The reduction of the size of the talon of the
incisor together with the loss of the specialized canines could be due to the food
becoming less hardy and fibrous and thus not necessitating piercing, prising and
tearing off. The improved cutting abilities of the heel would at the same time
be a better adaptation for the cutting of leafy vegetation.
I am thus of the opinion that the tapinocephalian dentition is a develop-
ment of that present in the Titanosuchia and that the Tapinocephalia hived off
from the Titanosuchia after these had acquired the typical talon and heel
incisors.
I have based the above account mainly on the South African material, but
hope to correlate with the Russian material in the near future.
108 ANNALS OF THE SOUTH AFRICAN MUSEUM
THE ARTICULATION AND MOVEMENTS OF THE LOWER JAW
(Figs. 39-41)
The quadrate has on its ventral surface a pair of convex condyles separated
by a groove and these fit fairly accurately into two concavities separated by a
ridge on the postero-dorsal face of the articular. The quadrate is firmly fixed
in the skull, being firmly clasped by the squamosal, braced by the quadrato-
Fic. 40. Jonkerta truculenta. S.A.M. 12030. xt.
A = posterior view of the jaw articulation.
B = posterior view of the lower jaw.
jugal, butted against by the paroccipital and held by the quadrate ramus of
the pterygoid.
The articular is also firmly wedged in between the angular and surangular
and the prearticular.
The only possible movement is thus at the joint between quadrate and
articular.
Fic. 39. Ventral view of the quadrate. x 4.
A = Parascapanodon avifontis S.A.M. 9127.
B = Parascapanodon sp. S.A.M. 12026.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS 109
ith the quadrate
= outer view.
iculating w
B
f the left mandible art
with the mouth closed.
.
Fic. 41. Jonkeria truculenta. S.A.M. 12030. }.
Inner View O
.
A=
IIO ANNALS OF THE SOUTH AFRICAN MUSEUM
The two condyles on the quadrate do not lie in the same plane—the outer
lying posterior to the inner. In section they present segments of circles with radii
of (in $.A.M. 12026) 26 and 32 mm. which are thus not co-centric. The radii
lie in a plane diverging from the sagittal plane in posterior direction. The result
of this structure is that when the jaws are opened the movement of the jaws is
a sliding one in posterior direction of at least 15 mm. and the backs of both jaws
at the same time move outwards for at least the same distance. At the same time
the lower edge of the jaw appears to move inwards. The latter two movements
are rendered possible by the fact that the two rami of the jaws are not firmly
fixed to each other at the symphysis.
What effects do these movements of the jaws have on the bite of a
titanosuchian ?
The forward movement of the jaws when the mouth is closed would push
the talons of the lower incisors from behind forwards between those of the upper
incisors and the heels of the lower incisors would slide under those of the upper
teeth thus executing a crushing and cutting action. The upper and lower canines
would move relatively as do the talons of the incisors, and the lower postcanines
would have a cutting action when moving past their upper antagonists.
When the jaws are closed the articulars move towards each other with the
result that the more posterior postcanines also move inwards and would thus
pass inwards of the upper postcanines leaving some space between the upper
and lower teeth.
If the lower mandibular edge, when the jaw is closed, moves outwards,
the alveolar edge would move inwards and this would still further increase the
space between upper and lower postcanines.
This would be functionally disadvantageous!
Faunistic
Hitherto the main elements of the fauna of the Tapinocephalus zone were
divided into two groups representing the opposing herbivores and carnivores
as follows:
herbivores: Pareiasauria, Tapinocephalia and Anomodontia.
carnivores: ‘Titanosuchia (including the Anteosauria), Therocepha-
lia and Gorgonopsia.
This presumed balance always appeared to be loaded in favour of the
carnivores, for not only were the Titanosuchia known to be large reptiles but
were in addition numerous, as judged by the frequency in which remains
(mostly poor, it is true) are encountered in the field.
In the new picture the herbivores are (and this is more natural) better
represented, the largest forms of the times being the herbivores. In order of size
the herbivores of this zone are: Titanosuchia, Pareiasauria, Tapinocephalia
and Anomodontia, and the carnivores: Anteosauria, Therocephalia and
Gorgonopsia.
THE DENTITION OF THE TITANOSUCHIAN DINOCEPHALIANS IST
The more marsh-dwelling herbivores were the Pareiasauria and the
Tapinocephalia; the more upland-dwelling forms the Titanosuchia and
Anomodontia. Of the carnivores the Anteosauria, with large heads, were more
of a slinking habit; the Therocephalia more active upland reptiles of prey, and
the Gorgonopsia just starting their role as important carnivores.
ACKNOWLEDGEMENTS
In the collection of the material here described I am pleased to be able to
express my gratitude to all those farmers in the Karoo who have so kindly
allowed me to work on their property, helped me in various ways, and always
been most hospitable. Mr. H. Zinn, technical assistant of the Museum, has
accompanied me on many of the collecting trips and has been most helpful.
In recent years C.S.I.R. has given me a grant which covered the current
expenses of the collecting trips undertaken.
Mr. C. Gow has during the last year assisted in the preparation of many
of the jaws studied. His painstaking work, chiefly with a vibro-tool, has produced
some really adequately prepared specimens.
The sections made of a number of specimens were rendered possible by the
use of a diamond-studded rotating saw purchased with a grant made by
Gs R.
The Trustees of the South African Museum gratefully acknowledge a
grant towards the cost of publishing this paper made by the South African
Council for Scientific and Industrial Research.
REFERENCES
Boonstra, L. D. 1936. The cranial morphology of some titanosuchid deinocephalians. Bull.
Amer. Mus. nat. Hist. 72, 99-116.
Boonstra, L. D. 1953. A suggested clarification of the taxonomic status of the South African
titanosuchians. Ann. S. Afr. Mus. 42, 19-28.
Boonstra, L. D. 1953. The cranial morphology and taxonomy of the tapinocephalid genus
Struthiocephalus. Ann. S. Afr. Mus. 42, 32-53.
Boonstra, L. D. 1954. The cranial structure of the titanosuchian: Anteosaurus. Ann. S. Afr.
Mus. 42, 108-148.
Broom, R. 1923. On the structure of the skull in the carnivorous dinocephalian reptiles. Proc.
zool. Soc. Lond. 1923, 661-684.
Broom, R. 1929. On the carnivorous mammal-like reptiles of the family Titanosuchidae. Ann.
Transv. Mus. 13, 9-36.
Broom, R. 1932. The mammal-like reptiles of South Africa. London: Witherby.
Erremov, J. A. 1940. Ulemosaurus svijagensis Riab- ein Dinocephale aus den Ablagerungen des
Perm der UdSSR. Nova Acta Leop. Carol. 9, 155-205.
Haueuton, 8S. H. 1915. On two new therocephalians from the Gouph. Ann. S. Afr. Mus. 12,
ama Ee
JANENSCH, W. 1959. Eine Jonkeria aus der Karru-Formation des Kaplandes. Paldont. £. 33, 22-49.
Ortov, J. A. 1959. [Carnivorous Deinocephalia (Titanosuchia) from the Upper Permian
deposits of the Middle Volga.] Trud. paleont. Inst. Acad. Sci. U.R.S.S. 72, 1-114 (in Russian).
Owen, R. 1879. Description of fragmentary indications of a huge kind of theriodont reptile
( Titanosuchus ferox, Ow.) from Beaufort West tract, Cape of Good Hope. Quart. F. geol. Soc.
Lond. 35, 189-199.
I12 ANNALS OF THE SOUTH AFRICAN MUSEUM
Van Hoepen, E. C. N. 1916. A new Karroo reptile. Ann. Transv. Mus. 5, No. 3, suppl. 3,1 p.
Watson, D. M. S. 1914. The Deinocephalia, an order of mammal-like reptiles. Proc. zool. Soc.
Lond. 1914, 749-786.
Watson, D. M. S. 1948. Dicynodon and its allies. Proc. zool. Soc. Lond. 118, 823-877.
Watson, D. M. S., & Romer, A. S. 1956. A classification of Therapsid reptiles. Bull. Mus.
comp. Kool. Harv. 114, 37-89.
-
Ann. S. Afr. Mus.
Plate XI
d reconstruction of the whole
-SIZE
fe
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ically in the South African Museum.
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repti
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrdége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, 9S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Hartman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuka, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
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by which a species has been previously designated (subsequent to 1758) are listed
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pagination and illustrations (plates and figures).
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Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 20, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XLVI
PART VII
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS
By
R. H. MiLztar
Marine Station, Millport, Isle of Cumbrae, Scotland
> ®
re > }
qu
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[CONTINUED ON INSIDE BACK COVER
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS
By
R. H. Mitiar
Marine Station, Millport, Isle of Cumbrae, Scotland
[Accepted April 1959]
(With 45 figures in the text)
CONTENTS
PAGE
Introduction . : : a hrs
Station list 3 : 4 fy ADORE
List of new species and forms ._ 119g
Description of species. § Lag
Distribution of species . oo e2t7
Summary . ; ; ; A esi)
References : ‘ : 220
INTRODUCTION
In a previous paper (Millar, 1955) I described a number of ascidians from
the coasts of South Africa, collected during an ecological survey carried out
by the Zoology Department of the University of Cape Town. A continuation of
this survey has produced more ascidian material, which forms a large part of
the subject of the present paper. The principal areas from which this new
material was collected (fig. 1) are: Langebaan Lagoon, Table Bay, False Bay,
Mossel Bay, Algoa Bay, and Mozambique. Small amounts of material were also
taken from several other places. The collecting was mainly from sub-littoral
areas, and this makes difficult a strict comparison with the faunistic results
obtained from the extensive surveys of Stephenson (1939, 1944, 1948), whose
studies refer to littoral areas.
Material is also included from the South African Museum, mainly
belonging to the Preter Faure collection, and some South African specimens from
the British Museum (Natural History), most of these latter being from Morten-
sen’s Java—South Africa Expedition.
Although most of the specimens formed the material of an ecological
survey, the present paper deals with systematics. I have had certain difficulties
owing to the still unsatisfactory state of the systematics in some ascidian genera;
this is particularly true of Polyclinum, Didemnum and Eudistoma. As I have
explained when discussing Dzdemnum stilense I believe that certain species will
be defined adequately only when a large series of specimens is available from
different areas and seasons, and when the biology is studied in addition to
113
Ann. S. Afr. Mus. 46 (7), 1962, 113—221, 45 figs.
ANNALS OF THE SOUTH AFRICAN MUSEUM
114
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FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS PES
preserved material. Such needs are, of course, difficult to meet, and impossible
in the course of a general ecological survey. For these reasons some of the
identifications are provisional, and some material has not been identified even
provisionally, as that course might lead to greater confusion.
I am indebted to Professor J. H. Day, of the Zoology Department of the
University of Cape Town, and to Dr. J. F. Croil Morgans, formerly of that
Department, for information regarding the specimens, and also to the Zodlogisch
Museum of Amsterdam for the loan of type specimens of many of Sluiter’s
South African species.
The Trustees of the Museum gratefully acknowledge the grant in aid of
publication of this paper received from the South African Council for Scientific
and Industrial Research.
STATION List
Most of the material belongs to the ecological survey of the University of
Cape Town, and to the Preter Faure collection of the South African Museum.
Details of stations relating to these two collections are given below. Collecting
details of other material mentioned in this report are given in the text.
UNIVERSITY OF CAPE TOWN ECOLOGICAL SURVEY
SALDANHA BAY
U.C.T., SB 1, Station 3, 13—7—46.
U.C.T., SB 91, Station 13, 14—7—46.
U.C.T., SB 92, Station 8, 14—7—46.
U.C.T., SB 94, Station 3, 13—7—-46.
U.C.T., SB 128, Salamander Bay, 7 m., 9-4—53, mud weed and sponge.
U.C.T., SB 130, off Donkergat Point, 7-9 m., 9-4—54.
U.C.T., SB 134, 4-3-37, from bottom of Szgnis.
LANGEBAAN LAGOON
U.C.T., LB 352, Oosterval, balanoid zone, 4—5—53, rocks.
U.C.T., LB 367, Skaapen Island, lower balanoid zone, 7—5—53, weed and
stones.
U.C.T., LB 374, south side of Skaapen Island, lower balanoid zone, 7—5-53.
U.G.T., LB 375, Skaapen Island, upper balanoid zone, 7—5-53.
U.C.T., LB 378, Langebaan jetty, 0-1 m., 7—-5—53, wooden pilings.
U.C.T., LB 384, Oosterval, L.W.N.T., 7-5-53, sandy limestone.
U.G.T., LB 472, 6-5-55.
TABLE BAY
U.C.T., TB 171, 33° 49°5’ S., 18° 27°5’ E., g-18 m., 4-8-46, shells or sand and
shells with occasional rocks and mussels.
U.G.T., TB 172, Station 3, 33° 50°35’ S., 18° 20’ E., 15:5 m., 3-7-47.
U.C.T., TB 173, Station 5, 35° 52-7’ S., 18° 26-8’ E., 20-5 m., 3-7—47.
116 ANNALS OF THE SOUTH AFRICAN MUSEUM
CAPE PENINSULA
U.C.T., CP 393, Oatland Point, intertidal, 27—9-54.
U.C.T., CP 425, Oatland Point, 14 inches above low water to below lowest
Poiniinceres. 12-10-54.
FALSE BAY
U.G.T., FAL 5, Station 3, just south of Seal Island, 35 m., 22-2-52, sand.
U.C.T., FAL 53, Gordon’s Bay, 18 m., 25-6—52.
U.C.T., FAL 108, Gordon’s Bay, 34° 09-3’ S., 18° 51’ E., 7-11 m., 23-1-53,
some sand.
U.C.T., FAL 109, Gordon’s Bay, approximately 34° 09°3’ S., 18° 51’ E., 7-11
M., 23-11-53.
U.G.T., FAL 117, Simon’s Bay, 23:5 m., 12-2—53, broken branched coralline.
U.C.T., FAL 121, Glencairn power station, rock terraces, 6 m., 17—2-53, scat-
tered rocks next to sand.
U.C.T., FAL 130, Oatland Point, 1-2 m. below L.W.S.T., scattered rocks,
26-2-53.
U.C.T., FAL 136, Oatland Point, o-2 m., 27—2-—53, vertical rock faces.
U.C.T., FAL 137, Gordon’s Bay quay, 0-4 m., 4-3—53, rock.
U.C.T., FAL 144, Oatland Point, bottom of Octomeris zone to 5 m. below,
9-3-53, rock face.
U.C.T., FAL 145, details as FAL 144.
U.C.T., FAL 152, Oatland Point, 4:5-5:5 m., 12-3—53, stones and rocks on sand.
U.C.T., FAL 158, Oatland Point, 0-3 m., 22—4-—53, rocks.
U.C.T., FAL 163, Oatland Point, S.W. face of Big Rock, 0-2 m., 23—5—53, area
free of Pyura community.
U.C.T., FAL 167, Oatland Point, S.W. face of Big Rock, area free of Pyura
community, 2-4 m. below Echlonia zone, 10—-6-53.
U.C.T., FAL 175, Oatland Point, S.W. face of Big Rock, 4—-6:5 m. below
Echlonia zone, 10—6—53.
U.C.T., FAL 176, Oatland Point, S.W. face of Big Rock, from top 2 m. of —
flourishing Pyura community, 10-6-53.
U.C.T., FAL 177, Oatland Point, 9-8-53.
U.C.T., FAL 181, from bottom of ship General Botha, Simon’s Town, 6—3-37.
U.C.T., FAL 182, Oatland Point, lagoon rocks, 7—9-53.
U.C.T., FAL 208, Africana, 34° 0-9’ S., 18° 42°4’ E., 36 m., rock, 10—9—53.
U.C.T., FAL 212, Africana, 34° 07:1’ S., 18° 35°6’ E., 21 m., 9—9—53, limestone
reefs and sand.
U.C.T., FAL 216, Africana, 34° 12°4' S., 18° 43°5' E., 42 m., 10-9-53, bottom
rocky.
U.C.T., FAL 221, Africana, N.W. of Seal Island, 34° 07-0’ S., 18° 32:5’ E.,
18 m., 9Q-9—-53, limestone reefs and sand.
U.C.T., FAL 225, off Miller’s Point, 34° 13-9’ S., 18° 31:6’ E., 40 m., 9-90-53,
coarse sand and finely broken shell.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 117
U.C.T., FAL 2209, off Cape Point, 34° 20-27’ S., 18° 31-8’ E., 64 m., 9-9-53,
coarse sand.
U.G.T., FAL 232, Africana, 34° 17°35’ S., 18° 31-4’ E., off Buffels Bay, 50 m.,
nO oS:
U.G.T., FAL 234, Africana, South Kogel Bay, 34° 15:35’ S., 18° 44:8’ E., 47 m.,
10—9—53, coarse sand with shell, pebbles and stones.
U.C.T., FAL 256, Oatland Point, 4:0-5:5m., 17-11-53.
U.C.T., FAL 259, Oatland Point, about 80 yards E. of Big Rock, 10:5 m.,
2I—11—53, coarse shelly sand.
U.C.T., FAL 265, half-way between Seal Island and Strandfontein, 15-25 m.,
4-4-54, rock.
U.C.T., FAL 272, Roman Rock, 100 yards ESE. of lighthouse, 14-17 m.,
18—9—-54, sloping granite rocks.
U.C.T., FAL 277, Roman Rock, 14-17 m., 21-9—54, rocks.
U.C.T., FB 1102, 21-4-47, 0°5 mile east of Seal Island, 27 m., rock.
U.C.T., FB 1105, 28-4-47, 34° 10’ S., 18° 28’ E., 24 m., sand.
U.C.T., FB 1106, 20-8—47, off Somerset Strand, 25 m.
MossEL BAy
Weer WMib 7. 34°04 17 9., 22> 13° 53” E., 19 m., 12-1—56, rock.
U.C.T., MB 14, 34° 11’ 04” S., 22° 10’ 09” E., 16 m., 13—1—56, rock.
U.G.T., MB 18, 34° 08’ 45” S., 22° 07’ 12” E., 13 m., 13—1—56, rock.
U.G.T., MB 22, 34° 08’ 50” S., 22° 07’ 20” E., 12°5 m., 13—1—56, rock.
U.C.T., MB 27, 34° 11’ 04” S., 22° 09 55” E., 19 m., 13—1—56, rock.
U.C.T., MB 38, 34° 10’ 07” S., 22° 07’ 46” E., 8-5 m., 16—-1-56, sand.
U.C.T., MB 39, 34° 10’ 08” S., 22° 08’ 00” E., 9 m., 16—1—56, rock.
U.C.T., MB 48, 34° 11’ 19” S., 22° 09’ 58” E., 10 m., 17—1—56, rock.
U.G.T., MB 53, 34° 10’ 57” S., 22° 09’ 55” E., 14. m., 17—-1-56, rock.
U-@:T., MB 55, 34° 10’ 42” S., 22° 00’ 38” E., 9 m., 17—1—56, rock.
U.G.T., MB 58, 34° 04’ 18” S., 22° 13” 32” E., 12-5 m., 18-1—56, rock.
U.C.T., MB 59, 34° 04’ 08” S., 22° 13’ 52” E., 11-5 m., 18-1—56, rock.
U.C.T., MB 60, 34° 04’ 18” S., 22° 14’ 10” E., 17-20 m., 18—1-56.
U.C.T., MB 65, 34° 04’ 47” S., 22° 13’ 06” E., 26 m., 18—1—56, rock.
U.G.T., MB 68, 34° 00’ 08" S., 22° 07’ 19” E., 13 m., 19—1—56, rock.
U.G.T., MB 69, 34° 08’ 35” S., 22° 07’ 20” E., 13°5 m., 19—1—46, rock.
U.C.T., MB 72, 34° 09’ 04” S., 22° 07’ 10” E., 12 m., 19—1—56, rock.
U.C.T., MB 84, 34° 11’ 26” S., 22° 10’ 8” E., 29 m., 21-1—56, rock.
ALGOA BAY
U.C.T., LIZ 1, Station 1, 33° 55-7’ S., 25° 37:2’ E., 8-5-10°0 m., 5-4-54, mud.
U.C.T., LIZ 2, details as Station 1.
U.C.T., LIZ 3, Station 2, 33° 56-1’ S., 25° 40-0’ E., 17-18 m., 5-4-54, sand.
U.C.T., LIZ 9, Station 3, 33° 58-1’ S., 25° 38-9’ E., 9m., 6—4—54, stones and rock.
118 ANNALS OF THE SOUTH AFRICAN MUSEUM
U.C.T., LIZ 11, Station 5, 33° 57-2’ S., 25° 38-0’ E., 9-10 m., 6-4-54, lime-
stone and grey clay.
U.C.T., LIZ 19, Station 8, 33° 58:5’ S., 25° 42:0’ E., 27 m., 7-4-54, sand and
shell.
U.C.T., LIZ 25, Station 12, 34° 00-4’ S., 25° 44°5' E., 39 m., 11-4-54, coarse
sand and shell.
U.C.T., LIZ 28, Station 13, 34° 00-8’ S., 25° 42-4’ E., 5-7 m., 11-4-54, rock.
U.C.T., LIZ 32, Station 3, 33° 58-1’ S., 25° 38-9’ E., 6—4—-54, stones and rock.
U.G.T., LIZ 34, Station 7, 33° 58:4’ S., 25° 40°5’ E., near charted ‘coral’
patch, 14 m., 7-4-54, stones.
U.G.T., LIZ 38, Station 8, 33° 58-5’ S., 25° 42-0’ E., 27 m., 6-4-54, sand and
shell.
MORRUMBENE ESTUARY
U.C.T., MOR 43, 20-1-54, channel at Linga-linga, 6-9 m.
U.C.T., MOR 50, 20-1-54, channel at mouth of Rio Coche, 3-5 m., sand.
U.C.T., MOR 76, 23-1-54, sand at L.W.S.T., Mongue.
U.C.T., MOR 77, 23-7-53, Linga-linga, 3 m., sand.
U.C.T., MOR o1, 12—7—54, channel at Linga-linga.
U.C.T., MOR 108, 14—7-54, edge of channel, Linga-linga, 1-5-4 m., sand.
U.C.T., MOR 109; 14-7-54, channel at Linga-linga, 4 m., sand.
U.C.T., MOR 122, 15—7—-54, off Linga-linga.
U.C.T., MOR 132, 16—7—54, channel opposite Rio Coche, 2 m., sand and shell.
U.C.T., MOR 187, 18—7—54, intertidal mud near L.W., N.W. of Mongue.
U.C.T., MOR 188, 18—7—54, stones at L.W.S.T., Mongue.
OTHER LOCALITIES IN MOZAMBIQUE
U.C.T., PEA 1, 26—7—53, attached to iron railings in bathing-pool.
U.C.T., PEA 3, Linga-Linga, 10 m., 23—7—-53, sand and weed.
U.C.T., PEA 4, Maxixe, 24—7-53, low water, sandbank and piles.
COMMERCIAL TRAWLERS
U.C.T., TRA 60, Kaffirkuils Bay, 34° S., 21° E., 62—70 m., 25-11-52.
U.C.T., TRA 62, 44 miles off Cape Barracouta, 63 m., 25-11-52, covering a
dromiid crab.
U.C.T., TRA 71, 32° 05’ S., 18° 14’ E., 66 m., 5—2-53, hard Polyzoa and rock.
SOUTH AFRICAN MUSEUM Pieter Faure COLLECTION
(denoted in text by PF)
PF 11, Mossel Bay, 0-55 m.
PF 586, 33° 50’ S., 25° 54’ E., 46-92 m.
PF 673, 33° 45’ S., 26° 44’ E., near Kowie, 73-79 m.
PE1704,°33° 58° 9.5125 51) Eo dom:
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS
PF 739, between Roman Rock and Cape Recife, 31 m.
PF 740, details as PF 739.
PF 895, 32° 47’ S., 28° 16’ E., 50-54 m.
PF 1095, between Bird Island and mainland, Algoa Bay, 18-29 m.
PF 1711, Cape St. Blaize, N. by E 2? E., 6 miles, 64 m.
PF 1864, Cape St. Blaize, N. 36 miles, 99 m.
PF 2326, Lion’s Head, N. 67° E., 25 miles, 248 m.
PF 2348, details as PF 2326.
PF 2361, details as PF 2326.
PF 2531, Lion’s Head, N. 63° E., 34 miles, 283 m.
PF 10165, Cape St. Blaize, N. by W. 4 W., 5 miles, 62 m.
PF 10477, Cape St. Blaize, N. by E., 8 miles, 72 m.
PF 10722, Cape Natal, W. by N., 6 miles, 91 m.
PF 12393, Itongazi River mouth, Natal, NW. 2 W., 3 miles, 46 m.
PF 12943, East London, low tide.
PF 13393, Cape Morgan, NW. £N., 11 miles, 160 m.
PF 13432, Sandy Point, NE. by E. 6 miles, 94 m.
PF 13481, Sandy Point, N. + E., 10 miles, 175 m.
PF 14560, Cape Point, N. 50° E., 18 miles, 341 m.
PF 14582, details as PF 14560.
PF 15584, Rocky Bank, False Bay, 31 m.
PF 15797, off Seal Island, False Bay, 18-19 m.
PF 15984, off Zwartklip, False Bay, 18-22 m.
PF 18785, Cape Seal, W. by N. 4 N., 7 miles, 72 m.
LisT OF NEW SPECIES AND FORMS
119
Synoicum capense sp. n. Botryllus anomalus sp. n.
S. australe sp. n. Metandrocarpa fascicularis sp. n.
Polycitorella pallida sp. n. Cnemidocarpa psammophora sp. n.
Tetrazona porrecta sp. n. Styela radicata sp. n.
Sigillina vasta sp. n. Boltenia africana sp. n.
Eudistoma digitatum sp. n. Halocynthia spinosa Sluiter f. defectiva n.
Didemnum (Polysyncraton) magnilarvum sp.n. Molgula cryptica sp. n.
Lissoclinum cavum sp. n. Eugyra myodes sp. n.
Ascidia stenodes sp. n. Eugyra macrentera sp. n.
DESCRIPTION OF SPECIES
Family Polyclinidae Verrill, 1871
Genus APLIDIUM Savigny, 1816
As I have pointed out (Millar, 1960) there seems to be no good reason for
separating the genera Aplidium Savigny 1816 and Amaroucium Milne Edwards
1842, and I am using Aplidium, which has priority.
I20 ANNALS OF THE SOUTH AFRICAN MUSEUM
Aplidium flavo-lineatum (Sluiter)
(Fig. 2)
Amaroucium flavo-lineatum Sluiter, 1898, p. 30, pl. 1, fig. 7; pl. 4, fig. 12.
Amaroucium astraeoides Sluiter, 1898, p. 33, pl. 1, fig. 9; pl. 5, figs. 2-5. Hartmeyer, 1912, p. 351,
pl. 44, fig. 5, text-fig. 10.
Amaroucium simplex Sluiter, 1898, p. 35, pl. 1, fig. 10; pl. 5, fig. 6.
Amaroucium erythraeum Michaelsen, 1934, p. 137, pl. 7, fig. 1. Millar, 1955, p. 270, fig. 2.
Known distribution
Cape Province.
A <>
OODOOD OC | EE
© 00000
ee Oo “o ya AY
: Se
ae 8h
ANIA Rad ; Ts
a SNORE NI Le aoee tee: ee ao
Fic. 2. Aplidium flavo-lineatum (Sluiter).
A, larva; B, stomach of type specimen of Amaroucium flavo-lineatum Sluiter ;
C, stomach of type specimen of Amaroucium astraeoides Sluiter; D, stomach
of type specimen of Amaroucium simplex Sluiter.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS I2I
Localities
able bays O.C.0., FB 171 B.
Cape Peninsula: U.C.T., CP 393 8; U.C.T., CP 425 R.
Feber payenu.cut., PAL 1902; U.GeghAL 163 Ns U.G.0, BAL 167 Y;
WEG yl PAL 175 K; U.Cov. FAL 162 f; Pl15797 A;. Mortensen’s
Java-South Africa Expedition, Station 65, 19-12-29, B.M.(N.H.)
TES. NO. 34.2.1 .4.
iMesh bay: W.-C... MB 18 P;°U.C:1., MB 48°A; U.C.1., MB 6o P;
Wee ahs MB 65 C.
arcnmnay a). Col LEZ 28K: U.Cil., LIZ. 32° T;. PF 704. .
Description
This species is often red in life, the collectors of the present material having
noted that the specimens were ‘red’, ‘blood orange’, or ‘scarlet’, but occasionally
‘colourless’. The variable amount of sand on the surface and within the colony
no doubt determines to a large extent its colour in the field. After preservation
the colour of the specimens may change, and the material in this collection
shows zooids with red, orange-yellow, sulphur-yellow, yellow-green and, in one
colony, bright blue colour in the body wall.
Most of the specimens in the present collection have quite conspicuous
small round or oval systems, each with a small common cloacal opening in the
centre.
I have examined many colonies and find that the number of folds on the
wall of the stomach varies from 19 to 33, but is usually between 25 and 30.
Larva
Larvae were present in colonies collected in January, February, April,
June, September, and December. They generally numbered from 1 to 4 ina
breeding zooid. Most larvae measure 0:80—1.00 mm. in length from the end of
the papillae to the base of the tail, but some were only 0-64 mm. The anterior
papillae have long, slender stalks. There is a variable number, generally 4-6,
of finger-like median ampullae, and many short lateral ampullae or large
vesicles.
Remarks
There has been much confusion in the past over this species, largely owing
to the inaccurate description of the type specimens given by Sluiter (1898).
Sluiter (1898) also described two other species, A. astraeoides and A. simplex,
the type specimens of which I have been able to examine along with those of
A. flavo-lineatum. In some important points I find Sluiter’s descriptions wrong,
and I have no doubt that the three species are synonymous, the name A. flavo-
lineatum having page priority. In particular the number of folds on the wall of
the stomach is not that given by Sluiter. Careful examination of the type
specimens (fig. 2, B-D), particularly transverse sections of the stomachs,
122 ANNALS OF THE SOUTH AFRICAN MUSEUM
showed that there are 23—26 narrow folds in each species, although Sluiter gave
the following descriptions :—
A. flavo-lineatum— to folds.
A. astraeoides—areolated stomach.
A. simplex—12 folds.
The discrepancy is difficult to understand, even assuming that Sluiter
made only a rough estimate of the total number from the number on one side
of the stomach.
This species has also appeared in accounts of South African ascidians
under the name A. erythraeum Michaelsen (Michaelsen, 1934; Millar, 1955),
but A. erythraeum was originally described from the Red Sea and I am now
doubtful if it is the same as the South African species, although anatomically
similar. Michaelsen (1934) also considers A. phortax (Michaelsen) from New
Zealand to be a synonym. If A. flavo-lineatum, A. erythraeum and A. phortax are
synonymous, then we are dealing with a species having an apparently very
curious and discontinuous distribution, and I think it more likely that the three
species are separate but anatomically similar. It might be possible to distinguish
them if the larvae of all three were known.
See below for remarks on the possible identity of A. circulatum (Hartmeyer)
and A. flavo-lineatum.
? Aplidium circulatum (Hartmeyer)
Amaroucium circulatum Hartmeyer, 1912, p. 349, pl. 39, fig. 2; pl. 44, fig. 6. Hartmeyer, 1913,
p. 130. Michaelsen, 1934, p. 130. Millar, 1955, p. 171.
Known distribution
Cape Province.
Locality
False Bay: U.C.T., FAL 234 W.
Description
The single colony is approximately mushroom-shaped, about 2-2 cm.
across the head, and provided with a short stalk. In the preserved state it is
buff-coloured, but in life, according to the collector’s note, the zooids are
‘brick-red’. There is sand within the colony but not on the surface.
The stomach of zooids in this colony has about 16 longitudinal folds, com-
pared with about 24 in the specimen which I previously described from False
- Bay (Millar, 1955). In his original account Hartmeyer (1912) described the
folds as numerous but did not state the number.
Remarks
This specimen agrees in most respects with Hartmeyer’s account of
A. circulatum. But, as Michaelsen (1934) has suggested, that species may be the
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 123
same as A. erythraeum (Michaelsen), and I have accepted A. erythraeum as a
synonym of A. flavo-lineatum.
The distinctions between A. circulatum and A. flavo-lineatum are not great,
the main one being the presence of a stalk on the colony. There may also be
fewer stomach folds. A much larger series of specimens will be needed, however,
to decide whether A. circulatum is a distinct species.
Aplidium pantherinum (Sluiter)
(Fig. 3)
Psammaplidium pantherinum Sluiter, 1898, p. 26, pl. 2, fig. 3; pl. 4, figs. 8, 9.
Psammaplidium obesum Sluiter, 1898, p. 28, pl. 1, fig. 6; pl. 4, figs. 10, 11.
Amaroucium unilarviferum Millar, 1955, p. 172, fig. 4.
B C
Fic. 3. Aplidium pantherinum (Sluiter).
A, larva; B, stomach of type specimen of A. pantherinum
(Sluiter); C, transverse section of stomach of type specimen
of A. obesum (Sluiter).
124 ANNALS OF THE SOUTH AFRICAN MUSEUM
Known distribution
Cape Province and Isipingo, Natal.
Localities
Table Bay: WU. LB 172;,0.C. 1, 2B 173 Ae
Off Cape of Good Hope: PF 2361 A.
False Bay: U.C.T., FAL 136 L; U.CG.T., FAL 145 A; Us@ol aie
152 N; U.G.T., FAL. 175°G; U/C. 2.) FAL 256'O:
Mossel Bay: U.C.T., MB 69 E.
Remarks
When I described a new species Amaroucium unilarviferum (Millar, 1955)
from Cape Province I had not examined Sluiter’s type specimens of South
African material. Having now done so I find such close agreement between
A. unilarviferum and two species which Sluiter named Psammaplidium pantherinum
and P. obesum that I must regard the three as synonymous. As Psammaplidium is
a synonym of Aplidium, the specific name becomes Aplidium pantherinum which
has page priority over A. obesum.
Only an examination of Sluiter’s type specimens shows the identity of these
species, as his descriptions are very misleading with regard to the stomach, a
key character in the determination of species of Aflidium. Careful examination
of isolated stomachs and of transverse sections of stomachs of the type specimens
of A. pantherinum shows 5 folds (fig. 3, B), although Sluiter stated that there were
12 folds. Five or 6 folds are also present in the stomach of the type specimens of
A. obesum (fig. 3 CG), which Sluiter stated had 8 folds.
The type specimens of A. pantherinum and A. obesum differ from each other
principally in external appearance, owing to the peculiar arrangement of sand
grains on the surface of the former. This is a character of little or no systematic
significance, and the identity of A. pantherinum, A. obesum and A. unilarviferum
is shown by the structure of the zooid and of the larva.
Larva (fig. 3 A)
Larvae range in length, measured from the end of the papillae to the base
of the tail, from 0-50 mm. (some larvae from type specimens of A. pantherinum
from Isipingo, Natal) to 0-80 mm. (some larvae from type specimens of A. obesum
from Sea Point, near Cape Town). The characteristic features of the larva, as
shown in all specimens, are the absence of lateral paired ampullae, and the
presence of a fringe of small epidermal vesicles round the anterior margin of
each side. From 1 to 3 larvae are present in each breeding zooid. The larva
which I figured under the name A. unilarviferum (Millar, 1955, fig. 4 D) was
evidently not fully developed, as the fringe of epidermal vesicles is not shown;
I have subsequently found the characteristic arrangement of vesicles in other
larvae from the same colony.
The larva can be used to distinguish between A. pantherinum and A. fuegiense
Cunningham, a South American species in which the stomach also has 5 or
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 125
6 folds. In A. fuegiense the larval trunk is deep, with prominent paired lateral
and unpaired median ampullae, but no vesicles (Millar, 1960), and is thus very
different from that of A. pantherinum.
Aplidium colelloides (Herdman)
(Fig. 4)
Amaroucium colelloides Herdman, 1886, p. 233, pl. 27, figs. 9-12.
Known distribution
Off Cape of Good Hope.
Locality
PF 18785 A.
Description
There are several colonies in the collection, all of a very characteristic
shape. The colony is divided into an oval, rounded, or almost triangular head
and a long narrow stalk (fig. 4 A). In one typical colony the head is 1-5 cm.
long and the stalk 6-0 cm. The head is smooth, free of sand, and dull orange-grey
in colour. The zooids are clearly visible and are arranged in long narrow oval
systems orientated so that the zooids appear to be in vertical double rows.
Common cloacal openings, which Herdman (1886) failed to find in the type
specimen, are narrow slits placed among the rows of zooids. The stalk is
sharply marked off from the head and has a uniform coating of sand grains.
In most specimens the lower end of the stalk is attached to a narrow stem-like
object which may be the tube of a polychaete.
The zooids (fig. 4 B) have a short thorax and abdomen which together
measure about 1-5 mm. in an average zooid. The post-abdomen is very long
and narrow and extends far down into the stalk of the colony. There are 6
pointed lobes on the oral siphon and the atrial opening is surmounted by a
triangular languet of moderate length. The body wall of the thorax is thin and
transparent, allowing the 9 rows of stigmata to be seen. Muscles on the thorax
are so poorly developed as to be scarcely visible. The abdomen is short and the
gut has a correspondingly short loop. There are 12 or 13 entire longitudinal
folds on the walls of the stomach. In some zooids the post-abdomen contains
an accumulation of reserve material, and in others a long series of testis follicles.
Although no ovary was seen several zooids were carrying larvae in the atrial
cavity.
Larva
The larva (fig. 4 C) measures about 0-70 mm. from the end of the papillae
to the base of the tail. The three papillae are borne on long narrow stalks.
Large numbers of small epidermal vesicles lie round the anterior end of the
126 ANNALS OF THE SOUTH AFRICAN MUSEUM
trunk in the region of the papillae. Both ocellus and otolith are present, the
ocellus, at least in the preserved state, being red, and the otolith black.
Remarks
This is an interesting species and the only previous record is that of Herd-
man (1886) who described the single type specimen. Herdman’s specimen was
collected by the Challenger from a depth of 150 fathoms, a few miles south of the
0.5 mm.
0.3 mm.
Fic. 4. Aplidium colelloides (Herdman).
A, colonies; B, zooid; C, larva.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS P27)
Cape of Good Hope. This colony was incomplete, and gave little idea of the
relatively great length of the stalk which apparently is usual. A long stalked
colony is exceptional in the genus Aplidium.
The present specimens are identified as A. colellordes mainly by the shape
of the colony, the proportions of the zooid, and the form of the larva, which
Herdman illustrated.
Aplidium retiforme (Herdman)
(Fig. 5)
Psammaplidium retiforme Herdman 1886, pp. 248-9, pl. 32, figs. 8-10.
Wauygunndt
hans ntuur’
prog
Fic. 5. Aplidium retiforme (Herdman).
A, colony; B, zooid.
Known distribution
mm..
Kerguelen.
Localities
False Bay: U.C.T., FAL 158 8; ?U.C.T.,
FAL 108 Z.
? Mossel Bay: U.C.T., MB 55 X.
? Algoa Bay: U.C.T., LIZ 9 J.
Description
The colony FAL 158 S consists of a
group of slender club-shaped lobes united
basally to a common stolon (fig. 5 A). Sand
and broken shell completely cover the surface
but do not penetrate within the lobes, where
the test matrix is clear. The largest lobes are
about 2:5 cm. long and o-8 cm. in greatest
diameter.
128 ANNALS OF THE SOUTH AFRICAN MUSEUM
Each lobe contains only a few zooids, probably constituting a single
system, but common cloacal openings were not seen.
The zooids (fig. 5 B) attain a length of 6 mm., or more if the post-abdomen
is very long. A layer of quite strong muscles is present on the thorax, which
consequently is usually contracted and bent, in the preserved condition. The
atrial opening is small and is surmounted by a short languet which has 3
shallow terminal lobes. There are 9 or 10 rows of stigmata. The stigmata are
moderately long and narrow, although Herdman (1886) describes those of his
specimens as ‘not large’ and shows them in his figure to be short. There are
about 16 undivided longitudinal folds on the wall of the stomach. The post-
abdomen contains neither ovary nor testes in the present specimens.
Remarks
This is one of those species which present a problem to the systematist.
On the one hand there is close agreement between the structure of the specimen
from False Bay and the type specimen from Kerguelen as described by Herdman, |
although it must be said that Herdman’s account lacks some important details.
On the other hand there is the great difference in temperature which must
raise doubts whether one species exists in both localities.
The specimens from the three queried localities have smaller colonies, but
otherwise are similar to the above description and may belong to the same
species.
Aplidium sarasinorum (Fiedler)
(Fig. 6)
Heterotrema sarasinorum Fiedler, 1889, pp. 859-78, pl. 25, figs. 1-14.
Known distribution
Ceylon.
Localities
False Bay: U.C.T., FAL 208 S.
Mozambique: U.C.T., PEA 3 K; U.C.T., MOR 77 K; MOR 108 L.
Locality unknown: S.A. Museum.
Description 7
Most of the colonies are growing on and round the stems of algae or
hydroids.
The specimen from False Bay is 4:2 cm. long, narrow, and flattened to a
leaf-like shape and divided for half its length into 2 broad lobes. The consistency
is firm and the surface almost smooth, but raised into small round swellings
which mark the positions of the zooids. The test is impregnated with sand.
Other colonies are more fleshy and less subdivided into lobes, but the shape
seems to depend largely on the form of the object to which the colony is
1
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 129
attached. Sand is generally present in the matrix of the colony, but is sparse or
absent on the surface.
In structure the zooids (fig. 6 A) resemble the type specimen described by
Fiedler (1889). The thorax is short, stout, and often bent in the preserved state,
owing to muscular contraction. The oral siphon is short and 6-lobed. The atrial
opening is a transverse slit with a wide 3-lobed languet (fig. 6 B). About 8
0.5 mm.
Fic. 6. Aplidium sarasinorum (Fiedler).
A, zooid; B, atrial languet.
powerful longitudinal muscles are present on each side of the thorax and con-
verge towards the ventral side of the abdomen. There is a band of shining pale
gold cells along each side of the endostyle, and although this feature was not
mentioned by Fiedler, it is present in all the South African specimens. The gut
has the usual subdivisions found in Aplidium. The stomach has 5-8 wide and
rather indistinct folds. ‘The post-abdomen is generally short, and only
indistinctly marked off from the abdomen. In some zooids there are no gonads
and in others a compact group of testis follicles immediately below the lower
bend of the intestine.
130 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
Fiedler (1889) founded his genus Heterotrema for a colony collected from
Ceylon, but Michaelsen (1923, 1924) pointed out the similarity between Hetero-
trema and certain species of Aplidium (Amaroucium), and Pérés (1952) regarded
H., sarasinorum as a variety of A. circumvolutum (Sluiter).
A. circumvolutum was originally taken from New Zealand waters and later
Pérés (1952) described specimens from Kerguelen as a variety of the same species,
A. circumvolutum var. kerguelenense. Pérés (1952) also described a new species
A, antarcticum from Kerguelen, but regarded it as very close to A. circumvolutum
and possibly identical.
I have no doubt that all the South African specimens listed here under
‘Localities’ are of the same species as Fiedler’s colony, but I do not follow
Pérés in identifying Heterotrema sarasinorum with Aplidium circumvolutum.
A. sarasinorum is characterized by the flat, expanded and lobed colony, with
zooids on both surfaces, the wide 3-lobed atrial languet, the stomach with about
7 folds, the very compact testis, and the short post-abdomen which is not
clearly marked off from the abdomen.
In A. circumvolutum the colony is more massive and broadly based, the atrial
languet long, the stomach smooth-walled (Sluiter, 1900) or with 5 or 6 folds
(Michaelsen, 1934), and the post-abdomen more sharply constricted and longer
than in A. sarasinorum.
In addition to these distinctions, which are admittedly rather slight, except
the different shape of the colony, there is the difference in distribution to
separate the species. It seems unlikely that the same species should be found at
Ceylon and at Kerguelen, and I believe that we are dealing with two species |
of similar structure, one distributed from South Africa across the Indian Ocean
(A. sarasinorum) and the other of wide distribution in southern waters including
New Zealand, Kerguelen, and also the Patagonian shelf, Graham Peninsula,
and South Georgia (Millar, 1960).
?Aplidium galeritum (Hartmeyer)
Amaroucium galeritum Hartmeyer, 1912, p. 344.
Known distribution
St. Francis Bay, Cape Province.
Locality
False bay OU: Cie Paleo re
Description
The specimen is slightly lobed and rather flattened, and measures 9:5 cm.
by 2:7 cm. by 1:0 cm. It is heavily encrusted on the surface and impregnated
throughout with sand and broken shell, so that the consistency is very firm.
There is no division of the colony into a softer head with zooids and firmer base
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 131
without zooids, which Hartmeyer (1912) found in his type specimen, but I do
not know if this is an important difference.
As regards the zooids, the structure agrees in most points with A. galeritum,
but in the specimen from False Bay there is no very distinct atrial siphon, and
there are only 10 to 12 folds on the stomach, whereas A. galeritum has a short
atrial siphon and 14 to 18 folds.
Larva
From 3 to 6 larvae are present in the atrial cavity of breeding zooids.
The trunk is about 0-4 mm. long, has the usual 3 papillae of the genus, and a
fringe of small epidermal vesicles round the anterior end. Both ocellus and
otolith are present. |
Remarks
I am not sure that this specimen agrees well enough with A. galeritum to
be placed in that species, but it may be within the variation to be expected,
and I hesitate to add another doubtful species to the South African list.
Genus syNoicuM Phipps, 1774
Synoicum capense sp. n.
(Fig. 7)
Diagnosis of species
Colony stalked, with a conical or hemispherical head. Surface of head and
stalk coated with sand and shell. Inner test matrix clear, without sand. A few
systems of zooids in each head. Zooids with 8-10 rows of stigmata, a very short
atrial siphon with moderately wide and long upper lip or languet. Stomach
smooth. Post-abdomen of variable length, with a long series of testis follicles.
Holotype
In the South African Museum, S.A.M. A25608 (U.C.T., FB. 1106C).
Localities
aiserpayen U,.G)t..F Bb ri0o,.Cs.U.C.L., FAL 53 L.
Description
The two colonies, which are stalked, are 1-3 cm. and 1°7 cm. long, the
head occupying less than half of the total length. The stalk is narrowest at the
base and gradually widens towards the upper end (fig. 7 A). The head is
almost twice as wide as the upper part of the stalk, and the larger colony is
o-6 cm. in greatest diameter. The head is widest at its junction with the stalk
and is either hemispherical or conical. Sand and broken shell fragments cover
the whole surface of the colony. Common cloacal openings are not visible, but
several must exist on each head, since dissection shows the zooids to be arranged
132 ANNALS OF THE SOUTH AFRICAN MUSEUM
common test, the inner test matrix of both head and stalk being clear.
The zooids (fig. 7 B) may reach 12 mm. in length, but some are shorter.
Only the thorax and abdomen are contained in the head of the colony and the
post-abdomen extends some distance down within the stalk.
.
in several systems. Sand and shell are confined to the surface layer of the :
0.5 cm.
Fic. 7. Synoicum capense sp. n.
A, colony; B, zooid.
mm
LEK) S°OIMSRagG
There are 6 short pointed lobes on the oral siphon. The atrial siphon is very
short with the upper margin drawn out into a short languet, which is either
quite simple or has 3 small distal lobes. About 6 slender longitudinal muscles
pass down each side of the thorax. Eight to 10 rows of stigmata are present in
the branchial sac, each row with about 12 stigmata. The oesophagus is of
moderate length, and the stomach is ovoid with smooth walls. The post-
stomach and the intestine are well-marked. There is little difference in diameter
between the abdomen and the post-abdomen, which may however be slightly
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 133
narrower. Some zooids of the larger colony have a fairly long series of testis
follicles in the post-abdomen, but none was found with an ovary. In the smaller
colony the zooids were without gonads.
Remarks
This new species perhaps most resembles S. adareanum (Herdman), but is
distinguished by the more slender club-shaped colony, the complete covering of
sand and shell, and the smaller number of rows of stigmata. From S. arenaceum
(Michaelsen) it differs in having several systems on each head and in lacking a
conspicuous rim round the cloacal openings.
Synoicum australe sp. n.
(Fig. 8)
Diagnosis of species
Colony rounded, smooth and moderately firm. Zooids arranged in small
round systems. Oral siphon with shallow lobes; atrial siphon tubular with
oblique opening and the dorsal margin slightly produced as a short lip. Longi-
tudinal muscles many and narrow. About 20 oral tentacles and 20 rows of
stigmata. Dorsal tubercle with a simple transverse oval opening. Oesophagus
longer than stomach. Stomach with smooth walls. Anus about half-way along
the thorax. Post-abdomen about equal in length to the thorax and abdomen
together. |
Holotype |
In the South African Museum. §8.A.M. A25609 (U.C.T., MB 27 H).
Locality
Mossel Bay: U.C.T., MB 27 H.
Description
There is only one specimen, a colony measuring 2-3 cm. by 1:6 cm. by
1-0 cm. The colony is slightly lobed, but there is no obvious area of attachment
and it is not apparent how the specimen was fixed to the substratum during life.
No foreign matter is present on the surface, which is quite smooth, and there is
also none within the colony. The colour, in the preserved state, is translucent
grey, with the systems of zooids showing through as paler stellate patterns.
The zooids (fig. 8) reach 6 mm. in length and are divided into a thorax
and abdomen of about equal length, and the post-abdomen which is as long
as the thorax and abdomen together. The thorax is wider than the other two
divisions of the body. The oral siphon is short with 6 shallow rounded lobes, and
the atrial siphon tubular with an oblique opening. No well developed languet
is generally present but the dorsal margin of the atrial opening is produced to
form a rounded lip of variable length. Numerous slender longitudinal muscles
pass along each side of the thorax, and circular muscle strands surround the
134 ANNALS OF THE SOUTH AFRICAN MUSEUM
oral siphon. About 20 tentacles stand at the base of the oral siphon. The dorsal
tubercle is small with a simple transverse oval opening. There are up to 20 rows
of stigmata, but the number in each row is difficult to count owing to contrac-
tion of the thorax.
The oesophagus is longer than the stomach, laterally flattened, and slightly
curved. It enters the oblique anterior end of the stomach. The stomach is
rounded with quite smooth walls. The post-stomach and intestine are together
0.3 mm.
Fic. 9. Polyclinum arenosum Sluiter. Larva.
1.5 mm.
0.2 mm.
Fic. 8. Synoicum australe sp. n. Zooid. Fic. 10. Polyclinum istpingense Sluiter. Larva.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 135
slightly longer than the stomach. The rectum extends about half-way along the
thorax, and ends in a 2-lipped anus.
None of the many zooids examined from this colony had any trace of
gonads. The post-abdomen contained numerous storage cells.
Remarks
Except for S. capense just described, no species of Synoicum has yet been
recorded from the coast of Cape Province. Michaelsen (1914, 1915) has
described Macroclinum angolanum from Angola, and this species should probably
be referred to the genus Synoicum. It differs in many respects from S. australe,
notably in its much smaller number of rows of stigmata, and much more
numerous oral tentacles. Two species described by Sluiter (1898) as Polyclinum
arenosum and P. insulsum were recorded respectively from Natal and Mozambique.
These were both regarded by Michaelsen (1919) as belonging to the genus
Macroclinum, which is now recognized as a synonym of Synoicum. I have examined
the type material and have no doubt that Sluiter was right in placing both
species in the genus Polyclinum.
In appearance of the colony the present species resembles S. gzard: (Herd-
man), an Antarctic species, but the conspicuously areolated stomach of the
latter readily distinguishes it. The zooid of S. australe is perhaps most like that
of S. adareanum (Herdman), a species of the high Antarctic, but the colony forma-
tion is quite different.
S. curcumvolutum Kott, from Kerguelen and Heard Island, is like S. australe,
but the colony is sometimes stalked and the zooids have only 12 rows of stigmata.
Another Antarctic species, S. minutum (Herdman), has a colony somewhat like
that of S. australe, but the zooids are irregularly scattered and have usually only
7 or 8 rows of stigmata.
Genus POLYCLINUM Savigny, 1816
Polyclinum arenosum Sluiter
(Fig. 9)
Polyclinum arenosum Sluiter, 1898, p. 20, pl. 4, figs. 1, 2. Millar, 1955, p. 174.
Known distribution
Isipingo, Natal; False Bay, Cape Province.
Locality
Algoa Bay: PF 1095 A.
Description
The deep division of the colony into long narrow lobes, which the specimen
shows, is typical of the species. The colony is considerably more massive than
Sluiter’s type specimen, as the lobes attain a length of 4 cm., compared with
I°4. cm. in Sluiter’s material.
136 ANNALS OF THE SOUTH AFRICAN MUSEUM
Larva
In Sluiter’s type specimens, which I have examined, larvae are present in
some zooids, but are in such a bad state of preservation that few details are visible.
They measure from 0-32 to 0:36 mm. in length, from the end of the papillae to
the base of the tail. Larvae (fig. 9) are also present in the new material from
Algoa Bay. These vary in length from 0-46 to 0-60 mm. They have the usual
3 anterior papillae, an ocellus and otolith. Small epidermal vesicles are grouped
along the dorsal and ventral parts of the trunk. Before the larva is fully developed
there are 4 pairs of lateral ampullae flanking the papillae, but later they sub-
divide to give rise to 8 pairs.
Polyclinum tsipingense Sluiter
(Fig. 10)
Polyclinum isipingense Sluiter, 1898, p. 21, pl 2, fig. 1, pl. 4, fig. 3. Hartmeyer, 1913, p. 83.
Millar, 1955, p. 175.
Known distribution
Isipingo, Natal; False Bay, Cape Province.
Localities
Morrumbene Estuary: U.C.T., MOR 50 J; MOR 76 F; MOR 108 M;
MOR 122 F; MOR 132 G; MOR 187 A.
Description
The colonies are of a rather solid appearance, and often reach 2—5 cm. in
greatest diameter. They are generally dull purple-brown in colour, and some
have a coating of sand and broken shell, but the surface of other colonies is bare.
Small common cloacal openings are scattered over the colony, but are not
raised on swellings as they were in the type material.
The zooids reach 5 mm. in total length. They are therefore shorter than
the zooids of the type material which Sluiter (1898) gives as 9 mm. long, but
the length of zooids in Polyclinum, as in most genera of the family, changes much
during the life of a colony and is a character of little systematic value. |
Larva
Larvae (fig. 10) are present in several colonies, and there are usually 2 in
each breeding zooid. They range in length from 0-32 to 0-40 mm., measured
from the end of the papillae to the base of the tail. There are 3 anterior papillae,
and 4 pairs of lateral ampullae. A dorsal and a ventral group of epidermal
vesicles is present on each side of the trunk.
Polyclinum neptunium Hartmeyer
(Fig. 11)
Polyclinum neptunium Hartmeyer, 1912, p. 331, pl. 38, fig. 9; pl. 44, figs. 2-4.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 137
Known distribution
Off Cape of Good Hope.
Localities
PF 2326; PF 2428; PF 2531; PF 14560.
Description
There are many colonies of this species in the Pieter Faure collection, all of
a dull purple-brown colour and fleshy appearance. Some are over 7 cm. in
greatest diameter and are thus rather larger than Hartmeyer’s type specimens.
The surface is generally smooth and free of encrusting matter, except on the
basal area of attachment where sand or small stones adhere to the test, but a
little sand may also be scattered on the surface. The zooids are arranged in oval
or elongated systems, but these are only sometimes visible externally.
In structure the zooids agree closely with Hartmeyer’s description.
Fic. 11. Polyclinum neptunium Hartmeyer. Larva.
Larva (fig. 11)
Larvae are present in several colonies. In length, measured from the end
of the papillae to the base of the tail, they range from 0:52 mm. to 0-80 mm.
The trunk is rather deep, dorsi-ventrally. The larva has the structure usual in
Polyclinum, with 3 anterior papillae, a sensory vesicle containing ocellus and
otolith, and well developed siphonal rudiments. The ampullary tissue well
illustrates the structure pointed out by Carlisle (1952) for Polyclinum aurantium
Milne Edwards; on each side a lateral ridge arises from the anterior end of the
trunk and sends one posterior branch dorsally and one ventrally. It is from
these ridges and their branches that the larger anterior ampullae and the
smaller dorsal and ventral vesicles arise; both ampullae and vesicles retain their
connection with the ridges by slender stalks.
138 ANNALS OF THE SOUTH AFRICAN MUSEUM
Genus PSEUDODISTOMA Michaelsen, 1924
Pseudodistoma africanum Millar
(Fig. 12)
Pseudodistoma africanum Millar 1954, p. 128.
Known distribution
Cape Province.
Localities
False Bay: U.C.T., FAL 158 L; U.C.T., FAL 175 F; PF 15584.
Mossel Bay: U.C.T., MB 65 D.
Algoa Bay: U.Q.T., LIZ 28 V.
Natal: PF 12393 A; off Durban, collection 8.8. Pickle, B.M. (N.H.) reg.
no. 1926., 6.1.4, 29° 91 077 5. 81,29 on) Eee am:
0.5 cm.
Fic. 12. Pseudodistoma africanum Millar.
Larva, possibly not fully developed.
Description
Most of the colonies agree well in shape and colour with the type specimens
(Millar, 1954). In the present collection the largest specimens are one of 8 cm.
length from Mossel Bay (U.C.T., MB 65 D), and a complex colony from Natal
(PF 12393 A) in which the head and stalk may reach 8 cm. The colony from
Natal is dull purple, unlike the other specimens which are some shade of yellow
or grey. The collectors have described the specimens variously as colourless or
pink in life.
In structure the zooids are generally typical of the species, and many have
a single very large egg in the post-abdomen. The zooids of a colony from Natal
(B.M. (N.H.) reg. no. 1926.8.1.4) have up to 20 longitudinal muscles on each
side, this being a greater number than in the type specimens.
Larva
Only one colony so far collected has larvae (fig. 12) and these were probably
not fully developed. This was the colony from Natal (B.M. (N.H.) reg. no.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 139
1926.8.1.4). In some zooids of this colony there was, in the atrial cavity, one
very large embryo or larva occupying the whole length of the thorax. The most
advanced larvae were about 1 mm. long from the end of the papillae to the base
of the tail. Unfortunately details were not clear, but there are 3 anterior papillae
in a vertical row, and both ocellus and otolith in the cerebral vesicle. A series
of ampullae is also present at the anterior end of the trunk.
Remarks
Pseudodistoma is somewhat intermediate between the families Polyclinidae
and Clavelinidae, having an abdomen and post-abdomen of the Aplidium-type
and a thorax of the Eudistoma-type. Such details of the larva as can be seen
suggest Aplidium rather than Eudistoma, thus confirming the position of the genus
in the family Polyclinidae.
Family Clavelinidae Forbes and Hanley, 1848
Genus CLAVELINA Savigny, 1816
Clavelina roseola Millar
Clavelina roseola Millar, 1955, p. 183.
Known distribution
False Bay, Cape Province; Red Sea (as Stomozoa murrayt, Kott 1957).
Localities
Mossel Bay: U.C.T., MB 60 Q; U.C.T., MB 65 E.
Description
One colony was collected at each station. The larger specimen is 5 cm.
long, 3°5 cm. wide, and 2°5 cm. high. The base of the colony is coated with
sand, but the heads are not. Some details may be added to those seen in the
type specimen. The musculature consists of numerous quite strong longitudinal
strands passing mainly from the intersiphonal region, down the sides of the
thorax to concentrate in two wide bands on the abdomen. In the new specimens
the folds on the stomach are few and have the appearance of artefacts. It 1s,
therefore, probable that the 11 or 12 longitudinal folds described in the type
material were also artefacts, and that the stomach in its natural state is smooth.
The anus is 2-lipped and lies opposite the 12th or 13th row of stigmata. Sub-
divisions of the gut below the stomach correspond closely to those in Kott’s
(1957) figure 3 of Stomozoa murrayt.
Remarks
Stomozoa murrayi Kott, collected from the Red Sea by the John Murray
Expedition, is evidently the same as Clavelina roseola and becomes a synonym.
The resemblances are very close, in the general form of the colony and zooid,
the gut, and particularly in the peculiar and characteristic siphonal lobes.
I40 ANNALS OF THE SOUTH AFRICAN MUSEUM
The only important difference is the large number of rows of stigmata in
S. murrayt (26) compared with C. roseola (17), but the greater development of
the branchial sac in the Red Sea specimens may be due to a more favourable
environment in warmer waters.
Kott (1957) accommodated her species in a new subfamily Stomozoinae,
but the siphonal lobes constitute the only character differing markedly from
the normal structure of Clavelina. The species seems to be no more than an
aberrant member of the genus Clavelina.
Clavelina steenbrasensis Millar
Clavelina steenbrasensis Millar, 1955, p. 185.
Known distribution
False Bay, Cape Province.
Localities
False Bay: U.C.T., FAL 175 E.
Mossel Bay: U.C.T., MB 18 Q; U.C.T., MB 38 D.
Description
There are several specimens in the collection, and these show that the
lateral compression of the colony noted in the type specimen is not a common
feature of the species. A well-developed colony is about 4 cm. in greatest
diameter and consists of a number of upright lobes, the largest about 2 cm.
long. The lobes are widest at the top and narrowest at the base where they are
united to a small mass of common test. The test is transparent and firm, the
dark blue zooids being visible from the outside. A section through the stomach
shows only 5 indistinct longitudinal folds, although in the type specimen there
were 8 folds. As in other species of Clavelina, however, it is often difficult to
decide whether or not some of the folds in fixed material are artefacts. Some
zooids have a slender posterior vascular process extending downwards from
the end of the abdomen.
Remarks
I have already pointed out the general similarity between this species and
C. gigantea Van Name, but the latter species even if it does not differ in the form
of the stomach as I previously believed, is distinguishable by its dark pigmented
test and apparently unpigmented zooids, and is recorded from the Gulf of
Mexico.
There are other specimens which I include in this species with some
hesitation. A single piece of a colony (U.C.T., FB 1106 D, 20—8—47) is solid,
smooth, and hyaline, and contains zooids like those of C. steenbrasensis but
without any pigment. There are also colonies from False Bay (S.A. Museum
Pieter Faure collection, no. 17458, 17-22 m.), and from an unknown locality
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS I41I
(S.A. Museum Preter Faure collection), which, although having a form of colony
resembling the typical specimens of this species, yet have a much softer and
more flexible test, and have no pigment on the zooids. It is possible that the
consistency of the test depends on the age of the colony, or the conditions of
preservation which may also affect the colour of the zooids.
Genus cystopiTEs Von Drasche, 1883
Cystodites roseolus Hartmeyer
(Fig. 13)
Cystodites roseolus Hartmeyer, 1912, p. 310.
Known distribution
Algoa Bay, Cape Province; Gulf of Guinea, west Africa; Seychelles.
Localities
Cape Peninsula: PF No. 2361 B.
Algoa Bay: between Bird Island and mainland, 18-30 m.
East of Algoa Bay: near Kowie, 33° 45’ S., 26° 44’ E., 73-78 m.
Description
The largest specimen is one from east of Algoa Bay, and measures 8 cm.
high and about 6 cm. across the base. This colony and the other almost as
large, from the same station, are roughly conical with a wide base of attachment
(fig. 13 A). All specimens are rather soft, with a smooth surface, and of a dull
pink or buff-pink colour. No common cloacal openings could be seen, and no
definite arrangement of the zooids, except in one colony from east of Algoa Bay,
in which the zooids appeared to be arranged in parallel spiral lines round the
colony. Hartmeyer (1912) was uncertain of the arrangement of zooids in his
type material, but described an apparently linear alignment of marks on the
surface of the colony, which may correspond with the appearance of the present
colony.
Spicules are generally absent from the most superficial layer of the common
test, but form a narrow layer a little below the surface, about the level of the
base of the thoraces. Below this there are scattered irregular masses of calcareous
matter, probably representing large groups of spicules. The spicules (fig. 13 B)
are mainly of a crude stellate shape with few and rather stout rays. Both the
central ball and the rays of the spicules are made up of many needle-like rods.
The spicules may attain 130u in diameter, and are therefore larger than the
spicules of Hartmeyer’s type specimen. There are also smaller lozenge-shaped
or rod-like spicules.
The zooids (fig. 13 C) are contracted and in this state measure about
2 mm. in length. In structure they are very like those of the type specimen, but
142 ANNALS OF THE SOUTH AFRICAN MUSEUM
have well-developed testes in the form of a rosette of follicles at the posterior end
of the abdomen. The branchial sac has narrow parastigmatic bars across the
rows of stigmata.
Larva
Larvae (fig. 13 D) are present in colonies from Cape Peninsula and Kowie.
They measure 0-8-1-2 mm. in length from the end of the papillae to the base of
the tail. The 3 vertically arranged papillae arise from within an elliptical band of
pO
0.2 mm.
Fic. 13. Cystodites roseolus Hartmeyer.
A, colony; B, spicules; C, zooid; D, larva.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 143
about 20 ampullae. In advanced larvae the rudiments of both oral and atrial
siphons are present. The sensory vesicle contains a cup-shaped ocellus which is
red in these alcohol-preserved specimens, and a spherical black otolith.
Remarks
This species is now known in South African waters from the Cape Peninsula
to Algoa Bay. It has also been described from the tropical waters of the Gulf
of Guinea and the Seychelles. The distribution thus given is a surprisingly
wide one, and the species appears to exist under the very different temperature
conditions of Cape Province and the Gulf of Guinea. In cases like this I wonder
if we are dealing with two species which are so similar structurally that they
cannot be distinguished, at least until many more specimens are available.
Cystodites dellechiajet (Della Valle)
Distoma dellachiajiae Della Valle, 1877, p. 40.
Known distribution
Very widely distributed in warm waters throughout the world, but
apparently not known from the West Indian Ocean.
Locality
_Morrumbene Estuary, Mozambique: U.C.T., MOR 43 W.
Description
There are several colonies, each investing a specimen of the crab Pseudo-
dromia integrifrons. The spicules form close-fitting capsules round the abdomens
of the zooids, but not round the thoraces. The convex disc-shaped overlapping
spicules of these capsules are about 200u in diameter and quite characteristic
of the species.
Genus POLYCITORELLA Michaelsen, 1924
Polycitorella pallida sp. n.
(Fig. 14)
Diagnosis of species
Colony cushion-like, broadly based, and dull white. Zooids closely spaced
and visible on the surface of the colony as small oval areas each with two
siphonal openings. Stellate and irregular spicules present in all layers of the
test. Zooids divided into thorax and abdomen. No lateral abdominal organs
visible. Siphons each with 6 lobes. At least 40 oral tentacles. Dorsal tubercle
simple. Thirteen to 15 rows of stigmata. Oesophagus long; stomach smooth-
walled; anus 2-lipped.
Holotype
In the South African Museum. S.A.M. A25610 (PF 739).
144 ANNALS OF THE SOUTH AFRICAN MUSEUM
Locality
Algoa Bay: PF 7309.
Description
The single colony is about 12 cm. long, 6-5 cm. wide, and 3°5 cm. thick.
It was evidently attached by most of the lower surface. In the preserved state
the colour is dirty white. The whole upper surface is marked by many closely
spaced low oval swellings, each marking the position of a zooid, and each having
the openings of the oral and atrial siphon. There is no encrusting material on
the colony.
Fic. 14. Polycitorella pallida sp. n.
A-C, spicules; D, zooid; E, dorsal
tubercle; ht., heart.
Spicules (fig. 14, A, B, C) are
numerous throughout all parts of the
test. Most of them are stellate with 5
to 10 rays in optical section. The rays
are sometimes conical and fairly smooth,
but usually can be seen to consist of
many needle-like rods which sometimes
produce an uneven or toothed end to
the rays (fig. 14 C). Groups of rods or
needles are also present, and form
irregular spicules. In diameter the
spicules range from 7p to 6op.
The zooids (fig. 14D) are divided
into a thorax 2-5 mm., and an abdomen
about 10 mm. long. The oral siphon is
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS I45
short and 6-lobed, and the atrial siphon, which originates a short distance from
the anterior end of the thorax, longer and more slender. Both siphons have
powerful circular muscles. Narrow closely-spaced longitudinal muscles cover
most of the thoracic region and, grouped together as a single narrower band,
continue down each side of the abdomen. Circular muscles are not conspicuous
on the thorax or abdomen.
The zooids generally show two constrictions, one between the thorax and
abdomen and the other about a quarter of the length of the abdomen behind
this. The latter constriction corresponds in position with a specially dense
accumulation of spicules and a region of tougher test forming a ring round the
abdomen at that point. This is the part of the abdomen where Michaelsen
(1924) found a pair of lateral abdominal organs in P. mariae, but I can find no
similar structures in the present species, although it is likely that this is a region
specially active in the formation of spicules.
There are at least 40 simple oral tentacles of different sizes. The tentacles
seem to be arranged in a single circle, but this is difficult to see. The dorsal
tubercle (fig. 14 E) is small with a narrow oval slit placed transversely. There
are 13 to 15 rows of stigmata in the branchial sac, each row with at least 12
rather wide and rectangular stigmata. The transverse bars are tall and narrow.
The dorsal languets are triangular and quite large. The oesophagus is narrow
and long, generally extending almost half the length of the abdomen. In outline
the stomach is ovate or somewhat rectangular. Its walls are smooth. The
remaining subdivisions of the gut are obscure, but behind the stomach there is
a clearly defined section which is probably the post-stomach, and another one
or possibly two indistinct chambers representing the mid-gut. The rectum is of
uniform diameter and ends in a 2-lipped anus about the level of the 8th row of
stigmata.
None of the zooids examined showed any trace of gonads. The heart is a
U-shaped tube beside the lower loop of the gut.
Remarks
This genus was established by Michaelsen (1924) for a species P. mariae
from the extreme northern end of North Island, New Zealand, and differs from
the genus Polycitor in having spicules in the test and lateral abdominal organs
on the zooids. Apart from the New Zealand species the genus probably contains
only one known species, described by Savigny (1816) as Eucoelium hospitiolum,
from the Gulf of Suez.
Savigny’s species differs from P. pallida in having few rows of stigmata, a
very short abdomen, and spicules confined to the surface of the colony.
P. mariae is much more like the South African species, but has a different form
of colony and has zooids with lateral abdominal organs. ‘The wide geographical
separation, with no intervening records, is also a reason for separating the
species.
146 ANNALS OF THE SOUTH AFRICAN MUSEUM
Genus TETRAZONA Michaelsen, 1930
Tetrazona porrecta sp. n.
(Fig. 15)
Diagnosis of species
Colony variable but usually long, narrow and lobed. Zooids not in systems.
Abdomen about 3 times as long as thorax. Both siphons short and 6-lobed.
About 16 oral tentacles. About 30 stigmata in each of the 4 rows. Oesophagus
long; stomach rectangular in outline. Testes below gut loop.
Holotype
In the South African Museum. 8.A.M. A25611 (PF 1095 B).
Locality
Algoa Bay: PF 1095 B.
A
|
|
4 cm.
mm.
Fic. 15. Tetrazona porrecia sp. n.
A, colony; B, zooid.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 147
Description
The colonies vary in size and shape a good deal, but are usually long and
relatively narrow with irregular lobes and swellings. As the colony generally
seems to have grown on the stem of an alga these swellings give some of the
specimens the appearance of a string of beads (fig. 15 A). Specimens of this kind
reach a length of at least 24 cm. The colony is firm, and the surface rough
owing to the presence of a uniform coating of sand grains. Fewer sand and shell
particles are present within the test.
The zooids (fig. 15 B) have an average length of about 4 mm., of which
the thorax occupies 1 mm. and the abdomen 3 mm. Both siphons open directly
to the surface, and both are short, with 6 sharp lobes. Muscles are not strong on
the thoracic wall and the zooids are consequently little contracted. The oral
tentacles, about 16 in number, are of moderate length, alternating in size, and
arranged in a single circle. Each of the 4 rows of stigmata has about 30 rather
narrow slits.
The abdomen is only slightly narrower than the thorax. The oesophagus
extends about half the length of the abdomen, and is quite wide, except where
it narrows before entering the stomach. In lateral outline the stomach is rect-
angular and in transverse section there is a suggestion of about 6 indistinct
folds, but these, or some of them, may be artefacts. The post-stomach is
narrower than the stomach, and the mid-gut, which forms the lower, horizontal
part of the gut loop, is wider. The rectum is of nearly uniform diameter.
Of the gonads, only the testis was present in the zooids examined, and this
consisted of a quite massive group of pear-shaped follicles lying immediately
below the lower bend of the gut loop, with the sperm duct passing straight
forwards to the atrial cavity.
Remarks
I am following Huus (1937) in accepting Michaelsen’s genus Tetrazona for
a small group of species differing from Cystodites essentially in lacking spicules.
The type species was described by Sluiter (1906) as Distoma glareosus, and the
genus evidently also includes Polycitor vitreus (Sars) and probably Polycztor
magalhaensis (Michaelsen). Removal of this well-defined group of species with
four rows of stigmata leaves the genus Polycitor with those species having a much
larger, but variable, number of rows of stigmata, with the type P. crystallinus
Renier.
Genus sycozoa Lesson, 1830
Sycozoa arborescens Hartmeyer
(Fig. 16)
Sycozoa arborescens Hartmeyer, 1912, p. 316.
Known distribution
Off Cape Town, South Africa.
148 ANNALS OF THE SOUTH AFRICAN MUSEUM
Localities
False Bay: U.C.T., FAL 234 S.
Mossel Bay: U.C.T., MB 48 B; U.C.T., MB 65 F.
Algoa Bay: PF 586 A.
Description
The colonies which come closest to Hartmeyer’s (1912) description of the
type specimen are two from Mossel Bay (MB 48 B). These colonies measure
respectively 2-0 and 2-2 cm. in greatest diameter (fig. 16 A, B). They consist of
a short basal column from the top of which spring several heads, each incom-
pletely divided into two. The heads are fan-shaped and laterally flattened. In
r52 99 9OO oe &
-°9¢¢g00 00
e
O55. emr:
Fic. 16. Sycozoa arborescens Hartmeyer.
A-C, colonies.
colour the colonies are pale grey with the zooids showing as darker marks. The
zooids are arranged, as in Hartmeyer’s specimen, in vertical double rows. They
agree with the description of the type material, and like it, lack gonads.
The three specimens from False Bay do not have the characteristically
branched colony. Two of them are single stalked heads (fig. 16 C), and one is
a pair of headless stalks joined basally. One of the stalked heads has the flattened
fan-shape which is found in the heads of more typical specimens of the species
and has the same double linear arrangement of zooids. Some of the zooids of
this colony have brood pouches containing a number of developing eggs, but no
larvae.
Other specimens, from Mossel Bay (MB 65 F) and Algoa Bay, are less
typical, but I am also including them in this species. They are single stalked
heads resembling the small specimen shown by Hartmeyer (1912) in his text-
figure 5.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 149
Remarks
The form of branching, which was more clearly developed in the type
specimen than in any of the present specimens, readily distinguishes this species
from others of the genus. It is the only Sycozoa known from South African waters.
Genus DISTAPLIA Della Valle, 1881
Distaplia capensis Michaelsen
(Fig. 17)
Distaplia capensis Michaelsen, 1934, p. 141.
Known distribution
False Bay and Still Bay, Cape Province.
Localities
Eemcenoy We eal., FAL 136M; U.C.T., FAL 158 N; U:C.T, FAL 175
NEU Cris BAL 182 W.
? Mossel Bay: U.C.T., MB 27 I.
0.5 mm.
0.3 mm.
Fic. 17. Distaplia capensis Michaelsen.
A, colony; B, zooid; C, larva.
150 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
The best-developed and most typical colonies are these from FAL 175 N
(fig. 17 A). In these the basal part forms a short squat column and the head,
which alone contains the zooids, is slightly wider than the column. The basal
column, which Michaelsen (1934) designates the stalk, is white and tough, and
the head has clear soft gelatinous test but is red-brown owing to the pigmentation
of the thoraces. In the collector’s notes the colonies are described as ‘purplish,
sloppy’, or ‘off-white, gelatinous’.
I can add little to the detailed description of the zooids, as the present
material confirms in every respect the structure of the type specimens. In all
zooids the folding of the stomach was distinct. In those sexually mature, only
the testis was developed; this consisted of a rosette of about 10 follicles and was
situated beside the intestinal loop. The position of the gonad, entirely within
the abdomen, is an important point of distinction between the present species
and D. skoogi (see figs. 17 B and 18 B).
There is one specimen from MB 27 I which I include, with some doubt,
in D. capensis. This is a single colony of soft consistency and pink-brown colour,
with small zooids having nothing to identify them, as the gonads are not
developed. The thorax is orange in colour, in the preserved state. Many of the
zooids have a single larva, and this seems to differ in some respects from that
of D. skoogi, in which species the specimen otherwise might be placed. The
length, from the end of the papillae to the base of the tail, is 0-8 to 0-9 mm.
Compared with the larva of D. skoogi the trunk is deep. The arrangement of
the papillae and their basal ampullae is similar. On each side of the postero-
ventral part of the trunk there is a curved flat structure which partly conceals
the oesophagus and rectum, only the stomach and part of the intestine being
clearly visible. No structures of this kind were seen in the larva of D. skoogi.
As far as could be determined these structures appear to be the thickened sides
of the abdomen. It is therefore likely that this colony is not of D. skoogi and
may be of D. capensis.
Distaplia skoogi Michaelsen
(Fig. 18)
Distaplia domuncula Michaelsen, 1923, p. 15.
Distaflia skoogi Michaelsen, 1934, p. 131.
Known distribution
Walker Bay and False Bay, Cape Province.
Localities
False Bay: U.C.T., FAL 232 E.
Off Cape St. Blaize: PF 1864.
Algoa Bay: U.C.T., LIZ 34 H.
? Morrumbene Estuary: U.C.T., MOR 43 X.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS I51
Description
Some of the colonies of this species in the present collection invested the
carapace of crabs, but others were attached to the stems of algae, and it appears
that the association with a crab, although common, is not obligatory. The
smallest specimen is one of 1:3 cm. diameter, attached to an algal stem, and the
largest specimen is 5:2 cm. in length, and is part only of a colony. The colonies
lcm
o)
Uw
3
3
C)
4, toaooooveve
90000890000"
< P)
eanngange?
Vg ggagoll?
ie:
0.5mm.
Fic. 18. Distaplia skoogi Michaelsen.
A, colony, B, abdomen of zooid; C, larva; D, ocellus and otolith of larva; l.c., lens
cells; oc., ocellus; ot., otolith.
are of fairly firm texture and all have clearly visible small round or oval systems
of zooids, generally closely crowded together. The common cloacal openings
are fairly conspicuous.
The zooids have already been described (Michaelsen, 1923; Millar, 1955).
There is rather more variation in the shape of the atrial opening and languet
152 ANNALS OF THE SOUTH AFRICAN MUSEUM
than the previous accounts suggest. In some zooids of some colonies the atrial
opening is relatively small and is surmounted by a long languet which has 3
distal lobes; in other colonies the zooids have a wide opening with a short, broad,
undivided languet. The most characteristic feature of the zooid is the position
of the gonad, which is accommodated in a small pouch projecting from the end
of the abdomen (fig. 18 B). This pouch often projects even farther than is shown
in Michaelsen’s figure.
The colonies from FAL 232 E and LIZ 34 H had fully developed larvae.
Not more than one larva was seen in each breeding zooid. The larva (fig. 18 C)
measures 1:0 to 1:1 mm. from the end of the papillae to the base of the tail.
There are 3 large papillae, two dorsal and one ventral, the ventral papilla
lying to the right of the sagittal plane. Each papilla has two round ampullae
at its base. The branchial sac and gut are well developed. Both ocellus and
otolith are present in the sensory vesicle, but they are so close together that
under low magnification it might be thought that only one pigmented organ is
present. The ocellus has 3 lens cells, apparently in linear arrangement (fig. 18 D).
Remarks
Michaelsen (1923) described a new species from South Africa and named
it Distaplia domuncula, and I used the same name for a later record (Muillar,
1955). The name, however, was pre-occupied by a species which Hartmeyer
(1913) had described, and Michaelsen (1934) gave his own species the new name
Distaplia skoogt.
Michaelsen (1934) believed Dvstaplia skooge Michaelsen and Dhistaplia
domuncula (Hartmeyer) to be separate. A comparison of the three species
recorded from Cape Province is given in Table 1.
TABLE I
Colony Kooids Gonads Stomach
D. capensis Slightly Unisexual. Beside gut loop; 14-16 folds.
stalked. 10-14 male fol-
licles.
D. skoogi Not stalked; Hermaphrodite. In sac projecting Smooth externally;
usually in- from abdomen; with lines _inter-
vesting crabs. about 5 male nally.
follicles.
D. domuncula Not stalked; Probably uni- Incompletely Smooth externally;
investing sexual. known. with lines inter-
crabs. nally.
D. capensis is evidently distinct from D. skoogi and D. domuncula, but the
latter two species are similar and might eventually have to be united.
Genus SIGILLINA Savigny, 1816
I am using the genus Szgi/lina in the wide sense employed by Michaelsen
(1930) and Hastings (1931) for the following single species, which does not
agree completely with any of the sub-genera which Michaelsen defined.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 153
Szgillina vasta sp. n.
(Fig. 19)
Diagnosis of species
Colony massive, with a system of cloacal canals below the surface, and
apparently a few common cloacal openings. Zooids with long oral and atrial
Fic. 19. Sigillina vasta sp. n.
A, zooid with testis; B, abdomen of zooid
with ovary.
mm.
mm.
154 ANNALS OF THE SOUTH AFRICAN MUSEUM
siphons, the latter at the posterior end of the thorax opening into a cloacal canal,
and usually backwardly directed. Three rows of stigmata. Abdomen about the
same length as thorax, and with a long narrow posterior vascular process.
Gonad beside intestinal loop, with the ovary sometimes at the extreme posterior
end of the abdomen.
Holotype
In the South African Museum. $.A.M. A25612 (U.C.T., TRA 60 E).
Locality
Kaffirkuils Bay: U.C.T., TRA 60 E.
Description
The specimen is part of a colony brought up in the catch of a commercial
trawler, and the whole colony was, according to the note accompanying the
specimen, a ‘large gelatinous mass approximately 3 by 8 by 10 inches (2-3 lbs.)’.
In the preserved state the specimen is purple-brown, with a very smooth surface.
There is no encrusting material. A few irregular slits were found on the surface,
and these are taken to be the external openings of the common cloacal canals
which lie a little below the surface. The oral openings are apparently scattered
without order and the zooids are therefore apparently not arranged in regular
systems.
The zooids (fig. 19 A) are quite large, and may reach 7 mm. in length, with
the thorax and abdomen of about equal length. The oral siphon is remarkably
long, and may equal or even exceed the thorax in length. Its opening has 6
lobes. The atrial siphon arises from the posterior end of the thorax and is about
the same length as the oral siphon. It is generally directed obliquely back,
instead of towards the surface of the colony, and leads into one of the common
cloacal canals. There are 6 lobes on the opening of the atrial siphon.
At the base of the oral siphon there are about 4 large tentacles and many
short ones occupying a zone anterior to the large ones. The dorsal tubercle is
small and its opening could not be clearly seen. Three rows of long narrow
stigmata are present with at least 12 stigmata per row. The oesophagus is quite
long, the stomach almost globular, and the post-stomach and intestine much as
in species of Eudistoma. Some zooids have a group of about 1o testis follicles
arranged in a rosette beside the gut-loop (fig. 19 A), with the sperm duct
passing straight forward. In other zooids the ovary is developed, consisting of a
small group of oocytes, or sometimes a single large egg projecting from the
posterior end of the abdomen (fig. 19, B). There is a posterior vascular process
from the end of the abdomen of very variable length.
Remarks
This remarkable species is difficult to place in its right genus. It would fit
Eudistoma were it not for the presence of common cloacal canals into which the
atrial siphons open. Another species, originally described by Sluiter (1895)
under the name Distoma deerratum and later by Hastings (1931) as Szgzllina
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 155
deerrata also has a long oral siphon, and a posterior atrial siphon opening into a
common cloacal canal. This species, however, which Hastings identifies with
Polycitor coalitus Sluiter, is known only from the tropical waters of Malaya and
north and north-eastern Australia, and in spite of structural similarities it is
very unlikely that the South African specimen is of that species.
There may be a case for recognizing a new subgenus of Szgzllina to accom-
modate the two species S. deerrata and S. vasta which differ from Eudistoma in
having the atrial siphon opening to a cloacal system.
Genus EUDISTOMA Caullery, 1909
Eudistoma digitatum sp. n.
(Fig. 20)
Diagnosis of species
Fully grown colony massive and firm with solid base, and apex divided
into a number of finger-like lobes which do not spread, but have a tendency to
grow inwards. Test fibrous, with large bladder cells which are few and widely
spaced. Zooids with a short wide thorax, a longer but narrower abdomen, and
a long posterior process of variable width. Gonads beside intestinal loop.
Holotype
In the British Museum (Nat. Hist.). Bowerbank, ‘Cape of Good Hope’.
Localities
Mossel Bay: PF 11 A; U.C.T., MB 48 C.
Algoa Bay: U.C.T., LIZ 32 P; B.M. (N.H.) collection, reg. no. 1852.3.12.
81, collected by J. Bowerbank; B.M. (N.H.) collection, reg. no.
1852.3.12.89, collected by J. Bowerbank.
‘Cape of Good Hope’: B.M. (N.H.) collection, no reg. no., collected by
J. Bowerbank.
Description
The colony, when large, has a very characteristic appearance, best shown
by the specimens collected by Bowerbank from ‘Cape of Good Hope’ (fig. 20 A),
and Algoa Bay. The 3 largest specimens have the following dimensions:
height (cm.) 140 7:6 10:0
width (cm.) 12°5 6-3 5:0
From a solid base there arises an upright portion which is usually stout
and columnar, but may be very short. The apex of this portion is divided into
a number, up to 12, of characteristic finger-like or wide tongue-like lobes.
These lobes are grouped close together and instead of spreading outwards have
a tendency to grow inwards at the top, towards the vertical axis of the colony.
The whole colony is very massive and firm, and the surface smooth without
156 ANNALS OF THE SOUTH AFRICAN MUSEUM
any encrusting matter. There is no sand within the colony and the common test
is fairly homogeneous, except for fibres and a few scattered bladder cells. In
well-preserved colonies the zooids are sometimes visible externally; they are
mainly confined to the lobes.
The zooids (fig. 20 B) consist of a short wide thorax often about 1 mm.
long, and a narrower abdomen about 2 mm. long. A long vascular process
A B
3 cm.
mm.
Fic. 20. Eudistoma digitatum sp. n.
A, colony; B, zooid.
projects from the end of the abdomen and extends, often for many mm.,
downwards towards the base of the colony. Generally this process is very slender,
but in some colonies it is almost as wide as the abdomen and contains large
quantities of opaque tissue, probably consisting of reserve food cells. The
siphons are both short, with 6 indistinct lobes, and open close together on the
surface of the colony. The thorax is usually widest at its anterior end. About
16 oral tentacles were counted, but others were present spread over a zone at
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 157
the base of the siphon. There are about 16 stigmata in each of the 3 rows.
The length of the oesophagus varies, but is usually greater than that of the
thorax. It is tapered at the lower end where it enters the round or wide smooth-
walled stomach. A narrow post-stomach leads to the curved intestine. The
rectum is of uniform diameter. In mature zooids there is a group of many small
testis follicles situated beside the lower part of the gut loop and sometimes
projecting slightly posterior to it. ‘The ovary is represented by one large oocyte,
no doubt with other smaller ones not visible.
Remarks
This species, which should not have been mistaken for others, does not
seem to have been described, although several species of Eudistoma are known
from South African waters. It is remarkable for the large and massive appearance
of the colonies.
Eudistoma coeruleum (Sluiter)
(Fig. 21)
Distoma coeruleum Sluiter, 1898, p. 14.
Known distribution
Natal; ?Still Bay, Cape Province; Mozambique.
Localities
Natal: Bluff, M.S. Evans, British Museum (N.H.) reg. no. 1876.9.3.4;
Scottburgh, S.A. Museum, low water level.
Fic. 21. Eudistoma coeruleum (Sluiter).
A, larva; B, C, sensory vesicle of different larvae to show variation in position of
ocellus and otolith.
158 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
This species is easily recognized by the club-shaped colonies of dark slate-
blue or purple colour. Zooids are confined to the expanded head of the colony,
and in the present material measured from 1-5 to 2-5 mm. in length; Sluiter
(1898) gives 4-5 mm. as the maximum length of zooids. In some of the new
material and especially that from Scottburgh, there are moderately powerful
longitudinal and transverse muscles on the thorax and the longitudinal ones
pass down towards the ventral side of the abdomen. In the branchial sac at
least 10 stigmata were counted in each row. The dorsal languets are long and
slender.
Larva
Colonies from both stations had larvae, only one to three in the atrial
cavity of each breeding zooid. The larva (fig. 21 A) measures 0-5 to 0-6 mm.
from the end of the papillae to the base of the tail. The trunk is rather long and
narrow, with the sensory vesicle about the middle of the dorsal side. Both a
black otolith and a paler ocellus are present, but their relative position varies
(fig. 21 B, C). There are three anterior papillae.
Eudistoma modestum (Sluiter)
(Fig. 22)
Distoma modestum Sluiter, 1898, p. 18.
Known distribution
Durban, Natal.
Localities
Algoa Bay: U.C.T., LIZ 1 Z; U.G.T., LIZ 9 M; U.CY., LiZ sai
PF 895.
Description
The colonies vary a good deal in appearance, but most are rather soft and
gelatinous in texture, and flattened, the flattening perhaps being an artefact.
The specimens do not have a definite stalk, as in Sluiter’s type material. As
Sluiter’s specimens did not exceed 1 cm. in diameter, however, they may not
have reached the full size of the species; it is possible that further growth would
have brought about a change in shape. The colour is brown or semi-transparent
with brown zooids showing through.
The zooids (fig. 22) are 2-4 mm. in length, and have a shorter waist than
is common in the genus. Although Sluiter described the waist as long and thin,
I have found, on examining the type specimens, that in many zooids it is only
slightly longer than the thorax. The longitudinal thoracic muscles in the
present material are slender and number up to 18. In some colonies the siphons
and the anterior end of the thorax are chocolate coloured, and brown spots also
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 159
mark the rest of the thorax. Zooids of the type material, which I have examined,
have the same parts marked with pink. The difference in colour may be due to
a difference in the method of preservation. In most other respects the structure
of the zooids is like that of the type specimens, but the oral tentacles number at
least 30, whereas Sluiter recorded only 16.
| mm.
Fic. 22. Eudistoma modestum (Sluiter).
Zooid.
Remarks
This is one of those cases in which the new specimens agree fairly closely
with type material but show such differences as might be expected to result
from their having been collected at a different age, or stage of maturity.
160 ANNALS OF THE SOUTH AFRICAN MUSEUM
Eudistoma reniert (Hartmeyer)
(Fig. 23)
Polycitor (Distoma) reniert Hartmeyer 1912, p. 309.
Known distribution
St. Francis Bay, Cape Province.
Localities
Noxth) of (Durban 2045974, 5..038 lo ake.
60 m., ‘Gilchrist Collection’, British Museum
(N.H.) reg. no. 1926.8.1.5.
Description
The single example of this species is a colony
of roughly cylindrical form, about 7-5 cm. long
and 3:0 cm. in diameter. It is very firm in con-
sistency, partly owing to the sand which encrusts
and impregnates the test. There is a small area
round each zooid free of sand, and the sur-
face of the colony consequently has a mottled
appearance.
The zooids (fig. 23) are about 5 mm. long,
and are similar in proportions to those described
by Hartmeyer (1912). According to Hartmeyer
the lower part of the gut has a twisted loop
characteristic of the species. This feature was
also noticeable in the material from the British
Museum (N.H.). A narrow process of variable
length arises from the lower end of the abdomen,
but Hartmeyer did not find a similar process in
his material.
] mm.
Remarks
The present record extends the known range Fic. 23. Eudistoma renieri
of this species into the warmer south-eastern (Hartmeyer).
waters off the coast of Natal. Zooid.
Eudistoma illotum (Sluiter)
(Fig. 24)
Distoma illotum Sluiter, 1898, p. 16.
Polycitor (Distoma) illotus (Sluiter). Hartmeyer, 1912, p. 303.
Known distribution
Cape ‘Town and Simon’s Bay, Cape Province.
3
i
|
|
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 161
Localities
False Bay: FAL 175 L.
Mossel Bay: ?MB7Q; ?MB 22 F; ?MB 27 J; ?>MB 48 D.
Description
Colonies are of rounded, lobed, upright or even slightly stalked, shape.
No sand is present either on the surface or within the colony. The test is of a
milky or semi-transparent appearance. Large vesicular cells are present in the
test and these were noted by Sluiter (1898) as a distinguishing feature of the
species.
a
0.3 mm.
Fic. 24. Eudistoma illotum (Sluiter).
Larva.
In the colonies from False Bay the zooids are 3-4 mm. in length, and
slender with a long waist as in the type specimens. ‘The zooids in the colonies
collected from Mossel Bay range from 2 to 6 mm. in length and have only a
short waist. I am therefore a little doubtful of the identification of the material
from Mossel Bay.
Larva
No larvae were found in the present material, but Sluiter’s type specimens,
which I have been able to examine, have larvae (fig. 24). These range from 0-96
to 1:20 mm. in length, from the end of the papillae to the base of the tail. There
are 3 anterior papillae with a pair of narrow lateral ampullae between each two
adjacent papillae, on each side. Both ocellus and otolith are present. Only one
larva was found in each breeding zooid, perhaps owing to the large size of the
larvae.
Reliable characters in the genus Eudistoma are few, there is considerable
162 ANNALS OF THE SOUTH AFRICAN MUSEUM
similarity in the zooids of different species, and the colony is often variable.
For these reasons the identification of a number of specimens is doubtful and
I include the following two species with hesitation.
?Eudistoma angolanum Michaelsen
Polycitor (Eudistoma) paesslerioides var. angolana Michaelsen, 1914, p. 430.
Known distribution
Gulf of Guinea (W. Africa); NW. and W. Australia.
Localities
False Bay: FAL 158 Q; FAL 208 T.
Mossel Bay : MB 7 Es MB GoIR:
Algoa Bay: LIZ 34 F.
Description
The colonies are usually pillow-shaped, rounded, or slightly lobed. They
are coated and impregnated with sand and are consequently hard.
The zooids seldom exceed 6 mm. in length, of which the thorax generally
occupies 1°0-1°5 mm. Muscles are well developed, consisting principally of
about 10 longitudinal strands on each side of the thorax, and a powerful ring
of muscles on each siphon. The waist of the zooids is long, containing the
correspondingly long oesophagus and rectum. A short vascular process some-
times extends below the abdomen. Gonads, present in specimens collected in
January 1958, consist of a group of 11-20 or more testis follicles beside the
lower part of the intestinal loop, and a small ovary.
Remarks
E. angolanum has not been recorded from closer than tropical west Africa,
but I can find no important differences between that species and the present
specimens. The identification remains doubtful, however, owing to the diffi-
culty of separating species of Ewdistoma, and to the wide geographical separation
of previous records.
?Eudistoma mobiusi (Hartmeyer)
Colella mobiusii Hartmeyer, 1905, p. 396.
Polycitor mobiust Hartmeyer, 1912, p. 305.
Known distribution
Cape Province and Madagascar.
Localities
False Bay: FAL 208 R.
Algoa Bay: LIZ 19 Z 1.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 163
Description
The single colony from False Bay and the two from Algoa Bay are all
about the same size, 1:-2—-1-4.cm. long. There is a rounded or flattened head, and
a narrower but slightly longer stalk. Both head and stalk are firm and pale
grey to dull yellow in colour. Zooids are visible on the head.
The zooids usually attain a length of up to 7 mm. The oral tentacles are
very small and arranged in several circles. There are at least 15 stigmata in
each of the three rows. The waist of the zooid is rather short, so that the
oesophagus and rectum are not so long as in some species of Eudistoma. This is
a feature, however, which might change at different periods in the life of the
colony. No gonads were present.
Family Didemnidae Giard, 1871
Genus DIDEMNUM Savigny, 1816
The genus Didemnum has given systematists great trouble and even the
most experienced workers have hesitated in deciding specific distinctions. Thus
Van Name (1945, p. 80) wrote in his account of the American species “The
treatment of the species that is possible with present information is necessarily
somewhat provisional and will doubtless require future revision, but the evidence
seems to be that instead of Didemnum being one of the largest genera of ascidians,
the species are in reality rather few, though in some cases widely distributed
geographically.’ And again in treating Didemnum candidum Savigny he states
(p. 86) that ‘This is the most difficult of all the American ascidians to deal with
from a taxonomic point of view. I am far from being able to overcome the fear
that I am confusing more than one species, but after the examination of a large
amount of material from various American localities I am at a loss to find a
reliable basis for dividing it by studying museum specimens.’ The situation is
similar with regard to the South African species. Michaelsen (1934, p. 147), in
his description of D. stilense from Still Bay, Cape Province, had to admit that
‘Because of our uncertainty as to the limits of variability and because of the
occurrence of so many species insufficiently described, it is hardly possible to
say whether D. stzlensis is indeed a new species, or whether it should be identified
with any older species.’
I believe that to try to identify preserved specimens, usually few in number,
collected at different seasons and states of their annual cycle, and generally
without any biological data, would only add further confusion. A proper
taxonomic study of the genus should involve regular collection of specimens
throughout the year, in large numbers, together with observation of biological
characters such as breeding season. For these reasons I have made a very
tentative identification of a number of specimens of the genus Didemnum, as
D. stilense.
164 ANNALS OF THE SOUTH AFRICAN MUSEUM
?Didemnum stilense Michaelsen
Didemnum stilense Michaelsen 1934, p. 146. Millar, 1955, p. 176.
Known distribution
Cape Province.
Localities
Langebaan Lagoon: U.C.T., LB 367 V.
False Bay: U.C.T., FAL 121 B; U.C.T., FAL 136 N; U.C.T., FAL 152 M;
U.G.T., FAL 158 X; U.C.T., FAL 163 P; U.C.T., FAL a aan.
U.C.T., FAL 176 R; U.C.T., FAL 182 U; U.C.0:) BAISorGeeaS
U.G.T.,. FAL 239 E; U.GT., FAL 272 F; U.C.1,, FAL Roy ae
U.C.T., FB 1106 B.
Mossel Bay: U.C.T., MB 18 R.
Algoa Bay: U.C.T:, LIZ 9 K; U.C.T., LIZ 28 S; UC. Re izreae
PF 586 B.
Description
Colonies are generally thin and encrusting but otherwise are of very
variable appearance. In the preserved state the colour is white, grey, buff,
pink or blue, and the collector’s notes show that living material is also of various
colours, but that these may change considerably on preservation. Spicules are
usually many, closely crowded, and present throughout the whole test. They
are generally of a regular stellate form with 6-10 conical rays as seen in optical
section. Sometimes the rays are more numerous, or small and rounded, or with
blunt ends. The larger spicules are generally 20—30p in diameter and occasionally
as large as 40 or even 70p.
The thorax of the zooids is rather smaller than the abdomen, and possesses
a slender retractor process which may be almost as long as the abdomen. The
atrial opening varies in size and depends at least partly on the degree of con-
traction of the thorax. The gut is divided as usual in the genus. A single large
egg generally represents the ovary when this is well developed. The testis has
a single large follicle with the proximal part of the sperm duct forming 8-10
coils on its surface.
Larva
Larvae were present in colonies from the following collections:
False Bay, 17-2-53, 20-8-47, 18-9-53.
Mossel Bay, 13-1-51.
Algoa Bay, 7—4-54.
The larva has the general structure commonly found in the family. There
are 3 anterior adhesive papillae, 4 lateral ampullae on each side, and a sensory
vesicle with ocellus and otolith. Larvae varied considerably in size, the smallest
having a length of 0.4 mm. (False Bay, 18-9-53) and the largest a length of
ie >
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 165
0:95 mm. (Algoa Bay, 7—-4-54). This is a large range within one species, but
Carlisle (19544, 19540) has shown that didemnid larvae vary in size according
to the size of the zooids, which itself is variable.
Remarks
It must be emphasized that the identification of the above specimens as
D. stilense is tentative.
Subgenus POLYSYNCRATON Nott 1891
Didemnum (Polysyncraton) magnilarvum sp. n.
(Fig. 25)
Diagnosis of species
Colony flat but rather fleshy. Spicules few, stellate, up to 75 in diameter.
Common cloacal openings numerous. Zooids with wide atrial opening, short
atrial languet, slender retractor process, testis of 8 to 10 follicles, and sperm duct
of 2 to 3 coils. Larva with trunk about 1-3 mm. long, having a collar of about
30 narrow ampullae surrounding the 3 papillae. Budding is precocious,
advanced larvae each having 2 buds.
Holotype
In the South African Museum. S.A.M. A25613 (PF 12393 B).
Locality
Natal: PF 12393 B.
Description
The single colony is 6 cm. by 3:7 cm. in diameter and 0-7 cm. thick. It is
flat but of a rather fleshy appearance. In the preserved state the colour is
greyish-pink. The upper surface is smooth without any encrusting matter, and
the whole colony is soft and flexible. Certain areas of the colony are paler owing
to the presence of greater numbers of spicules, which, however, are scarce in
most places. The common cloacal openings are numerous and scattered
irregularly over the surface. Spicules are nowhere numerous in the common test,
but are most plentiful near the upper and lower surfaces and scarcest in the
central layers. ‘They are stellate and the largest ones reach 75u in diameter, but
most are smaller.
The zooids (fig. 25 A, B) do not show any obvious arrangement in systems.
They are generally about 1 mm. long, the thorax being only slightly longer than
the abdomen. The oral siphon is tubular, with 6 rather shallow lobes. Theatrial
opening is wide and exposes quite a large part of the branchial sac. A short atrial
languet is present; it has a narrow base and slightly wider free end which is
sometimes indented, giving the languet two shallow lobes. No trace was found
of lateral thoracic organs, and there were no accumulations of spicules beside
166 ANNALS OF THE SOUTH AFRICAN MUSEUM
the thorax such as generally mark the position of these organs in species which
possess them.
There are 4 rows of stigmata, and a long slender retractor process from the
base of the thorax. The oesophagus is of moderate length, leading to the ovate
or almost globular stomach. There is a short post-stomach, and simple intestinal
loop. The testis consists of a rosette of 8 to 10 follicles, and the sperm duct makes
about 24 spiral turns before passing forward towards the rectum.
Larva
Numerous larvae (fig. 25 CG) were embedded in the common test. When
fully developed these have a trunk about 1-3 mm. long, measured from the end
of the papillae to the base of the tail. The 3 papillae project from the centre of a
0.5 mm.
eed
0.5 mm.
Fic. 25. Didemnum (Polysyncraton) magnilarvum sp. n.
A, thorax of zooid; B, abdomen of zooid; C, larva.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 167
collar of about 30 narrow finger-like ampullae, formed by the subdivision of the
paired ampullae generally present in didemnid larvae. The sensory vesicle has
both ocellus and otolith. In addition to the branchial sac and gut of the oozoid
there is, on each side of the larval trunk, an equally well developed branchial sac
and gut representing the first two blastozooids. A similar precocious development
was noted by Lahille (1890) in Dizdemnum (Polysyncraton) lacazi. In the Didem-
nidae this is characteristic of Diplosoma but I do not know of other references to
the phenomenon in Polysyncraton. ‘The larvae of only a few species are known,
however, and I believe that the precocious appearance of blastozooids, together
with the subdivided anterior collar, may be found characteristic of larvae in the
sub-genus Polysyncraton.
Remarks
Few species of the sub-genus are known from the waters of South Africa
or the western Indian Ocean. D. (Polysyncraton) spongiotdes (Hartmeyer) and
D. (Polysyncraton) chuni (Hartmeyer), both from 318 m. off Cape of Good Hope,
appear to be very closely related to each other, if distinct, but they are clearly
distinguished from D. (Polysyncraton) magnilarvum by their small number (3 or 4)
of testis follicles. D. (Polysyncraton) tubtporae (Michaelsen) from Zanzibar is
distinguished by its characteristic lateral thoracic organs with adjacent masses
of spicules. D. (Polysyncraton) paradoxum Nott var. mahenum Michaelsen is
described from the Seychelles and the species originally from New Zealand.
The variety seems to be rather like the present species, but has much smaller
spicules and usually only 4 testis follicles. D. (Polysyncraton) amethysteum (Van
Name) is known from tropical Atlantic America, tropical west Africa and the
eastern Mediterranean; it is therefore a species of widespread occurrence in
warm waters and might be expected to occur in the Indian Ocean. It resembles
the new species but has smaller spicules, fewer testis follicles and a larva only
half as large.
Genus LEPTOCLINIDES Bjerkan, 1905
Leptoclinides capensis Michaelsen
Leftoclinides capensis Michaelsen 1934, p. 151.
Known distribution
Preekstoel, near Still Bay, Cape Province.
Localities
able Bay WG eb 171 As
Balsebayen) Cube EAL dig:
Description
The specimen from False Bay is a colony 1 cm. in diameter, which had
been growing on the carapace of a small dromiid crab, according to the
168 ANNALS OF THE SOUTH AFRICAN MUSEUM
collector’s note, but the specimen from Table Bay is incomplete and the sub-
stratum unknown. In the colony from False Bay there is a single central common
cloacal opening; Michaelsen (1934) also apparently found only one opening in
his type specimen. I have found the structure of the common test to agree in
general with Michaelsen’s description, except in the absence of bladder cells
from the whole superficial layer, the type specimen having a narrow subcortical
layer without bladder cells. Spicules are few, being concentrated almost entirely
near the opening of the oral siphon and at the lateral thoracic organs. The
zooids agree well with those of the type specimen, but have rather larger lateral
thoracic organs. Six spiral turns of the sperm duct lie over the single testis
follicle.
The colony from Table Bay, collected on 8-4-1946, had developing larvae
in the common test, but none fully formed.
Genus LissocLinum Verrill, 1871
Lissoclinum cavum sp. n.
(Fig. 26)
Diagnosis of species
Colony consisting of an upper and a lower layer of common test, separated
by a large common cloacal space across which the zooids are stretched. Spicules
up to 40u in diameter. Zooids with a very large atrial opening, and two groups
of spicules over the postero-ventral corners of the thorax. Testis with about
5 follicles. Sperm duct not coiled.
Holotype
In the South African Museum. S.A.M. A 25614 (U.C.T., LIZ 32 V).
Localities
False Bay: U.G.T., FAL 234 Ut.
Algoa Bay: U.C.T., LIZ 32 V.
Description
The specimens from both stations are thin irregular sheets, of a dirty yellow
or brown colour. Small darker spots on the surface mark the oral openings.
The quite numerous round or oval common cloacal openings are scattered over
the surface. The colony consists of an upper thin layer of common test and a
lower rather thicker layer. Between these layers is the very large common cloacal
cavity. Spicules (fig. 26 A) are numerous and closely packed in the upper and
lower layers of the colony, and are also present in the test which covers the
abdomen. They are stellate but somewhat irregular with a tendency for the
rays to be arranged in pairs, or to be reduced to round swellings. ‘The maximum
diameter found was 40u. In addition there is an ovoid mass of spicules at the
postero-ventral corners of the thorax, presumably marking the position of the
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 169
lateral thoracic organs. The zooids (fig. 26 B), each enclosed in a thin sheath of
test material, are suspended between the two layers of test. Most zooids are
between 1 and 2 mm. in length. The thorax is generally slightly longer than
the abdomen. The oral siphon is tubular, of variable length, and ends in 6
indistinct lobes. Much of the dorsal surface and sides of the thorax are exposed
Sy
30 u
0.5 mm.
| a ee |
0.2 mm.
Fic. 26. Lissoclinum cavum sp. n.
A, spicules; B, zooid; C, larva.
by the very large atrial opening, the anterior border of which has a short
projecting lip or languet. There are about 24 long slender oral tentacles which
meet across the base of the oral siphon. Each of the four rows has about 14
stigmata. The abdomen lies almost at right angles to the thorax, thus displacing
the gut from the attitude normal in didemnids. The stomach, which is ovoid
or cylindrical, is followed by a short post-stomach, and an intestine and rectum
of almost uniform diameter. No ovary was present in any of the zooids examined.
The testis consists of a rosette of 5 or 6 wedge-shaped follicles, and the sperm duct,
originating from the centre of the rosette, does not make any spiral turns, but
is gently curved near its point of origin.
Larva
The specimen from Algoa Bay, taken on 6-4-1954, contained larvae (fig.
26 C). The trunk of the larva is about 0-6 mm. long from the end of the papillae
170 ANNALS OF THE SOUTH AFRICAN MUSEUM
to the base of the tail. The papillae project from the centre of a group of
irregular ampullae. Both ocellus and otolith are present in the sensory vesicle.
Remarks
Lissoclinum bilobatum Millar is perhaps the South African species most like
L. cavum, but is distinguished by having only 2 testis follicles.
Genus TRIDIDEMNUM Della Valle, 1881
Trididemnum cerebriforme Hartmeyer
Trididemnum cerebriforme Hartmeyer, 1913, p. 139.
Known distribution
Cape Province.
Localities
Saldanha Bay: U.C.T., SB 130 U.
Langebaan Lagoon: U.C.T., LB 352 U; U.C.T., LB 374 F; U-CT EB
Bye on OP Cal MA Heys Sms es UN Col bee Fis aig (Ge
False Bay: U.C.T., FAL 136 8; U.C.T., FAL 144 Z; U.C-T,, FA or6us:
U.G.T., FAL 225 P; U.C.T., FAL 234 U2; U.C.i., FA aoees
Mortensen’s Java—South Africa Expedition 1929-30. Station 61, B.M.
(N.H.) reg. no. 34.2.1.3; Mortensen’s Java—South Africa Expedition,
1929-30. Station 65, 19.12.29, B.M. (N.H.) reg. no. 34.2.1.5.
Mossel Bays U.C.1T., MB 59 K; U:C/E. MBS. S:
Algoa Bay: U.G.T., LIZ 9 L; U.C.T.; LIZ 28 T; U.C i) Eiaieeae
UW. Gol EYZ oak
Commercial trawlers: U.C.T., TRA 71 F.
Description
The largest of the three specimens on which Hartmeyer (1913) based this
species had a much-folded surface but in the smallest the surface was almost
plain. The brain-like folding cannot therefore be regarded as a diagnostic
character. Among the numerous specimens in the South African collections
that I have examined there are many colonies with a smooth surface and only
a few which are much convoluted. One of the latter kind, taken by Mortensen’s
expedition, is folded over so that the anatomically lower surfaces are almost in
contact, leaving an extensive but narrow space into which open slits which I
take to be the common cloacal openings. ‘These openings are therefore on the
under surface of the colony, a condition thought to exist (Michaelsen, 1930) in
Leptoclinides brasiliensis, but otherwise very rare in the Didemnidae. In all other
respects this South African specimen agrees with more typical examples of
T. cerebriforme. The zooids of various colonies show much variation in the
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS El
amount of pigment on the thorax. In some there is no pigment, in others a
small dark spot over the anterior end of the endostyle, and in many specimens
the thorax has a great quantity of dark blue pigment.
Genus DipLosomMA Macdonald, 1859
Diplosoma listerianum Milne Edwards
Leptoclinum listerianum Milne Edwards, 1841, p. 84.
Diplosoma listerianum Michaelsen, 1919, p. 42.
Known distribution
West coast of Europe; Mediterranean; Adriatic; Sargasso Sea; west
Africa; Luderitz Bay, South West Africa to Durban, Natal.
Localities
Saldanha Bay: U.C.T., SB 92.
Malse Bay: U.C.0., FAL 109 Y; U.C.T., FAL 272 F.
misoa bays, U.C. 1. LIZ 1 Y.
Remarks
I can find no constant differences between these specimens and colonies
of D. listerianum from British waters. There is some doubt, however, regarding
the validity of certain species of Diplosoma, and it may prove that D. modestum
described by Michaelsen (1920) from the Seychelles and Zanzibar is identical
to D. listerianum. The Atlantic American D. macdonaldi Herdman also appears
to be very similar anatomically, but Van Name (1945) was unwilling to
identify it with D. listerianum without having examined European material.
This is a genus in which anatomical features of colony and zooid are very
uniform and specific differences may have to be sought in larval or biological
characters.
Family Cionidae Lahille, 1887
Genus cCIONA Fleming, 1822
Ciona intestinalis (Linnaeus)
Ascidia intestinalis Linnaeus, 1767, vol. 1, pp. 2, 1087.
Ciona intestinalis Fleming, 1822, p. 512.
Known distribution
West coast of Europe, Mediterranean, tropical west Africa, east and west
coasts of north America, Malay Archipelago, Japan, Australia, South Africa.
Localities
Saldanha Bay: U.C.T., SB 94; U.C.T., SB 134 A.
Description
The South African specimens are typical of the species as found in European
waters. I was able to confirm the distribution of the longitudinal muscles to the
siphons, as previously described (Millar, 1953).
172 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Ascidiidae Adams, 1858
Genus ascip1a Linnaeus, 1767
Ascidia stenodes sp. n.
(Fig. 27)
Diagnosis of species
Anterior part of body narrow and elongated, with terminal oral siphon.
Atrial siphon posterior to middle of body. Body wall with a sharply defined
band of transverse muscles along the dorsal side. Gut small, mainly in the
posterior part of the body, and forming a single narrow loop with the anus near
the oesophagus.
Holotype
In the South African Museum. S.A.M. A25615 (U.C.T., FAL 259 Q).
Localities
False Bay: U.C.T., FAL 5 J; U.C.T., FAL 259 QO; U.G. 9.) FB vex
B
Fic. 27. Ascidia stenodes sp. n.
A, specimen with test removed, seen from the left; B, dorsal tubercle;
C, region of atrial opening, an., anus; atr.op., atrial opening; ovd., oviduct.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 173
Description
There are 6 specimens in the collections, the largest being about 5:5 cm.
long, and the smallest 1-0 cm. long. The surface of all specimens is thickly
coated with small stones, pieces of shell and sand which obscure the shape of
the body. The test, when freed from adhering particles, is transparent and not
very thick. When the body is removed from the test it is seen to consist of a
long tapering anterior part terminating in a short narrow oral siphon, and a
wider posterior part which accommodates the gut (fig. 27 A). The short incon-
spicuous atrial siphon is near the posterior end of the body and in young
specimens very near it. Most of the body wall of the left side is thin and trans-
parent, without muscles, but a thick narrow band of transverse muscles runs
along the dorsal side and extends only a short distance on to the left and right
sides; the margins of this band are sharply defined. The left side of large
specimens also has a ventral band of transverse muscles, but this is absent in
small specimens.
The oral tentacles number about 30 in a specimen of body length 1-3 cm.,
about 40 in one of 1-7 cm. and 60 in one of 5:5 cm. In two small specimens
examined, the opening of the dorsal tubercle is a slightly curved longitudinal
slit with the concavity to the left (fig. 27 B). In large specimens it is an irregular
C-shaped slit. The small ovoid ganglion is immediately posterior to the tubercle.
The branchial sac extends back to the posterior end of the body. The dorsal
lamina is a moderately wide smooth-edged membrane. The longitudinal bars
are of uniform thickness and number about 40 in the specimen 1-7 cm. long.
There are 2—7 stigmata in each mesh according to the size of the specimen.
Rather small papillae are present at the intersections of transverse and longi-
tudinal bars, but no intermediate papillae.
The gut is rather small. It starts with the short curved oesophagus lying
across the body and leading to the ovoid stomach which passes obliquely
forward. A few longitudinal folds are present on the wall of the stomach. The
intestine makes a simple loop and the rectum ends near the base of the atrial
siphon, in a small two-lipped or simple anus.
The gonad, as usual in the genus, is spread over a large part of the intestinal
wall, and the ducts lie beside the intestine and rectum (fig. 27 C).
Remarks
The diagnostic features of this species have already been noted. To them
might be added the covering of shell fragments or stones, which was found on
all specimens, but this is a character which may depend on the nature of the
substratum.
According to the collector’s note the specimens FAL 259 Q were lying
unattached on the bottom and this may be the usual condition in the species.
Ascidia sydneiensis Stimpson
Ascidia sydneiensis Stimpson, 1855, p. 387.
For synonymy see Kott 1952, pp. 310, 311.
LA. ANNALS OF THE SOUTH AFRICAN MUSEUM
Known distribution
West Indies; Hawaii; Malay Archipelago; Amboina; Japan; Australia;
South Africa.
Localities
False Bay: U.C.T., FAL 158 V; U.C.T., FAL 175 Q; U.C.T., FAL 181 A;
from catch of a trawler, S.A. Museum, April 1904.
Mossel Bay: U.C.T., MB 18S.
East London: S.A. Museum no. 12943, low tide.
Algoa Bay: U.Q.T., LIZ 32 S.
S.A. Museum (no details).
Description
These specimens agree closely with those which I have already described
from South African waters (Millar, 1955). The largest of the present animals
is 13 cm. long.
Remarks
Except for the South African records A. sydneiensis is known only from
warm waters where, however, it is very widely distributed throughout the
world.
I am not sure that A. sydneiensis, A. incrassata Heller, and possibly A. multi-
tentaculata Hartmeyer, all recorded from South African waters, might not prove
to represent a single species.
Family Corellidae Lahille, 1887
Genus CORELLA Alder & Hancock, 1870
Corella eumyota ‘Traustedt
Corella eumyota Traustedt, 1882, pp. 271, 273.
For synonymy see Kott, 1952, pp. 318, 319.
Known distribution
Antarctic; Subantarctic; South Africa; St. Paul (Indian Ocean); New
Zealand; Australia.
Localities
Langebaan Lagoon: U.C.T., LB 367 U; U.C.T., LB 374 E; U.C.T., LB
375 X.
Family Agnesiidae Huntsman, 1912
Genus AGNESIA Michaelsen, 1898
Agnesia glaciata Michaelsen
Agnesia glaciata Michaelsen, 1808, p. 370.
Agnesia krausei Michaelsen, 1912, p. 181, figs. 24, 25.
Agnesia capensis Millar, 1955, p. 191, fig. 19.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 175
Remarks
No specimens of A. glaciata are present in this collection, but material
which I previously examined from South Africa contained one specimen of a
species which I described as a new species A. capensis (Millar, 1955). During
examination of the Discovery collections I found another specimen, also from
False Bay, the same locality as that of the type specimen of A. capensis. The
Discovery specimen was intermediate between A. capensis and A. glaciata, and I
have given reasons (Millar, 1960) for regarding A. capensis as a synonym of
A, glaciata.
Family Styelidae Sluiter, 1895
Subfamily Botryllinae Adams, 1858
Genus BOTRYLLUs Gaertner, 1774
Botryllus magnicoecus (Hartmeyer)
Botrylloides nigrum var. magnicoecum Hartmeyer, 1912, p. 271.
Botryllus magnicoecus (Hartmeyer). Michaelsen and Hartmeyer, 1928, p. 331.
Known distribution
South Africa; Australia; New Zealand.
Localities
False Bay: U.C.T., FAL 109 Z; U.C.T., FAL 158 U; U.C.T., FAL 175 GC;
U.G.T., FAL 225 M; PF 15797 B; PF 14582.
Table Bay: Woodstock Beach, S.A. Museum, 19-6—-14, washed up on
beach, collected by K. H. Barnard.
Saldanha Bay: U.C.T., SB 128 P.
Algoa Bay: U.G.T., LIZ 9 H.
Botryllus anomalus sp. n.
(Fig. 28)
Diagnosis of species
Atrial openings of the zooids lead directly to the surface of the colony,
instead of into common cloacal chambers. Zooids with 8 rows of stigmata,
5 or 6 folds on the stomach, and a short straight pyloric caecum. A gonad on
each side of the body, each gonad consisting of a group of 6 or 7 testis follicles
and a single large ovum.
Holotype
In the South African Museum. S.A.M. A25616 (Saldanha Bay, 5-9-12).
Localities
Saldanha Bay: S.A. Museum, level of low water, 5—9-12, collected by
K. H. Barnard.
Table Bay, Woodstock Beach: S.A. Museum, washed up on beach, 19—6-14.
collected K. H. Barnard.
% -
i a —
176 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
There are several colonies of this species in the collection. A large specimen
is 9 cm. long, 3 cm. wide, and less than 1 mm. thick. All specimens have been |
growing on thin flat algal fronds. ‘The surface of the colony is smooth and even, ‘
without any foreign matter, and the colour of the specimens, in the preserved
state, a dull pink. There are no common cloacal cavities in the test and the
Fic. 28. Botryllus anomalus sp. n.
A, surface view of part of colony to show arrange-
ment of zooids, and vascular ampullae; B, zooid,
seen from the dorsal side; C, transverse section of
stomach.
0.5 mm.
zooids are not arranged in systems, but are closely and regularly spaced over
the colony (fig. 28 A). Each zooid (fig. 28 B) has a pair of small openings on
the dorsal side, marking the oral and atrial siphons, both of which open directly
on the surface of the colony. Rounded vascular ampullae are present between
the zooids.
The zooids are small, rarely exceeding 1 mm. in length. Near the anterior
end of the branchial sac is the round oral opening, through which can be seen
the 8 tentacles. The branchial sac has 3 inner longitudinal bars on each side,
and 8 rows of stigmata along the length of the sac. The digestive system consists
of a short oesophagus leading from the posterior end of the branchial sac to the
short barrel-shaped stomach. This has 5 or 6 complete folds on the walls
4
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 7/7)
(fig. 28 C), and a short straight pyloric caecum with slightly swollen distal end.
The intestine makes a sharp bend to the left and upwards to the short rectum
which ends in the simple anus below the atrial opening. On each side of the
body is a single gonad, each gonad consisting of a group of 6 or 7 pear-shaped
testis follicles and one large dark brown ovum. The left gonad is slightly anterior
to the right one. No larvae were present in the colonies.
Remarks
This species is chiefly remarkable for the opening of the individual atrial
siphons of the zooids directly on the surface of the colony, and the corresponding
absence of common cloacal chambers. The same features characterize Botryllus
primigenus Oka which, however, is distinguished by the presence of only 4 rows
of stigmata in the branchial sac, and the rather larger, curved pyloric caecum.
Genus BOTRYLLOIDES Milne Edwards, 1841
?Botrylloides leacht (Savigny)
Botryllus leachii Savigny 1816, p. 199.
Known distribution
Western Europe; Mediterranean; Australia; New Zealand; South Africa.
Localities :
Saldanha Bay: U.C.T., SB 91.
Remarks
This species has already been recorded by Michaelsen (1934) from Table
Bay and False Bay. No gonads are developed in the present specimen, and
I am identifying it as B. leacht because it agrees with that species in other
characters. It must be admitted, however, that the distinctions are not entirely
satisfactory in a number of species in both Boérylloides and Botryllus.
A single colony (U.C.T., SB 89, 13—7—46) from the same area is similar
but has more circular systems and I am not sure if it is of this species, or possibly
Botryllus schlosseri (Pallas).
Botrylloides mgrum Herdman var. giganteum Pérés
Metrocarpa nigrum var. giganteum Pérés, 1949, p. 205.
Known distribution
(Of variety) Senegal, west Africa; Durban, Natal and Knysna, Cape
Province.
Locality
Morrumbene Estuary: U.C.T., MOR 122 G.
178 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
The colony is dark purple in the preserved state. The zooids are large, with
a long narrow thorax containing about 16 rows of stigmata. There is a small
pyloric caecum. The gonad consists of an anterior testis with about 12 follicles,
and a posterior ovary containing one large ovum.
Subfamily Polyzoinae Hartmeyer, 1903
Genus GYNANDROCARPA Michaelsen, 1900
Gynandrocarpa unilateralis (Michaelsen)
(Fig. 29)
Gynandrocarpa placenta (Herdman) var. unilateralis Michaelsen 1900, p. 29.
Gynandrocarpa placenta (Herdman). Michaelsen 19048, p. 30.
[non] Goodsiria placenta Herdman, 1886, p. 328.
Gynandrocarpa domuncula Michaelsen, 19044, p. 247.
Known distribution
South Africa; south-east Africa.
Localities
False Bay: U.G. 0), PAL 265, 8; U: Cale WANG oa):
Mossel Bay: U.C.T., MB 53 Q; U.C.T., MB 65 G; PF 1711; PF 10165.
Algoa Bay: PF 1095 C; Port Elizabeth, B.M. (N.H.) registered no. 1871-
5-12-1; U.C.T., LIZ 28 N.
Description
The colonies are of two types: (1) a flattened upright disc with basal stalk
attached to one edge of the disc (fig. 29 A), and (2) a hollow dome-shaped
plate with no stalk (fig. 29 B). The stalked type of colony appears to have been
attached, during life, to a firm object on the bottom of the sea, but the dome-
shaped specimens were taken from the carapace of crabs. ‘These latter colonies
were originally considered to be a separate species, G. domuncula Michaelsen,
but Michaelsen (1934) later regarded them merely as colonies which are
modified in shape by their attachment to the crab. Having examined several
specimens of both types I entirely agree with this view, as I find no differences
in the zooids or larvae of the two types.
Preserved specimens are generally dull buff, grey, or pink-grey in colour,
but in some colonies the zooids are quite bright pink, which may have been the
colour of the colony in life. One specimen is described in the collector’s note
as ‘strawberry red’.
Michaelsen (1904a) has dealt thoroughly with the structure of the zooids
of this species under the name G. domuncula. The most interesting feature is the
gonad (fig. 29 C), which is single and on the right side. Many zooids of the new
material had a well-developed gonad. This consists of a sac-like ovary flanked
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 179
on each side by a rounded or ovoid testis. The oviduct is short and has its distal
part expanded to form a brood pouch which lies against the outer surface of
the branchial wall. The brood pouch opens into the cavity of the branchial sac
by means of an oval slit in the branchial wall, which at this place is devoid of
stigmata. In the present material the brood pouch of breeding zooids never
contained more than 2 or 3 embryos or larvae. One zooid contained a straight-
tailed fully developed larva in the branchial sac, thus confirming that larvae
pass from the brood pouch into the branchial sac before escaping to the
exterior.
Each testis has many small pear-shaped follicles, the ducts from which unite
to form one duct from each follicle, and these join the single sperm duct. The
emb.
_—————E— Eee
]} mm.
Fic. 29. Gynandrocarpa unilateralis (Michaelsen).
A, stalked colony; B, stalkless colony; C, gonad; br.p., brood pouch; emb., embryo; 1.b., longitudinal
branchial bars; ovy., ovary; sp.d., sperm duct; te., testis; D, larva.
180 ANNALS OF THE SOUTH AFRICAN MUSEUM
sperm duct opens into the peribranchial cavity. Michaelsen has pointed out
that, with the oviduct opening to the branchial sac and the sperm duct to the
atrial cavity, there may be obligatory cross-fertilization between zooids.
Larva
The larva (fig. 29 D) is large, having a trunk 1-0 to 1:1 mm. long and a
tail of 2-0 to 2:1 mm. Epidermal ampullae almost completely cover the anterior
two-thirds of the trunk. There are 2 dorsal and one ventral papillae. The
rudiments of the oral and atrial siphons are already present in the larva. A
sensory vesicle is present but contains neither ocellus nor otolith. In shape and
organization this larva is very similar to that of Dextrocarpa solitaris (Millar,
1955), but is more than twice the size. Larvae obtained from the stalked type
of colony and from the stalkless (“(domuncula’) type are indistinguishable.
Remarks
There has been confusion between this species and another South African
species of similar shape, Polyandrocarpa placenta (Herdman), as I have already
pointed out (Millar, 1955). Both the form of the colony and the general structure
of the zooids are alike in the two species, which are most clearly separated by
the gonads. In P. placenta there are several polycarp-type gonads, a condition
very different from that described above in G. unilateralis. It is remarkable,
however, that the larvae of P. placenta have, according to Herdman (1886), an
‘arrow shaped’ trunk about 1-0 mm. long and a tail 2-5 mm. long. They there-
fore agree well with the larvae of G. unilateralis, and also appear to lack both a
pigmented ocellus and otolith. But the larvae of P. placenta were present in the
peribranchial cavities and not, as in G. unilateralis, in the brood pouch or
branchial sac.
Genus DEXTROCARPA Millar, 1955
Dextrocarpa solitaris Millar
(Fig. 30)
Dextrocarpa solitaris Millar, 1955, p. 200.
Known distribution
False Bay, Cape Province.
Localities
PF 18785 B.
Description
Each specimen consists of a group of pear-shaped individuals joined by a
narrow stalk to a basal stolon (fig. 30 A). They therefore have a social rather
than a colonial or solitary organization. The whole test is heavily coated with
sand and shell fragments. The stalk uniting the individuals with the stolon
varies in length, being in some cases much shorter, and in others longer, than
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 181
the body. Within the stalk there is a short extension of the innermost layer of
test which surrounds the body (fig. 30 B, t), but no extension of the body wall,
and the individuals therefore do not appear to be in organic connection with
each other. In the type specimens I found no connexion between individuals,
which were therefore regarded as quite solitary. The present specimens show a
0.3 cm.
Fic. 30. Dextrocarpa solitaris Millar.
A, a group of individuals; B, specimen with test
removed, seen from the left; t., test.
condition intermediate between a fully social organization with zooids in
organic connexion, and a quite independent organization. It may be that in
this species budding is confined to relatively young individuals, and in older
ones complete separation may occur, or the apparently social organization may
result from the fusion of the test of neighbouring but separate individuals.
I have little to add to my earlier account of the species. A large individual
dissected had about 38 oral tentacles, 19 folds on the stomach, and the following
arrangement of longitudinal bars in the branchial sac:
dorsal line o (8) 2 (7) 3 (8) 3 endostyle.
ee
182 ANNALS OF THE SOUTH AFRICAN MUSEUM f
Breeding individuals had up to 110 eggs each, in various stages of development,
contained in the brood pouch. The larvae are as previously described. i
Genus ALLOEOCARPA Michaelsen 1900 i
Alloeocarpa capensis Hartmeyer
(Fig. 31)
Alloeocarpa capensis Hartmeyer, 1912, p. 261. é
Known distribution :
St. Francis Bay, Cape Province.
Localities
Algoa Bay: PF 740; PF 2348.
Fic. 31. Alloeocarpa capensis Hartmeyer.
A, gut; B, gonads; C, larva.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 183
Description
The collection contains 4 colonies, or pieces of colonies, from Algoa Bay,
the largest of which is 10 cm. by 4:5 cm. in diameter. In external appearance
these specimens are very like the type material illustrated by Hartmeyer (1912,
plate 38, fig. 7). The colonies are attached to shells or algal stems. They are of
a dirty grey-brown colour and the surface has low but conspicuous swellings
over the zooids.
The specimen collected off Lion’s Head has zooids which are more or less
free, being united only to a basal sheet of test by their narrow posterior ends.
The zooids, when extracted from the common test, are dull red.
Hartmeyer’s (1912) description of the zooids leaves little to be added, but
I have found the arrangement of the gonads to be different. According to
Hartmeyer the male gonads are on the left half of the body and the female on
the right. In all the mature zooids which I have examined, from colonies at
both collecting stations, the testes and ovaries were on the left, the testes in the
anterior part and the ovaries far back in the posterior part (fig. 31 B). Itis a
little difficult to determine the exact position of the ovaries relative to the
middle line of the body, but careful examination shows them to lie to the left
of the membrane which unites the endostyle to the ventral body wall. I have
seen no gonads male or female to the right of this membrane, in any of the
many zooids dissected. The position of the ovaries is confirmed by the location
of larvae, which are in the left peribranchial cavity.
Larva
Only a few zooids had larvae (fig. 31 C), and there were never more than
two in a breeding zooid. The larva is large, the trunk being 0-9 to 1:0 mm. long,
and the tail about 1-3 mm. long. The trunk is brick-red. There are 3 anterior
papillae and, a short distance behind these, a complete ring of about 24 narrow
ampullae round the trunk. When the larva has been dehydrated and cleared a
single spherical black body is seen, in the sensory vesicle, but is quite hidden by
the red pigment of the trunk in the untreated larva. This body is presumably a
‘photolith’, the compound sensory structure of larvae in several genera of the
subfamily Polyzoinae.
Genus METANDROCARPA Michaelsen, 1904a
Metandrocarpa fascicularis sp. n.
(Fig. 32)
Diagnosis of species
Zooids separate, with a narrow stalk joining them to the basal common
test. About 30 oral tentacles in two circles. Dorsal tubercle with a straight
oblique slit. Branchial sac with 7-9 longitudinal bars on each side. Stomach
with about 16 folds and a hooked pyloric caecum. Ten to 20 testes on each side,
on the posterior part of the body wall. Two to 5 ovaries on each side, on the
anterior part of the body wall.
184 ANNALS OF THE SOUTH AFRICAN MUSEUM
Holotype
In the South African Museum. S.A.M. A25617 (PF 10477).
Locality
Near Mossel Bay: PF 10477.
Description
There are 4 colonies the largest of which is 3:5 cm. by 2:0 cm. in diameter
and 1-0 cm. thick. The colonies are attached to pieces of sand-encrusted tubes
2mm.
|
Fic. 32. Metandrocarpa fascicularis sp. n.
A, colony; B, zooid, with test removed, seen from the left; C, oral tentacles; D, dorsal
tubercle; E, a testis; F, larva.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 185
probably the tubes of polychaete worms. The zooids of a colony are closely
crowded together, giving the appearance of a bunch of grapes (fig. 32 A).
Large zooids (fig. 32 B) reach a length of 4 or 5 mm. The body of the zooid is
pear-shaped, with the lower narrow end produced into a short slender stalk
which unites the zooid with the basal creeping stolon of the colony. In some
zooids the body is almost sessile on the stolon and in others the stalk is well
developed. The oral and atrial openings are small and inconspicuous, situated
quite close together on the upper end of the body. In the preserved state the
specimens are pale brown, and have no encrusting material, except on the
stolon.
The test is thin but semi-opaque. The body wall is dull red, but sufficiently
thin and transparent to allow the internal organs to show through. Muscles
are poorly developed. The oral and atrial siphons are very short and end in
simple round openings. There are about 30 oral tentacles arranged in two circles,
the outer one consisting of about 16 large tentacles and the inner one of smaller
tentacles (fig. 32 C). The dorsal tubercle is small and has a narrow straight slit-
like opening which is obliquely placed (fig. 32 D).
In the branchial sac of large zooids there are 9 longitudinal bars on each
side and smaller zooids have 7 or 8. ‘There are no folds. The dorsal lamina is a
narrow plain-edged membrane. The oesophagus is narrow. The stomach is of
a long barrel-shape, with about 16 longitudinal folds and a narrow, hooked
pyloric caecum. The intestine forms a flat bend parallel to the stomach, and
the rectum ends in a smooth-edged anus.
The female gonads are confined to the anterior part of the body and the
male gonads to the posterior part. On each side of the endostyle there are 2 to
5 large spherical ovaries of a dark brown colour. These seem to produce only
one mature egg at a time, since ovaries are often seen with a single developing
embryo. Embryonic development takes place within the ovary or oviduct
instead of freely in the atrial cavity.
The testes are 10 to 20 small pear-shaped glands on the left and right sides
of the posterior part of the body, each testis with a very slender sperm duct
(figs92) 1):
Larva
Zooids were never found with more than two larvae. The larva (fig. 32 F)
is reddish brown and has a trunk 0-7 to 0-8 mm. long and a tail of about 2:0 mm.
There are 3 anterior papillae arranged in a triangle, and a circle of about 24
narrow ampullae round the trunk. From each of these ampullae a line of dark
pigment extends back along the trunk. The single black sensory spot (“photo-
lith’) can be seen only after dehydration and clearing.
Remarks
No species of Metandrocarpa has yet been described from the waters of
South Africa or the western Indian Ocean.
186 ANNALS OF THE SOUTH AFRICAN MUSEUM
There are other specimens in the collection which I cannot definitely
identify, but which may belong to this species (e.g. PF 2348).
Genus POLYANDROCARPA Michaelsen, 19044
Polyandrocarpa anguinea (Sluiter)
Styela (Polycarpa) anguinea Sluiter, 1808, p. 52.
Known distribution
Knysna and False Bay, Cape Province.
Localities
Mossel Bay: U.C.T., MB 18 T; U.C.T., MB 22 G; U.C.T., MB 53 R;
U.G.T., MB 55 U; U.C.T., MB 60 S.
Algoa Bay: U.C.T., LIZ 28 Q.
Morrumbene Estuary: U.C.T., MOR 108 N; U.C.T., MOR 132 H.
Remarks
This species has hitherto been placed in the genus Polycarpa and was
considered as a simple form which grows in close aggregations of individuals.
A study of the more numerous specimens now available in the present collection
makes it clear that the species is a colonial styelid, to be placed in the genus
Polyandrocarpa. It is very similar to P. lapidosa (Herdman) from south-eastern
Australia, but appears to be specifically distinct, and I shall compare the two
species in another publication.
Subfamily Styelinae Herdman, 1881
Genus POLYCARPA Heller, 1877
Polycarpa rubida (Sluiter)
(Fig. 33)
Styela (Polycarpa) rubida Sluiter, 1898, p. 53.
Polycarpa rubida (Sluiter). Hartmeyer 1909, p. 1364. Michaelsen, 1918, p. 37.
Known distribution
Mozambique.
Locality
Morrumbene Estuary: U.C.T., MOR 132 1.
Description f
The single specimen is 2:3 cm. long and 1-4 cm. wide. It had been basally
attached, and is upright with the oral opening terminal and the atrial opening
about one-third of the body length from it. Both openings are almost sessile,
the siphons being very short. The test is grey with a pink-brown tinge, and has
irregular furrows and mounds, but is not further divided into small fields as in
the type specimen described by Sluiter (1898). Small rose-coloured marks on
the surface of the test may indicate the colour in life; Sluiter’s specimen was
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS I 87
evidently red and the preserving alcohol had extracted this colour. The test is
moderately thick and is tough. Within, it is pearly white. The body wall is dull
red-grey, the red colour being most marked round the oral and atrial openings.
Circular muscles form a continuous sheet and below these is a layer of longi-
tudinal muscles, the whole set of muscles rendering the body wall thick and
opaque.
There are 24 oral tentacles of alternating lengths. The dorsal tubercle is
a fairly prominent pad with an angular U-shaped opening facing obliquely
forward and to the left (Fig. 33A). Four well-
developed folds are present on each side of
the branchial sac, with the following arrange-
ment of longitudinal bars:
dorsal line 2 (15) 2 (10) 2 (10) 1 (9)
1 endostyle.
The dorsal lamina is a plain membrane.
Most of the gut is situated in the posterior
part of the body (fig. 33 B). ‘The oesophagus
is curved and the stomach barrel-shaped with
about 20 indistinct longitudinal or slightly
oblique folds and a hooked pyloric caecum.
The intestine and rectum form an S-shaped
bend and the anus has about 15 shallow lobes
In Sluiter’s specimen the anus was some dis-
tance behind the atrial opening, but in the Fic. 33. Polycarpa rubida (Sluiter).
present specimen it lies close to it. iy dorsal cubercles By eur:
The gonads are small and numerous, about 40 being present on the left
side of the body. They are of the type usual in the genus and are scattered over
the inner surface of the body wall. Between them are numerous transparent
flattened endocarps. A few endocarps are also present within the intestinal loop.
Remarks
This new specimen differs from the type specimen principally in having
fewer longitudinal bars between the branchial folds, a character which may be
related to the much smaller size of the specimen. The opening of the dorsal
tubercle is also different, but as Sluiter remarked, the S-shaped slit of the type
specimen is easily derived from a horse-shoe-shaped slit.
Genus CNEMIDOCARPA Huntsman, 1913
Cnemidocarpa asymmetra (Hartmeyer)
Tethyum (Styela) asymmetron Hartmeyer, 1912, pp. 253-257.
Styela asymmetra (Hartmeyer). Michaelsen, 1915, pp. 394-398.
Cnemidocarpa asymmetra Hartmeyer, 1926, pp. 180-183.
[non] Styela asymmetrica Sluiter, 1904, p. 87.
188 ANNALS OF THE SOUTH AFRICAN MUSEUM
Known distribution
Cape Town, Cape Province and Luderitz Bay, South West Africa.
Locality
Langebaan Lagoon: U.C.T., LB 378 Q.
Description
The larger of the 2 specimens in the collection is 5-2 cm. high and 3:0 cm.
wide, and the smaller specimen 4:0 cm. high and 3:0 cm. wide. The body is
erect with both siphons projecting from the upper end. The test is slightly
wrinkled, rather thin but tough, and semi-transparent. In the larger specimen
the opening of the tubercle is horse-shoe-shaped with both horns turned out-
wards, and in the smaller specimen similar but with only the right horn turned
outwards. The following is the arrangement of longitudinal bars on one side
of the smaller specimen:
dorsal lamina o (8) 3 (11) 3 (10) 2 (8) 2 endostyle.
There are 10-14 stigmata per mesh. The gonads form the most characteristic
feature of the species. In the present specimens the shape of the gonads does not
differ much from Hartmeyer’s (1912) original description. It is remarkable,
however, that in the new specimens the testis lobes occupy a central position,
being largely embedded within the ovary, whereas Hartmeyer (1912) gives the
opposite arrangement (‘die innere Partie wird vom Ovarium, die aussere von
den Hoden gebildet’). ‘The gonoducts are arranged in pairs and are distributed
over the inner surface of the gonads. The larger specimen from Langebaan has
a total of 7 of these pairs.
Cnemidocarpa psammophora sp. n.
(Fig. 34)
Diagnosis
Surface coated with sand or shell. Test with short hair-like processes over
most of the surface and one or a few long root-like basal processes. Many closely
spaced flat oral tentacles. Dorsal tubercle with a C-shaped or simple longitudinal
slit. Branchial sac with 4 rather low folds. Stomach with about 20 folds and a
short, curved pyloric caecum. Five to 8 gonads on the left side and 10 or 11
on the right.
Holotype
In the South African Museum. S.A.M. A25618 (U.C.T., LIZ 25 L).
Localities
Alsoa Bay: U-C.0* LIZ 19:2 90 Ci ei Zios ale:
Description
All three specimens in the collection have an almost globular body
(fig. 34 A, B). In two the diameter is 10 mm. and in the third the body is
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 189
15 mm. long and 12 mm. wide. These measurements were made over the
coating of sand grains and broken shell which completely covers the body.
Two simple openings fairly close together on the upper side mark the positions
of the oral and atrial siphons which do not project from the outline of the body.
Among the sand grains and broken shell are many short tag-like processes of
the test, which are generally distributed over the surface. At the base of the
5mm.
Fic. 34. Cnemidocarpa psammophora sp. n.
A, B, two intact specimens; C, specimen with test removed, seen from the right; D, E, dorsal
tubercles of two specimens; F, part of stomach and intestine, to show pyloric caecum;
G, gonads; ovd., oviduct; sp.d., sperm duct.
body a few of these tags have been greatly developed as stout rooting processes.
Generally about 6 of these are present, and one of them is longer and thicker
than the rest. In the largest individual the main process is 24 mm. long.
When the test is removed the body is seen to be almost globular or a little
elongated, with the siphons projecting as short conical tubes with shallow ter-
minal lobes (fig. 34 C). The body wall is semi-opaque although the muscles
190 ANNALS OF THE SOUTH AFRICAN MUSEUM
are rather poorly developed. On the inner surface of the body wall are quite
numerous small endocarps. The base of the oral siphon is provided with many
closely spaced slender and flattened tentacles. In two specimens these numbered
at least 60 and 80 respectively. ‘The opening of the dorsal tubercle in the largest
specimen is a simple narrow longitudinal slit (fig. 34 D), and in one of the
smaller specimens dissected a C-shaped slit facing forward, with slightly
inrolled horns (fig. 34 E). The dorsal lamina is a moderately wide membrane
with smooth margin. There are 4 rather low folds on each side of the branchial
sac with the following arrangement of longitudinal bars in the largest specimen:
dorsal lamina 1 (18) 2 (9) 3 (20) 3 (16) 2 endostyle.
The gut consists of the curved oesophagus, the barrel-shaped stomach with
about 20 folds and a short pyloric caecum (fig. 34 F), and the S-shaped intestine
and rectum. The border of the anus is cut into a large number of very small
lobes.
Gonads are present on each side of the body and number on the left from
5 to 8, and on the right 10 or 11. The gonads are tubular and nearly straight,
of the kind that characterizes the genus (fig. 34 G). Most of the gonad is occupied
by a long tubular ovary, and closely applied to each side of this is a series of
small pear-shaped testis follicles. The individual sperm ducts join the common
sperm duct which passes along the mesial surface of the gonad. Both the sperm
duct and the oviduct project slightly from the end of the gonad.
Remarks
There is no west African or South African species with which C. psammophora
is likely to be confused. The only species of the genus recorded from the western
Indian Ocean is C. madagascariensis Hartmeyer, a species incompletely described
as the only specimen was damaged. From what is known, however, it is evident
that several characters distinguish it from C. psammophora. C. madagascariensis has
a test divided into areas separated by furrows, and lacks the coating of sand
and the tags and rooting processes of C. psammophora. Hartmeyer’s species also
has many fewer oral tentacles and fewer gonads.
Several species of Cnemidocarpa are adapted to life on sandy bottoms.
C. psammophora is one of these and shows the adaptive features of this ecological
group, namely, small size, spherical form, and rooting processes of the test.
Genus sTYELA Fleming, 1822
Styela angularis (Stimpson)
Cynthia angularis Stimpson, 1855, p. 387.
Tethyum costatum Hartmeyer, 1911, p, 564.
Known distribution
South Africa.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS I9gi
Localities
False Bay: U.G.T., FAL 158 W; U.C.T., FAL 216 G; U.C.T., FAL 225 L;
Weir BAL 220M:
Mossel Bay: UW:C.T., MB 7 R; U.C.T:, MB 14 Y; U.C.T., MB 18 U;
UC. MBr438 By U-C:l) MB se7S; UC. 1, MB 55 Ve U.Grr:
MB 68 F; U.C.T., MB 72 U.
Cape Morgan: PF 13393; PF 13432; PF 13481.
Description
The characteristic shape of this species renders it unmistakable (see Hart-
meyer, 1911, Millar, 1955). In the present collections specimens range from a
very small size up to 9°5 cm. long.
Styela marquesana Michaelsen
Styela marquesana Michaelsen, 1918, p. 27.
Known distribution
Lourengo Marques, Mozambique and Durban, Natal.
Localities
Morrumbene Estuary: U.C.T., MOR 132 K.
Cape Natal: PF 10722.
Description
Michaelsen’s (1918) detailed description leaves little to be added. In the
present material specimens have from 22 to 26 folds on the stomach, and the
branchial bars vary between the following limits:
dorsal lamina 4 (10) 1 (10) 1 (10) 2 (6) 3 endostyle, and
dorsal lamina 10 (12) 13 (16) 15 (15) 10 (9) 3 endostyle.
Styela radicata sp. n.
(Fig. 35)
Diagnosis of species
Surface coated with sand or shell. Test with one or a few basal processes.
Dorsal tubercle with a simple transverse slit. Stomach with about 15 folds, and
a small pyloric caecum. One gonad on each side, with tubular ovary and lobed
testis follicles close to the sides of the ovary.
Holotype
In the South African Museum. 8.A.M. A25619 (U.C.T., MOR 132 J).
Locality
Morrumbene Estuary: U.C.T., MOR 1932 J.
Description
The single specimen (fig. 35 A) is almost spherical, but slightly narrowed
at the base, and measures 1-1 cm. in height and 1-0 cm. in width. It is com-
192 ANNALS OF THE SOUTH AFRICAN MUSEUM
pletely covered with sand grains and fragments of shell. The oral and atrial
openings are on the upper side and not very far apart. They scarcely project
from the surface. On the lower side there is a root-like process of the test about
4 mm. long, and a few much shorter tags. The test is thin, but very firm perhaps
owing to the closely adhering sand and shell.
Fic. 35. Styela radicata sp. n.
A, intact specimen; B, specimen with test removed, seen from the left; C, dorsal tubercle;
D, anus; E, gonad.
Muscles are not strong and consist of radial strands surrounding the two
siphons, and a series of circular muscles mainly round the siphons but also
spreading downwards over the upper half of the body (fig. 35 B). The body
wall is thin and delicate. The internal siphons are very short, and are not lobed.
There are 32 oral tentacles, of which about 16 are quite small. The dorsal
tubercle is small, with a simple transverse slit (fig. 35 C). The branchial sac
is delicate, with 4 well developed folds having the following arrangement of
longitudinal bars:
dorsal lamina 2 (16) 4 (7) 4 (11) 5 (7) 2 endostyle.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 193
The dorsal lamina is narrow, with a plain margin. Each mesh of the branchial
wall contains an average of 4 stigmata, which are long, narrow and regular.
The short curved oesophagus leads to the barrel-shaped stomach which
has about 15 undivided longitudinal folds and a very short rounded caecum.
The intestine and rectum make a simple S-shaped bend ending in a narrow
anus with minutely lobed margin (fig. 35 D).
One gonad is present on each side of the body. On the left side it is anterior
to the intestinal loop and directed obliquely back towards the atrial opening.
The right gonad occupies a corresponding position. Each gonad (fig. 35 E)
consists of an almost straight tubular ovary ending in a short simple oviduct,
and a series of 8 to 10 lobed testis follicles along the whole length of each side
of the ovary.
No endocarps are visible on the inner surface of the body wall.
Remarks
This species is quite unlike any other known from the waters of west,
South or east Africa. Its closest resemblance is to S. schmitti van Name from the
southern part of the east coast of South America. ‘The small size, the shape, the
coating of foreign particles, and the basal process are similar in the two species,
but §. radicata differs from S. schmitt2 in having only one gonad on each side
and in the structure of the branchial sac. The external similarities probably
indicate adaptive convergence rather than systematic affinity.
Family Pyuridae Hartmeyer, 1908
Genus PpyuRA Molina, 1782
Pyura stolonifera (Heller)
(Fig. 36)
Cynthia stolonifera Heller, 1878, p. 92.
Pyura stolonifera (Heller) Hartmeyer, 1911, p. 554.
Known distribution
South Africa;? east and south Australia.
Localities
Hascbay Cle Wh AlsrAg Ye Go. FAL 163 R; U.C:i., FAL 177.N;
U.C.T., FAL 221 8; PF 15984 A.
Mossel Bay: U.C.T., MB 18 V; U.C.T., MB 53 T; PF 11 B.
AlgoaiBay, ULGil Lie wx UE. l., LIZ 2) B; PE rog5 D:
East London: S.A. Museum (no details).
Description
P. stolonifera is a species which may have to be split, if a critical examina-
tion is made of enough material from different places. Most of the specimens
194 ANNALS OF THE SOUTH AFRICAN MUSEUM
listed above are undoubtedly of this species, and have the dorsal tubercle
basically in the form of a C opening posteriorly, but with the horns spirally
inrolled and contorted to an extent that increases with the size of the animal
(fig. 36 A-D). This is the arrangement found by Hartmeyer (1911) both in the
specimens taken by the German South-polar Expedition in Simon’s Bay, and
in Heller’s type specimens which Hartmeyer re-examined. Specimens from
A B | G
————EeEe
0.5 mm.
Fic. 36. Pyura stolonifera (Heller).
A-D, dorsal tubercles respectively of small (A, B), medium-sized (C)
and large (D) specimens; E, spicules.
Australia, however, which have been assigned to this species (Hartmeyer and
Michaelsen, 1928) have a dorsal tubercle facing forward. I believe that we may
be dealing with two species which are structurally similar except in some
characters like the dorsal tubercle. It would be surprising if P. stolonifera occurred
in Cape Province, South Africa, and on the south coast of Australia but not
between these places. Kott (1952), in dealing with the Australian records of the
species, admits that ‘the nomenclature of the group is still very confused’.
Some, but not all, of the specimens from False Bay and Mossel Bay have
:
:
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 195
spicules (fig. 36 E) in various body tissues, but not in the test. The spicules are
most abundant in one large specimen from Mossel Bay (MB 18 V), in which the
walls of the endostyle, the branchial folds, the oral tentacles and the dorsal
tubercle have many closely packed simple or branched spicules. Spicules of this
kind have been recorded in certain tissues of Cynthiopsis valdiviae Michaelsen
and spicules of a simple shape in C. herdmanni (v. Drasche), and both of these
species are regarded as representing P. stolonifera (Heller) (Hartmeyer, 1911).
Pyura capensis Hartmeyer
(Fig. 37)
Pyura capensis Hartmeyer, 1911, p. 561.
Known distribution
False Bay, Cape Province.
Fic. 37. Pyura capensis Hartmeyer.
A, intact specimen; B, oral tentacle; C, dorsal tubercles of two specimens;
D, gut; E, one of the gonadial sacs.
196 ANNALS OF THE SOUTH AFRICAN MUSEUM
Locality
Mossel Bay: U.C.T., MB 18 W.
Description
The largest of the 3 specimens is 2-5 cm. long and 1:6 cm. wide, and the
smallest is 1-4 cm. long and 1:3 cm. wide. In all 3 the body is upright and the
siphons fairly close together at the upper end. One of the characteristic features
of this species, which readily distinguishes it from P. stolonifera, is the subdivision
of the test into hexagonal or irregular plates, each plate having a red or brown
central mark (fig. 37 A). This feature was noted by Hartmeyer (1911) and is
distinct in the new specimens. The test is thin but tough.
The oral tentacles (fig. 37 B) number about 16 in the largest specimen.
They have slender primary branches with few and small secondary branches.
In the two specimens dissected the opening of the dorsal tubercle is a U-shaped
slit with a few short straight branches (fig. 37 C). The tubercle illustrated by
Hartmeyer (Taf. 57, fig. 8) is more complex but has the same essential features.
The 7 branchial folds have the following arrangement of longitudinal bars:
dorsal line 1 (10) 3 (10) 2 (15) 2 (14) 2 (16) 2 (12) 3 (9) 2 endostyle-.
A long row of dorsal languets is present.
The gut of the new specimens agrees closely with Hartmeyer’s description.
The margin of the anus, which Hartmeyer appears not to have seen clearly, is
smooth.
In most species of Pyura each gonad consists of a compact double row of
sacs with central common ducts, but in P. capensis the individual gonadial sacs
are numerous and scattered so that the gonads have a more diffuse appearance
than is usual in the genus.
Remarks
This is a distinctive species, and the present material, the first since the
type specimens were described, confirms the account given by Hartmeyer.
Genus microcosmus Heller, 1878
Microcosmus oligophyllus Heller
(Fig. 38)
Microcosmus oligophyllus Heller, 1878, p. 101.
Known distribution
Cape Province.
Localities
Mossel Bay: U.C.T., MB 22 H.
Saldanha Bay: Hoedjies Bay, low water, S.A. Museum.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 197
Description
This species has already been adequately described (Heller, 1878; Hart-
meyer, 1912; Millar, 1955), and only a few points about the new specimens
need be noted.
There are many specimens, 17 having been col-
lected at low water from Hoedjies Bay alone. The
dorsal tubercle shows some variation in the shape of
its opening. In the type specimen re-examined by
Hartmeyer (1912) the opening was U-shaped with
the right limb longer than the left and slightly inrolled. Fic. 38. Microcosmus
Another specimen from the same collection as the type eae
had the right limb more markedly inrolled and the oe
left one only slightly so. In the new material this difference in the limbs is
sometimes even more obvious, and in some specimens results in the two limbs
having well-formed spirals in opposite directions, giving the opening a modified
horizontal S-shape (fig. 38).
Microcosmus exasperatus Heller
Microcosmus exasferatus Heller, 1878, p. 99.
For synonymy see Van Name, 1945, pp. 346, 347.
Known distribution
Formosa; China Sea; Malay Archipelago; West Indies; Venezuela;
Colombia; Natal; Mozambique.
Localities
Natal: PF 11001.
Mozambique: U.C.T., PEA 1 B; U.C.T., PEA 4 Z; U.C.T., MOR 43 M;
U.C.T., MOR 108 O; U.C.T., MOR 109 R; U.C.T., MOR 122 H;
U.C.T., MOR 188 M.
? Mossel Bay: U.C.T., MB 22 I.
Remarks
This is a well-known species in many warm areas throughout the world.
It seems to be common in coastal waters of Mozambique. There is one doubtful
record from Mossel Bay; this specimen is covered with sand, and has only 7
folds on each branchial wail, but in other respects it agrees well with
M. exasperatus.
Maicrocosmus pedunculatus Pérés
(Fig. 39)
Maicrocosmus pedunculaius Pérés, 1951, p. 1062.
Known distribution
Senegal and Sierra Leone, west Africa.
198 ANNALS OF THE SOUTH AFRICAN MUSEUM
Localities
Mossel Bay: U.C.T., MB 72 V.
Morrumbene: Estuary: U.Q.T., MOR 91 R; U.C.T., MOR 132 L.
Description
A specimen of average size is 1-6 cm. long and 1-4 cm. across, slightly
compressed laterally and triangular in outline with the lower, narrow end
produced into one or several root-like processes (fig. 39 A). These processes
are whitish and free of sand but the rest of the body is completely coated with
sand grains. A narrow ridge which joins the siphons is continued down the
sides of the body towards the base.
The internal structure (fig. 39 B) is in complete agreement with the
descriptions already published of this species (Pérés, 1951; Millar, 1956).
Ds
8
CS a ah es
i :
4 4s oy paren SAE
pt) VN? 2.
- ba TS ee it o
. S x = ase PONS
- co Lhd
Fic. 39. Microcosmus pedunculatus Pérés.
A, intact specimen; B, specimen with test removed, seen from the left.
Remarks
This species apparently presents a remarkable case of discontinuous
distribution, since the only previous records are from tropical west Africa north
of the equator. The identification of the South African specimens, however, can
scarcely be doubted on anatomical grounds. This is either a case of structural
convergence of two species, or true discontinuous distribution, or continuous
distribution of M. pedunculatus masked by the accidental absence of specimens
from collections in intermediate areas.
Genus BOLTENIA Savigny, 1816
Boltenia africana sp. n.
(Fig. 40)
Diagnosis of species
Body erect, not stalked, completely covered with sand and shell. About
12 compound oral tentacles. Dorsal tubercle with a straight, C-shaped or
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 199
S-shaped slit. Branchial sac with transverse stigmata, and with 6 folds on each
side. Stomach with longitudinal folds, and a compact lobed mass of glandular
tubules. Intestine and rectum of uniform diameter. Anus with a plain margin.
One gonad on each side, the left one in the secondary intestinal loop.
Holotype
In the South African Museum. 8.A.M. A25620 (PF 15984 B).
2 cm.
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Fic. 40. Boltenia africana sp. n.
A, intact specimen; B, specimen with test removed, seen from the right; C, dorsal tubercles of
three specimens; D, part of branchial wall; E, gonad.
Localities
False Bay: U.C.T., FAL 212 T; PF 15984 B.
Mossel Bay: U.C.T., MB 55 Y.
Algoa Bay: U.C.T., LIZ 38 G.
Description
A number of the specimens collected from False Bay and from Mossel Bay
had been growing closely together and formed compact masses. Large indivi-
200 ANNALS OF THE SOUTH AFRICAN MUSEUM
duals reach 4 cm. in length and 3:5 cm. in breadth. The shape is variable,
being cylindrical, squat or irregular (fig. 40 A). The two siphons generally
project from the upper end but are sometimes scarcely visible. Sand, broken
shell and other debris completely cover the surface and no doubt render the
animals very inconspicuous in nature. his may explain why the species has
not previously been described. The test is only moderately thick, but is tough.
When removed from the test the body is seen to be red-brown. The siphons are
quite long, and conical. A set of strong muscles radiates from each siphon over
the sides of the body, and circular muscles pass round the siphons. The body
wall is rather thin, so that the gut and gonads are visible (fig. 40 B).
There are 11-14 oral tentacles, of alternating sizes. ‘The largest of these are
tri-pinnate. The dorsal tubercle varies a good deal, the opening being a simple,
almost straight longitudinal slit, or C-shaped, or in the form of a horizontal S
(fig. 40 CQ). On each side of the branchial sac there are 6 folds, of which the
ventral 2 or 3 are smaller than the more dorsal ones. A long row of dorsal
languets is present. In a specimen of moderate size, and in a large one, the
longitudinal bars of the branchial sac had respectively the following arrange-
ment:
dorsallme 4 (8). 1 (14) 3 (15) 3 (2)) 4 (6) 5 G6) S¥endoseyic
dorsal line 15 (15) 10 (33) 8 (26) 12 (23) 12 (20) 12 (5) 12 endostyle
The stigmata, as usual in the genus, are transversely placed in longitudinal
rows (fig. 40 D). A longitudinal bar passes across the centre of each row of
stigmata.
The oesophagus is narrow, curved, and rather short. The stomach is wide
at the oesophageal end and tapers towards its junction with the intestine, where
there is a conspicuous lobed mass of glandular tubules. Numerous longitudinal
folds are present on the walls of the stomach. The intestine forms a flat, hori-
zontal, almost closed loop and the rectum bends upwards and forwards towards
the base of the atrial siphon, where it ends in the plain-edged anus. There is
little variation in diameter along the whole length of the intestine and rectum.
One gonad is present on each side of the body, that of the left side lying
just above the dorsal limb of the intestine, that is in the secondary gut loop
(fig. 40 B). On the right side the gonad is in a corresponding position. Each
gonad is a long, rather bulky, and slightly curved body (fig. 40 E). The ovary
occupies the mesial side and the testis is confined to the parietal side, next to
the body wall. Indentations on the margins of the ovary give it a slightly lobed
appearance. The testis follicles are round bodies which form a white mass
visible from the outside, through the body wall. Both the sperm duct and the
oviduct are at the posterior end of the gonads, from which they project only
very slightly.
Remarks
No species of Boltenia, as characterized by the transverse arrangement of
stigmata, has been known hitherto from west, south, or east Africa.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 201
Genus HALOCYNTHIA Verrill, 1879
Halocynthia spinosa Sluiter f. defectiva n.
(Fig. 41)
Species Halocynthia spinosa Sluiter, 1905, p. 16.
Diagnosis of form
As the typical form of the species, but having 2 gonads on the left side and
none on the right.
Holotype
In the South African Museum. 8.A.M. A25621 (U.C.T., MB 55 W).
Known distribution
(Of species) Gulf of Aden; Red Sea.
Localities
alse bay; W.C.W., FAL 137 N.
Mossel Bay: U.C.T., MB 55 W; U.C.T., MB 68 G.
Description
The largest specimen in the collection is 4:5 cm. long and 5:0 cm. across.
The body is rounded with a narrow base, or roughly hemispherical and broadly
based (fig. 41 A). On the upper side the two siphons project prominently. In
colour the specimens vary from pale grey to brown and generally have a pink
tinge or quite bright red suffusion, especially on and near the siphons. Spines
are present on almost all parts of the test usually in groups of one long central
spine and several short encircling spines (fig. 41 B). The spines are larger on
the siphons than elsewhere, but not so large as those shown by Sluiter (1905)
in his figure of.the species. The test is fairly thick and is tough. Stout longitudinal
and circular muscles render the body wall thick and opaque.
The oral tentacles number 12 to 14 in these specimens, and are tri-pinnate.
In form the dorsal tubercle is constant, having a C-shaped slit facing forward
and with the horns spirally inrolled (fig. 41 D). The dorsal languets form a long
series of slender projections from the roof of the branchial sac. Nine or 10 tall
narrow folds are present on each side of the branchial sac, with up to 26 longi-
tudinal bars on the largest folds.
The gut forms a wide loop of almost uniform diameter, except for the
stomach which is short and wide with longitudinally pleated walls and a lobed
mass of glandular tubules (fig. 41 E).
The gonads constitute the main peculiarity of these specimens. There are
two on the left side (fig. 41 C) and none on the right. The gonads are large, and,
instead of being parallel are curved so that they meet and fuse at their dorsal
and ventral ends, forming what appears to be one ring-shaped body with two
pairs of gonoducts.
202 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
In the type material Sluiter (1905) described the gonads as a lobed mass
on each side, but Michaelsen (1919) noted, as an abnormality, the complete
absence of gonads on the right side. It is interesting that Van Name (1921)
found a similar condition in a specimen of H. pyriformis (Rathke) from the
Bahamas, although that species normally has 4—7 gonads on each side. There
Fic. 41. Halocynthia spinosa Sluiter f. defectiva n.
A, intact specimen; B, spines of test; C, specimen with test and right body wall removed, seen
from the right; D, dorsal tubercle; E, gut.
seems to be a tendency to eliminate the gonads of the right side in Halocynthia,
This tendency has apparently become fixed in the population of H. spinosa
inhabiting the waters of the Cape Province, and it is perhaps best to recognize
the situation by describing that population as a forma of the species. Neverthe-
less, if further collections from intervening waters of the western Indian Ocean
make it clear that there is discontinuous distribution of the species, it may be
necessary to recognize f. defectiva as a new species.
6A
ee
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 203
Family Molgulidae Lacaze-Duthiers, 1877
Genus MOLGULA Forbes and Hanley, 1848
Molgula falsensis Millar
Molgula falsensis Millar, 1955, p. 217.
Known distribution
False Bay, Cape Province.
Localities
alse Bay 0.C.0., FAL) 136 R; U.C.T., FAL 158 Q; U.C.T., FB 1105.
Description
A few specimens of this species are present in the collections. They all
confirm the characters which I have described in the type material (Millar,
1955). In some of the new specimens, however, the oviduct although turned up
at right angles to the gonad, is quite short and in others it is not bent up from
the gonad. In the type specimens I could not see the sperm duct, but in the
new specimens it is situated about half-way along the gonad and is a distinct,
free, finger-like projection.
The test of some of the new specimens is thicker and much firmer than in
the original specimens and approaches in appearance and consistency the
condition found in many species of Ascidia. Short test projections or hairs are,
however, present.
Molgula conchata Sluiter
Molgula conchata, Sluiter, 1808, p. 58.
Ctenicella conchata (Sluiter). Hartmeyer, 1913, p. 128.
Known distribution
Knysna, Cape Province.
Localities
Mossel Bay: U.C.T., MB 7S; U.C.T., MB 39 M; U.C.T., MB 58 M;
U.C.T., MB 60 T; U.C.T., MB 72 W.
Alison bays (O.C.m., LIZ 11 Zi.
Description
Many specimens are contained in the collection, including large ones over
4 cm. long. All of them show the characters which readily distinguish this from
other South African species of Molgula, namely: the coating of sand, slight
lateral compression of the body, horizontal S-shaped slit of the dorsal tubercle,
_and the form of the gonad and its position relative to the intestinal loop.
Remarks
The present records extend the known range of this species westwards to
Mossel Bay and eastwards to Algoa Bay. The geographical distribution never-
204 ANNALS OF THE SOUTH AFRICAN MUSEUM
theless seems to be very restricted, and M. conchata is replaced by allied species
in neighbouring areas.
Molgula scutata Millar
Molgula scutata Millar, 1955, p. 215.
Known distribution
Langebaan, Cape Province.
Localities
Saldanha Bay: U.C.T., SB 1.
PAlgoa Bay: U.C.T., LIZ 28 U.
Description
The dorsal tubercle of this specimen has an oblique and almost straight
slit; the previous examples had a C-shaped or an S-shaped slit.
One specimen from Algoa Bay, which I have provisionally included in
this species, is similar in all respects to typical specimens except that each gonad
is bent downwards at its dorsal end. This may be an individual peculiarity.
Remarks
This, the second record of the species, scarcely extends its known range.
Molgula cryptica sp. n.
(Fig. 42)
Diagnosis of species
Surface covered with sand. About 18 oral tentacles. Dorsal tubercle with
C-shaped opening usually facing to the right. Seven branchial folds, each with
3 longitudinal bars. Primary intestinal loop closed; secondary loop semi-
circular. Left gonad in secondary loop. Each gonad consisting of a sausage-
shaped ovary and a testis of radially disposed follicles surrounding the blind
end of the ovary.
Holotype
In the South African Museum. 8.A.M. A25622 (U.C.T., FAL 158 R).
Locality
False Bay: U.C.T., FAL 158 R.
Description
The body is generally ovoid, with the short siphons close together at the
upper end. Most of the surface is covered with a thin layer of adhering sand
grains. The largest specimens in the collection are about 3 cm. long, including
the siphons. The test is rather thin and, but for the coating of sand, would be
transparent. |
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 205
When the test is removed the body is seen to be elliptical or almost round
in outline, and the tubular siphons close together (fig. 42 A, B). Each of the 6
oral lobes and the 4 atrial lobes has a series of narrow projecting teeth forming
a fringe round the openings. The body wall in the preserved state is pale brown
but is sufficiently transparent to allow the gut and gonads to be seen. Muscles
are mainly confined to the siphons, where circular and longitudinal ones exist,
and to the dorsal half of the body which has one set of longitudinal muscles
radiating from the base of each siphon.
The oral tentacles generally number about 18 and are of alternating sizes.
They are tri-pinnate, but are not particularly bulky (fig. 42 C). The dorsal
tubercle is small, with a C-shaped or V-shaped opening. The C-shaped slit is
more common and usually faces to the right (fig. 42 D).
Ossie
Fic. 42. Molgula cryptica sp. n.
A, B, specimens with test removed, seen from the right and left respectively; C, oral
tentacle; D, dorsal tubercle; E, branchial fold in transverse section to show the three
longitudinal bars (in black); F, gonad.
206 ANNALS OF THE SOUTH AFRICAN MUSEUM
Seven low folds are present on each branchial wall, with a constant arrange-
ment of 3 longitudinal bars on the folds; this appears to be characteristic of the
species. On the ventral face of each fold near the base, there is a single bar, and
two bars are present at the summit of the fold (fig. 42 E). The dorsal lamina is
quite long and wide with a plain margin which is sometimes inrolled. The
stigmata are subdivided into oval or long narrow straight slits which, as they
tend to be arranged in regular transverse rows, obscure the spiral pattern.
The oesophagus is short, and it narrows towards its junction with the
stomach. A large mass of tubular glandular tissue surrounds and hides the
stomach. The intestine and rectum are narrow throughout their length. The
primary loop is quite closed and the secondary loop forms a semicircle (fig.
42 B). No incisions are present on the margin of the anus.
On the left side the gonad lies in the secondary intestinal loop, which it
largely fills, and on the right side it is immediately dorsal to the renal sac
(fig. 42 A, B). The ovary is sausage-shaped or somewhat pear-shaped, with a
rather short narrow oviduct directed towards the atrial siphon. Round the
ventral, blind end of the ovary the many small testis follicles are radially
arranged to form a compact crescentic or rosette-like testis (fig. 42 F). The
common sperm duct is wide and extends about half-way along the inner face
of the ovary to end in a remarkably large opening. The opening of the sperm
duct is, in fact, wider than that of the oviduct.
The renal sac is a large curved body in the lower half of the right side.
Larva
Larvae are present in the atrial cavity of some specimens. The larvae have
a trunk about 0-2 mm. long and a tail of about 0-7 mm. There is a single black
sensory pigment spot.
Remarks
M. cryptica is most like M. pulchra, which has been recorded from sub-
Antarctic South America and adjacent parts of the Antarctic. M. pulchra,
however, has a large number of oral tentacles, and the sperm duct has a
characteristic convoluted shape. It seems likely that these two species have
diverged from a common ancestral stock.
I have brought together, in fig. 43, those characters of gut and gonad
which best distinguish the four species of Molgula found in the present South
African collections.
Genus EUGYRA Alder and Hancock, 1870
Eugyra myodes sp. n.
(Fig. 44)
Diagnosis of species
Test provided with fine processes, and covered with sand grains. Body wall
thin and transparent. Muscles consisting mainly of (1) short transverse bands
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 207
D
Fic. 43. Comparison of gut loop and gonads of A, Molgula falsensis ; B, Molgula scutata;
C, Molgula cryptica; and D, Molgula conchata.
©). nak B
G
Fic. 44. Eugyra myodes sp. n.
A, B, specimen with test removed seen
respectively from the right; and left C, oral
tentacle; D, dorsal tubercle; E, gonad.
208 ANNALS OF THE SOUTH AFRICAN MUSEUM
in a line between the siphons and extending down the anterior and posterior
margins of the body, and (2) a V-shaped series of bands on each side of the
body. About 8 tri-pinnate oral tentacles. Dorsal tubercle with an almost straight
oblique slit. Branchial sac with 7 longitudinal bars and regular double spiral
stigmata. Primary intestinal loop open at the bend. Right gonad above renal
sac. Left gonad in secondary intestinal loop. Gonad consisting of a central sac-
like ovary surrounded by lobed testis follicles.
Holotype
In the South African Museum. 8.A.M. A25623 (U.C.T., LIZ 3 Q).
Locality
Algoa Bay: U.C.T., LIZ 3 Q.
Description
The larger of the two specimens is about 1:5 cm. in greatest diameter, and
the smaller specimen about 1-1 cm. In outline both are almost circular, and at
least in the preserved state the body is laterally compressed to some extent.
The siphons are not visible externally and the whole body is covered with
sand grains. Numerous fine hair-like processes project from the surface of
the test. The test, when freed of adhering sand grains, is thin, flexible and
transparent.
The internal siphons are separated by a distance equal to about half of the
body diameter (fig. 44 A, B). They are short and their margins bear narrow
teeth, which on the oral siphon number about 24 and on the atrial siphon
about 20. Circular muscles surround each siphon but do not extend beyond
their bases. The remaining muscles of the body wall consist of 2 sets. The first
set is of short transverse fibres arranged in a line which runs between the siphons
and extends in the median plane down the anterior and posterior sides of the
body (fig. 44 A, B). The second set is a V-shaped line, on each side of the body,
of longer fibres. The body wall is very thin and transparent.
There are 8 or 9 tri-pinnate oral tentacles of alternating sizes (fig. 44 CQ).
The dorsal tubercle is small, with a straight or very slightly curved slit (fig. 44
D). Behind the dorsal tubercle and to the right of the ganglion lies the small
branched neural gland.
The branchial sac is delicate. There is quite a long dorsal lamina which
increases in width towards the posterior end, and has the margin rolled in.
Seven longitudinal bars, only slightly curved, are present on each side, and
below each bar a row of stigmata in double spirals. Each component slit of the
double spiral makes about 3 coils.
The oesophagus is curved. The stomach is not large and has on its right
wall a glandular diverticulum with longitudinally folded walls. The primary
loop of the intestine is open in its anterior part but closed in the middle region.
The rectum bends up and forward and ends near the base of the atrial siphon
in the plain-edged anus (fig. 44 B).
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 209
The left gonad lies in the secondary intestinal loop and the right one above
the renal sac. In each the ovary is sac-like with a short terminal oviduct. Round
the whole margin of the ovary and applied to its mesial face is a series of about
20 lobed testis follicles (fig. 44 E). The ducts of these lead to a main duct which
passes along the face of the ovary, to open by a few short slender sperm ducts
projecting into the atrial cavity.
On the posterior part of the right side is the small straight renal sac.
Remarks
Few species of Eugyra have been recorded from west, South or east African
waters, and of those few none is likely to be confused with the present species.
From Walvis Bay, South West Africa, Michaelsen (1914, 1915) has described
E. woermanni which, having a gonad only on the right side, properly belongs to
the sub-genus Gamaster, and is very distinct from E. myodes. No species of Eugyra
is known from Cape Province or the African waters of the eastern Indian
Ocean. FE. kerguelenensis Herdman is known from Kerguelen, the coastal waters
of Argentina, and also the Patagonian Shelf and South Georgia (Millar, 1960).
That species bears the closest resemblance to E. myodes. It also has bands of
short muscles on the body wall but these are differently arranged. The position
of left gonad mainly within the primary intestinal loop also distinguishes E.
short muscles on the body wall but these are differently arranged. The position of
the left gonad mainly within the primary intestinal loop also distinguishes £.
kerguelenensis, as does the C-shaped opening of the dorsal tubercle. Nevertheless
the similarities are enough to suggest a close relationship between the two species.
Eugyra macrentera sp. n.
(Fig. 45)
Diagnosis of species
Test with hair-like processes, and covered with sand grains. Muscles
confined to the siphons, which have circular and radial strands. Eight bipinnate
oral tentacles. Dorsal tubercle with a C-shaped opening. Branchial sac with 7
longitudinal bars. Stigmata in uninterrupted double spirals. Gut long, with the
primary loop almost completely closed, and the secondary loop also very
narrow. Left gonad in secondary intestinal loop. Right gonad oblique. Renal
sac small.
Holotype
In the South African Museum. 8.A.M. A25624 (U.C.T., LIZ 11 Za).
Locality
Alcoa pays WG... IZ 11 2.2.
Description
There is only one specimen and this measures 1-2 cm. by 0-6 cm. The
body is oval and is covered with a rather thick layer of sand grains. It appears
210 ANNALS OF THE SOUTH AFRICAN MUSEUM
to have been attached to the frond of an alga, part of which is still adhering to
the specimen. The test is thin and its surface has numerous fine hair-like
processes.
The body wall, in spite of some brown pigmentation, is sufficiently trans-
parent to allow the gut and gonads to be seen when the test is removed. Narrow,
closely spaced circular muscles surround the siphons and stouter radial muscles
spread out for a short distance from their bases (fig. 45 A, B). The remainder
of the body wall, however, lacks muscles. Narrow teeth form a fringe round the
margins of both siphons.
(ONS) inn
Fic. 45. Eugyra macrentera sp. n.
A, B, specimen with test removed, seen respectively from the left and right; C, oral
tentacle; D, dorsal tubercle.
Eight bipinnate tentacles are situated at the base of the oral siphon (fig.
45 C). The dorsal tubercle is small, with a C-shaped opening facing towards the
right (fig. 45 D). The dorsal lamina is rather narrow but long, and has a plain
margin. Seven tall longitudinal bars are situated on each side of the branchial
sac. They are less curved than in many species of Eugyra. Below each of these
bars is a row of infundibula, each with a pair of spiral stigmata. There are, in
addition, some smaller accessory infundibula in the spaces between the main
ones. Narrow transverse bars separate adjacent transverse rows of infundibula.
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 2II
The oesophagus is short, narrow and curved, and the small stomach bears
a lobed glandular diverticulum. The intestine is long and folded into a flat
double loop, making an almost closed primary loop and a narrow secondary
loop. The rectum bends forward and upward, to end in a plain anus near the
base of the atrial siphon.
In this specimen the gonads are not well developed, and seem to be in a
post-spawning state. They are long and rather narrow, consisting of a central
tubular ovary fringed by testis lobes. No further details of the gonads could be
seen. The left gonad is in the secondary intestinal loop and the right gonad lies
obliquely across the centre of the right side, pointing towards the atrial siphon.
The renal sac is a small semi-circular body about the middle of the right
side.
Remarks
E. macrentera is remarkable for the length of the intestine and the way in
which it is folded. A few species of Molgula, notably M. platei Hartmeyer and
M. contorta Sluiter, have a somewhat similar intestinal loop, an arrangement
which allows a long intestine to be accommodated in a small space.
ANNALS OF THE SOUTH AFRICAN MUSEUM
22
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FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS
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FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 24107)
DISTRIBUTION OF SPECIES
Geographical components of the ascidian fauna
Stephenson (1944) has analysed the littoral fauna and flora of the South
African coast between Port Nolloth on the west and Durban on the east, and
recognizes the following components:
(1) the warm-water component,
(2) the cold-water component,
(3) the ubiquitous component,
(4) the south coast component,
(5) the more local components.
The ascidians described in the present paper (Table 2) and in a previous
one (Millar, 1955) can be grouped into the first four components defined by
Stephenson. Excluding doubtful records and insufficiently known species, the
ascidians can be divided as follows:
The warm water component:
Aplidium sarasinorum
Polyclinum isipingense
P. arenosum
P. constellatum
Pseudodistoma africanum
Cystodites dellechiajer
Eudistoma coeruleum
E.. modestum
E. reniert
Ascidia pygmaea
Botrylloides nigrum var. giganteum
Symplegma viride
Polyandrocarpa anguinea
Polycarpa rubida
Styela plicata
S. aequatorialis
S. marquesana
Microcosmus exasperatus
M. pedunculatus
The cold water component:
Aplidium flavo-lineatum
A. pantherinum
Polyclinum neptunium
Cystodites roseolus
Trididemnum cerebriforme
Leptoclinides capensis
218 ANNALS OF THE SOUTH AFRICAN MUSEUM
Corella eumyota
Botryllus magnicoecus
Cnemidocarpa asymmetra
Microcosmus oligophyllus
Molgula scutata
The ubiquitous component:
Diplosoma listerianum
Ciona intestinalis
The south coast component:
Aplidium circulatum
A. retiforme
Clavelina roseola
Sycozoa arborescens
Distaplia capensis
Ascidia sydneiensis
Agnesia glaciata
Gynandrocarpa unilateralis
Dextrocarpa solitaris
Alloeocarpa capensis
Styela angularis
S. pupa
Pyura capensis
P. stolonifera
Boltenia africana
Molgula falsensis
M. conchata
Table 3 shows the number of species in each of the four main components,
as a percentage of the total number of ascidian species collected, and also shows
the comparable percentages obtained from Stephenson’s (1944) figures of all
species of littoral animals.
TABLE 3
Percentage of total species
Ascidians All littoral animals
Warm water component 39 52
Cold water component 22 18
Ubiquitous component 4 8
South Coast component 35 13
FURTHER DESCRIPTIONS OF SOUTH AFRICAN ASCIDIANS 219
It should be pointed out that the figures are not strictly comparable
because the ascidians include both littoral and sublittoral material.
The ascidian fauna compared with the littoral animals as a whole, shows
a warm water component almost as important, and a relatively more important
south coast component. The cold water component and the ubiquitous com-
ponent are about the same relative size in ascidians as in the total of littoral
animal species in South Africa. There is therefore quite good agreement
between ascidians and other groups.
The distribution of South African species in other regions shows some
interesting features. Particularly notable is the high proportion of the ascidian
fauna which appears to be endemic. Care must be used in comparing the
results in the present paper with those of other authors dealing with different
areas, owing to the persisting difficulties of ascidian systematics, and the con-
sequent possibility of confusion of species. Nevertheless it appears that more
than half of the species described in the present account are endemic. As
pointed out by Ekman (1953), however, the faunas of the neighbouring regions,
South West and south-east Africa, are very imperfectly known; it is therefore
likely that the list of endemic species will shorten as knowledge of these areas
increases.
The wider affinities of the South African ascidian fauna appear to be with
the adjacent areas of the warmer Indian Ocean. Only Apflidium retiforme,
Corella eumyota, and Agnesia glaciata are found also in any of the cooler waters of
more southerly regions. In this respect the South African ascidian fauna con-
trasts markedly with that of southern South America where many species are
shared with the Subantarctic and Antarctic regions (Millar, 1960). The tem-
perature regimes of southern South America and southern South Africa are, of
course, very different, but it seems likely that the greater geographical isolation
of the latter may also have been important in preventing fusion of the faunas of
South Africa and more southerly areas.
In other respects also the ascidians of South Africa and South America are
dissimilar. Several South American ascidians reach a very great size, as for
instance Distaplia cylindrica (Lesson), and the molgulids Ascopera gigantea Herd-
man and Paramolgula gregaria (Lesson). No species of a comparable size is known
from South Africa, except Szgillina vasta sp. n., which is known only from the
incomplete type specimen. Differences in the abundance of phytoplankton may
be responsible for the different sizes of the ascidians in the two areas.
SUMMARY
1. A systematic account is given of the ascidians from South African waters
contained in several collections.
2. Sixty-four species are identified, and five are given provisional identifi-
cations. Of the total, seventeen are described as new species, and one as a new
form of a known species.
3. The distribution of the species is briefly discussed.
220 ANNALS OF THE SOUTH AFRICAN MUSEUM
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Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
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Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuka, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
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Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.—All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name... .
B. Form of bibliographic references to synonymic names.—The first reference following
any name in the synonymy should be to the earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XLVI
PART VIII
DACTYLISPA CAPICOLA (PERINGUEY) UND VERWANDTE
(203. Beitrag zur Kenntnis der Hispinae (Coleoptera, Chrysomelidae))
Von
EricH UHMANN
Stollberg-Erzgebirge, Lessingstr. 15. Deutschland, DDR
Ss. GIESRAS
een
“ee oe ~
ISSUED FEBRUARY 1962 PRICE 17c.
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TRUSTEES OF THE SOUTH AFRICAN MUSEUM
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[CONTINUED ON INSIDE BACK COVER
DACTYLISPA CAPICOLA (PERINGUEY) UND VERWANDTE
(203. Beitrag zur Kenntnis der Hispinae (Coleoptera, Chrysomelidae))
Von
ErIcH UHMANN
Stollberg-Erzgebirge, Lessingstr. 15. Deutschland, DDR
[Angenommen Juli 1961]
(Mit 2 Abbildungen)
Dactylispa capicola (Pér.) 1898
In dem zahlreichen siidafrikanischen Dactylispa-Material, das mir zur
Bearbeitung vorgelegt worden ist, fanden sich immer wieder Stiicke, die zur
Beschreibung der D. capicola passten, aber immer wieder waren die Stiicke
untereinander verschieden. Durch das freundliche Entgegenkommen der
Herren Dr. Hesse und Dr. Andreae vom South African Museum erhielt ich
jetzt das gesamte capicola-Material des Museums zur Begutachtung. Fast alle
10 Stiick weichen mehr oder weniger voneinander ab. Herr Kollege Andreae
war so liebenswiirdig, mir ausftihrlich tiber das Material zu schreiben.
Zunachst hier die Urbeschreibung Péringuey’s (1898, p. 121).
,,dispa capicola n. sp.
Linear; black, opaque, nearly glabrous; antennae moderately slender,
basal joint very thick, the five following narrow, third one a little longer than
the fourth, the five ultimate ones moderately thickened and pubescent; discoidal
raised parts of the thorax smooth and very distinct; elytra deeply foveate, and
with three dorsal series of short spines on each side, and a marginal one, the
spines of which are not longer than the dorsal ones; the first interval along the
suture has a slightly raised, somewhat spinose line, the spines of which are more
developed in the posterior part. Length 44-54 mm.; width 2-2} mm. Hab. Cape
Colony (Cape Town, Stellenbosch, Worcester).”’
Im Schliissel (Péringuey, 1898, p. 119) steht die Art neben D. gracilis (Pér.),
sie wird von ihr getrennt:
,¢2. Black, opaque, nearly glabrous; elytra costulate . . . capicola
cl. Black, opaque, pubescent; elytra not costulate . . . gracilis’.
Wie aus Gréssen- und Patriaangabe hervorgeht, haben Péringuey mehrere
Stiicke vorgelegen, seine Beschreibung wird sich also nicht nur auf ezn Stiick
beziehen. Im Material ist nur eines vorhanden, von Cape Town, von Péringuey
223
Ann. S. Afr. Mus. 46 (8), 1962, 223-230, 2 figs.
224 ANNALS OF THE SOUTH AFRICAN MUSEUM
selbst mit capicola bezettelt. Dieses Stiick hat als Typus, Lectotypus, zu gelten.
Leider fehlen ihm die Fihler. Von den beiden anderen Fundorten ist kein
Stiick vorhanden, wohl aber 1 Stiick von Durban, Natal (Py typ.), 2 Stiicke
von Douglas, Cape Province (Py), 1 Stiick von Malvern, Natal (Barker) (Py),
1 Stiick von Natal (Barker), 1 Stiick von Durban, Natal (——), 2 Stiicke von
Eshowe, Zululand (——), 1 Stiick von George, Cape Province, 7:86 (—).
Py” bedeutet: Das Stiick wurde von Péringuey bezettelt und mit seinem
Signum versehen, ,,typ’’ bedeutet nicht ,,type’’, sondern ,,typical’’, denn der
Fundort ist nicht in der Beschreibung genannt. Sie wurden augenscheinlich
erst spater von ihm bestimmt.
Alle diese Stiicke und die ahnlichen meiner Sammlung sind vom Typus
mehr oder weniger verschieden. Am ahnlichsten ist der Typus der D. bodongi
Uhm. aus Beira, Mocambique, die ich (1930, p. 130) ausfithrlich beschrieben
habe. D. bodongi hat kurze, starke Dornen auf der Deckenscheibe, die dichten
Seitenranddornen sind kurz, gerade, die 4 Spitzenranddornen sind etwas
langer als die des Seitenrandes, aber immer noch kurz und kraftig. Decken
kurz aber deutlich behaart. In meiner deutschen Beschreibung der D. bodongi
ist zu berichtigen auf Zeile 3: ,,3. Fiihlerglied 14 mal so lang wie breit”, auf
Zeile 4: ,,so lang wie breit’’.
Der Typus von D. capicola hat dieselbe Anordnung der Deckenelemente,
die Spiesse der Dornen sind aber alle diinner und erscheinen dadurch langer,
die Seitenranddornen sind auch diinner, anscheinend etwas langer und nach
hinten gekriimmt, die 5 Spitzenranddornen sind auch diinner, deutlich langer
als die des Seitenrandes. Decken anscheinend kahl, die Behaarung k6nnte
aber abgerieben sein. In 1931 (32) spreche ich von einem typischen Stiick
der D. capicola aus dem Briisseler Museum, das nicht so deutlich behaart ist
wie D. bodongi. Die Dornen erscheinen dadurch etwas langer.
Dass dem Typus die Fiihler mit Ausnahme der beiderseitigen beiden
Basalglieder fehlen, ist zu bedauern. Es muss aber darauf hingewiesen werden,
dass die Lange der einzelnen Fihlerglieder bei vielen Dactylispa- (und tiberhaupt
Hispini-) Arten nicht unbedingt als Artmerkmal gelten kann.
Bei 50 X sieht man, dass der Rest der Behaarung, besonders an der
Deckenspitze, aus kurzen, niederliegenden Harchen besteht, nicht aus ziemlich
langen, aufrechten Haaren wie bei D. bodongz.
BESCHREIBUNG DES LECTOTYPUS
Langlich, parallel, schwarz, Decken dunkelblau-metallisch. Stirn nieder-
gedriickt, runzlig, mit Mittelfurche, Hals matt, schagriniert. Kopfschild breit,
glanzend, sehr fein niederliegend behaart, vor den Fihlern spitz. Halsschild
so lang wie breit, Bewehrung kraftig, Vorderdornen weit gespreizt, mit sehr
schragem Vorderast und fast senkrechtem, kiirzerem, gebogenem Hinterast,
Seitendornen fast 3, 0, der vorderste am langsten, schrag nach vorn gerichtet,
der mittlere fast gerade abstehend, der dritte gekriimmt, erst nach hinten,
hy
»
Scie ee
DACTYLISPA CAPICOLA (PERINGUEY) UND VERWANDTE 225
dann nach aussen gerichtet, so lang wie der mittlere. Beide Halsschildflachen
durch eine tiefe Mittellinie getrennt, seitlich steil abfallend, glatt, schwach
gewolbt, schwach quergerunzelt, vorn neben der Mittellinie schmal, aber
ziemlich tief ausgerandet (Abb. 1). Antebasalrand gut entwickelt, scharf.
Die vertieften Stellen der Scheibe mit deutlich behaarten Kérnchen. Vorder-
ecken vollig verrundet, mit je einem deutlichen Borstenzylinder, Hinterecken
seitlich vorgezogen. Decken mit parallelen Seiten, die 10 Punktreihen durch die
Basen der Dornen gest6rt, 9. und 10. in der Mitte vereinigt, ,, Punkte’’ viereckig
(Griibchenpunkte). Zwischenraume schmal, unregelmdssig. Dornen der
Decken ziemlich kurz, mit starker Basis und schlankem Spiess, die Spiesse
aN
ABB. I
Abbildung der Halsschildflachen der D. capicola (Pér.).
Die Abbildung zeigt die kleinen Ausschnitte vorn an den Flachen. Wenn diese grésser werden,
k6énnen die schmalen Leisten neben der Mittelrinne abgetrennt werden. Die Furche wird viel
feiner, und es entsteht eine vertiefte Mittelflache wie bei gracilis.
vorm Hinterrande kraftig. Naht mit Schliessddrnchen, erst auf dem Abfall
mit einigen Dérnchen. I. Zwischenraum mit einer Reihe von Dérnchen (von
Péringuey erwahnt); II. Zwischenraum mit II 1-5, mit II 1b, II 5 stark;
IV. Zwischenraum mit IV 1b, IV 1, 2, beide etwas hinter II 2 und II 3,
mit IV 3-IV 5, diese Dornen schrag vor den entsprechenden von II;
VI. Zwischenraum mit 4 Schulterdornen, VI 2 hinter IV 1, VI 3 schrag
hinter IV 2, VI 4 am Ende des kaum angedeuteten Eindruckes, neben IV 4;
VIII. Zwischenraum mit einer Reihe van Dérnchen, davon 2 in der Schulter-
kehle, VIII 5 nicht betont. Zusatzbedornung gering, ausser den erwahnten auf I
und im Eindruck, einige auf VIII und vor der Deckenspitze. Behaarung
siehe oben. Fiir ein gutes Unterscheidungsmerkmal halte ich die Bedornung
des Seitenrandes im Verhaltnis zum Spitzenrande. Die Dichte der Bedornung
(ideale Bedornung, siehe unten) bleibt sich gleich, Seitenranddornen nur
etwas langer als die der Scheibe, nach hinten gekriimmt, am vdéllig verrundeten
Hinterrand kiirzer, am Spitzenrande deutlich langer als an der Seite. Spitzen-
randdornen etwa 4, sie gehen allmahlich in die Seitendornen wber. Mittel-
226 ANNALS OF THE SOUTH AFRICAN MUSEUM
schenkel deutlich bed6rnelt, Mittelschienen schwach gekriimmt. Rand der
Epipleuren hinten gezahnelt. Lange fast 6 mm. (Kopf geneigt). Schulterbreite
fast 2 mm.
Dactylispa subcapicola n. sp.
Differt a D. capicola (Pér.) elytris evidenter pilosis, spinis lateralibus
elytrorum longioribus, fere rectis (spinae apicales spinas laterales parum
superantes), femoribus intermediis exilissime granulatis.
Zum Holotypus wahle ich ein Stiick, das von Péringuey selbst bezettelt
worden ist: ,,Hispa capicola Pér. typ’’. Fundortszettel: Durban, Natal. C. N.
Barker, 1898. Da Péringuey in seiner Beschreibung Natal nicht als Patria
aufgefiihrt hat, kann das Stiick nicht als ,,Cotype’’ (jetzt ,,Paratypoid’’)
aufgefasst werden, sondern ,,typ’’ muss heissen ,,typical’’, wie ich schon oben
ausgefiihrt habe. |
Der Holotypus und seine Paratypoide sind dem Typus von D. capicola
sehr a4hnlich, unterscheiden sich von ihm vor allem durch deutliche Behaarung
auf den Decken und die Bedornung der Deckenrander: Dornen des Seitenrandes
langer, so lang wie die der Scheibe, fast gerade; Dornen des Spitzenrandes nur
wenig langer als die des Seitenrandes, nicht so auffallend langer wie bei D.
capicola.
Holotypus. Gestalt, Farbe und Glanz wie bei D. capicola. Stirn eingedriickt,
langsrunzlig, hinten mit einer kleinen Grube. Augenrander mit diinnem
Harchensaum. Fiihler kurz, gerade bis zur Schulter reichend, Schmal- und
Breitseite nicht ausgepragt, Glied 7-11 schwach verdickt, Glied 1 dick, schrag
abgeschnitten, unten vorgezogen; Glied 2 rundlich, viel kiirzer und diinner
als Glied 1; Glied 3 abgerundet-zylindrisch, 14 mal so lang wie 2; 4-6 einander
fast gleich, jedes kiirzer als 3; 6 kiirzer als 5; 7 zylindrisch, so lang wie 3;
8-10 einander gleich, zylindrisch, fast quer; 11 um die Spitze langer als Io.
Glied 2-6 fein behaart, die folgenden tomentiert. Halsschild nur wenig
breiter als lang. Bewehrung kraftig, Vorderdornen im rechten Winkel gespreizt,
Vorderast nur wenig langer als der hintere. Seitendornen 2, 1, der freie Dorn
etwas langer als die beiden mit gemeinsamer Basis, alle an der Spitze etwas
zuruckgekriimmt. Scheibe mit 2 glatten, scharf abgesetzten Flachen, die durch
eine feine Mittelfurche getrennt sind. Vorn sind beide neben der Mittelfurche
schmal ausgeschnitten, doch so, dass eine feine Leiste sie von der Mittelfurche
trennt wie bei D. capicola. Antebasalrand scharf. Die vertieften Stellen mit
deutlich behaarten Kérnchen. Vorderecken vollig verrundet, mit deutlichen
Borstenzylindern. Hinterecken seitlich vorgezogen. Decken mit parallelen
Seiten, die 10 Reihen durch die Basen der Dornen gestért, 9. und 10. in der
Mitte vereinigt. Die Punktgriibchen unregelmassig viereckig. Dornen der
Decken etwas langer und mit kraftigeren Spiessen als bei D. capicola. Spiesse
der Dornen auf dem Abfall ziemlich lang. Bedornung wie bei D. capicola,
aber die Deutung der Dornen durch viele den Hauptdornen fast gleiche
Zusatzdornen unsicher. Behaarung fein, deutlich, aufrecht. Deckenrander
a
DACTYLISPA CAPICOLA (PERINGUEY) UND VERWANDTE 227
dicht bedornt, zu jedem Querleistchen zwischen den einzelnen Punkten gehort
ein Randdorn (ideale Bedornung). Weiteres siehe bei D. capicola. Rand der
Epipleuren fein bedérnelt. Mittelschenkel fein bedérnelt (50 x), die anderen
Schenkel nicht nennenswert skulptiert. Mittelschienen schwach gebogen,
einfach. Lange 5.5 mm., Breite 24 mm.
MATERIAL
Holotypus, siehe oben.
1. Paratypoid. Durban, Natal (C. N. Barker), capicola, Uhmann det.
2. Paratypoid. Durban, Natal (Barker), Hispa capicola Pér.
3. Paratypoid. Durban, Natal (Barker 1898), capicola, Andreae det.
4., 5. Paratypoid. Douglas, Cape Province, Dactylispa ?capicola Pér.
In coll. Uhmann:
6. Paratypoid. Malvern, Natal (Barker), Dactylispa ?capicola Per.
Einschaltung der D. subcapicola n. sp. in meinen ,,Schliissel der Dactylispa-
Arten Afrikas .. .”’ (1955, p. 160):
8 (11) Seitenranddornen kurz.
g (10) Deckendornen schlank, so lang oder langer als die sehr kurzen
des Seitenrandes. Halsschild nicht breiter als lang.
ga (gb) Oberseite deutlich aufstehend behaart. Bewehrung kurz. Flachen
des Halsschildes vorn ziemlich breit ausgeschnitten, dort mit
schmaler, vertiefter Mittelflache (Abb. 1: Legende). Decken
mit vielen Zusatzdérnchen. In allen Dingen zierlich.—typ!—
4°5 mm. Natal, Sambesia, Zululand (neu) .. . gracilis Péringuey
gb (ga) Oberseite auf den Decken kaum sichtbar behaart. Bewehrung
kraftiger. Flachen des Halsschildes vorn schm4ler ausgeschnitten,
ohne schmale, vertiefte Mittelflache (Abb. 1). Die scharfe
Mittelfurche bis tiber den Ausschnitt verlangert. Decken mit
weniger Zusatzdornchen. In allen Dingen kraftig. Der D. bodongi
ahnlich.—typ!—6 mm. Cape Town.......... capicola Péringuey
11 (8) Seitenranddornen langer.
12 (13) 3. Fihlerglied kurz, nur wenig langer als die Nachbarglieder.
Seitenranddornen etwa halb so lang wie eine Decke breit (in
der Quermitte). Seitenbewehrung halb so lang wie der Halsschild
breit. Seitenflachen gross, breit, vorn ausgeschnitten, mit
durchlaufender Mittelfurche wie bei capicola. Decken deutlich
aufstehend behaart, mit vielen Zusatzdérnchen.—5.5 mm.
Durban, Natal; Douglas, Cape Province . . . subcapicola n. sp.
228 ANNALS OF THE SOUTH AFRICAN MUSEUM
d
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NV Yo eevee Ph ev vies
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—-*-+2=<5
ABB. 2
Oberseite der Puppenhaut von Dactylispa callosa Uhm
Halsschild. Die 5 Erweiterungen des Halsschildes sind hier nach oben gebogen. Er ist in der
Vorderhalfte in der Mitte aufgerissen.
Mes = Mesothorax; Met = Metathorax,
Pi— Prothorax:
DACTYLISPA CAPICOLA (PERINGUEY) UND VERWANDTE 229
Dactylispa callosa Uhm. 1935, 1948
Neanotypus (Puppen-Exuvie). Pearly Beach near Quoin Point, District
Bredasdorp, Aug. 1958. Zwischen Grasstengeln. (Abb. 2.)
Puppenhaut dunkelbraun, ob durch dussere Einfliisse? 4:25 mm.
Unterseite. Fithlerscheide schwach segmentiert. Deckenscheide mit
angedeuteten Rippen, ohne Zahnchen oder Borsten am Rande. Knie ohne
Borsten. Bauch sehr fein schagriniert, scheinbar fein punktiert. An Stelle der
Querlinien von Kérnchen oder Héckerchen finden sich auf dem 4. bis 7. Sternit
(1. bis 3. nicht sichtbar) Querleisten, die wohl aus der Verschmelzung der Basen
von Erhabenheiten entstanden sind. Die Leiste des 4. Sternites lasst noch
einzelne Hockerchen erkennen, an den anderen Sterniten sind die Hockerchen
mehr oder weniger erloschen. Vor dem eigentlichen Hinterrande mit starker,
nach hinten steiler Querfalte, auf der beiderseits jeder Querleiste 2 sehr kleine
Zahnchen stehen.
Oberseite. Kopf durch das Schliipfen zerstort. Halsschild quer, beiderseits
am Rande mit 5 Zapfen. Der zweite von vorn nur kurz, die anderen fast
gleich; der 1. in den Vorder-, der 5. in den Hinterecken. Scheibe durch das
Schlipfen vielleicht verandert: mit stark vertiefter Mittellinie, beiderseits mit
einer erhabenen Beule, die durch eine vertiefte Furche rings von der Scheibe
abgesetzt ist.
Mesonotum quer, doppelt so breit wie lang, mit S-formig geschwungenen
Seiten und vorgezogener, abgestumpfter Spitze, beiderseits etwas eingedriickt.
Metanotum glatt, beiderseits zu einer schwachen Langsfurche vertieft.
Seiten des 1. Tergites einfach, die des 2. bis 4. Tergites etwas lappenartig
erweitert und dort mit je einem kurzen, feinen Zylinder. Endborsten sind auch
bei 100facher Vergrésserung nicht zu erkennen gewesen. In 1958 (p. 215) sprach
ich die Vermutung aus, dass die Anhange der Segmente 2-5 vielleicht immer
in der Dreizahl! auftreten wiirden. Das kann ich an der vorliegenden Puppen-
haut wegen der dunklen Farbung und den mir zu Gebote stehenden Mitteln
nicht einwandfrei nachweisen. Tergit 5 hat keinen Zylinder beiderseits. Alle
vorhandenen Zylinder sind sehr fein und diinn, man kann sie wohl als
,,ninfallig’’ bezeichnen. Sonst lasst sich an den Tergiten 2-5 je ein sehr kurzes,
feines Bérstchen nachweisen, das man wohl als 2. Anhang bezeichnen kann.
Der 3. Anhang, wenn vorhanden, miisste ausserordentlich klein sein. Tergit 6
hat einen einfachen Rand, Tergit 7 besitzt am Rande einen grossen,
geschwungen-zugespitzten Zahn, der aber anscheinend ein Produkt der Seiten
der Scheibe ist. Tergit 8 + 9 tragt 4 gleiche dreieckige Zahne in gleichem
Abstande. Auf der linken Seite steht zwischen den Zahnen 1 und 2 ein kurzer
Zylinder, der rechts fehlt. Diese 4 Zahne sind etwas nach unten gebogen.
Stigmen auf sehr kurzen Zylindern, die des 5. Tergites zahnartig spitz
ausgezogen.
Tergit 2 bis 6 mit zunehmend starker Querfalte (auf 4 und 6 ist der Abfall
nach hinten recht steil). Jede biegt sich vor dem Seitenrand riickwarts um.
230 ANNALS OF THE SOUTH AFRICAN MUSEUM
Auf Tergit 2 bis 4 liegt das Stigma vor der Falte, auf 5 sitzt es auf ihr. Auf 6
befindet sich auf dem riickw4rts gerichteten Aste der Falte ein kleines, spitzes
Zahnchen auf verdickter Basis. Tergit 7 mit 2 einfachen Querfalten, Tergit
8 + 9 uneben.
Auf Tergit 2 bis 6 liegen die oft beobachteten beiden Querreihen von
kleinen Erhabenheiten, aber die Reihen sind nicht so regelmassig und deutlich
ausgepragt und voneinander geschieden. Sie werden gebildet durch sehr viele
winzige Erhabenheiten, die ein helles Schiippchen tragen. Auf dem 6. Tergit
scheint die dem Hinterrande nachste Reihe aus kleinen, durchsichtigen,
stumpfen Zahnchen zu bestehen. Die Hinterreihen sind breit. Tergit 7 ohne
die kleinen Rauhigkeiten, dafiir sind die Querlinien deutlich.
LITERATURVERZEICHNIS
PERINGUEY, L. 1898. Catalogue of the South African Hispinae (Coleoptera), with descriptions
of new species. Ann. S. Afr. Mus. 1, 113-1930.
Unmann, E. 1930. Hispinen des Deutschen Entomologischen Institutes in Berlin-Dahlem.
II. Teil. Ent. Bl. 26, 125-136.
Unmann, E. 1931(32). Katalog der afrikanischen Hispinen des Berliner Zoologischen Museums
Mitt. zool. Mus. Berl. 17, 836-8091.
Unumann, E. 1935. Afrikanische Hispinen aus dem Naturhistorischen Reichsmuseum in
Stockholm. Ent. Tidskr. 56, 121-126.
Uumann, E. 1948. Die Holotypen einiger meiner Hispinae-Arten im Naturhistoriska Riks-
museum in Stockholm. Ark. Zool. 41 B, no. 3, 1-2.
Unmann, E. 1955. Schliissel der Dactylispa-Arten Afrikas ohne Madagascar. Rev. Zool. Bot.
afr. 51, 147-180.
Unmann, E. 1958. Hispinae aus Siidafrika. III. Teil. 7. ent. Soc. S. Afr. 21, 214-226.
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Bettrdge zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Hartman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-500.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. and ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.—All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name... .
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should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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ANNALS
POUT HvArFRICAN, MUSEUM
ON THE DENTITION AND TOOTH REPLACEMENT
IN TWO BAURIAMORPH REPTILES
By
A. W. CromMPTon
South African Museum, Cape Town
be LIBR At 2
ae aeeipeieetatien
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[ CONTINUED ON INSIDE BACK COVER
ON THE DENTITION AND TOOTH REPLACEMENT
IN TWO BAURIAMORPH REPTILES
By
A. W. CROMPTON
South African Museum, Cape Town
[Accepted August, 1961]
(With 3 plates and 1o figures in the text)
CONTENTS
MM ErOCUCHORM tate feat PA Mace ode HE et ee) por
Dentition and tooth replacement in Ericiolacerta parva Watson . 234
Dentition and tooth replacement in Bauria cynops Broom . . 241
Discussion . ee od ek ie Stok eae ae . 250
Summary . Alb ar sl Siete Ns a ue as rar MCS.
References : : Ns hye Bi saz ee ee seis e254
ieyato) the) lettermeyol the figures, 5. ees ls 255
INTRODUCTION
The bauriamorphs are an advanced group of mammal-like reptiles which
have independently developed many mammalian features otherwise found only
in the cynodonts and mammals. Broom (1911) suggested that because of their
less specialized nature the bauriamorphs are probably more closely allied to
the group from which mammals arose than the cynodonts are. He suggested
(1932) that the ictidosaurs (including Diarthrognathus) probably arose from a
small bauriamorph that lived during Cynognathus zone times. The structure of
the skull of Diarthrognathus (Crompton, 1958) confirms this view.
Very little detailed information on the dentition of bauriamorphs is
available and the purpose of the present paper is to record information on the
dentitions of two, Ericiolacerta and Bauria.
Recently a detailed study of the dentition of Dzarthrognathus, has been
completed by the author; this will be published in the near future. The present
study will therefore enable the possible relationships of this advanced mammal-
like reptile to be discussed with greater confidence.
Information reported here on Ericiolacerta parva is based on the type and
only known specimen of this animal and is in addition to that published by
Watson (1931) in his excellent description of this animal. As many teeth as
possible were exposed and the left mandible was separated from the skull.
The present description of the dentition of Bauria cynops is based upon the
type skull (S.A.M. 1333) described by Broom (1909). The exposed surface of
the skull had been damaged but fortunately little attempt was made to prepare
231
Ann. S. Afr. Mus. 46(9), 1962, 231-255, 3 pls., 10 figs.
232 ANNALS OF THE SOUTH AFRICAN MUSEUM
individual teeth and consequently much useful information could still be
obtained. In order that the teeth could be studied in detail the lower jaw was
freed from the skull. Unfortunately the matrix does not respond to acetic acid
and it was therefore necessary to prepare the teeth mechanically.
Short descriptions of the dentition of Bauria have been given by Broom
(1909, 1932, 1937), Boonstra (1938) and Brink (1953); these are based upon
three skulls and an isolated mandible. Considerable diversity of opinion exists
on several important aspects of the dentition. Little detailed information is
available on the dentitions of the other bauriamorphs Bauroides, Melinodon,
Sesamodon and Watsoniella. Seeley (1895) has given a description of the post-
canines of Microgomphodon and Mucrohelodon. Unfortunately it is difficult to
follow, and no illustrations accompany the description.
The importance of Ericiolacerta is that it is the earliest theriodont that is
known in which the apices of the crowns of marginal teeth occlude and were
worn to horizontal or spherical surfaces. In the scaloposaurids and early
cynodonts the postcanines are longitudinally ovate and sheared past one another
to achieve a scissor-like cutting action. In the Cynognathus zone bauriamorphs,
the gomphodont cynodonts, and their descendants the tritylodontids, the cheek
teeth are transversely widened and occluding crowns met one another.
Ericiolacerta is therefore an ideal animal to study the shift from a shearing to a
crushing or grinding tooth action. A similar shift took place in the early
mammals (Patterson, 1956), and an attempt will be made in this paper to
compare the jaw action of early mammals with those of some mammal-like
reptiles in order to determine whether or not grinding was achieved in the
theriodonts with transversely ovate postcanine teeth.
Tooth replacement in the mammal-like reptiles and the origin of the
mammalian type of tooth replacement has not received a great deal of
attention. Watson (1931) described as unique among mammal-like reptiles
the replacement of a maxillary tooth in Ericiolacerta vertically from above. He
concluded from this study that all the functional teeth belonged to one set and
that the replacing tooth observed indicated that tooth change affecting allthe
teeth was about to set in and that it probably only occurred once. This would
mean that in Ericiolacerta a mammalian type of replacement had almost been
achieved. Boonstra (1938) showed that in both the maxilla and dentary of
Bauria replacing teeth lay lingual to the functional postcanines, but unfortunately
he gave no detailed description. Parrington (1936) showed that in Dimetrodon,
Thrinaxodon and Tribolodon replacement was alternate and that it was possible
to divide the postcanine teeth into two series that are situated alternately in
the jaw. He expressed the view that the number of replacements of each series
was reduced in Thrinaxodon. He accepted as possible Bolk’s theory for the origin
of the mammalian type of replacement, viz. that as the teeth crowns became
more complex there was no space for teeth of one series to erupt between
those of the other and that instead one series replaced the other to form the
deciduous and permanent dentitions of mammals.
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 233
Romer and Price (1940) confirmed that replacement was alternate in
Dimetrodon, but that in addition a wave of replacement passed along each
alternately numbered series. These waves passed from back to front.
Tooth replacement in gomphodont cynodonts has been described and
discussed by Broom (1913), Brink (1955, 1956) and Crompton (1955b, 1958).
It is at present also being studied by Mr. S. Fourie. Brink is of the opinion that
although forms such as Diademodon from the Lower Trias do not have a dental
succession identical to that of placental mammals, the incisors and anterior four
postcanines were replaced only once, the 12th to 16th teeth never, and the 5th
to 11th certainly not more than once. The canine, Brink feels, was probably
replaced more than once. This would suggest that gomphodont cynodonts were
tending towards a mammalian dental succession. Unfortunately Brink has
not described the material upon which these conclusions are based.
Gomphodont cynodonts from the Middle Trias, such as Scalenodon, do
not confirm this view (Crompton, 1955b, 1958). Here there appears to have
been frequent replacement of the incisors and canines, whereas in the postcanine
series replacement is limited and is recorded only from the posterior end of the
dentition where sectorial teeth were replaced by transversely ovate teeth. With
growth, teeth are added to the postcanine row posteriorly, while teeth are lost
at the front of the postcanine row. Recent unpublished work on Dzademodon
by Mr. Fourie confirms that postcanine teeth are lost anteriorly and added
posteriorly. Similar observations have been made on Oligokyphus (Kiihne, 1956).
The dental succession of gomphodont cynodonts therefore appears to be
fundamentally different from that of placental mammals, and the origin of
the mammalian pattern must be searched for in other mammal-like reptile
groups.
Kermack (1956) has reported on tooth replacement in the Therocephalia
and the Gorgonopsia. He claims that alternate replacement (his ‘functional
distichical replacement’) has been lost in the postcanine series in these families.
His material was insufficient to enable him to determine the order of replacement
except in the case of the canines.
Recently Edmund (1960) has surveyed in great detail tooth replacement
in the lower vertebrates. Although mammal-like reptiles are not treated above
the pelycosaur level, Edmund’s contribution is a most valuable basis for the
study of tooth replacement in mammal-like reptiles. In order to explain
mammalian tooth replacement, which is so different from the alternating
replacement typical of most reptiles, Edmund has suggested that the deciduous
and permanent dentitions represent individual zahnrethen.
It is hoped that a description given in this paper of the conditions in
Ericiolacerta and Bauria will contribute towards the solution of the problem
of the origin of the mammalian dental succession.
I am deeply indebted to Professor D. M. S. Watson for his very kind
permission to undertake additional preparation on the type of Ericiolacerta parva,
and to Dr. K. A. Kermack and Mrs. F. Mussett for their valuable assistance
234 ANNALS OF THE SOUTH AFRICAN MUSEUM
to me during my stay in London. I wish also to record my thanks to Dr. F. R.
Parrington for his helpful criticism and encouragement.
DENTITION AND TooTH REPLACEMENT IN ERICIOLACERTA
It is difficult to determine the number of teeth present in the type because
the tip of the snout is slightly damaged. Watson (1931) claimed 6 teeth in the
premaxilla and 8 in the maxilla and that no canine could be distinguished.
Although a total of 14 teeth is probably correct it is difficult to be certain of the
exact number in the maxilla and premaxilla.
Cheek teeth in both mandible and maxilla are preserved in various
stages of the replacement cycle. These can be conveniently divided into four
stages (fig. 1). In stage I a freshly erupted tooth was loosely held in an enlarged
1 T IV
long. gr.
Fic. 1.—Ericiolacerta parva. Four stages in the replacement cycle of cheek teeth.
(Key to lettering, p. 255.)
alveolus. In a later stage (II) attachment bone (at. bo.) was built up around the
neck of the tooth above the longitudinal groove (long. gr.) which extended the
length of the tooth row on the lingual side in the maxilla and dentary. Next,
replacement teeth developed in pits in the base of the longitudinal groove
(stage III), and as the replacing tooth increased in size, it migrated labially
and resorbed first the alveolar wall separating it from the alveolus and second
the root of the functional tooth (stage IV).
‘For description purposes the teeth are numbered consecutively from the
front because no distinct canine is present.
Lower dentition (figs. 2 A, B, 4 C)
The 15th tooth from the front has a well-preserved unworn tricuspid
crown. The central cusp is slightly higher than the accessory cusps. The outer
surface of the tooth is convex, but due to a slight swelling on the inner surface
of the crown a short distance below the apex of the main cusp the upper
portion of the inner surface of the upper half of the crown is slightly concave
(figs. 2 B, 14). The posterior lower teeth of Ericiolacerta behind the 8th all have
this characteristic shape. The tooth is firmly held and the longitudinal groove
does not extend as far posteriorly as this point. The 14th tooth is less firmly held.
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 235
The crown is unworn and in contrast to the 15th is terminated by four small
cusps. The second cusp from the anterior border is the largest. Their apices
do not lie in the same plane but are arranged to form an arc. A wide gap
separates the 12th and 14th teeth. The functional tooth appears to have been
lost, although this could not be confirmed beyond doubt. A pit in the base of the
longitudinal groove containing a replacing tooth (fig. 2 A, rep. t. 13) is
w.fac.h.
B .fac.o.
14 lO
Fic. 2.—Ericiolacerta parva. A, medial view of posterior portion of the left upper and lower
dentitions, and B, posterior view of 14th and 1oth teeth. Both x to. (Key to lettering, p. 225.)
present lingual to this gap. It has been interpreted as a replacing tooth for the
13th.
The 12th tooth is partially erupted (stage I). The accessory cusps have
been lost as a result of damage but there is no sign of wear on the remaining
central cusp. The 11th is fairly tightly held (stage II) and the longitudinal
groove is present lingual to its base. An oblique wear facet sloping inwards
and downwards has obliterated all trace of the anterior accessory cusps. The
10th is tightly held and a large pit is present lingual to it in the base of the
longitudinal groove (stage III). The matrix was not completely removed from
this pit and it presumably contained a replacing tooth in life. The crown is
considerably worn and two distinct facets can be identified (fig. 2 B 10): a
236 ANNALS OF THE SOUTH AFRICAN MUSEUM
horizontal facet (w. fac. h.) on the labial side, and an oblique facet (w. fac. o.)
on the lingual. The portion of the crown supporting cusps was completely worn
away. The goth tooth is loosely held by its alveolus (stage I). It shows no signs
of wear and is terminated by three small cusps. The upper portion of the
crown is slightly narrower antero-posteriorly than the lower, and the tooth
therefore has a dumb-bell appearance. The 8th tooth is tightly held. A small
pit is present in the longitudinal groove lingual to its base (stage III). The
crown is worn to two distinct facets, as is the roth.
It is difficult to interpret the 7th tooth. It is very small; the crown is
extensively worn and tightly held by attachment bone. The crown surface is
considerably lower than adjacent teeth. No pit is visible at the base of the
longitudinal groove. The 7th tooth in the upper jaw is also small and has been
considered by Watson (1931) as the first maxillary tooth. No. 6 in the lower jaw
is loosely held by its alveolus, indicating that fossilization took place soon
after it had erupted (stage I). It is sharply differentiated from the unworn
teeth in the posterior portion of the jaw in that the crown is terminated by a
single sharp cusp. In view of this it is possible to interpret it as a canine and the
7th as the first postcanine. The crown of the 5th tooth also consists of a single
cusp, but it is more tightly held by attachment bone than the 6th. A fairly wide
gap separates the 5th and 6th teeth.
The anterior mandibular teeth are only well preserved on the right,
where six alveoli, some with and some without teeth, could be counted. The
6th tooth is well worn. The 5th is unworn and, in contrast to conditions on the
left, is terminated by 3 cusps as in the left 9th and 15th teeth. The crown of the
4th tooth is extensively worn to two wear facets as in the left 1oth. The 3rd
alveolus is empty, the 2nd tooth is not well preserved, and the Ist appears to
be a replacing tooth.
Upper dentition (Plate XII A: figs. 2 A, 3 A, B, C, 4 A, B)
The teeth of the maxillae are shown in lateral view in Plate XII A, medial
view in figs. 2 A and 3 A, and in crown view in fig. 3 B. They vary greatly in
size, and as in the lower jaw large spaces separate the individual teeth.
The maxilla medial and posterior to the 14th tooth is damaged, but it
appears that this tooth was tightly held and that a large pit (fig. 3 B p.) was
present lingual to it (stage III). A single flat wear-facet dipping slightly upwards
in a lingual direction is present. The 13th is preserved as it was after it erupted
and is loosely held in a large alveolus that extends lingually to the longitudinal
groove (stage I). The apex of the crown has unfortunately been damaged and
lost, but no signs of wear could be detected on the remaining portion. The
crown of the 12th tooth is well worn to a single facet (fig. 3 C 12). The tip
of the crown of a replacing tooth is present in a pit lingual to its base (stage ITT).
The 11th tooth and its replacing tooth have been described by Watson (1931).
This is the only position where stage IV of the replacement cycle is preserved,
viz. in which a replacement tooth is preserved shortly before it was to have
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 237
erupted. The lingual portion of root of the functional tooth (rem. f. 11) has
been resorbed and the crown lost. The crown of the replacing tooth (rep. t. 11)
fits closely against the remnant of the root of the functional tooth. The pit
containing the replacing tooth is extremely large and confluent with the
alveolus. The crown of the replacing tooth is terminated by a single cusp. The
tooth as seen in posterior view is illustrated in figure 3 C, 11. A small replacing
tooth (rep. t. 10) is present medial to the roth tooth (stage III). The crown
rem.f. 1.
Fic. 3.—Ericiolacerta parva. A, medial view of upper right dentition; B, crown view of upper
left dentition; C, posterior view of 13th, 11th and 8th, roth, 12th teeth and D, occlusion of
upper and lower teeth. All x 10. (Key to lettering, p. 255.)
238 ANNALS OF THE SOUTH AFRICAN MUSEUM
is extensively worn to a nearly horizontal wear-facet (fig. 3 C 10). The 9th tooth
is preserved as it was shortly after it erupted. The apex of the crown has been
damaged but no signs of wear could be detected on the remaining portion.
The 8th tooth is much smaller than the oth, is tightly held and extensively
worn, but the matrix medial to it and the three anterior teeth could not be
removed without difficulty to determine details of replacement. The 7th tooth
(Watson’s first maxillary tooth) is smaller than the 8th and has a slightly
worn crown. It is separated by a large gap from the 8th. It could not be
ascertained whether a tooth has been lost from this gap. No. 6 is sharply
pointed and shows no signs of wear whereas the apex of No. 5 is worn to a
horizontal facet (Plate XII). No. 6 could possibly be interpreted as a small
canine, and this would confirm the findings in the lower jaws.
The crowns of the teeth in the right maxilla (fig. 3 A) could only be seen in
lingual view because the right cheek is covered by an articulated hand. Nos. 14,
12 and 8 are preserved as they were shortly after they erupted and no signs of
wear could be detected in these teeth (stage I). The 9th and 11th teeth have
small pits for replacing teeth at their bases, and the crown of the oth is worn
to an oblique wear-facet. The 13th is preserved with the crown of a well-
developed replacing tooth situated in a pit medial to it (stage III). The 1oth
is very tightly held by attachment bone and has a well-worn crown. The
replacement tooth lingual to it is the largest in the right maxilla and at death
was presumably about to erupt.
Tooth replacement and tooth wear in ERICIOLACERTA
The teeth are preserved in different stages of replacement, and it is clear
that the functional teeth do not belong to one series all of which were about to
be replaced by a second set, as suggested by Watson (1931). It is also clear from
the above description that replacing teeth developed initially in shallow pits
at the base of the longitudinal groove, i.e. lingual to the functional teeth. The
longitudinal groove probably supported the dental lamina. As the tooth germs
increased in size, a pit developed at the base of the groove to support them. As
the tooth enlarged it migrated labially, absorbing first the lingual alveolar
wall and later the root of the functional tooth. Only in its final stages was
replacement vertically from above or below in the upper and lower jaw
respectively.
It remains now to determine the order of replacement. In fig. 4 the various
stages of replacement in the left and right maxillae and the left mandible are
shown diagrammatically. In the maxilla the order of replacement was clearly
alternate. In the left maxilla the odd-numbered teeth, 9 and 13, have unworn
crowns and are preserved as they were shortly after they erupted. Replacement
of the 11th lagged slightly behind the other odd-numbered teeth, but its replace-
ment tooth is large and in life would have been the next tooth to erupt. On the
other hand the even-numbered teeth 14, 12, 10 and 8 all have worn crowns, are
firmly held by their alveoli, and have small pits for replacing teeth lingual to
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 239
them. In the right maxilla the even-numbered series consisting of the 8th,
12th and 14th teeth are preserved recently after they erupted. Eruption of the
roth replacement tooth would have lagged slightly behind these teeth as in
the case of the 11th in the left maxilla. It therefore appears that on the left
the odd teeth were erupting whereas on the right the even teeth were erupting.
This may indicate that the teeth have been incorrectly numbered, but not
necessarily because a similar phenomenon was encountered by the author in
early cynodonts. A small difference in the degree of development of the
Fic. 4.—Ericiolacerta parva. Diagrammatic representation of the order of replace-
ment of the posterior teeth. A, left maxilla; B, right maxilla and C, left dentary.
replacing teeth is also preserved in the odd-numbered series. The replacing
tooth for the 13th is larger than those for the 9th and 11th.
Except for these minor differences it appears that alternate teeth developed
and erupted simultaneously. Similar results were obtained in Thrinaxodon
(Parrington, 1936). This is in contrast to early Cistecephalus zone cynodonts
(Crompton, unpublished MS.) and other early therapsids (Romer and Price,
1940), where a wave of replacement passes from back to front along each
alternately numbered series.
It is difficult to interpret the order of replacement in the mandible.
Replacement does not appear to be of the regular alternate type observed in
the maxilla. Over certain short distances there is evidence of alternate replace-
ment, e.g. 10 and 8, while over others there is not. Unfortunately, the right
lower dentition could not be studied.
Wear of the teeth
The wear surfaces indicate that the upper and lower teeth must have
occluded crown to crown, and opposing upper and lower teeth did not shear
past one another. Teeth preserved recently after they erupted show no signs
of wear whereas teeth about to be replaced appear to have lost approximately
half of their crowns as a result of wear.
240 ANNALS OF THE SOUTH AFRICAN MUSEUM
It is difficult to interpret the wear-facets of occluding teeth. Upper teeth
have a single flat wear-facet sloping upwards and inwards, whereas lower teeth
have two distinct facets, a labial horizontal facet and a flat inner facet sloping
inwards and downwards. Sharp angles are formed where the facets meet the
edges of the crowns and one another (fig. 3 D). It is difficult to determine the
jaw actions causing these wear-facets. It is unlikely that a single dorso-ventral
DR
ES
> i a
Y WW
—AIMAX CS
€\ 7 : Int.c
ay Ws “@O.
NS
Fic. 5.—Bauria cynops. Ventral view of palate x 14.
(Key to lettering, p. 255.)
movement of the lower jaw would account for their shape. It is also unlikely
that the abrasive action of food particles would produce such constant and
sharply defined wear-facets. It is possible, therefore, that in Ericiolacerta more
complex jaw movements were possible.
Watson (1931) has described upper cheek teeth as . transversely
widened, the high original cusp passing on its lingual side into a ridge which
separates two concave areas into which grind the posterior and anterior cusps
of the two lower molars between which it fits’. The present study does not
confirm this arrangement. The wear patterns clearly show that the labial cusps
of the uppers did not occlude outside the opposing lowers. On the contrary,
the apices of the crowns when present must have met one another. The teeth are
not transversely expanded to the extent suggested by Watson, and at their
bases the crowns are circular in cross-section.
6
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 241
THE DENTITION OF BAURIA CYNOPS
The palate (Plate XIII A and fig. 5) has been well described by Boonstra
(1938) and Broom (1937). The secondary palate (sec. p.) is formed only by the
premaxillae and maxillae. It is narrowest across the posterior border which lies
adjacent to the 5th postcanines. Anteriorly to the posterior border of the
secondary palate the maxillae curve sharply upwards to their contact with the
premaxillae (P.M.). The premaxillae have a broad tongue-shaped process
which extends backwards in the mid-line to meet the maxillae. Two ovoid
foramina (p. for.) are present between the lateral edges of this structure and
the maxillae. These foramina were not reported by Boonstra or Broom. From
the anterior edge of these foramina an interdigitated suture between the
premaxillae and maxillae passes outwards through the diastema between the
last incisor and the canine. No evidence for Broom’s view that part of the
vomer is visible between the premaxillae and maxillae could be found.
Upper dentition (Plates XII B, XIII, and figs, 5, 6, 8)
There are 4 incisors, 1 canine and 10 postcanines in the upper jaw. This
is in agreement with the findings of Broom (1909), Boonstra (1938) and Brink
and Kitching (1953). The roof of the diastema between the incisors and canines
has the form of a shallow pit and receives the lower canine. A thin strip of the
maxilla forms a labial border to this pit and therefore the tip of the lower
canine was not visible when the jaws were closed. The crowns of all the incisors
and both canines were damaged, but the postcanines are fairly well preserved,
except for damage to their outer surfaces. Because of this damage it was essential
to prepare both left and right dentitions. The postcanines of each side are
arranged to form a curve. In the region of the 5th postcanines the opposing
curves are closest to one another, and anterior and posterior to this point they
diverge away from each other (fig. 5). Figures of the length of the postcanine
series are of little value unless it is clearly stated how the measurements were
taken. Measured along the lingual surface of the upper postcanine series they
measure 4-1 cm.
In crown view (fig. 6) the postcanines are oval with the lingual side
of the crown considerably wider than the labial. Brink and Kitching (1953)
claimed that in the third specimen of Bauria the crowns of adjacent teeth
expand so that they come into contact with one another, but that their necks
are separated from one another. It is clear in the type, however, that the
majority of the postcanines were separated by narrow gaps with an average
width of 0-6 mm., although in some cases teeth have been slightly displaced
with the result that consecutive crowns touch one another.
The form of the crowns of the teeth is of great interest because of the
marked changes they undergo as a result of wear. These changes are essential
for the interpretation of jaw action and tooth replacement, and will therefore
be described in detail. Interpretation of the wear pattern is complicated by
242 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 6.—Bauria cynops. A, B and C, reconstructed sections through the 4th, 5th and 7th
right upper postcanines and D, crown view of the upper postcanine dentition. All x 3.
(Key to lettering, p. 255.)
ORR le mse ce ee ae ee ee
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 243
the fact that the occluding tooth rows do not lie parallel to one another, but
at a small angle to one another. The majority of the upper postcanines lie
slightly further lingually to the lowers, but the anterior upper postcanines and
canines lie slightly labially of the corresponding lower teeth.
In order to understand the shape of the crowns, outline drawings of the
posterior view have been given in addition to the crown views (fig. 8). The 5th
postcanine (fig. 6 B, rep. t. 5) on the right side was preserved shortly before
it was due to be replaced, and both the replacing and functional teeth are in
good condition (Plate XII B). Contrary to all previously published reports, the
crown of an unworn upper tooth is not flat. On the contrary, it supports a
prominent cusp (lab. c.) on the narrow labial edge of the crown while the
wide lingual edge of the crown supports a crenulate ridge. In posterior view
the crown has a concave centre, the centre point of the concavity lying towards
the labial side of the crown. A broad, ill-defined transverse ridge extends a
short distance across the crown from the apex of the main cusp. On either side
the crown falls slightly away. On the left side the replacing tooth of the 5th
postcanine was preserved shortly after it had fully erupted and become the
functional tooth; it has the same crown structure as the incompletely erupted
counterpart on the right.
The oth, 7th and 4th are little worn, and a labial cusp and prominent
lingual edge can still be recognized. The 2nd upper postcanine is also little
worn and consists of a high labial cusp and small, ill-defined lingual ridge at
a much higher level than the labial cusp. Wear is greater on the lingual side
in this tooth, whereas in the posterior teeth wear is greatest on the labial side.
This is due to the crossing of upper and lower postcanine rows. In the 8th and
6th the crowns show advanced stages of wear. Two prominent wear-facets
are visible. The labial cusp has been worn away completely to a flat wear-facet
extending lingually and slightly upwards to the centre of the crown where it
meets a flat oblique wear-facet extending upwards and inwards from the
lingual edge of the crown. The orientation of these wear-facets is best seen in
the posterior profiles and stereo-photographs.
In the functional 5th postcanine on the right, wear had proceeded further.
The labial wear-facet extends further across the crown. The prominent ling11
edge and oblique lingual wear-facet have been considerably reduced in size.
In the 3rd tooth the crown is worn to a practically horizontal plane. The ist
and 1oth postcanines are considerably smaller than the other postcanines and
consist of a high labial cusp lingually of which the crown falls away rapidly.
The measurements of the individual postcanines in mm. are as follows:
WIGH SANE: POSte | 2-2 VR 2eAy OR WOO) | 90 | 29) | 321 || 2:7 | 297 2
Wax vine apy o°On SO) Asti) 4e7 | 5S. lea-7 | 51 14s | 2:9 2
244
ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 7.—Bauria cynops. A, crown view of lower postcanines and B and C, sections
through mandible to show the roots of the 6th and 7th postcanines. All x 3.
(Key to lettering, p. 255.)
ee ee
’
eS a on
< ee en ee eee ee eee Oe ee)
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 245
Lower dentition (Plate XIV; figs. 7, 8, 9)
There are three lower incisors. This confirms the findings of Brink and
Kitching (1953). It is easy to understand why Broom (1909) considered that
the type had four. A narrow diastema separates the 3rd lower incisor from the
canine. The crown of the 4th upper canine fits into this diastema. The anterior
region of the snout is damaged in the type in such a way that the crown of the
upper 4th incisor is preserved fitting into this diastema, and is broken off at the
UPPER POSTCANINES. hoes
Gr, LA | a fa
LOWER POSTCANINES.
(oobi aa aemm
Fic. 8.—Bauria cynops. Posterior views of upper and lower postcanines. All x 3.
same level as the remaining lower incisors so that it can easily be mistaken for a
lower incisor.
The postcanine series follow immediately after the canine without an
intervening diastema. As in the case of the upper jaw the postcanines of both
sides are arranged to form an arc, and the two arcs diverge away from one
another in front and behind the 5th postcanine. Eleven postcanines are present
on both sides. In crown view the postcanines are transversely ovate but have a
greater eccentricity than the corresponding uppers.
The roth left postcanine shows the least signs of wear (Plate XIV B; fig. 8).
It consists of a prominent labial cusp. The remainder of the crown is flat except
246 ANNALS OF THE SOUTH AFRICAN MUSEUM
for a faint ridge along the lingual edge and a prominent medium transverse
ridge. This ridge is drawn upwards to form an ill-defined cusp in the centre
of the crown. Faint ridges extend from the apex of the main cusp along the
anterior and posterior borders of the crown. On the right side the 1oth post-
canine is an old tooth which had not been replaced before death, and is greatly
worn.
In the oth left postcanine the main cusp has been lost, but the medium
transverse ridge, although slightly worn, can still be recognized.
In the 4th, 5th and 7th postcanines initial wear stages are present. The
central ridge has been worn away and a basin is present in the centre of the
crown lingually to the main cusp. This basin is flanked in front and behind by
the two ridges extending inwards from the apex of the main cusp. In the 8th
tooth wear is further advanced and this basin has deepened considerably.
Consequently the main cusp is more prominent in this tooth than in unworn
teeth. In the 6th postcanine wear has advanced further. The main cusp is low
and an oblique wear-facet is present on the lingual surface of the crown. The
crowns of the 1st and 3rd teeth are smaller than adjacent teeth and are worn
to a flat horizontal surface. The 11th postcanine is much smaller than the
remaining postcanines; it consists of a prominent labial cusp lingually of
which the crown falls rapidly away.
As in the upper postcanines, a small gap separates consecutive crowns.
The gaps are larger anteriorly than posteriorly. The measurements of the
individual postcanines, given in mm., are as follows:
Max. ant.-post. | 2-2 | 2-5 >|) 2:2.) 9°@ | 8:0) | 9-9 | 9-4) 9-0 || 9-9 sia males
Max. ling.-lab. | 3-2 | 4-2 | 3-4.) 4:8 | 4:8 | (4°83 | 5-4 | 4:85) 5-1 eee
Relationship between the wear pattern of upper and lower postcanines
An outstanding characteristic of the dentition is that in the initial wear
stages the upper teeth are subject to wear mostly on their labial edge and
centre of the crown, whereas in the lower postcanines wear is greater in the
centre of the crown and on the lingual edge. It is only in very advanced wear
stages that postcanines tend to acquire nearly flat occluding surfaces.
Although an attempt has been made to illustrate (fig. 10) the three progres-
sive stages of wear of Bauria postcanines, it is naturally difficult to determine
from one specimen the exact changes the postcanine teeth underwent as a result
of wear. Careful investigation of other bauriamorph specimens will be necessary
to confirm these findings. The initial wear (fig. 10 A, I) in the centre of the lower
postcanines, and wearing away of the upper labial cusps, is apparently due
to the upper labial cusp meeting the lower postcanine lingually of the lower
labial cusp. The lingual edge of the upper tooth occludes lingually of the
la
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 24°7
lingual edge of the lower teeth and this results in the formation of the oblique
wear-facet extending upwards and inwards from the lingual edge of the upper
postcanines. In a more advanced stage (II) an oblique wear-facet is developed
on the lingual edge of the lower teeth where the oblique wear-facet of the
uppers meets the lowers. Consequently both upper and lower are worn away
on their lingual sides during this phase. In an advanced wear stage (III) all
B
unworn
Fic. 9.— Bauria cynops. A, medial view of the right lower postcanine series and B,
graph to illustrate decrease in extent of wear of odd and even-numbered post-
canines. Wear stages I, II, and III correspond to those shown in Fig. 10.
A X 3. (Key to lettering, p. 255.)
the prominent features of the crown tend to be worn away so that only flat
occluding surfaces remain.
The height of the crown above the alveoli borders varies greatly in consecu-
tive postcanines. As a general rule the height is greater in unworn teeth than
in worn teeth. The result is that wear-facets of consecutive teeth are seldom on
the same horizontal plane (fig. g A).
The correct interpretation of the degree of wear is of importance in
determining the order of tooth replacement in the postcanine series.
Roots of postcanines and alveoli
Several teeth are loosely held in enlarged alveoli whereas others are
tightly held by their alveoli. In the latter the alveoli are built up on the lingual
248 ANNALS OF THE SOUTH AFRICAN MUSEUM
side of the tooth in such a way that a horizontal shelf is formed at the base of
the lingual side of the postcanines. This is more marked in the upper jaw than
in the lower. As a general rule worn teeth are tightly held and unworn teeth
loosely.
A fracture through the right ramus of the lower jaw has exposed the
roots of the 6th and 7th postcanines (fig. 7 B, C). The single root of the 7th
is substantial and tapers gradually to a point. There is no indication of any
division of the root. The alveolar bone is closely apposed to the root. A narrow
unworn
ul Mt
A
Fic. 10.—Bauria cynops. A, occlusal relations of unworn and three wear stages
typical postcanine teeth and B, three wear stages compared with unworn upper
and lower postcanines. All x 3.
bony transverse septum separates successive teeth. The remnants of the septum
between the 6th and 7th obscure part of the root of the 6th postcanine. Preserva-
tion of the dentary surrounding the root is not exceptional, but it is possible
to recognize large cavities and canals lingually to the apex of the root. In
addition the root is slightly resorbed on its lingual side.
Unfortunately no sections through the root of an upper postcanine exist.
Reconstructed sections based on external features only are given in fig. 6 A, B,
C. Features that could not be observed are shown in dotted lines. Posterior to
the secondary palate a thick mass of bony tissue constituted by the maxilla
and a thin plate of the palatine (PAL) lies lingually to the last six teeth (fig. 6 C).
The medio-ventral edge of this bony mass is drawn downwards to form a
prominent ridge so that a narrow groove separates the outer surface of this
ridge from the teeth. A longitudinal section was cut through the bony mass.
Numerous cavities and canals are present in it, but their shape and course
could only be accurately determined if serial sections were made. Anteriorly
this lingual mass of bone is obscured by the presence of secondary palate, but
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 249
the ridge on the lingual edge of the lingual mass continues forward close to the
lingual surface of the teeth.
Tooth replacement
Only in one case are a functional tooth and its successor preserved and
visible in a single alveolus. This supplies important information on how tooth
replacement actually takes place, and details on the structure of an unworn
crown.
The tooth preserved in the act of replacing is the 5th right upper (fig. 6 B).
Its alveolus is greatly enlarged. The functional tooth lies hard against its
labial edge. The crown of a replacing tooth is situated lingually and slightly
behind the functional tooth. The lingual edge of the replacing tooth is preserved
at the same level as the edge of the alveolus. The replacement tooth is obliquely
orientated and does not, as would be expected, erupt vertically downwards,
but rather downwards and inwards. The lingual edge of the root of the
functional tooth has been resorbed and the labial edge of the replacing tooth
is closely apposed to it. After the functional tooth was lost, the crown of the
replacing tooth would presumably have moved in a labial direction to the
vertical position. On the left side replacement was in advance of that on the right
because the old functional tooth has been shed and an unworn replacement
tooth is present. Although the alveolus is still greatly enlarged the new tooth
is vertically orientated. Alveoli size is therefore important in determining the
order of tooth replacement. Although horizontal sections through the bony
mass lingual to the last 7 upper postcanines revealed no additional replacing
teeth, the wear patterns of adjacent teeth appear to indicate that active
replacement of the postcanines was probably still taking place in both upper and
lower jaws although in no case in the lower jaw are both the replacing and old
functional teeth preserved in a single alveolus. A probable explanation for this
is that the replacing teeth developed initially deep in the maxilla or dentary
lingually to the apex of the roots of the functional teeth. An empty cavity in this
position alongside the root of the greatly worn 6th lower postcanine appears to
confirm this view. Replacing teeth would therefore only become visible lingually
to the functional teeth shortly before eruption. It was considered inadvisable to
section the type to confirm this view. Boonstra (1938) mentioned a series of
replacing teeth lingual to the functional postcanines exposed by grinding in
the American Museum specimen, but did not give a detailed description.
In Ericiolacerta the replacing teeth are visible throughout their develop-
ment, and the order of replacement is easily determined. In the presentspecimen
of Bauria the order of replacement has to be determined by other criteria such
as the degree of wear of the postcanines, size of alveoli, and size of individual
teeth. These alone are insufficient because, for example, if replacement in both
jaws is not accurately synchronized, occluding teeth of different ages may well
produce different wear patterns in teeth of similar age.
It seems clear from the presence of unworn teeth in the postcanine series
250 ANNALS OF THE SOUTH AFRICAN MUSEUM
and the actual replacement observed, that tooth replacement is still taking
place throughout the postcanine series. The available evidence suggests that
teeth replace alternately in Bauria as in Ericiolacerta. For example, in Bauria,
in the right mandible (fig. 9 A), the 6th, 8th and roth postcanines are slightly
smaller, are more tightly held by their alveoli, and are lower in height above
the alveolar border than the 5th, 7th or gth. Wear is certainly further advanced
in the former group. Similar conditions can be observed in the left mandible.
The gth, 7th and 5th postcanines are larger, more loosely held in their alveoli,
and higher than the 6th and 8th. In the left mandible, replacement is in
advance of that of the right.
In the left maxilla the 5th and 7th are large, unworn, loosely held in their
alveoli, and high-crowned in contrast to the 6th and 8th.
Replacement in the right maxilla also lagged slightly behind that in the
left, and the replacing tooth for the 5th had not yet erupted before death.
Alternate teeth throughout the series did not erupt simultaneously. For example,
the right 7th upper postcanine already showed signs of wear although the 5th
had not yet erupted. On the left the 5th alveolus is larger than the 7th. The
10th postcanine in the left mandible had been replaced only shortly before
death, yet the 8th is well worn and in life would presumably have been the next
to be replaced. This evidence suggests that a wave of replacement moved
forwards along the odd- and even-numbered postcanines. In the case of the
maxilla, the wave that affected the odd-numbered postcanines was in advance
of that which affected the even-numbered, but the waves are spaced so that
alternate replacement is present over short distances.
An expected result of replacement waves moving in an anterior direction
would be that the wear of alternate teeth would diminish in an anterior direc-
tion. There is some evidence of this (fig. 9 B). In the right mandible the 1oth
postcanine is greatly worn and was about to be replaced. The 8th and 6th
teeth are well worn but the 4th is less worn. The 2nd tooth appears to have
erupted shortly before death and shows few signs of wear. The gth, 7th and 5th,
on the other hand, appear also to have erupted shortly before death and are
not much worn, whereas the 3rd is small, greatly worn and loosely held in its
alveolus, and presumably would have been the next to be replaced. The three
wear stages upon which fig. 9 B is based correspond to those shown in fig. 10.
Similar facts were observed in the maxilla.
Tooth replacement in occluding upper and lower jaws appears to have
been fairly closely correlated, with the result that a recently erupted and unworn
tooth of the upper jaw would have met a recently erupted and unworn tooth
of the lower jaw. For example, in both upper and lower jaws on the right side the
gth, 7th and 5th postcanines appear to have been replaced shortly before death.
DIscussION
In both Ericiolacerta and Scaloposaurus there was a marked reduction in the
size of the canines. The postcanine teeth of both genera are similar. Both are
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 251
circular in cross-section at the alveolar border and in both there is a tendency
to develop small cusps although these are limited to the lower jaw in Ericiolacerta.
In Scaloposaurus the shearing action of the postcanine teeth has been retained,
but in Ericiolacerta the reduction of the canines has apparently made it possible
for the apices of the upper and lower postcanines to meet despite the limited
area of contact, and the apices of the crowns are completely worn away in
old teeth.
It is not possible to determine whether the shearing action was retained in
Ericiolacerta or whether a limited amount of lateral and propalinal jaw move-
ment was possible. One of these three movements may account for the wear-
facets on the postcanines of Ericiolacerta which could not be accounted for only
by a simple dorso-ventral movement of the lower jaws relative to the uppers.
In Bauria and all bauriamorphs from the Cynognathus zone the teeth have
expanded transversely to increase the occlusal areas. The crushing action of
Bauria is therefore much more efficient than that of Ericiolacerta, and the latter
appears to be very close to ancestral forms which deviated away from a simple
sectorial action of apposing dentitions.
A similar shift from a sectorial dentition to a crushing dentition took place
in some early mammals (Patterson, 1956), e.g. the dryolestids. Here, too, there
has been a reduction in the size of the canines. The development in the
bauriamorphs paralleled that of some of the later mammals and consequently
there is no question of the known bauriamorphs being ancestral to any known
Jurassic or Rhaetic mammal. A possible exception is Dvarthrognathus. ‘This
form may have developed from early bauriamorphs, and because it possesses
a squamoso-dentary articulation may therefore be classified as a mammal.
The late cynodonts also developed a crushing dentition, but a reduction
of the size of the canines does not appear to have triggered off this development.
In cynodonts such as Diademodon the sectorial action was retained and a
crushing part of the tooth added lingually to the original labial shearing cusps,
the apices of which never met.
It has been tacitly assumed that the flat crowns of Bauria are the result
of a grinding action of the jaws. Because of the mode of replacement in Bauria
the crowns of the postcanine teeth are of varying heights. Consequently, for
individual teeth of the upper and lower jaws to have met it is necessary for the
teeth to intermesh. This would appear to have ruled out any significant
propalinal movement of the mandible. The large upper canine, the pit for the
lower canine, the large lingual cusps of the lower postcanines, and the transverse
processes of the pterygoid would probably eliminate any significant lateral
movement of the mandible. For this reason it is unlikely that true grinding was
developed in Bauria but that the upper and lower teeth could only crush food
as the result of a simple pounding action. Unfortunately details of the nature
of the mandibular joint in Bauria are not available. Recent work to be published
shortly on the jaw action of gomphodont cynodonts has indicated that it is
unlikely that any propalinal movement was present in these forms and that the
252 ANNALS OF THE SOUTH AFRICAN MUSEUM
wear pattern of the postcanines which suggest propalinal movement are due to
other factors.
In Ericiolacerta and Bauria it has been shown that the teeth replaced
alternately. There is no question, therefore, that a mammalian type of dental
succession was present in either of these bauriamorphs. In Friciolacerta it has
been shown that the replacing teeth developed initially in distinct pits in the
dentigerous bones and lie lingually to the functional dentition. The pits were
connected by a shallow longitudinal groove, which presumably housed the
dental lamina. As the replacing teeth increased in size their pits enlarged and
became confluent with the alveoli of the functional teeth, and the replacing
teeth moved labiad. Recent work to be published in the near future has shown
that in the Cistecephalus zone cynodonts, in Thrinaxodon and in Tribolodon, the
replacement teeth developed in a similar fashion. This type of replacement
was not confined to mammal-like reptiles. In his excellent résumé on tooth
replacement Edmund (1960) has cited the relevant literature and has pointed
out that in Wothosaurus and Pleisiosaurus replacing teeth developed in an almost
identical fashion to that described above. Edmund (1957) has shown that in
ornithischian dinosaurs a groove for the dental lamina connected a series of
foramina which penetrated the dentigerous bone and opened into the alveoli
of the functional teeth. Tooth germs budded off from the dental lamina and
passed through these foramina into the base of the alveoli.
In Bauria the teeth did not develop in superficial pits, but apparently deep
in the dentigerous bones lingually to the base of the roots of the functional
teeth. No indication of a groove for the dental lamina or any foramina (guber-
nacular canals) which connected the developing teeth with the oral epithelium
could be observed, but better material is necessary to confirm this point. This
may be interpreted to mean that tooth replacement has ceased in the particular
specimen of Bauria described above, but this is unlikely. To obtain the type of
replacement found in Bauria and in mammals it was necessary for the dental
lamina or remnants of it to lie not superficially but to be buried within the
dentigerous bones. A similar development has taken place in the crocodiles
(Edmund, 1960), where remnants of the dental lamina remain in each alveolus
and retain their capacity to form teeth. The reason for the shift in Bauria
to a deep site for the development of the replacing teeth was presumably
correlated with the transverse widening of the teeth in this form. If the develop-
ing teeth were to have developed in superficial pits they would have projected
into the oral cavity and either been damaged by or hindered mastication.
A similar shift from a superficial to deep-lying position took place in other
groups which have transversely widened teeth, e.g. Diadectes, Watson (1954).
In Diademodon replacing teeth also developed deep in the dentigerous bones
below the functional teeth, but the groove for the dental lamina remained in
the superficial position lingually to the bases of the postcanines. A similar shift
from a superficial site for replacing teeth to a site deep in the dentigerous bones
probably took place in the ancestors of mammals. In mammals, in contrast to
DENTITION AND TOOTH: REPLACEMENT IN TWO BAURIAMORPH REPTILES 253
Bauria and Diademodon, developing permanent teeth retain their contact with
the oral epithelium through their gubernacular canals.
The origin of the mammalian dental succession from the types of dental
succession reported in mammal-like reptiles remains an unsolved problem.
Bauria shows that alternate replacement was present in advanced mammal-like
reptiles and that teeth did replace alternately even though expanded transversely.
It is pertinent to mention that alternate replacement was also still present in the
advanced mammal-like reptile (or early mammal) Diarthrognathus from the late
Trias. This does not support the view of Bolk (1922) that as the postcanines
became more complex they would no longer erupt alternately but that one of
the alternate-numbered series would actually replace the other series, one
becoming the deciduous and the other the permanent. The specialized condi-
tions in gomphodont cynodonts do not help to solve the problem of the origin
of the mammalian succession. An alternate explanation for the mammalian
dental succession has been suggested by Edmund (1960). Edmund has shown
that in early tetrapods such as Captorhinus the teeth of a single zahnreithe erupt
as a unit and are ankylosed to the dentigerous bones. Edmund has suggested
that in mammals the deciduous teeth represent the remnants of one zahnrethe
and permanent dentition a second anterior zahnreihe. This theory also explains
the front-to-back order of replacement in mammals and, because of its
simplicity, is extremely attractive. But certain problems present themselves.
Edmund made no attempt to correlate his theory with palaeontological
evidence. Eruption and functioning of teeth in a complete zahnrethe is known
only in a few specialized groups such as the captorhinids. In mammal-like
reptiles alternate tooth replacement similar to that of typical reptiles is present
except in the specialized gomphodont cynodonts, where the position is difficult
to ascertain.
Edmund’s theory would imply that in the late mammal-like reptiles there
should be a tendency to reduce the number of teeth to two zahnreihen, but no
tendency in this direction has been reported.
Although these facts alone do not disprove Edmund’s view they certainly
do not confirm it. An alternative explanation for the origin of the mammalian
conditions is worth while considering. It is possible to derive the mammalian
succession from forms in which the time lag between the budding-off of the
even- and odd-numbered tooth germs from the dental lamina is reduced so
that odd- and even-numbered teeth can erupt consecutively and function
simultaneously. Even in mammals consecutive teeth in the series do not erupt
one after the other in an orderly fashion; e.g. in man the first deciduous molar
erupts before the deciduous canine, and the first molar erupts before the second
premolar. Compared with the pelycosaurs there appears to be a tendency to
reduce the relative time-interval between the eruption of alternate teeth in
later mammal-like reptiles. In pelycosaurs there is a tendency for the odd- and
even-numbered teeth not to function simultaneously. This has been termed
‘functional distichial replacement’? by Kermack (1956). He has pointed out
254 ANNALS OF THE SOUTH AFRICAN MUSEUM
that functional distichial replacement is not present in the therocephalians
and gorgonopsians studied by him. On the other hand, although alternate
tooth replacement is present in Thrinaxodon and Bauria, the functional periods
of odd- and even-numbered teeth overlap considerably. A reduction of the
time-lag between the formation of odd and even teeth together with a lengthen-
ing of the time between eruption and replacement of individual teeth would
be expected if the form were moving towards the mammalian type of replace-
ment. It is difficult because of lack of reliable evidence on tooth replacement in
mammal-like reptiles to determine a trend of this nature, but tooth replacement
was apparently far less frequent in a form such as Bauria than in Enriciolacerta.
A similar reduction of the frequency of replacement can be traced in the later
cynodonts compared with those of the Cistecephalus and Lystrosaurus zones.
What is urgently required is further growth series of individual mammal-like
reptiles.
SUMMARY
(1) The dentition, tooth replacement and wear stages of the postcanine
teeth of the type specimens of Ericiolacerta parva and Bauria cynops are described.
(2) In both genera alternate tooth replacement was present. In Enciolacerta
the replacing teeth developed initially in pits in the base of the superficial
groove which supported the dental lamina. In Bauria replacing teeth appear
to have developed deep in the dentigerous bones. The origin of the mammalian
order of replacement is briefly discussed.
(3) Ericiolacerta represents an early stage in the transition from a sectorial
to a pounding or crushing dentition. This transition was probably correlated
with reduction of size of the canines in this form. In Bauria a crushing dentition
is better developed than in Ericiolacerta.
(4) It is doubted whether a true grinding action was present in Bauria.
Wear of the postcanines could be accounted for solely by a simple pounding
action.
REFERENCES
Bok, L. 1922. Odontological essays. Fifth essay. 7. Anat. Lond. 57, 55-75.
Boonstra, L. D. 1938. On a South African mammal-like reptile, Bauria cynops. Palaeobiologica
6, 163-183.
Brink, A. S. 1955. A study of the skeleton of Diademodon. Palaeont. Afr. 3, 3-39.
Brink, A. S. 1956. Speculations on some advanced mammalian characteristics in higher
mammal-like reptiles. Palaeont. Afr. 4, 77-06.
Brink, A. S. & Kircuine, J. W. 1953. On some new Cynognathus zone specimens. Palaeont.
Afr. 1, 29-48.
Brom, F. & ScHRO6DER, J. 1935. Beobachtungen an Wirbeltieren der Karrooformation.
VII. Ein neuer Bauriamorphe aus der Cynognathus-Zone. S.B. bayer. Akad. Wiss. 1935, 21-36.
Broom, R. 1905. Preliminary notice of some new fossil reptiles collected by Mr. Alfred Brown
at Aliwal North, S. Africa. Rec. Albany Mus. 1, 269-278.
Broom, R. 1909. Notice of some new South African fossil amphibians and reptiles. Ann. S. Afr.
Mus. 7, 270-278.
Broom, R. 1911. On the structure of the skull in cynodont reptiles. Proc. zool. Soc. Lond. 1911,
893-925.
DENTITION AND TOOTH REPLACEMENT IN TWO BAURIAMORPH REPTILES 255
Broom, R. 1913. On evidence of a mammal-like dental succession in the cynodont reptiles,
Bull. Amer. Mus. nat. Hist. 32, 465-468.
Broom, R. 1932. The mammal-like reptiles of South Africa. London: H. F. & G. Witherby.
Broom, R. 1937. On the palate, occiput and hind foot of Bauria cynops Broom. Amer. Mus.
Novit. No. 946, 1-6.
Crompton, A. W. 1955a. A revision of the Scaloposauridae with special reference to kinetism
in this family. Navors. nas. Mus., Bloemfontein 1, 149-183.
Crompton, A. W. 1955b. On some Triassic cynodonts from Tanganyika. Proc. zool. Soc. Lond.
125, 617-6609.
Crompton, A. W. 1958. The cranial morphology of a new genus and species of ictidosaurian.
Proc. zool. Soc. Lond. 130, 183-216.
Epmunp, A. G. 1957. On the special foramina in the jaws of many ornithischian dinosaurs.
Contr. R. Ont. Mus. Zool. Palaeont. No. 48, 1-14.
Epmunp, A. G. 1960. Tooth replacement phenomena in lower vertebrates. Contr. R. Ont. Mus.,
Life Sci. Div. No. 52, 1-190.
Kermack, K. A. 1956. Tooth replacement in mammal-like reptiles of the suborders Gorgonopsia
and Therocephalia. Phil. Trans. (B) 240, 95-133.
Ktune, W. G. 1956. The Liassic therapsid OLt1cokyPHus. London: British Museum.
ParRRINGTON, F. R. 1936. On tooth replacement in theriodont reptiles. Phil. Trans. (B) 226,
121-142.
PATTERSON, BRYAN. 1956. Early Cretaceous mammals and the evolution of mammalian molar
teeth. Fieldiana, Geol. 13, 1-105.
Romer, A. S. & Price, L. W. 1940. Review of the Pelycosauria. Spec. Pop. geol. Soc. Amer.
No. 28, i-x, 1-538.
SEELEY, H. G. 1895. Researches on the structure, organization and classification of the fossil
reptilia. Part ix. On the Gomphodontia. Phil. Trans. (B) 186, 1-57.
Watson, D. M. S. 1914. Notes on some carnivorous therapsids. Proc. zool. Soc. Lond. 1914,
1021-1038.
Watson, D. M. S. 1931. On the skeleton of a bauriamorph reptile. Proc. zool. Soc. Lond. 1931,
1163-1205.
Watson, D. M. S. 1954. On Bolosaurus and the origin and classification of reptiles. Bull. Mus.
comp. Zool. Harv. 111, 299-449.
Key To LETTERING
at. bo. Attachment bone.
c. Canine.
can. Canal in dentary.
DEN. Dentary.
Lt25 Functional 5th postcanine.
lab. Labial.
lab. c. Labial cusp.
Jeena Nh oF Pit for lower canine.
ling. Lingual.
long. gr. Longitudinal groove.
MAX. Maxilla.
p. Pit for replacing tooth.
PAL. Palatine.
p. for. Palatal foramen.
rem. f. 11 Remnant of 11th functional tooth.
rep. t. Replacing tooth.
rep. t. 11 11th replacing tooth.
sec. p. Secondary palate.
V: Vomer.
w. fac. Wear-facet.
w. fac. h. Horizontal wear-facet.
w. fac. o. Oblique wear-facet.
Ann. S. Afr. Mus., Vol. XLVI Plate XII
A (top): Ericiolacerta parva. Left maxilla. B (bottom): Bauria cynops. Detail of replacement of
5th postcanine.
fl
;
; a:
i
;
i
= — ™
i
i
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5
}
Tie
%
¥ =
~
z “— Le Aba
; 2 ink Sar ok SE he ia ee
1 r a
” ~ y
Ann. S. Afr. Mus., Vol. XLVI Plate XIII
A (top): Bauria cynops. Ventral view of palate. B (bottom): Bauria cynops. Right postcanine
dentition.
Ann. S. Afr. Mus., Vol. XLVI Plate XIV
A (top): Bauria cynops. Crown view of left postcanine series. B (bottom): Bauria cynops. Detail of
posterior three postcanines.
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, 8. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Hartman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-500.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
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by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name... .
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should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 20, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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ANNALS
OF THE
Pee ine AERICAN MUSEUM
VOLUME XLVI
PART X
NOMENCLATORIAL CHANGE FOR A LONG-KNOWN
SOUTH AFRICAN FISH
By
J. L. B. Smira
Research Professor and South African Council for
Scientific and Industrial Research Fellow in Ichthyology,
Rhodes University, Grahamstown
jen Hdd ty ate
in Wh UE oe i
ISSUED FEBRUARY 1962 PRICE toc.
PRINTED FOR THE
TRUSTEES OF THE SOUTH AFRICAN MUSEUM
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[CONTINUED ON INSIDE BACK COVER
NOMENCLATORIAL CHANGE FOR A LONG-KNOWN
SOUTH AFRICAN FISH
By
Jj. L. B. Surry
Research Professor and South African Council for
Scientific and Industrial Research Fellow in Ichthyology,
Rhodes University, Grahamstown
[Accepted September 1961]
(With 1 plate)
More than sketchy knowledge of the shore fishes of southern Africa came
only long after that of most other parts. This was a result of various unusual
circumstances, stormy seas, poor harbours and hostile natives, all of which
retarded European settlement, so that long after early voyagers and travellers
had gained fairly extensive knowledge of the fishes of parts as remote as
Australia, Cape fishes remained poorly known. While certain early travellers
and collectors visited the Cape, more active and able exponents operated in
the Mauritius and Madagascar area, with the result that many species from
there were described and named in quite early times. As it has been found
that many fishes of the Indo-Pacific cover vast areas, one result has been that
certain species named as new in the Pacific I have since found to have been
described before from Mauritius and other near-by parts of the western Indian
Ocean.
Family Pomadasyidae
A well-known South African fish whose habits and nature have proved
most interesting from many aspects (Smith, 1935, p. 201; 1950) is the so-called
Spotted Grunter, long known as Pomadasys operculare Playfair, 1866, the type
from Zanzibar. This fish is widely distributed in the Indian Ocean, being
common about India and Madagascar, where all available evidence indicates
clearly that in both countries it has been caught by and was well known to
the natives from quite early times. In Madagascar it is one of the best-known
inshore fishes (I have photographs of catches by native fishermen there) and
as it is not only of striking appearance but an excellent table fish, it seemed
likely that it must have been noticed by earlier travellers and collectors in the
Mauritius-Madagascar area. I therefore set out to see if it had not been
25/7
Ann. S. Afr. Mus. 46 (10), 1962, 257-260, 1 pl.
258 ANNALS OF THE SOUTH AFRICAN MUSEUM
described before Playfair, and soon discovered that it had. There is no doubt
that Playfair’s name must yield by priority to commersonni Lacépéde, 1802,
the type from Madagascar, collected and described by Commerson in the
previous century. Lacépéde (18024, p. 431), on a drawing by Commerson of
a fish from Madagascar, describes as follows (names as in original, description
translated): “Le LABRE COMMERSONNIEN (Labrus commersonnit). Nine sharp rays
and 16 articulated rays in dorsal fin. Teeth in both jaws about equal. One
sharp ray and 17 articulated rays in anal. The back and a great part of the
sides of the fish sprinkled with equal round small spots.’ Then Lacépéde
(1802a, in pl. 23, fig. 1, LABRE commersonnien) shows D IX 15, and anal with
one spine and about 16 rays, pelvics with at least 7 soft rays, and deeply lunate
caudal. Lacépéde’s artist Des¢ve was often impressionistic and inaccurate by
present standards, but the drawing could have been based on the species now
under review.
Lacépéde (1802), p. 181, no. 26) also describes LUTJAN MICROSTOME
(Lutjanus microstomus), also on a drawing by Commerson of a Madagascar fish, as
follows: ‘Nine spines and 16 articulated rays in the dorsal: the anal sickle-
shaped; the head conical and long: the opening of the mouth small: a denticula-
tion (serration) near the nape: the pectoral narrow: a great number of dark
spots, irregular and small, on the body and tail.’ Also Lacépéde (1802a,
pl. 34, fig. 2) shows LUTJAN MICROSTOME with D IX 16, anal total about 16,
and a serrated preopercle margin; the body with small spots, and unmistakably
the species here under review. Lacépéde (1802), p. 216) states: ‘Nobody
has yet published the description of (LUTJAN) muicrostome, a drawing of which
we have seen among the manuscripts of Commerson, and which lives in the
great Equinoxial Ocean or in parts of the Great Ocean near the tropics. The
teeth of microstome are small and slender and the anus is nearer the head than
the tail.’
Cuvier & Valenciennes (1830, p. 252) describe ‘Le PRISTIPOME COMMER-
SONIEN (Pristiboma Commersonit, nob.; Labre Commersonien et Lutjan microstome,
Lacép.)’ as follows (names as in original, description translated) :
‘Among the papers of Commerson is a drawing which has been shown
twice in the work of M. de Lacépéde, the first time (vol. III, pl. 23, fig. 1)
with the name Labre Commersonien (pp. 431 and 477)’ (to which Cuvier &
Valenciennes give the footnote: “The characters of Labre Commersonien
([Lacépéde] p. 431) and notably the 17 rays in the anal, are taken from the
figure ([Lacépéde] pl. 23) which has been distorted by [the artist] Deséve.’) ;
‘the second time (vol. ITI, pl. 34, fig. 2) with the name Lutjan microstome (vol. IV,
pp. 181 and 216)’ (to which Cuvier & Valenciennes give the footnote: “The
characters of Labre microstome ({Lacépéde] p. 181) are taken from the other copy,
also made by Deséve, and conform better with the original, but again he
distorts the anal.’). ‘.. . It [is a fish that] was found, says the traveller [Commer-
son], in the mouth of small rivers in the south of Madagascar. The specimen
described weighs about 5-6 ounces, and is about 8-9 inches long, but it grows
NOMENCLATORIAL CHANGE FOR A LONG-KNOWN S.A. FISH 259
larger and reaches about 2 pounds in weight. The colour is a blue silvery white,
covered with brown-black spots. The dorsal, anal and caudal are brownish,
the other fins whitish or transparent. The teeth are very small, like a file.
The margin of the preopercle is denticulate, no marked spine on opercle.
The dorsal is deeply cleft. ‘The pectorals are long and pointed. The first anal
spine is very short, the second abruptly longer, very strong, the third much
less. The caudal is forked. B. 7; D. 10/16; A. 3/9; p. 17; V. 1/5.’
In view of this unmistakable designation of the species, it is plainly the
fish that has been known all this time under the name of P. operculare Plyfr.
It is curious that Lacépéde’s name has not before been revived. That this
has not been done appears to be due chiefly to Giinther, who in 1859
(p. 289) synonymized commersonni Lacépéde, 1802, microstoma Lacépéde, 1802
and commersoni as described by Cuvier & Valenciennes (1830, p. 252) all with
hasta Bloch, 1790. Giinther’s commanding position caused this opinion to be
accepted, uncritically, by all later workers on the Indian Ocean fauna. The
following table shows the difference between hasta Bloch and operculare Playfair,
and the correspondence of the latter with commersonni Lacépéde.
hasta operculare commersonni
Dorsal D XII-XIII 13-15 D X (-XI) 13-16 X 16
Anal A III 6-7 A III 8-10 III 9
Gillrakers 12-13 15-17 a
Markings Interrupted crossbars Many small dark spots Many small dark spot
In order to make quite certain I wrote to the Natural History Museum in
Paris to ask if they had Commerson’s Madagascar specimen. Dr. M. Bauchot
replied to say that the specimen (like most of Commerson’s fishes) cannot
be found, and that Commerson’s original drawing is apparently lost, but on
going through Cuvier’s papers there was found a copy by Cuvier’s artist of
Commerson’s original 1768 drawing with notes copied from Commerson’s
manuscript; of this they kindly sent me a photograph (shown here). Also
pinned to the drawing was a copy of Lacépéde’s illustration. This figure leaves
no doubt about the identity of Commerson’s fish from Madagascar, and the
name Pomadasys operculare Playfair, 1866 used for close on a century must yield
by priority to Pomadasys commersonni Lacépéde, 1802.* A photograph of a
specimen from the Transkei of Pomadasys commersonni of size comparable with
Commerson’s fish is shown here (pl. XV).
I wish to record my gratitude to the South African Council for Scientific
and Industrial Research for financial assistance, and to Dr. M. Bauchot of
Paris for valuable assistance rendered in searching their archives, and for the
photograph reproduced in Plate XV.
* The original spelling with double “‘n’”’ by Lacépéde is used here.
260 ANNALS OF THE SOUTH AFRICAN MUSEUM
REFERENCES
Biocu, M. E. 1790. Naturgeschichie der auslandischen Fische. 4. Berlin.
Cuvier, G. L. C. F. D., & VALENCIENNES, A. 1830. Histoire naturelle des poissons. 5. Paris: Levrault.
GUNTHER, A. 1859. Catalogue of the acanthopterygian fishes in the collection of the British Museum.
1. London: British Museum.
LACEPEDE, B. G. E. de la V. 1802a. Histoire naturelle des poissons. 3. Paris: Plassan.
LAcEPEDE, B. G. E. de la V. 1802b. Histoire naturelle des poissons. 4. Paris: Plassan.
Piayrair, R. L. 1866. The fishes of Zanzibar: Acanthopterygi. London: van Voorst.
SmiTH, J. L. B. 1935. New and little known fishes from South Africa. Rec. Albany Mus. 4, 169-235.
SMITH, J. L. B. 1950. Pomadasys operculare Playfair, and the South African seas. Ann. Mag. nat.
Hist. (12), 3, 778-785.
Ann. S. Afr. Mus. Vol. XLVI Plate XV
Above: Pomadasys commersonni (Lacépéde). 150 mm. (Transkei).
Below: A copy by Cuvier’s artist of a drawing by Commerson of a fish about 8 inches in length
from Madagascar. Below, left, pinned to the drawing, is a copy of Lacépéde’s drawing published
in 1802, III, pl. 23, fig. 1. Also notes copied from Commerson’s original manuscript are written
above, indicating Fort Dauphin, Madagascar, as locality. (Photo: Museum, Paris.)
Ul pp 9 POOF
jis ea chy Pree hei . esewass
jor
4 EFS,
1 a ae SRB)
is
ek
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, 8. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Hartman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
I—262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
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by which a species has been previously designated (subsequent to 1758) are listed
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arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 250.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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ANNALS
OF THE
SOWPH «AFRICAN MUSEUM
VOLUME XLVI
PART XI
BEET HYDROZOA OF THE SOUTH AND WEST COASTS OF
SOUTH AFRICA
PART I. THE PLUMULARIIDAE
By
N. A. H. MiLitarp
Koology Department, University of Cape Town
\G we Ss eu
x |
: oS
Ra
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[ CONTINUED ON INSIDE BACK COVER
THE HYDROZOA OF THE SOUTH AND WEST COASTS OF
SOUTH AFRICA
PART I. THE PLUMULARIIDAE
By
N. A. H. Mittarp
Koology Department, University of Cape Town
[Accepted April, 1961]
(With 12 figures in the text)
CONTENTS
PAGE
Introduction . Le we 200
Station list . 3 ‘ 203
Remarks on classification . 267
List of species : : 6 BB
Systematic section fh OTA:
Summary . Sn. ge Salads uae Bits)
Neferences, 1%: ‘ : 5 Stik}
INTRODUCTION
This paper is the continuation of a series on the systematics of the hydroid
coelenterates of the South African coasts. The hydroids of False Bay, near
Cape Town, and those of the east coast (Natal and Portuguese East Africa)
have already been described (Millard, 1957, 1958, 1959a), as have the hydroids
from ships’ hulls in Table Bay (Millard, 1959b). This leaves the species from
the west and south coasts to be dealt with, the latter including very rich collec-
tions from the Agulhas Bank. It was found that it is no easy matter to decide
on a line of demarcation between the west and south coasts when one is dealing
with material dredged off the shore, and also that the distribution of the species
does not justify such a division. These coasts are thus dealt with together.
However, the hydroids of False Bay, which effectively separates the two coasts
as far as the littoral material is concerned, have already been described. This
paper, therefore, deals with the west and south coasts from South West Africa
on the west to the southern border of Natal on the east, with the exception of
False Bay. In a few instances recently collected specimens from False Bay or
Natal have been mentioned when they have something of importance to add
to the description. These records are quoted within brackets.
It was also found impracticable to separate littoral and estuarine material
from that of deeper waters, as the Hydroida are essentially a sublittoral group
and only a very small percentage of the population extends its range upwards
into the littoral area. The term ‘littoral’ is used here in preference to ‘inter-
261
Ann. S. Afr. Mus. 46 (11), 1962, 261-319, 12 figs.
262 ANNALS OF THE SOUTH AFRICAN MUSEUM
tidal’, and is taken to include forms within reach at low water of spring tide
even though they are not exposed and thus are not strictly between tide-marks.
Since the scope of the paper is thus very large and deals with an enormous
amount of material, it is proposed to divide it into parts, this, the first,
dealing with the Plumulariidae. In toto it should complete the survey of
the coast.
An analysis of the geographical distribution is being left to a later date
when the descriptions are complete and more records are available, as also
keys to the identification of species.
The material described has been obtained from a number of different
sources and I am indebted to many bodies and individuals for their co-operation
in building up the collection.
Some of the material dates from the time when the late Professor
T. A. Stephenson held the chair of Zoology at the University of Cape Town,
and was collected by him and his associates during his intertidal survey of the
coast. ‘This material is catalogued with a single reference letter.
The bulk of the material has been collected since that date by members
of the Zoology Department under the direction of Professor J. H. O. Day, on
expeditions to estuaries round the coast, on instructional student camps, and
particularly by cruises of the University research vessel, the John D. Gilchrist.
The Division of Fisheries has contributed material dredged by the r.s. Africana,
and Messrs. Irvin and Johnson material brought up by commercial trawlers.
Finally there is a fairly bulky collection which was submitted to me for
identification by the South African Museum and which was dredged by the
s.s. Pieter Faure 50-60 years ago. Much of this material has deteriorated during
the years, but since most of the Plumulariidae can be identified on their skeletal
parts it has proved very useful and provided a number of valuable records,
particularly from those regions which were only scantily covered by the
John D. Gilchrist.
The details of the collecting stations, including date, latitude and longitude,
depth, and nature of bottom, are given in the station list in the following
section, and only the catalogue numbers are quoted under individual species.
In the description of the species the full synonymy has not been repeated
where it has been quoted in previous papers of the series. Instead the latter are
referred to.
A visit to the British Museum of Natural History in the latter half of 1960
enabled me to examine many type specimens and confirm identities of various
South African species. For this privilege I am most grateful to the director of
the museum and to Dr. W. J. Rees and members of his department. It has
also made possible the solution of a number of problems of synonymy. I am
also indebted to the director of the Munich Museum for the loan of slides of
Stechow’s collection from South Africa. |
Various bodies have contributed either directly or indirectly to the
expenses involved, including the South African Council for Scientific and
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 263
Industrial Research, the Carnegie Corporation, the Staff Research Fund of
the University of Cape Town, and the Publications Fund of the University.
The type specimens of new species will be deposited in the South African
Museum, and in these cases the registered museum number is quoted as well
as the University catalogue number.
The Trustees of the South African Museum acknowledge with thanks
receipt of a grant from the University of Cape Town for this publication.
STATION LisT
A. Littoral material from Oudekraal on the west coast of the Cape Peninsula.
Wate 119594) bositions 39-50-5192) bo.22-2: E.
AFR. Material dredged by the government research vessel, R.s. Africana.
Date Position Depth (m.) Bottom
AFR 729 15.8.4.7 31°22-8’S./16°20-2’E. 366 bk M, R
AFR 736 17.8.4.7 30°42°4’S./15°59°2’E. 201 co gn S, Sh
AFR 865 8.1.48 34°35°5'9./19°18°2’E. 27 R,S
AFR 945 19.3.48 36°25’S./21°8’E. 177 S,R
AFR 994 19.4.48 34.°34°5/S./21°22°5’E. 68 co S, Sh
B. Littoral material from Lambert’s Bay on the west coast. Date: 30.7.48.
Position: 32°5'S./18°14’E.
BMR. Bushmans River Estuary, on sand and muddy banks. Date: 10.9.50.
Position: 33°41'S./26°40’E.
CP. Littoral material from various localities on the west coast of the Cape
Peninsula.
Date Locality Position
(Oe | 29.4..38 Kommetje 34°8°5'S./18°19°4’E.
CP 328 31.12.48 Sea Point 33°55°2'S./18°22-6’E.
CP 335 12.9.49 Hout Bay 34°3’S./18°21’E.
CP 646 5.12.60 Oudekraal 33°58°5'9./18°22-2’E.
CP 650 1.2.61 Bakoven 33°57°0'S./18°22°3’E.
CPR. Material from various localities in the Cape Province.
Date Locality Position Depth (m.)
CERO GI 14.1.49 Cape Agulhas 34.°50'S./20°1’E. littoral
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 44 —.1.58 Port Nolloth 29°15'S./16°52’E. littoral
CPR 46 20.6.59 Umgazi Bay 31°43'S./29°26’E. 27
E. Littoral material from Port Elizabeth on the south coast. Date: July 1936.
Position: 33°56'S./25°36’E.
FAL. False Bay.
Date . Position Depth (m.) Bottom
FAL 326 10.9.57 34°7°7'S./18°26:0’E. 3-5 R
KNY. Knysna Estuary, on the south coast. Position: 34°5'S./23°4’E. (average).
Date Depth (m.) Bottom
KNY 22 16.7.47 1-4. M
KNY 28 W747 littoral wooden pole
KNY 30 16.7.47 5-7 ,M
IKNY (57 20.7.47 115 R
KNY 70 15.7.47 2-6 Ss
KNY> 71 15.7.47 7] Sh, S, M
KNY 127 15.4.49 O-I M
KNY 165 Q.7.50 littoral R, Buoy
264
L. Littoral material from East London, on the south coast. Date: July 1937.
ANNALS
Position: 33°1'S./27°54’E.
LAM. Dredged in Lambert’s Bay, west coast.
LAM 2
LAM 7
LAM 9
LAM 13
LAM 16
LAM a1
LAM 26
LAM 27
LAM 30
LAM 35
LAM 43
LAM 45
LAM 46
LAM 51
LAM 59
LAM 62
126
1asy
190
257
367
378
392
4.72
51 I
513
LIZ 7
LiZn3
LIZ 16-17
LIZ 27
MB. Dredged in Mossel
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NM
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OF THE SOUTH AFRICAN MUSEUM
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Date Position Depth (m.)
16.1.57 32°4:5'S./18°18-3’E. 17
18.1.57 32°5’S./18°17°9’E. 29
Dele ay 32°4°7'S./18°17°7' EB. 23
19.1.57 32°4/S./18°18-1E. 18
17.1.57 32°4:8'S./18°18-2’E. II
17.1.57 32°7-5/8./18°17-6'E. 20
18.1.57 32°4°9/S./18°17°5E. 27
16.1.57 32°4°1'S./18°18-4’E. 16
19.1.57 32°5°1'S./18°1 7-7 E 20
19.1.57 92°5:5/9./18°17°7' E 27°5
Q1.1.57 32°4:9'S5./18°18-2’E 13°5
21.1.57 32°5/S./18°18-2’E 8
CIEE | B9°APAS./ TO yey ku. 23
23.1.57 32°6°5 S./1O 17-7 E- 16°5
23.1.57 32°9’S./18°18’E. 16
23.1.57 32°1°5/S./18°18’E. 25
LB. Material from Langebaan Lagoon, on west coast.
Date Position Depth (m.)
15.7.46 BQC7 hos) LO a1 ki. 4
16.7.46 33°6"3'9./16 3:3. 7
26.4.49 33°11°3/S./18°5°5/E. O-2
2.5.51 Bin 7 Teh Ike) Gh lee littoral
7.5.53 33°6/S./18°1°5’E. littoral
7.5.53 33°6/S./18°9’E. O-I
8.5.53 93°7-O1S.|1O 2° 1 Le. 2°5
6.5.55 33-7°4'S./18°2°5’E. 3-4
23.9.57 33°6/S./18°1'5/E. littoral
5.1.58 33°6-5/S./18°2’E. littoral
LIZ. Dredged in Algoa Bay, Port Elizabeth, south coast.
Date Position Depth (m.)
6.4.54 33°58°1’S./25°38-9’E. 9
6.4.54 33°58°2’S./25°38-8’E. 7-8
7-454 33°58°4'S./25°40°5’E. 14
11.4.54 34.°0°8'S./25°4.2°4'E. 5-7
Bay, south coast.
Date Position Depth (m.)
12.1.56 34.°8:0'S./22°8°5’E. QI
12.1.56 BY PGy STIG ECY 1B 19
13.10.56 BAL l= 95/225 LOntgis. 16
13.1.56 BA iE (94) 2270;0781. 19
13.1.56 34°11.1'S./22°10°1'E. QI
15.1.56 34.°8°3/S./22°9°4’E. 31
16.1.56 34°9°3/S./22°10:0'E. 31
16.1.56 34°10°1'S./22°8-0’E. 9
16.1.56 34°8-5/S./22°8-8’E. 25
17.1.56 34°11°3/9./22°10°:0 EB. 10
17.1.56 34°11:0'S./22°9-9'E. 14
17.1.50 34°10°7'S./22°9'6’E. 9
18.1.56 34°4°3/S./22°13°5’E. 12°5
18.1.56 34.°4:1'S./22°13-9'E. 115
18.1.56 34.°4°3/S./22°14°2’E. 17-20
19.1.56 34°8-9’S./22°7-0’E. 18
19.1.56 84 °O°1'S./22° 7°21. 12
20.1.56 34°5:0'S./22°11°8’E. 19
20.1.56 34.°6-2’S./22°10°9’E. 27°5
HYDROZOA OF SOUTH AND WEST COASTS
BAett-619./22> 10-2) 1. 29
9A°T1-4/9./22°10°1 BE. 29
MB 83 21.1.56
MB 84 21.1.56
MB 88 18.1.56
34.°4°8/S./22°13°1’E.
26
OF SOUTH AFRICA
S, Sh, R
R
co S, Sh, R
265
N. Littoral material from Port Nolloth, west coast. Date: 11.11.35. Position:
ZOO.) TO 2ues
NAD. Dredged off Natal, east coast.
NAD 1
NAD 9
NAD a1, 22
Date
17.5.50
23.4.50
12.8.58
29°46’S./31°17’E.
Position Depth (m.) Bottom
30°47°1'S./30°29°1’E. 44. St
110-130
49
29°58’S./31°2’E.
PP. Littoral material from Paternoster, west coast. Date: 24.9.57. Position:
32°43'5./17°55 E.
S. Littoral material from Still Bay, south coast. Date: January, 1932. Position:
BAe 2 038/21. 20) Er.
SAMH. Material dredged by the s.s. Pieter Faure and lodged in the South
African Museum. (The 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 given to the nearest minute.)
Date Position Depth (m.) Bottom
SAMH 148-153 15.7.1898 34°8’S./22°16’E. St
SAMH 163 28.10.1898 33°54'S./25°47'E. 40 d 8
SAMH 165-169 11.11.1898 33°4.9'S./25°56’E.
SAMH 190-194 3=.28.12.1898 33°9/S./28°3’E. 86 S, Sh, R
SAMH 195, 196 15.2.1899 33°53'S./25°45 E. 33-42
SAMH 197-201 7.3.1899 33°59/S./25°51 EE. 24-27
SAMH 210 15.3.1899 33°4'7'S./26°19'E. 18-29 8S, Sh, St
SAMH 212 24.3.1899 33°50'S./26°35’E. gI M
SAMH 215-225 19.6.1899 34°26'S./21°42’E. £S
SAMH 238-241 ~—=11.10.1900 34°8/’S./22°59°5'E. 73 S, Sh, Crl
SAMH 248 24.10.1900 94°29°5./22°01E:: ai R
SAMH 249 15.7.19O1 33°13°5/S./27°58’E. 89 brk Sh
SAMH 254-277 17.7.1901 9307S (27° 47°5 E. {S$
SAMH 278-280 25.7-1901 32°50'S./28°18°5’E. 86 brk Sh
SAMH 284 6.8.1901 33°5'S./27°50’E. 7 R, brk Sh
SAMH 285-203 13.8.1901 32°45/S./28°26’E. 53 brk Sh, St
SAMH 297-208 13.8.1901 32°4.7'S./28°28’E. 82 brk Sh
SAMH 299-304 15.80.1901 32°42’S./28°26’E. 31 R
SAMH 305-309 10.9.1901 33°54’S./26°51’E. 120 brk Sh, St
SAMH 319, 320 23.9.1901 34°5'S./26°34’E. 115 S, Sh, bk Spks
SAMH 322-333 22.9.1904 34°12’S./22°15°5’E. 51 £S
SAMH 344-347 19.10.1904 34°15'5/9./22°14’E. 64 M
SAMH 349 15.8.1905 34.°8’S./23°32’E. 73 M
SAMH 353 28.8.1906 33°54°5'9./26°28’E. 104 M
SAMH 355 —.1.1913 29°55/S./31°14’E. littoral
SAMH 360 25.7.1901 32°50’S./28°18-5/E. 86 brk Sh
SB. Saldanha Bay, west coast.
Date Position Depth (m.) Bottom
SB 141 2.5.57 33°2°5'S./18°2’E. littoral R
SB 150 22.09.57 33°2°5S./18°2’E. littoral R
SB 168 25.9.57 Coie Dixie fl eine a oe littoral R
SB 178 28.4.59 33°3°6/S./18°0-4’E. 15 Sh, kh 8, R
SB 194 1.5.59 33°3°5'S./17°59°2’E. 20 Rea
SB 196 1.5.59 33°4°4S./17°56-4E. 35,
266
ANNALS OF THE SOUTH AFRICAN MUSEUM
SCD. Dredged off the south coast.
SCD 5
SCD 20
SCD 22
SCD 26
SCD 29
SCD 33
SCD 36, 37
SCD 50
SCD 52
SCD 61
SCD 67
SCD 69
SCD 75
SCD 79-81
SCD 82
SCD 84, 85
SCD 94
SCD 96
SCD 103
SCD 104
SCD 108
SCD 117
SCD 122
SCD 126
SCD 138
SCD 141
SCD 152
SCD 153
SCD 169
SCD 175
SCD 186-8
SCD to1
SCD 195
SCD 199
SCD 201
SCD 206
SCD 219
SCD 234
SCD 239
SWA. Littoral material from rocky shore at Luderitz, South West Africa.
Date
19.4.58
26.5.58
26.5.58
23.5.50
22.6.58
21.5.50
19.5.58
18.5.58
20.8.58
15.08.58
4-7-59
5:7-59
16.7.59
16.7.59
17-7-59
17-7-59
20.7.59
20.7.59
22.7.59
23-7-59
23-7-59
14.2.60
14.2.60
3.6.60
28.6.60
28.8.60
2.6.60
25.11.60
24.11.60
30.11.60
30.11.60
29.11.60
29.11.60
30.11.60
29.11.60
30.11.60
29.11.60
30.11.60
29.11.60
Position
34°15'S./25°5'E.
34°9°3'S./23°23°8'E.
34°26-7'S./23°26-0’E.
33°47'5./26°4’E.
33°38-6’S./26°54°7’E.
35°3'9./27°56:2'E.
32°15:2'S./28°57°7'E.
31°38:8'S./29°34°4’E.
34° 1'S./25°45°5 E.
32°17°7'S./28°54°5'E.
34°20'S./24°22’E.
33°31'S./27°14°5E.
32°39'S./28°38’E.
32°43'S./28°28’E.
Bo 82709. /2q 540 gts.
33°3'9./27°55 E.
33°55°5 9-/25°51'E.
34°21'S./25°41'E.
35-7'9./22°15/E.
34°26°5/S./21°48’E.
34°35'S./21°56’E.
34°46’S8./22°5/E.
34°55'S./21°26’E.
34°3’S./25°59'E.
33°58°9/S./25°41°4’E.
34°20'S./23°31’E.
34°10'S./23°32 EB.
34°4°3'S./23°25°8'E.
34°7°5'S./23°31°7'E.
34°10'S./23°32’E.
34°5'8'S./23°23°2’E.
34°51’S./23°41'E.
34°2:0'5./23°28-4’E.
34°51'S./23°41E.
34°2:0’8./23°28-4’E.
Date: —.7.57. Position: 26°33'S./15°9’E.
TB. Material dredged from Table Bay.
TRA 35
TRA 36
TRA 38
TRA 42
Depth (m.)
49
Date Position Depth (m.)
11.2.47 33°47°5'S./18°24°3’E. 19-20
25.10.46 33°52°5'9./18°26’E. 22
4.8.46 33°49°5'9./18°27°5/E. 9-18
11.2.4.7 33°48'°3'S./18°24’E. 10-12°5
Tle 33°4.7°5 9./18°24°3’E. 19-20
26.6.4.7 33°52°7'9./18°28-7’E. 9
BOTA 33°52°7'9./18°26°8’E. 20°5
15.12.57 33°48-6/S./18°24-6’E. 15
TRA. Material collected by commercial trawlers.
Date Position Depth (m.)
21.1.50 34°34’S./20°50’E. 70
21.1.50 34°34’S./20°50’E. 73
—.7.50 34°30'S./20°56’E. 73
—.7.51 34.°30’S./20°55’E. 70
24.9.52 31°50’S./16°50’E. 330
TRA 47
Botiom
R
Sh
M, Sh
R
S, Sh
R
R
R
2S
2S
Ss, M
St, Sh
br S, Sh
R
bk M, S
Sh
S
co S, brk Sh
co S, Sh, St
R
kh §
Bottom
Sh. ot
iy
DAD DNA
RNANNN
GP jor Ge fer
DN
ct
DN
St
ior
a)
~
DN
Se
DN
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 267
TRA 56 28.11.52 34°40’S./21°35’E. 70-77) Ree S
TRA 86 23.3.53 32°41°7'S./17°58°5 E. 9 )
TRA 92 —.1.54 35,°3’8./21°50'E. 110 Ss
TRA 99 18.1.56 34°25°5'S./21°50°2’E. 60 S
TRA 134 23.2.57 34°19'S./18°30’E. 52 S, Sh, R
TRA 151 6.3.58 34°51'S./19°55E. 22 R
TRA 156 15.10.58 BA 1249. / lo. 220E, 18-27 Cable
TRA 159 6.7.58 33°560'S./25°36’E. Turtle’s back
WCD. Dredged off west coast.
Date Position Depth (m.) Bottom
WCD 7 24.359 SABO P3095) LO ale]: by bu 43 R
WCD i2 24.3.59 34°9°4'S./18°16°5’E. 75 R
WCD 18 ~—_.29..4.59 33°5°618./17.54°5 E. 64. kh M
WCD 20 30.4.59 eGR ts) iyo ay ae 86 R
WCD 32 15.12.59 34°10'S/18°16’E. 03 R
WCD 34 ~—:115.12.59 34°11-2’S./18°20°2’E. 27 R
WCD 56 21.9.60 32°4:6'S./18°18’E. 18 Saple
WCD 81 15.9.49 34°5/S./18°21’E. II N)
REMARKS ON CLASSIFICATION
In the taxonomy of the Plumulariidae three problems are of recurrent
interest, namely:
(1) The validity of fixed or movable nematothecae as a basis for subdivision;
(2) the presence or absence of cauline hydrothecae as a useful diagnostic
character, and
(3) the value of the method of branching in the limitation and grouping of
genera.
In 1883, Allman, using the type of nematotheca as a basis, divided the
Plumulariidae into two large sections: the Eleutheroplea and the Statoplea.
However, the presence of certain genera such as Heteroplon Allman 1883 and
Heterotheca Stechow 1921, with two kinds of nematothecae, made this simple
division impracticable. In 1923b Stechow made use of 4 sub-families, but also
used as his basis the type of nematotheca. Stechow’s subfamilies included:
Kirchenpaueriinae Stechow 1921. Nematothecae reduced.
2. Plumulariinae Kiihn 1913 (= Eleutheroplea). Nematothecae 2-chambered
and movable.
Acladiinae Stechow 1923b. Intermediate forms.
4. Aglaopheniinae Stechow 1911 (= Statoplea). Nematothecae 1-chambered
and immovable.
eS
The Kirchenpaueriinae and the Aglaopheniinae appear to be valid sub-
families representing natural assemblies of genera (in fact the latter is raised to
family rank by some systematists), but the other subfamilies need further
consideration.
The Acladiinae was created for intermediate forms with fixed mesial nema-
tothecae and fixed or movable lateral nematothecae. It was an unsatisfactory
group, as Stechow himself realized, for he abandoned it in 1925, adding its
genera to the Plumulariinae. It has in fact been increasingly recognized that
268 ANNALS OF THE SOUTH AFRICAN MUSEUM
in this group the type of nematotheca does not form a good basis for family,
or even generic, diagnosis. ‘This was pointed out as early as 1913 by Billard
and in 1915 by Bale, and most systematists have dropped the genera Heteroplon,
Heterotheca and Antennellopsis Jiderholm 1896 for this reason.
There is, however, a group of genera which are linked by the presence of
cauline hydrothecae, including Thecocaulus Bale 1915, Halopteris Allman 1877,
Gattya Allman 1886 and Schizotricha Allman 1883, genera which Stechow dis-
tributed between the Plumulariinae and Acladiinae. Bale was the first to
realize the importance of cauline hydrothecae as a diagnostic character
when he established the genus Thecocaulus in 1915 for ‘forms in which hydro-
thecae are borne on the rachis as well as on the pinnae’. Such genera possess
stem internodes which are essentially similar to those of the hydrocladia, and
thus differ from the ‘Plumularia’ type where the stem internodes lack hydro-
thecae and thus differ from those of the hydrocladia. This difference is a funda-
mental one exhibited in the earliest stages of growth, influencing, as it does,
the nature of the first upright stem. It is a sound character for generic diagnosis,
and under no circumstances should species with cauline hydrothecae be
included in the same genus as species without, as has been done by Billard
(1913) (i.e. in the genus Plumularia). It is also considered to be a sound character
on which to group genera, and a new subfamily, the Halopterinae, is proposed
for forms with cauline hydrothecae. (Stechow’s Acladiinae cannot be retained
since Acladia is a synonym for Halopteris.) The Plumulariinae can be retained
in Stechow’s sense, though excluding certain genera dealt with below.
Stechow implies that the Kirchenpauerinae is the most primitive subfamily
of the Plumulariidae because of the poorly developed nematothecae, but it is
maintained here that the arrangement of hydrothecae and hydranths is of
far greater fundamental significance. It is rational to assume that a primitive
form would be one in which a functional feeding unit (i.e. a portion of stem
and a hydranth) is established as rapidly as possible in early development and
before budding commences, as in fact occurs in the sympodial growth of
primitive Calyptoblast families such as the ‘Campanulinidae’ and Haleciidae.
This arrangement is possible in the Halopterinae but not in the other Plumu-
lariidae where a stem must develop hydrocladia before the formation of
hydranths is possible. The Halopterinae is thus considered to contain the
primitive stock of the Plumulariidae.
Genera of Halopterinae
The genus Halopteris was established by Allman in 1877 for H. carinata,
which is the type species. Totton in 1930 showed that Plumularia catherina
Johnston 1833 (the type species for Thecocaulus) is congeneric, and thus sinks
Thecocaulus in the synonymy of Halopteris.
Although Allman’s conception of Halopteris (and also Stechow’s) was the
presence of fixed nematothecae, its main diagnostic characters are now con-
sidered to be the presence of cauline hydrothecae, a pinnate stem with
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 269
unbranched hydrocladia and an untoothed thecal margin. The genus as thus
envisaged includes the following species:
Plumularia alternata Nutting 1900, P. buski Bale 1884, P. campanula Busk 1852,
Halopteris carinata Allman 1877, Plumularia catherina Johnston 1833, P. concava
Billard 1911, Halopteris constricta Totton 1930, Plumularia crassa Billard 1911,
P. diaphana Heller 1868, Halopteris gemellipara n. sp., Aglaophenia glutinosa
Lamouroux 1816, Thecocaulus heterogona Bale 1924, Plumularia liechtensternit
Marktanner-Turneretscher 1890, Thecocaulus minutus Trebilcock 1928, Plumu-
laria polymorpha Billard 1913, P. sulcata Lamarck 1816, P. tuba Kirchenpauer
1876, Heteroplon valdiviae Stechow 1923, Plumularia zygocladia Bale 1914b,
and possibly also Plumularia diaphragmatica Billard 1911 and P. jedani Billard
1Q13.
In addition to the characters mentioned above, these species usually have
other features in common. Thus, the colonies as a whole tend to resemble
Aglaophenia in their general appearance rather than Plumularia, with robust
stems and close-set hydrocladia and hydrothecae. Most of the species possess
hinge-joints near the base of the stem or its branches. There is a tendency
towards the production of opposite hydrocladia—in H. gemellipara, H. catherina
and H. zygocladia the hydrocladia are opposite throughout all or most of the
length, and in practically all other species opposite hydrocladia are borne by
the first or second thecate internodes. Internodal and intrathecal septa are
typically absent. Each hydrocladial internode typically has one median inferior
nematotheca and one or two pairs of laterals, and sometimes one or more
supracalycines. The cauline internodes have a similar arrangement, though they
may possess supernumerary supracalycines as well. The nematothecae may be
I- or 2-chambered, movable or immovable. The gonothecae, where known, are
dimorphic, typically with the female seated on the stem and the male on the
hydrocladia of the same colony, and they bear nematothecae, at least in the
female.
It is not suggested that all these characters are essential for the inclusion
of a species in Halopteris (in fact there are exceptions as regards most of the
characters listed), but that the common possession of most of them links
together a group of species which are closely related and can be considered as
constituting a single genus. Few hard and fast rules can be laid down, as many
of the species appear to be in a state of flux and show surprising variability
of structure, even within a single colony. It has already been demonstrated
how the segmentation of the hydrocladia, and the structure and arrangement
of nematothecae, may vary in H. glutinosa (see Millard, 1958, and also Broch,
1933).
It is proposed to include in the same subfamily (i.e. Halopterinae) the
genera Antennella, Schizotricha, and Monostaechas, which are clearly related to
Halopteris, and differ only in their method of branching.
270 ANNALS OF THE SOUTH AFRICAN MUSEUM
The genus Antennella Allman 1877 was created for A. gracilis Allman 1877,
a form with simple stems similar to hydrocladia, and this is the type species.
Schizotricha Allman 1883 was created for forms with pinnate stems and branch-
ing hydrocladia, and the type species is S. unifurcata Allman 1883 according to
Totton 1930. Finally Monostaechas Allman 1877 was created for M. dichotoma
Allman 1877 (= M. quadridens (McCrady) 1857) in which the hydrocladia
branch sympodially from the posterior surface, the resulting ‘stem’ being formed
from the proximal parts of successive hydrocladia (cf. also Billard, 1913, p. 14).
The type of branching, however, forms no inflexible demarcation between
the four genera so far mentioned. It is known, for instance, that many species
of Halopteris may exist in a simple form, and some species of Antennella may
produce pinnate stems. Thus Broch (1933) has shown that Antennella secundaria
can produce pinnate stems although the simple form is the more common,
and that Halopteris diaphana can exist in the simple form although the pinnate
form is more common. H. campanula, H. catherina and H. constricta can all exist
in the simple form (Bale, 1913, Hincks, 1868, and Millard, 1957, respectively).
Accordingly Broch has sunk Thecocaulus (= Halopteris) in the synonymy of
Antennella, and this practice has been followed by most Continental syste-
matists. But it has been shown recently (Millard, 1958) that Monostaechas
natalensis, which has a completely different method of branching, can also
exist in the simple form, and Billard 1913 has shown that A. secundaria can
branch in the Monostaechas way. To follow Broch’s system we should then sink
Monostaechas also in the synonymy of Antennella. It appears to the present author
that within this group of species the simple form is the most primitive type and
that from it have led two main evolutionary lines, one leading to Halopteris
with the tendency towards pinnate branching, and one to Monostaechas with
branches arising from the posterior surface of the stem, and that since these
genera represent grades of evolution it is wise to keep them separate. The
placing of intermediate types will always be a matter of opinion, though in
general a species should be named according to its dominant form, thus
Antennella secundaria but Halopteris diaphana.
Passing on to forms with branching hydrocladia, these surely represent a
further stage of the evolutionary line leading through Halopteris, and if we are
to retain Halopteris as separate from Antennella then we must also retain
Schizotricha, in spite of intermediate forms.
Another genus to be included in the Halopterinae is Gattya Allman 1886
(= Paragattya Warren 1908, see p. 281), which was created for Gattya humilis
Allman 1886, a form with a toothed thecal margin. Whether a toothed hydro-
theca is a sufficient character to distinguish a genus is a matter of opinion.
In most of their characters the species are closely related to the Antennella-
Halopteris line (for instance, G. humilis possesses hinge-joints and occasionally
branching hydrocladia, and the first thecate internode of the stem bears a pair
of opposite hydrocladia). Possibly they represent a deviation from the Halopteris
stock, which is already foreshadowed in H. carinata and H. valdiviae which have
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 271
a sinuated hydrothecal margin. At any rate the character is a definite one and
useful in distinguishing species, and the genus may be allowed to stand.
The genera NEMERTESIA and ANTENNOPSIS
The genus Antennopsis was founded by Allman in 1877 for A. hippuris,
the features distinguishing it from Nemertesia Lamouroux 1812 being the
‘scattered disposition’ of the hydrocladia, and the uncanaliculated coenosarc.
The former character has generally been accepted as worthless, and the latter
character has also been discredited by many authors (see Bedot, 1921). The
SCHIZOTRICKHA
SA
GATTYA
HALOPTERIS
MONOSTAECHAS =e
Lat:
ANTENNELLA
Fic. 1. Diagram showing possible evolutionary trends in the Halopterinae.
25 7/2 ANNALS OF THE SOUTH AFRICAN MUSEUM
author considers that the type species of Antennopsis (A. hippurts Allman 1877) is
congeneric with the type species of Nemertesia (Sertularia antennina Linnaeus
1758) and consequently sinks Antennopsis in the synonymy of Nemertesia.
There are, however, certain species (Antennularia fascicularis Allman 1883
and Antennopsis scotiae Ritchie 1907) included by Bedot in Antennopsis which
differ markedly from other species of the composite genus Nemertesia and from
the type species of this genus. In Nemertesia the hydrocladia arise from an unfas-
cicled stem, or from the central tube of a fascicled stem, either irregularly or in
verticils. In the two species mentioned the hydrocladia arise, quite irregularly,
from the peripheral tubes of the stem, and there is no recognizable central or
main tube. It is just as though the hydrorhizal tubes of a colony of Antennella
had become bound together to form a fascicled stem, each branching or
separating from the others independently to give hydrocladia. This character
is, I feel, of generic value, and I propose a new genus Corhiza for the reception
of the two species mentioned and two new species described below, with
Antennopsis scotiae Ritchie 1907 as the type species (i.e. Corhiza scotiae (Ritchie)
1907). |
These four species are also linked by certain less important characters
which do not occur in Nemertesia and which emphasize the cleavage between -
the two genera, namely the large hydrothecae, the absence of a mamelon on
the base of the hydrocladium, the position of the lateral nematothecae on long
supporting processes at the sides of the hydrotheca, and the gonothecae which
are borne on the hydrocladia and themselves bear nematothecae.
Further, the author feels that the sum total of the characters of the genus
Corhiza indicates a closer relationship with the Halopterinae than with the
Plumulariinae, and proposes to place it in the former subfamily. The stem of
course bears no cauline hydrothecae, but if the stem is considered as a con-
glomeration of hydrorhizal tubes this becomes understandable.
The genus Corhiza appears to be closely related to Antennella and Mono-
staechas. In A. africana the hydrorhizal tubes often rise up from the substratum
in bunches, giving rise to clumps of simple stems (i.e. hydrocladia). A further
development of this process would produce a colony of the Corhiza type. A
fascicled stem similar to that of Corhiza is found in some members of Mono-
staechas (M. natalensis Millard 1958 and WM. faurei Millard 1958), and MM.
natalensis appears to occupy an intermediate position between the two genera,
where the branching of the hydrocladia has not developed to the same extent
as in other species of the genus.
The evolutionary relationships between the genera of Halopterinae as
visualized by the author are illustrated in figure 1.
The subfamilies of the Plumulariidae are summarized as follows.
Subfamily 1. Halopterinae nov. subfam. Hydrocladia arising independently
from hydrorhiza, from a pinnate stem, or from the superficial tubes of a
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 273
compounded stem. Pinnate stems bearing cauline hydrothecae. Paired
lateral nematothecae present, of variable structure.
South African genera: Antennella Allman 1877
Corhiza nov. gen.
Gattya Allman 1886
Halopteris Allman 1877
Monostaechas Allman 1877
Schizotricha Allman 1883
Subfamily 2. Kirchenpauerinae Stechow 1921. Stem simple or pinnate,
the latter without cauline hydrothecae. No paired lateral nematothecae.
Median nematothecae poorly developed, seldom 2-chambered, often
rudimentary and sometimes represented by naked sarcostyles only.
South African genera: Airchenpaueria Jickeli 1883
Oswaldella Stechow 1919
Pycnotheca Stechow 1919
Subfamily 3. Plumulariinae Kihn 1913. Stem upright, giving off hydro-
cladia pinnately or in verticils. No cauline hydrothecae. Paired lateral
or supracalycine nematothecae present. | Nematothecae 2-chambered,
movable, free from hydrotheca.
South African genera: WNemertesza Lamouroux 1812
Plumularia Lamarck 1816
Subfamily 4. Aglaopheniinae Stechow 1911. Stem upright and pinnate.
No cauline hydrothecae. Hydrocladia with close-set hydrothecae generally
facing towards the anterior surface. Paired lateral nematothecae normally
present. Nematothecae 1-chambered and immovable, usually adnate to
hydrotheca.
South African genera: Aglaophenia Lamouroux 1812
Cladocarpus Allman 1874
Halicornaria Allman 1874
Lytocarpus Allman 1883
Thecocarpus Nutting 1900
List OF SPECIES
Subfamily 1 Halopterinae
Antennella africana Broch 1914 Halopteris gemellipara n. sp.
Antennella secundaria (Gmelin) 1788-1793 Halopteris glutinosa (Lamx.) 1816
Corhiza bellicosa n. sp. Halopteris tuba (Kirch.) 1876
Corhiza pannosa n. sp. Halopteris valdiviae (Stechow) 1923
Corhiza scotiae (Ritchie) 1907 Monostaechas natalensis Millard 1958
Gattya humilis Allman 1886 Schizotricha simplex Warren 1914
Halopteris constricta Totton 1930
274 ANNALS OF THE SOUTH AFRICAN MUSEUM
Subfamily 2 Kirchenpaueriinae
Kirchenpaueria pinnata (Linn.) 1758 Oswaldella nova (Jarvis) 1922
Kirchenpaueria triangulata (Totton) 1930
Subfamily 3 Plumulariinae
Nemertesta ciliata Bale 1914b Plumularia pulchella Bale 1882
Nemertesia cymodocea (Busk) 1851 Plumularia setacea (Ell. & Sol.) 1755
Nemertesia ramosa Lamx. 1816 Plumularia spinulosa Bale 1882
Plumularia filicaulis Kirch. 1876 Plumularia wasini Jarvis 1922
Plumularia lagenifera Allman 1886
Subfamily 4 Aglaopheniinae
Aglaophenia late-carinata Allman 1877 Halicornaria exserta n. sp.
Aglaophenia pluma (Linn.) 1758 Halicornaria ferlusi Billard 1901
Cladocarpus leloupi n. nom. Lytocarpus filamentosus (Lam.) 1816
Cladocarpus lignosus (Kirch.) 1872 Thecocarpus flexuosus (Lamx.) 1816
Halicornaria arcuata (Lamx.) 1816 Thecocarpus formosus (Busk) 1851
SYSTEMATIC SECTION
Subfamily Halopterinae
Antennella africana Broch, 1914
Antennella africana. Millard, 1957, p. 226.
Records. West coast: TB 1A, 17B. TRA 86N, 156E. LAM 30], 35D. SB 196L.
CP 650F. WCD 56K.
South coast: AFR 994L. TRA 38N. MB 8R (pp), 12X, 47S, 52E, 58D,
59E, 60K. LIZ 7V. SCD 5J, 22B, 36U, 81Q, 96G, 152Y, 153E, 175F. SAMH
241, 279; 293, 309.
Remarks. This material includes one specimen (SAMH 309) in which all the
dimensions are about double the normal.
The species is common on the west and south coasts of South Africa (from
Luderitz Bay on the west (Broch, 1914) to approximately 29°E. on the south)
and occurs in the intertidal region down to a depth of 120 m.
Antennella secundaria (Gmelin), 1788-1793
Antennella secundaria. Millard, 1958, p. 199.
Records. South coast: MB 8R (pp), 47S. SAMH 153, 165, 191, 212, 277, 360.
LIZ 16G, 27K.
Description. Colonies reaching a height of 1:2 cm. Stems and hydrothecae
sturdy and of greater dimensions than the material from Inhaca (Millard, 1958).
Athecate internodes comparatively short and only rarely exceeding the neigh-
bouring thecate internodes in length, bearing one or two nematothecae.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 275
Measurements (mm.)
* Inhaca Mossel Bay
Hydrocladium, length of basal part 0°99-2°75 1°73-3-27
thecate internode, length 0-26-0°34. 0°32-0°43 |
athecate internode, length 0:26-0:38 0°22-0°45
athecate internode, maximum diameter 0-04—0°05 0:08—0-12
Hydrotheca, length abcauline O-15-0'19 0°20—0:225
diameter at margin O-15-0'19 0°27—0°32
‘Gonotheca, male, length (without Dedicely 0°32—0°40
maximum diameter te 0°22—0°29
Gonotheca, female, length Guithout Sedicel) 0-61—0-66
maximum, diameter 0°41—-0°48
Remarks. In poor material it is difficult to distinguish A. secundaria from A.
africana. ‘The number of nematothecae on the intermediate internodes and the
proportionate length of the latter are no criterion, as both are variable charac-
ters. The only sure criterion is the presence of supplementary lateral nematothe-
cae in A. africana and of a median supracalycine nematotheca in A. secundaria,
and these are often lost. There appears to be no difference in the structure of
the gonophores of the two species.
In South Africa both species are known only in the simple form. A. secun-
daria occasionally gives off one or two branches, but only from the back of the
basal part of the stem, as mentioned also by Billard (1913, p. 8). True pinnate
branching never occurs.
A. secundaria occurs on the south and east coasts of South Africa (from
approximately 19°E. on the south coast (Stechow, 1925) to Inhaca on the east
coast), in the littoral region, and down to a depth of 100 m. (Stechow, 1925).
Genus CorHIZA nov. gen.
Type species: Antennopsis scotiae Ritchie, 1907
Diagnosis. Plumulariidae with an upright and fascicled stem composed of a
number of interwoven and intercommunicating tubes of equal diameter and
importance. Hydrocladia arising from the component tubes in a completely
irregular fashion, not rebranching. Hydrotheca with untoothed margin.
Nematothecae (so far as is known) all bithalamic and movable, including
paired laterals. Gonothecae borne on the hydrocladia, bearing nematothecae.
Corhiza bellicosa nov. sp.
(Fig. 2 A—E)
Types and records. Holotype: SCD 84S (South African Museum registered num-
ber = SAMH 364). Paratype: SCD 153J. (Both from south coast.)
* Including material described by Millard, 1958.
276 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description of holotype. Several short and thick fascicled stems reaching a maxi-
mum height of 5:0 cm., a few of which branch irregularly, giving rise to numer-
ous hydrocladia in an irregular fashion from the component tubes in the distal
region. The individual tubes of the stem are connected to one another by
communicating pores and branch irregularly. In the distal region the tubes
usually separate from one another in clumps (the branches), and each one
terminates in a hydrocladium. The hydrocladia of a clump or branch tend to
face in the same direction, giving a ‘Monostaechas’ appearance to the
colony.
Hydrocladia unbranched, with proximal region athecate, divided by an
irregular number of transverse internodes and bearing a double series of
nematothecae. Remainder divided into thecate internodes by oblique
nodes.
Hydrotheca borne on a projection near base of internode, with depth
exceeding diameter, swollen in basal half, then narrowed slightly and expanding
again to margin. Margin smooth, at right angles to internode. Mostof adcauline
wall adnate, but a small distal portion free.
Nematothecae generally 13 to an internode, including: 1 median inferior
borne close to proximal end of internode and not reaching base of hydrotheca;
1 pair lateral inferior borne on base of internode slightly above level of median
inferior and just reaching base of hydrotheca; 1 pair lateral borne on long
processes arising at the side of the hydrotheca and not quite reaching thecal
margin; 1 pair lateral borne on outer surfaces of bases of above processes;
1 pair lateral borne at the side of the hydrotheca about midway between above-
mentioned process and thecal margin, overtopping hydrotheca; generally
2 pairs supracalycine borne above thecal margin (1 pair close to the midline
and another pair further to the sides and at a slightly higher level).
There is, however, much variation in the number and position of the
supracalycine nematothecae, and as few as 2 and as many as 9 have been
observed, generally in pairs but often quite irregularly arranged.
All nematothecae bithalamic and movable. Nematotheca large, expanding
towards margin, with sides of distal chamber cut right away to level of dia-
phragm resulting in a margin of 2 flaps—a high abcauline (i.e. outer) one,
and a lower adcauline (i.e. inner) one.
Gonophores absent.
>
A-E. Corhiza bellicosa n. sp.
A. Two stems from the holotype, SCD 84S. There has probably been some distortion of
the hydrocladia during preservation.
B-D. Two lateral views and an anterior view of portions of hydrocladia, from the holotype.
B is taken from near the proximal end, and C from near the distal end, of hydrocladia.
E. Different views of the lateral nematothecae, the top one from SCD 153J, the other two
from the holotype.
F. Monostaechas natalensis Millard. Different views of the nematothecae from SCD 153D (large
form).
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 7]
Fic. 2. Corhiza bellicosa and Monostaechas natalensis.
278 ANNALS OF THE SOUTH AFRICAN MUSEUM
Measurements (mm.)
SCD 848 SCD 153]
(Holotype)
Internode length (on posterior surface) 0°4.9—0°94. 0°55-0°80
diameter (above hydrotheca) 0-13-0'18 0-14—0°20
Hydrotheca, height .. 0°22—0'27 0°23-0°31
diameter at margin O°1'7—0°22 0°16—-0:24,
diameter /height 0:67-0:85 0-70—-0'84.
Nematotheca, height 0:09—-0°15 O-11-0-16
Remarks. This species is very close to Monostaechas faure: Millard 1958, differing
from it in the unbranched and longer hydrocladia, in the presence of 2 extra
pairs of nematothecae to each internode, in the greater measurements of all
individual parts, and in the shape of the hydrotheca where the proportion of
diameter/height is greater. ‘The shape of the nematotheca is very characteristic
of the species.
Corhiza pannosa nov. sp.
(Fig. 3 A-B, D-G)
Types and records. Holotype: SCD 5H (South African Museum registered num-
ber = SAMH 362). Other records: SCD 61H, 67M, 186%. SAMH 292.
WCD 12G.
Description of holotype. A bushy colony reaching a height of 5:3 cm. Hydrorhiza
a matted reticulum. Stem strongly fascicled, branching irregularly, and giving
off hydrocladia from its component tubes in a completely irregular fashion.
Tubes of stem not canaliculated.
Hydrocladia borne on short apophyses of stem, reaching a maximum
length of 2 cm. and bearing up to 20 hydrothecae. Basal 4 or 5 mm. devoid of
hydrothecae, but divided by transverse nodes into 2 or more rather long inter-
nodes bearing a double series of nematothecae. Remainder of hydrocladium
divided into thecate internodes by oblique nodes, though in some parts the
distal end of an internode may be cut off by a transverse node.
Hydrotheca large, with almost parallel sides, standing away from hydro-
cladium at an angle of about 50°, with about half adcauline wall adnate to
internode.
Nematothecae generally 7 to each internode, namely: 1 medial inferior;
2 pairs laterals (1 seated on a process arising next to the hydrotheca and not
>
A-B, D-G. Corhiza pannosa n. sp. A, the whole colony, and B, the lateral and supplementary
lateral nematothecae in adcauline view, from the holotype SCD 5H. D-G, portions of
hydrocladia from different samples to show variation. (D, the holotype. E, WCD 12G.
F, SAMH 292. G, SCD 61H). F shows a gonotheca.
C. Corhiza scotiae (Ritchie). The lateral and supplementary lateral nematothecae from TRA
35B as a comparison with B.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 279
Fic. 3. Corhiza spp.
280 ANNALS OF THE SOUTH AFRICAN MUSEUM
quite reaching margin, and 1 seated in the angle between this process and the
internode) ; 2 (or occasionally 1 or 3) median superior.
Nematothecae all bithalamic and movable. Lateral nematotheca large
and of characteristic shape: terminal chamber cup-shaped and deep, and with
nearly parallel sides which turn in slightly at margin, with a distinct, deep
excavation in adcauline (i.e. mesial) wall. Median nematotheca smaller, with
very short adcauline wall.
Gonophores absent.
Measurements (mm.)
SCD 61H SAMH 292 SCD186X WCD12G SCD5H
(holotype)
Internode length eons distal
end when cut off). . . 0°62-0:88 0:62-0:68 0:77-0:95 0:80-1:09 0°70—-1°17
diameter, maximum .. O'0Q-O°12 O*1I—O°13 009-0714 O°13-O0°17 O°18—0:23
Hydrotheca, height abcauline . 0°26-0°38 0:34-0:38 0°35-0°46 0:30-0:36 0-34-0°46
height adcauline, free pans . O°16—0°245 0:20-0:24. 0°20—-0°:22 0°14-0°20 0-16—-0°25
diameter at margin .. 0°32-0°34 0°345-0°39 0°31-0°43 0:36-0:39 0°37-0-41
free part/abcauline height | . 0°58-0°71 0°54-0°66 0°50-0°57 0°42-0°58 0°41-0°555
diameter/abcauline height . 0°84-1°31 O°QQ-1'I5 0°72—-1:06 1:00-1:23 0°87—1-09
Nematotheca height, lateral . O1O-O'II OtII-O'I14 O'II-O°13 OTI-O'16 O°15—O°19
supplementary lateral . 0°03-0°08 0:045-0:06 0:07-0:09 0°07—0°12 0*09-0°15
Gonotheca, height 0:80 0:96-1:07
breadth 0°33 0:36-0:59
Remarks. In this species the form of the colony resembles closely that of Corhiza
fascicularis (Allman) 1883 from Tristan da Cunha, the type material of which
apparently no longer exists. It can be distinguished from other species of the
genus in particular by its distinctive lateral nematothecae, which resemble
most those found in Halopteris glutinosa (cf. Millard, 1958, figure 10). ‘The
details of the hydrocladia resemble those of Monostaechas natalensis Millard 1958,
from which it is distinguished by its larger hydrothecae which are less adnate,
and by the nematothecae.
Remarks on ‘other records’. 1 have included with this species one stem fragment
bearing hydrocladia (SCD 67M) exactly like the holotype, and 4 samples of
solitary hydrocladia which appear to represent simple forms of the same species.
The latter agree in all important features and particularly in the shape of the
lateral nematothecae which are identical. A few minor differences do, however,
occur. Thus the transverse nodes separating off intermediate internodes are
better defined and almost invariably present, and in 3 of the samples (SCD 61H
and 186X, and SAMH 292) the processes supporting the lateral nematothecae
arise at a slightly higher level on the internode. Also in these simple forms most
of the individual parts are smaller, particularly the lateral and supplementary
lateral nematothecae. Yet the range of variation in the measurements of the
different samples, and the overlapping which occurs, make it impossible to
distinguish two species on these grounds.
Two of the simple forms (SAMH 292 and SCD 186X) bear male gonothe-
cae. Each gonotheca arises immediately below a hydrotheca, is roughly
pear-shaped but curved towards the hydrocladium, and bears a round
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 281
operculum at the wide distal end and two large nematothecae near the
base.
These simple forms can always be distinguished from South African species
of Antennella by the characteristic shape of the lateral nematothecae.
Corhiza scotiae (Ritchie), 1907
(Fig. 3 C)
Antennopsis scotiae Ritchie, 1907, p. 543, pl. 3 (fig. 3). Ritchie, 1909, p. go, fig. 8. Millard, 1957,
p- 235. .
Records. West coast: WCD 7L, 12F, 18M.
South coast: TRA 35B, 38C, 92D. SCD 26B, 67L, 84R, 153G, 175D,
191Q, 195K. SAMH 307, 323, 331, 345:
Remarks. ‘This endemic species has a somewhat restricted distribution round the
South African coast, being found between the lines of 17° and 28°E. longitude
and in a depth-range of 18-120 m. It has never been found in the littoral area,
and is typical of the Agulhas Bank.
Gattya humilis Allman, 1886
Gattya humilis Allman, 1886, p. 156, pl. 24 (figs. 5-7).
Paragattya intermedia Warren, 1908, p. 323, pl. 47 (fig. 27), fig. 16. Mi€ullard, 1957, p. 230.
Millard, 1958, p. 209.
Records. West coast: PP 4Z. LAM 27G, 43H. SB 168D (recorded by Day, 1959,
as Paragattya intermedia). CP 646D, 650E.
South coast: MB 15D, 52G, 60M, 88K. LIZ 27G. SCD 36R, 84Q.
SAMH 275.
Remarks. Examination of Allman’s type material of Gattya humilis in the British
Museum of Natural History (reg. no. 86.2.19.41), and a comparison with
Warren’s cotypes of Paragattya intermedia, show that these species are synony-
mous. The material figured by Allman (pl. 24, fig. 6) includes 3 young stems,
but in the same colony branching stems are also present similar to those
described by Warren. Allman failed to observe the median superior nemato-
theca. Allman’s material is labelled ‘Port Elizabeth (?), Cape of Good Hope’,
and is almost certainly South African in origin. The genus Paragattya Warren
1908 thus becomes sunk in the synonymy of Gattya Allman 1886, with Gattya
humilis Allman 1886 as the type species.
The following points may be added to the published descriptions. Firstly
the first two oblique nodes of the stem are extra well defined as hinge-joints,
and between them arise not one, but a pair, of opposite hydrocladia. Secondly
the hydrocladia occasionally rebranch, the branch being of similar construction
to the hydrocladium and bearing 1-3 hydrothecae. Both these features are
also characteristic of the closely related genus Halopteris.
282 ANNALS OF THE SOUTH AFRICAN MUSEUM
Halopteris constricta ‘Totton, 1930
(Hing G)),
Halopteris constricta. Millard, 1957, p. 227, fig. 14 A.
Records. South coast: LIZ 27J. MB 55H.
(Additional from False Bay: FAL 326D.)
Description. Two small colonies including simple and pinnate forms. Structure
as previously described, except that in some stems the first (and sometimes
also the second) segment after the hinge-joint gives rise to a pair of hydrocladia
arising one on each side of the cauline hydrotheca.
Female gonothecae (not previously described) borne on thecate inter-
nodes of stem, each on a very short pedicel which arises on one side of the
internodal projection which supports the hydrotheca. Gonotheca strongly
curved towards the stem, with a wide distal aperture facing the stem, bearing
2 bithalamic nematothecae on basal part. (These gonothecae were not present
in the samples from the south coast, but on new material from False Bay.)
Halopteris gemellipara nov. sp.
(Fig. 4. A-F)
Types and records. Holotype: SAMH 308. Other records: (NAD 22B). SCD 36P,
153L.
Description. Stem unfascicled, reaching a maximum height of 5:5 cm., bearing
pinnately arranged hydrocladia. Basal part without hydrothecae or hydro-
cladia, divided by irregular transverse nodes, and bearing a median series of
nematothecae on the anterior surface. No hinge-joints. Remainder divided
into rather long internodes by oblique nodes; each internode bearing a hydro-
theca near the base on the anterior surface, and a pair of opposite hydrocladia
arising one on each side of the hydrotheca.
Hydrocladium borne on a short apophysis of the stem and separated from
it by a transverse node, which, however, is often not clearly visible. Remaining
nodes oblique. First internode athecate, bearing one median nematotheca near
distal end. Remaining internodes thecate, but with the distal end sometimes
cut off by a transverse node situated just above the attachment of the hydro-
theca. Hydrothecae not exactly on the distal surface of the hydrocladium, but
shifted slightly towards the anterior surface.
>
A-F. Halopteris gemellipara n. sp. A and B, two hydrocladial internodes from SAMH 308 (the
holotype) and NAD 22B respectively. C, the whole stem; D, a cauline internode in anterior
view; E, a lateral nematotheca; F, the median inferior nematotheca, all from the holotype.
G. Halopteris constricta Totton. A portion of the stem in lateral view showing female gonothecae,
from FAL 326 D.
H, J. Halopteris valdiviae (Stechow). H, part of a hydrocladium with 2 male gonothecae; J, a
female gonotheca. Both from SAMH 349.
K. Halopteris glutinosa (Lamx.), female gonotheca, from SAMH 278.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA
ee
A—.B
Dl! : 5S mm.
Fic. 4. Halopteris spp.
283
284 ANNALS OF THE SOUTH AFRICAN MUSEUM
Hydrotheca cylindrical, expanding towards margin; adnate for about 2
height, then free; abcauline height exceeding (or rarely equal to or less than)
width at margin; margin forming an angle of 55° or more with internode.
Each thecate internode bearing 3 nematothecae, one median inferior
seated well below the base of the hydrotheca, and 2 laterals seated on short
processes one on each side of the hydrotheca and not reaching margin. A
single median supracalycine nematotheca sometimes present on hydrocladial
internodes, and 2-4 on cauline internodes. _
Median nematotheca short and broad, probably immovable, 2-chambered ;
with small basal chamber and larger distal chamber separated by transverse
septum; with distal chamber cut away on adcauline side and margin facing
towards internode. Abcauline wall strongly convex in supracalycine nemato-
thecae, the nematotheca of the first hydrocladial internode, and the median
inferior nematothecae on the stem; only slightly convex in the median inferior
nematothecae on the hydrocladia.
Lateral nematotheca minute, 1-chambered, immovable, with adcauline
wall cut away and margin facing towards internode.
Gonothecae absent.
Measurements (mm.)
SAMH 308 NAD 22B- SCD 36P
Stem, internode length .. se .. 1°24-1°69 1:18-1:96 1-00—1°63
diameter ais he be .. 0°28-0'41 0°21-0°32 =0°27—0°365
Hydrocladium, internode length, first
athecate Ags be: i .. 0°34-0°55 0°43-0°48 0°39-0°53
normal thecate ms Hes .. 0°55-0°71 0:66-0-80 0°53-0-70
Hydrotheca, height abcauline .. .. 0°29-0:38 0°40-0°50 024-050
height, adcauline, free part .. O'10-0°16 = =0°13-0:18 —0:08-0°15,
diameter at margin .. yl .. 0°29-0°33 0:24-0:29 0°26-0°295
free part/abcauline height .. .. 0°29-0°46 0:30-0:36 0:24-0°47
diameter/abcauline height .. .. 0°82-1:02 0°51-0°60 0°54-1°17
Nematotheca, height, median cauline .. 0:095-0:18 0:13-0:16 0-12—0°16
hydrocladial, median inferior .. O'0Q-O'II O'115—-0°135 0:09—-0°13
lateral Sif a we .. 0°035-0:05 0:05-0:06 0:03-0:06
Remarks. In the measurements and the figure a sample from Natal (NAD 22B)
is included, as it illustrates the variation possible in the species. It differs from
the holotype in several particulars, mainly in the proportions of the hydro-
thecae, which are deeper and narrower. As a result the diameter at the margin
is little more than half the abcauline height, whereas in the holotype the
diameter is practically equal to the height. Further, the axis of the hydrotheca
is more nearly parallel to the hydrocladium so that the margin forms an angle
of about 70° with it as against about 55° in the holotype (fig. 4B). Intermediate
internodes are also more common on the hydrocladia, especially towards their
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 285
distal ends. Sample SCD 36P contains both forms, though on separate stems.
Since the points mentioned are all differences of proportion and degree, and
all structural details are exactly the same, and bearing in mind the tendency
towards variation in Halopteris, I hesitate to separate these two forms as separate
species, at least until more material is available.
One stem of the holotype shows a rather unusual abnormality and bears
a pair of hydrocladia on the posterior surface in addition to those on the anterior
surface.
H. gemellipara resembles H. catherina (Johnston) 1833 and H. zygocladia
Bale 1914a in the possession of opposite hydrocladia, but differs from both in
the monothalamic lateral nematothecae. In this respect, and in other structural
details, it is very similar to Antennellopsis integerrima Jaderholm 18096, from Japan,
differing from it in the longer free part of the hydrotheca and the oblique
hydrocladial internodes.
FHlalopteris glutinosa (Lamx.), 1816
(Fig. 4 K)
Heteroplon pluma Allman, 1883, p. 32, pl. 8 (figs. 1-3).
Plumularia glutinosa. Billard, 1910, p. 36, fig. 16. Stechow, 1925, p. 502.
Plumularia alternata (Nutting), 1900. Jarvis, 1922 (pp), p. 345, pl. 25 (fig. 16).
Halopteris glutinosa. Millard, 1958, p. 200, fig. 10 A—-D.
Records. South coast: LIZ 16E, 27H. MB 8S, 15E, 24V, 47M, 52D, 6o0J.
SCD 36Q, 84P, 153K, 169U. SAMH 103, 197, 225, 276, 278, 298. TRA 92G.
Description. Colonies altogether more robust than those on the Natal coast
(Millard, 1958), most of them reaching a height of 2-3 cm., but three large
colonies reaching 10, 12 and 18 cm. (TRA 92G, SCD 84P, and SAMH 278
respectively), thus resembling more closely the colonies described by Allman
and Stechow. In the larger colonies the lower regions of the stem have no
visible nodes, the hydrocladia are usually lost, and the hydrothecae tend to be
eroded and shallow. All measurements exceed those of the east coast material,
though the proportions and appearance of the hydrothecae are similar. Hydro-
cladia may bear up to 15 hydrothecae. Lateral nematothecae with terminal
chamber deeply cut away on adcauline (1.e. inner) side.
Stem internodes may bear as many as 3 extra pairs of nematothecae
above the margin of the hydrotheca, and occasionally the 1-chambered
supracalycine nematotheca is paired in the lower part of the hydrocladium.
Female gonothecae (not previously described) borne on the stem, each
on a short pedicel of one segment arising next to a hydrotheca. Very large,
flattened, bluntly rounded at distal end, bearing 2-5 2-chambered nematothe-
cae near base, reaching a length of 3-74 mm. and a diameter of 1°70 mm.
Remarks. The material described by Jarvis (1922) from East Africa, and assigned
by her to Plumularia alternata Nutting, is present in the British Museum of
Natural History. On examination the samples from Zanzibar and Wasin
286 ANNALS OF THE SOUTH AFRICAN MUSEUM
/
both proved to be mixed, with some plumes (those illustrated in pl. 25, fig. 16,
of Jarvis) resembling exactly Halopteris glutinosa and attributable to this species.
In other plumes, however, the stems were more slender and geniculate and
possessed athecate intermediate internodes, and the hydrothecae had sinuated
margins, each with one distinct anterior lobe and 2 lateral lobes. It was
undoubtedly these plumes which led Jarvis to assign the material to P. alternata,
although the stems do not possess the delicacy of Nutting’s material. They
possibly represent a separate variety of H. glutinosa or a separate species
altogether.
Halopteris tuba (Kirch.), 1876
(Fig. 5)
Plumularia tuba Kirchenpauer, 1876, p. 44, pl. 1 (fig. 2), pl. 4 (fig. 2).
Acladia africana Marktanner-Turneretscher, 1890, p. 261, pl. 5 (figs. 11, 11a).
Heteroplon jaederholmi Stechow, 1912, p. 366, figs. F, G.
Plumularia (Heteroplon) africana. Stechow, 1925, p. 500, figs. 44, 45.
Thecocaulus tuba. Leloup, 1939, p. 12, fig. 8.
Records. South coast: SCD 5M, 52L, 79F, 82K, 84N, 1175, 153F, 175A, 186W,
219U, 234Q, 239A. SAMH 194, 238, 249. AFR 865.0.G.
Description. Colonies of unfascicled, unbranched stems reaching a maximum
height of 22 cm.
Basal part of stem without hydrothecae or hydrocladia, with a few irregular
transverse nodes and a double row of movable nematothecae on the anterior
face. ‘This part usually terminated by 1 or 2 oblique hinge-joints. Where 2
hinge-joints occur the segment between them bears a single median hydrotheca
on the anterior face and a pair of hydrocladia arising one on each side of it,
also 1 medial inferior nematotheca and 4-8 movable nematothecae rather
irregularly arranged above the hydrotheca.
Remainder of stem bearing hydrothecae and hydrocladia on anterior
surface, unsegmented except for extreme distal end, where regular, slightly
oblique nodes may occur. The first hydrotheca is median, and from the same
level arises a pair of hydrocladia, one on each side of it. Thereafter the hydro-
thecae are displaced alternately to the right and left of the mid-line and the
hydrocladia arise singly, alternately from the right and left sides of the hydro-
thecae (from the right side of a hydrotheca displaced to the right and vice versa).
The hydrothecae and hydrocladial origins thus form 2 longitudinal rows on the
anterior surface of the stem. Very rarely a pair of hydrocladia arises from the
same level. Corresponding to each hydrotheca are 5 cauline nematothecae:
1 inferior, 2 lateral, and 2 minute supracalycine. Of these the inferior nema-
totheca is not situated immediately below its hydrotheca, but is displaced
towards the opposite side of the stem.
Hydrocladia borne on short apophyses of the stem arising next to the cau-
line hydrothecae (the hydrothecae thus appear to be seated on the apophyses),
divided into internodes by distinct oblique nodes. Apophysis without nema-
tothecae. First internode short, athecate, with 1 median movable nematotheca.
a
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 287
o:-S5mm. as Se i eae
Fic. 5. Halopteris tuba (Kirch.).
A. The lower part of the stem in anterior view, showing hinge-joints and arrangement of
hydrocladia and cauline nematothecae. From SAMH 238.
B, C. Cauline hydrothecae from SCD 5M to show shape of hydrotheca and supracalycine
nematothecae—B from upper region, and C from lower region, of stem. Aperture for
lateral nematotheca shown on left of diagram.
D. The first 2 hydrocladial internodes, from SCD 5M.
E-G. Stages in the development of the female gonophore, from SCD 5M.
288 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remaining internodes (up to 12 in number) each with 1 hydrotheca, 1 median
inferior nematotheca, 2 lateral nematothecae, and 1 supracalycine naked
sarcophore. Old colonies may have weak internodal septa at the proximal
and distal ends of the internodes.
Hydrotheca shallow, with width exceeding (or, rarely, equal to) depth,
and with untoothed margin. Hydrocladial hydrotheca adnate to internode to
within a short distance of margin, then free. Cauline hydrotheca with abcauline
wall greatly thickened, and margin eroded, particularly on adcauline and
lateral sides, so that no free part remains. Hydropore very small, circular,
close to abcauline side.
Nematophores of 4 types:
(i) Lateral nematothecae: slender, elongated, and movable. Bithalamic,
with funnel-shaped distal chamber which is not cut away on adcauline
(i.e. inner) side. Those on the hydrocladia arise from short processes of the
internodes next to the hydrotheca and usually reach well above the margin.
Those on the stem are much larger, and are not seated on processes, but
emerge directly from oval openings in the perisarc. They are not con-
tiguous to the hydrotheca but separated from it by a distance equal to
about half the thecal width. The median nematotheca of the first hydro-
cladial internode is very similar, but smaller and with a wider base—
it appears to be intermediate between this and the following type.
(ii) Median nematothecae: short, hook-shaped and immovable. With a
transverse septum and thus bithalamic. Distal chamber with adcauline
wall completely cut away. Situated well below hydrotheca and not reach-
ing to base.
(iii) Supracalycine nematothecae: immovable, hook-shaped, minute, mono-
thalamic. Situated immediately above the eroded adcauline wall of the
cauline hydrothecae.
(iv) Naked sarcophores: situated in the angle behind the free part of the
adcauline wall of the hydrocladial hydrothecae. It is possible that a
nematotheca may be present and fused with the thecal wall, but it is not
visible as a separate entity.
Male and female gonothecae (not previously described) borne on the
same colony and often on the same hydrocladium with the male more distal
than the female.
Female gonotheca arising from a small triangular pedicel attached to
the first thecate internode of hydrocladium just below the hydrotheca and
slightly to one side. Very large, flattened, rounded distally and tapering
proximally. Bearing 3 nematothecae near base on surface facing hydrocladium,
these being similar in size and appearance to the laterals of the stem. Containing
a single embryo between the arms of a bifurcating blastostyle.
Male gonothecae borne on internodes of hydrocladia other than the first.
Smaller than female, elongated-oval, with abcauline side more convex than
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA
289
adcauline, with distal aperture. (Appearance similar to male gonothecae of
H. glutinosa (see Millard, 1958, fig. 10).)
Measurements (mm.)
SCD 5M SCD 79F
Hydrocladial internode, length on posterior sur-
face, athecate 0-2I—0°30 0°27—0°30
EhWecate .. 0-50—-0'64 0°43-0°53
Hydrotheca, cauline, heen abeauliive 0-19—0°22
width at mouth ee 0°22-0°27
hydrocladial, height abeaulinies 0-12—0°16 0:16-0:20
width at mouth 0°20-0°27 0-20-0'25
Nematotheca, cauline, lateral 0-26—0°30
median inferior O-12—0°13
supracalycine 0:04-0:06
hydrocladial, lateral 0-16—-0-21 O-1I-O°17
median inferior (first) O-II-O'15 O-1I-O'14
median inferior (normal) 0:08-0°15 0:065-0°12
gonothecal : 0°24—0°30 0-19—-0°26
Gonotheca, female, length Patou pedicel 2°95-3°36 2°83-3-11
maximum diameter I*51—-1'Q4. 1°40—-1°54.
Gonotheca, male, length 0°36-0°44.
maximum diameter O°15-0°25
Remarks. 1 am satisfied that Marktanner-Turneretscher’s species Acladia
africana (redescribed by Stechow, 1925) is a synonym for Kirchenpauer’s
species Plumularia tuba, and have accordingly combined the two. Kirchenpauer’s
material was re-examined by Leloup in 1939, who testifies to the excellence of
Kirchenpauer’s description and adds a more accurate diagram of a hydrocladial
internode.
The only features distinguishing the two species are (i) the segmentation
of the stem which is said to be absent in A. africana and present in P. tuba, and
(ii) the hydrotheca, which is said to be completely adnate in A. africana and
only partially so in P. tuba.
Kirchenpauer’s type material has unfortunately been lost, but I have
seen a prepared slide of Stechow’s material of A. africana, and have found that
the upper part of the stem is in fact segmented, and that the hydrocladial
hydrothecae do have a short region which is not adnate.
There is an apparent variability in the length of the lateral nematothecae
on the hydrocladia, which sometimes just overtop the thecal margin (as in
Leloup’s figure) and sometimes greatly overreach it (as in Stechow’s figure,
1925). This appears to be a result of erosion of the thecal margin in the latter
case (in deep hydrothecae the margin is sinuated and in shallow hydrothecae
the margin is even and worn). Both types may occur on the same stem, but the
lateral nematothecae always reach approximately the level of the next node.
290 ANNALS OF THE SOUTH AFRICAN MUSEUM
A full description of the species has been given, since none of the published
descriptions is quite complete. The arrangement of the cauline nematothecae
was found to be exactly similar in Stechow’s material.
It is evident that, as in all typical species of Halopteris, the arrangement of
the cauline segments is essentially the same as that of the hydrocladia, differing
only in the tendency for nodes to disappear, the lateral displacement of the
hydrothecae and the nature of the supracalycine nematophores.
This endemic South African species occurs on the Agulhas Bank east of
19°E., extending round the coast to Natal waters. It is found in depths of
II-130 m.
Halopteris valdiviae (Stechow), 1923
(Fig. 4 H, J)
Thecocaulus(?) valdiviae. Stechow, 1925, p. 495, figs. 42-43.
Halopteris valdiviae. Millard, 1957, p. 228, fig. 14 B.
Records. South coast: SAMH 1096, 248, 324, 332, 346, 349. SCD 79G, 169W.
West coast: AFR 736X. WCD 7M.
Description. ‘Two magnificent tree-like colonies reaching 35 and 37 cm. in
height (SAMH 248 and 349), and a number of smaller colonies and detached
portions. A few details may be added to previous descriptions.
Hydrorhiza forming a dense mat 5 cm. wide by 3 cm. high. Stem strongly
fascicled, reaching 1 cm. in diameter near base, branching irregularly. Stem
and branches giving off hydrocladia-bearing pinnae from the deeper tubes in
an irregular fashion and from all surfaces.
Pinnae unfascicled, or very lightly fascicled near base, with basal part
terminated by 2 oblique hinge-joints, but occasionally as many as 4, possibly
due to regeneration. Segment between hinge-joints bearing 1 median cauline
hydrotheca, and 1 pair of opposite hydrocladia on anterior surface. First
segment beyond hinge-joints similar, with median hydrotheca and 1 pair of
hydrocladia. Remaining segments each bearing 1 cauline hydrotheca, and 1
hydrocladium arising alternately from the right and left of the hydrotheca.
Hydrocladia as described by Stechow (1925) and Millard (1957), bearing up
to 9 hydrothecae.
Nematothecae as previously described, except that it is now possible to
distinguish a pair of minute, monothalamic supracalycine nematothecae behind
the free part of the adcauline wall of each cauline hydrotheca.
Gonothecae (not previously described) dimorphic, male and female on
same colony, each borne on a small triangular pedicel of 1 segment which
arises immediately below a hydrotheca and slightly to one side.
Male gonothecae borne on thecate internodes of hydrocladia, pear-shaped
but slightly asymmetrical with the greatest curvature on side away from hydro-
cladium, with flattened distal end, bearing 2 bithalamic nematothecae near
base on side facing hydrocladium.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 291
Female gonothecae borne on internodes of pinnae, very much larger than
male, flattened in a plane at right angles to pinna, expanding distally to a wide
aperture, bearing 3 or 4 bithalamic nematothecae on side facing pinna.
Measurements (mm.)
Gonotheca, male, length ne Ae ae aH, a .. 0°58-0°83
maximum diameter a = ne a .. 0°20—0°:39
Gonotheca, female, length .. ae Ss Sy af 2. 1°24-1°55
maximum diameter ae e. bs mS eis -2) W:O1—1-22
The measurements of the trophosome agree well with those given by
Millard (1957), except that the range is in some cases greater.
Remarks. It is now obvious that the material described by Millard (1957)
consisted of 2 detached pinnae bearing hydrocladia.
This endemic South African species occurs on the west and south coasts
as far east as 29°E., and in depths of 33-201 m. It is remarkably similar to
H. tuba in the arrangement and structure of its hydrocladia, differing from it
in its fascicled stem, and proportions of hydrotheca in which the depth is
almost invariably greater than the width.
Monostaechas natalensis Millard, 1958
(Fig. 2 F)
Monostaechas natalensis Millard, 1958, p. 206, fig. 12.
Records. South coast: GPR 46J. SCD 153D.
Description. The first colony is of small size and similar to the holotype in its
appearance and measurements, but the second colony (SCD 153D) is altogether
of larger proportions. The fascicled stems branch irregularly and reach a
maximum height of 7-6 cm., and individual hydrocladia reach a length of
3-4 cm. All the minute measurements are larger. The proportions of the parts
are the same except that a greater part of the adcauline thecal wall is free
from the internode. Towards the ends of the hydrocladia there is a tendency
for the distal parts of the internodes to be cut off by transverse nodes. No
branching hydrocladia were observed—any branching which may occur is
obscured in the fasciculation of the stem. The general appearance of the
colony is very similar to that of Corhiza pannosa n. sp., but it is clearly distin-
guished from it by the nature of the lateral nematothecae. With the larger
size the structure of the latter could be more exactly ascertained. Each nema-
totheca has the sides deeply cut away to the level of the diaphragm, with a
high, bonnet-shaped abcauline (i.e. outer) wall and practically no adcauline
(i.e. inner) wall at all. It resembles that of Corhiza bellicosa n. sp., except that
the adcauline wall is more reduced. In the holotype the structure of the nema-
tothecae was not easy to determine because of the small size and the opacity
of the contents, but a re-examination of the material with a phase-contrast
292 ANNALS OF THE SOUTH AFRICAN MUSEUM
microscope shows that they are of a similar nature and that the diagram given
by Millard in 1958 (fig. 12E) was not strictly accurate. The measurements of
the larger colony are given for comparison with the holotype.
Measurements (mm.) SCD 153D
Internode length (on posterior surface) .. aoe ue .. O'7I-1°05
diameter (above hydrotheca) - ie HY .. OTI4-0°19
Hydrotheca, height .. vs ie Br oe ae ». 0°34-0°45
height of free part a: a: ae si x .. | OFFG=0794
free part/total length .. ae a oS if ..> 0°44-0°65
diameter at margin... oe a oe ne ~. 0°27-0:34
Nematotheca, length ae in es a: wr .. 0°09—0°17
Remarks. In view of the larger measurements sample SCD 153D might well
be included in a separate variety, but one is reluctant to establish a new variety
on size alone when little is known about the variation within the species.
Schizotricha simplex Warren, 1914
Schizotricha simplex Warren, 1914, p. 83, pl. 6 (figs. 1-19), figs. 1-4.
Records. S 65B (recorded by Stephenson, Stephenson and du Toit, 1937).
L 430. CPR OA.
Description. Colonies reaching a maximum height of 1-4 cm., agreeing exactly
with the beautiful description by Warren, bearing male and female gonophores.
In the larger stems practically all the ‘pinnae’ bear 3 thecate internodes and
sometimes even 4, and many of the ‘pinnules’ bear 2.
Subfamily Kirchenpaueriinae
Kirchenpaueria pinnata (Linn.), 1758
Kirchenpaueria pinnata. Millard, 1957, p. 233. Millard, 1959b, p. 252.
Records. West coast: CPR 44A. SB 141U. LB 126, 190Q, 257P, 367X, 378E,
392R, 472H, 513J (recorded by Day, 1959). WCD 18N.
South coast: S 65A (recorded by Stephenson, Stephenson and du Toit —
(1937) as K. unilateralis). KNY 710, 165A (recorded by Day, Millard and
Harrison, 1952). E 83. LIZ 7X, 27F. E54, 317, 3360. (CPR 7Daoms
Kirchenpaueria triangulata (Totton), 1930
(Fig. 6 E-J)
Plumularia triangulata Totton, 1930, p. 225, fig. 61.
Records. South coast: SCD 175G.
Description. Colony growing on the stems of Nemertesia ciliata Bale, and including
both simple and pinnate forms.
2
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 293
Pinnate stem reaching a length of 1 cm., and consisting of long internodes
separated by transverse nodes which may be indistinct in the lower region.
Each internode giving rise to a hydrocladium-bearing apophysis from its
distal end. Each internode bearing a well-developed mamelon on the upper
surface of the apophysis, and 2-4 cauline nematothecae. The latter including 1
on proximal end immediately above apophysis of last internode, rarely 1 near
distal end below the apophysis, and 1 (or rarely 2) on upper surface of apophy-
sis distal to the mamelon.
Hydrocladia alternate, the two rows in one plane or shifted very slightly
to one side of the stem. Hydrocladium normally homomerous and consisting
of long thecate internodes separated by transverse nodes, but intermediate
athecate internodes commonly occur and in quite an irregular fashion. Thus
there may be 1-4 or none at all before the first thecate internode, and 1 or 2
of irregular length between successive thecate internodes.
Hydrotheca borne on the proximal half of the internode, which thus has
a long distal region; flaring slightly to margin, completely adnate or with a
very short free portion, with diameter exceeding depth.
Nematothecae: 1 median inferior and one median supracalycine; with
convex abcauline wall and practically straight adcauline wall and oblique
margin; movable; monothalamic, or with a very thin septum close to base.
Simple stem reaching a length of about 4 mm., similar to hydrocladium,
borne on a long apophysis of hydrorhiza. Measurements of parts slightly
smaller than corresponding ones of pinnate stem.
Gonothecae (male) borne by pinnate stem on hydrocladial apophyses to
one side of mamelon, elongated, tapering below, truncated above, triangular
in section. Pedicel of 2 segments.
Measurements (ram.)
Pinnate Simple
stem stem
Stem internode, length 0°48—-0°75
diameter bs iy Re: 0:07-0:16
Hydrocladium, thecate internode, length 0:66-0:85 0°59-0°81
diameter at distal end 0:055-0°075 0:05-0:07
Hydrotheca, height abcauline 0:07—0:095 0:07-0:08
height of free part 0:00-0:01 0:00-0:01
diameter at mouth O-10-0'12 0:095-O0'1I
Nematotheca, length 0:05-0:08 0:05-0:07
Gonotheca, male, length 1°74-2°46
maximum. diameter 0:80-0:85
Remarks. The author agrees with Bedot (1923) that the main diagnostic charac-
ter separating Azrchenpaueria from Plumularia should be the absence of paired
nematothecae. Although the nematothecae in KAirchenpaueria as a whole are
characteristically reduced, any attempt to separate the two genera on the
294. ANNALS OF THE SOUTH AFRICAN MUSEUM
W
O'lmm,.
0°'3 mm.
Fic. 6. Oswaldella nova (Jarvis) and Kirchenpaueria triangulata (Totton).
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 295
structure of the nematothecae alone will fail because of the presence of inter-
mediate forms. Thus a group of species with but one superior nematotheca
(including P. halecioides Alder 1859, P. bonneviae Billard 1906, P. inermis Nutting
1900, and others) has been assigned by Bedot (1923) to Kirchenpaueria, by
Stechow (1923b) to a new genus Ventromma, and by most other authors to
Plumularia. To this category also belong Plumularia irregularis Millard 1958
and P. triangulata Totton. Most of these species have monothalamic superior
nematothecae, and are thus intermediate between a typical Plumularia and a
typical Kirchenpaueria. In K. triangulata, however, a poorly developed septum
is often visible in the nematotheca, and in K. irregularis it is quite distinct, so
that in these genera the nematothecae are strictly speaking bithalamic and
verging on the Plumularia type.
The South African material of K. triangulata agrees closely with Totton’s
description and resembles it in the epizootic habit of growth. The material
shows much evidence of regeneration after injury, and this probably accounts
for the presence of the many intermediate internodes, which were not men-
tioned by Totton. In one case the end of a stem has regenerated as a
hydrocladium.
The species is closely related to K. irregularis (Millard) 1958, differing
from it in the longer distal part to the hydrocladial internode and in the
arrangement of the cauline nematothecae.
It is the first record of the species from South Africa.
Oswaldella nova (Jarvis), 1922
(Fig. 6 A-D)
Plumularia nova Jarvis, 1922, p. 347, pl. 26 (fig. 20).
Kirchenpaueria adhaerens Millard, 1958, p. 203, fig. 13 F, G.
Records. South coast: MB 47Z. SCD 84T. TRA o2H.
West coast: WCD 7N.
(Additional from Natal: NAD 1P.)
Description. A number of samples of epizootic colonies growing on other hydroids
(Halopteris glutinosa (Lamx.)), H. valdiviae (Stechow) and Gattya humilis Allman).
coed
A-D. Oswaldella nova Jarvis.
A. Part of colony on back of stem of Halopteris glutinosa, from TRA 92H (hydrocladia of
host cut off short).
B-D. Portions of hydrocladia from different colonies to show variation in size and shape:
B from TRA 92H, C from NAD 1P, and D from MB 47Z.
E-J. Kirchenpaueria triangulata (Totton), from SCD 175G.
E. A colony including a pinnate stem bearing gonothecae, and several simple stems,
removed from host.
F. A portion of a pinnate stem showing arrangement of nematothecae.
G, H. Hydrocladial internodes from a simple stem and a pinnate stem respectively.
J. A single nematotheca with diaphragm.
296 ANNALS OF THE SOUTH AFRICAN MUSEUM
Hydrocladia arising direct from adherent stem, usually unbranched, but in
one sample (TRA 92H) often forked. The division occurs immediately above
the origin of a hydrotheca; both limbs are of equal thickness, and their hydro-
thecae do not face one another, but lie in the same plane as those of the original
hydrocladium. Unbranched hydrocladia bearing up to 14 hydrothecae.
Hydrotheca variable in size and shape, with diameter generally exceeding,
but occasionally equal to, height; usually free from internode for a small
proportion of its length, but occasionally completely adnate.
Gonothecae absent, but scars probably of old ones present below hydro-
thecae and slightly to one side.
Measurements (mm.)
Type
MB 47Z NAD 1P. TRAo2H — material
Internode length (posterior
surface) if .. 0°29-0°31 0°35-0°41 0:42-0:57 0:°39-0°48
diameter near distal
end Me .. 0:06-0:07 0:065-0:08 0:08-0:11 0:04-0:05
Hydrotheca, length abcau-
late see a .. 0°05-0:08 0:075-0'10 0:08-0'12 0:06—-0:075
length of free part .. 0:01-0:03 0:00-0:02 O-0I-0:04, *O-01 ~
free part/abcauline
length. .. O°125-0°43 0:00-0:25 O°10-0°40 *0-125
diameter at mouth .. 0:°105-0°135 O*10-O‘II O:II-O°13 0:07-0:08
Remarks. Examination of the type material of Jarvis’s species Plumularia nova
in the British Museum of Natural History (slide no. 23.2.15.283 from Zanzibar)
has established the fact that this species is synonymous with Kzirchenpaueria
adhaerens Millard. The correct measurements are quoted above and replace
the incorrectly calculated ones given in 1958. The type material is scanty and
damaged, and no hydrocladia bear more than 2 hydrothecae. The median
inferior nematothecae are missing, though their orifices remain. In the only
hydrotheca lying in a suitable position there is a short free part to the adcauline
wall. The remains of one very damaged gonotheca are present, but it is impos-
sible to determine the shape.
The discovery of forked hydrocladia in the new material necessitates the
transfer of the species from Kirchenpaueria to Oswaldella, to which genus all its
structural details indicate a closer affinity. Further knowledge of the growth-
habits may indicate that it is merely an epizootic form of O. bifurca (Hartlaub)
or O. antarctica (Jaderholm).
* One measurement only.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 297
Subfamily Plumulariinae
Nemertesia ciliata Bale, 1914b
(Fig. 7 E-G)
Nemertesia ciliata Bale, 1914b, p. 170, pl. 36 (fig. 1). Bale, 1915, p. 298. Briggs, 1914, p. 307,
pl. 10 (fig. 3). Jaderholm, 1919, p. 23.
Records. West coast: TRA 47N. AFR 729H.
South coast: TRA 92F. AFR 945L. SCD 5N, 175C. SAMH 3209.
Description. Several colonies, the largest reaching a height of 24:5 cm. Stem
and larger branches fascicled, smaller branches unfascicled, branching always
in one plane. Hydrocladia given off in whorls of 2, 3, or 4, but irregular in
lower parts of branches; consisting of alternate thecate and athecate internodes,
of which the first is always thecate.
Appearance and structure as described by Bale and Briggs, with the
following minor points of difference.
The branching is not so luxuriant as in the specimen figured by Briggs
and the branches are more widely spaced, with intervals in the neighbourhood
of 5-10 mm. between them. In branches with two hydrocladia to a whorl it
sometimes happens that the alternate pairs are not set at right angles to each
other but compressed in one plane, forming angles of 40-60/120-140° with
each other, and giving a flattened appearance to the branch. The same ten-
dency may be found in stems with 3 hydrocladia to a whorl, as in NV. japonica
(Stechow) 1909, but it is not constant within a colony. The hydrocladia are
often longer than in the type material, reaching 5-7 mm., and the internodes
are in general somewhat longer, but the measurements of the hydrothecae
are very similar.
Gonothecae as described by Bale.
Measurements (mm.)
Hydrocladium, internode length, thecate oe Af .. 0°29-0°51
athecate ate in he ae = a .. 0*14-0:28
diameter at distal end of internode .. ot ao .. 0°04—0°095
Hydrotheca, height abcauline a Ne ot a .. 0°02—0:07
diameter at margin .. ie bs M nie .. 0°06-0:085
Nematotheca, lateral, height a of ae ia -. 0°05-0-10
Gonotheca, length .. ie $4 i a reaching 0°49
maximum diameter .. oy - oh reaching 0°24
Remarks. This is the first record of the species from South Africa. So far it is
known only from Tasmania and Japan.
I have not separated a var. cruciata as was done by Bale (1915), although
both this and the normal form are present, as the tendency to produce different
numbers of hydrocladia to a whorl almost certainly depends on the growth
and age of the colony as has been shown for other species of the genus.
298 ANNALS OF THE SOUTH AFRICAN MUSEUM
rl F i 50 5mm.
Fic. 7. Nemertesia spp., portions of hydrocladia showing variation.
A-D. Nemertesia ramosa Lamx.
A. The typical appearance from TRA 125K. No superior medial nematotheca.
B. An example with smaller hydrothecae, longer internodes, and superior medial nema-
tothecae, from NAD oK.
C, D. Hydrocladial internodes from old and young regions respectively of the same colony,
from TRA 1348S. C with internodal septa and thickened perisarc.
E-G. Nemertesia ciliata Bale.
E. The typical appearance, from TRA 47N.
F, G. Hydrocladial internodes from old and young regions respectively of the same colony,
from SAMH 329.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 299
Nemertesia cymodocea (Busk), 1851
Nemertesia cymodocea. Billard, 1910, p. 39. Millard, 1957, p. 234. Millard, 1961, p. 207.
Records. West coast: SB 178B. TB 3, 4, 5, 18, 21A. TRA 156A.
South coast: MB 81N. AFR 865R. TRA 92K. SCD 26C, 36T, 52M,
153H, 175E, 219V. SAMH 148, 240, 347.
Description. The length of the hydrocladial internodes is variable, sometimes
being almost double that quoted by Billard (1910, p. 40), though the size of
the hydrotheca remains fairly constant.
Observations suggest that during the growth of a colony the stems first
produce alternate hydrocladia, one to an internode, and that later the decussate
arrangement develops at the tip of the stem, and still later the hexastichous
arrangement. Thus young colonies bear only alternate hydrocladia and stems
may reach a length of 10 cm. (and bear gonophores) before the decussate
arrangement starts. The hexastichous arrangement is characteristic of the
distal ends of old colonies, though by this time the proximal hydrocladia may
be lost or buried by the peripheral tubes of the stem. In young colonies, too,
the peripheral tubes bear a thick coating of nematothecae, which appear to
fall off with age.
Nemertesia ramosa Lamouroux, 1816
(Fig. 7 A—-D)
Antennularia ramosa, var. plumularioides Billard, 1906, p. 215.
Nemertesia ramosa. Millard, 1957, p. 235. Millard, 1961, p. 206.
Nemertesia ramosa, var. plumularioides (Billard), 1906. Vervoort, 1959, p. 293, figs. 46 b, 47.
Records. West coast: WCD 12D, 32H.
South Coast: TRA 92E. SAMH 333. SCD 5P, 96A, 153N, 175B.
(Natal: NAD 9K.)
(Additional from False Bay: TRA 1348.)
Description. A number of colonies of varying size, the largest reaching 28-2 cm.
Considerable variation in appearance and structure occurs, much of which is
probably due to growth and ageing of the colony.
The number of hydrocladia produced in a verticil appears to increase
with age. In young colonies the hydrocladia have an alternate arrangement,
I or 2 pairs to an internode. As growth proceeds the young shoots produce
2, 3, 4, or even 5 hydrocladia to a verticil, members of one verticil alternating
with those of the next resulting in as many as 10 longitudinal rows. In old
colonies only the verticillate arrangement is visible since the lower hydrocladia
with alternate arrangement tend to fall off and their apophyses to be buried
by the peripheral tubes of the stem.
The thickness of the perisarc also appears to increase with age and is
possibly influenced by other factors as well. In young colonies the perisarc is
thin, with the hydrocladia white and the stem a pale yellow or straw colour.
300 ANNALS OF THE SOUTH AFRICAN MUSEUM
In old colonies the thickened perisarc gives to the stem and bases of the hydro-
cladia a dark brown colour. Associated with the thickening of the perisarc
the nodes of the stem become indistinguishable and the hydrocladial apophyses
very thick and long, athecate internodes become more numerous in the
hydrocladia, and in the apophyses and basal parts of the hydrocladia appear
internodal septa (up to 12 to an internode). The appearance of these regions
is thus very different from that in the growing tips of the stems and the distal
ends of the hydrocladia where the structure is similar to that usually described
for the species.
Measurements (mm.)
Hydrocladium, internode length, thecate oo ae .. 0°34-0°86
diameter at distal end of internode .. as ais .. O°0G=O-%2
Hydrotheca, height abcauline ve “ ae ce .. 0:06-0°14
diameter at margin... = es xe at .. 0°07-O°14,
Nematotheca, lateral, height ay ae oe a .. O:07-O785
Gonotheca, length .. x an Be fy reaching 0:94
maximum diameter .. ae of f, reaching 0°43
Remarks. Billard (1906) has created a var. plumularioides for forms of NV. ramosa
with alternate hydrocladia, but if, as seems practically certain, the arrangement
of hydrocladia is a matter of age, the retention of this variety is unnecessary.
Vervoort (1959) has described specimens with alternate hydrocladia reaching
lengths of up to 25 cm. from tropical west Africa, but he himself mentions that
the hydrocladia may be arranged in whorls of 3 in the upper parts of the larger
colonies.
Plumularia filicaulis Kirchenpauer, 1876
Plumularia filicaulis. Millard, 1958, p. 209, fig. 13 D, E.
Records. West coast: WCD 34D, 81K.
Plumularia lagenifera Allman, 1886
Plumularia lagenifera. Millard, 1957, p. 230.
Records. West coast: TRA 156D. LAM 32], 7L, 9V, 13H, 26Q, 46N, 590A, 62T.
PP 1W. LB511E (recorded by Day, 1959). A382. WCD 34C, 81H. CP 650C.
South coast: KNY 300 (recorded by Day, Millard and Harrison (1952)
as P. setacea).
Plumularia pulchella Bale, 1882
Plumularia pulchella. Millard, 1957, p. 232.
Records. South coast: SCD 37S. SAMH 290, 302. MB 8Q, 47Q, 52F, 55G,
60L. KNY 71B, 165B (recorded by Day, Millard and Harrison, 1952).
LIZ 27C.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 301
Description. Colonies reaching a maximum height of 1-1 cm. (KNY 71B),
but usually between 0-3 and 0-7 cm.
Gonothecae ovate, truncated distally, with wide distal aperture, 5-54
times height of hydrotheca.
Plumularia setacea (Ellis and Solander), 1755
Plumularia setacea. Millard, 1957, p. 232. Millard, 1958, p. 212. Millard, 1959b, p. 252.
Records. West coast: WCD 18R, 2o0F, 34E. ‘TRA 156B. B 114D. LAM arT,
27Z, 35C, 45X. SB 150A. LB 152, 367W (recorded by Day, 1959). TB 2,
21H. CP 640C.
South coast: SCD 5K, 52S, 169V. TRA 159A. SAMH 108, 224. CPR 1,
46H. MB 817, 24X, 47R, 55F. E 170 (recorded by Stephenson, Stephenson
and Bright, 1938). LIZ 17H, 27E. L 57, 145 (recorded by Eyre, Broekhuysen
and Crichton, 1938).
Plumularia spinulosa Bale, 1882
Plumularia spinulosa. Bale, 1884, p. 139, pl. 12 (figs. 11-12). Bale, 1888, p. 783, pl. 19 (figs. 11-13).
Warren, 1908, p. 320.
Monotheca spinulosa var. obtusa Stechow, 1923b, p. 224.
Plumularia spinulosa var. obtusa. Millard, 1957, p. 232.
Plumularia spinulosa var. typica Stechow, 1923b. Millard, 1958, p. 212.
Records. South coast: SCD 29G, 36W, 75G, 85A. SAMH 199, 291, 303.
MB 47P, 58C. LIZ 13D, 27D. L 111 (reported by Eyre, Broekhuysen and
Crichton, 1938).
Remarks. I have no longer attempted to distinguish var. obtusa from var. typica
as I have found that the length of the terminal spine of the hydrocladium varies
not only in different samples but also within the same colony. The origin of
the hydrocladium, whether from the middle or distal end of a stem internode,
is also a variable feature.
Plumularia wasim Jarvis, 1922
(Fig. 8)
Plumularia wasini Jarvis, 1922, p. 349, pl. 26 (fig. 23), fig. 1.
Records. South coast: CPR 46G.
Description. Stem reaching a height of 1-6 cm., unfascicled, unbranched, divided
into internodes of irregular length by transverse nodes; basal part short and
without hydrocladia; no hinge-joints; distal part giving rise to alternate hydro-
cladia, 1-3 per internode, the 2 rows in the same plane. Intervals between
hydrocladia not regular, some being short and others long. Stem bearing 2
rows of cauline nematothecae (2-14 per internode), sometimes on the lateral
sides but often shifted on to the anterior or even the posterior surface.
Hydrocladia borne on very short apophyses of stem, divided into alternate
302 ANNALS OF THE SOUTH AFRICAN MUSEUM
athecate and thecate internodes terminated by oblique and straight nodes
respectively. Apophysis without nematothecae or mamelon.
Hydrotheca large, with even or slightly sinuated margin; free from inter-
node for more than half height; with diameter always exceeding depth. Hydro-
thecae not strictly on upper surface of hydrocladium but shifted slightly onto
anterior surface.
Nematothecae: 1 (or occasionally 2) on each athecate internode; 4 on
each thecate internode, including 1 median inferior not quite reaching base of
hydrotheca, 1 pair of laterals seated on processes next to hydrotheca and not
quite reaching its margin, and 1 superior in angle behind adcauline thecal wall.
All bithalamic and movable except superior which is monothalamic.
Gonothecae borne on thecate hydrocladial internodes, each on a short
pedicel of 2 segments, and each bearing 2 or 3 large nematothecae on basal
region. ‘wo types present: the larger (probably female, but empty) on first
thecate internode of hydrocladium only, pear-shaped, with large operculum
at distal end; the smaller (male) usually on thecate internodes other than the
first, more rounded and without operculum.
Measurements (mm.)
Stem internode, length se te x oA 7 .. 0°34-0°99
diameter bye ate af: ck ie ie .. O°145—-0°24 .
Hydrocladium, internode length (on posterior surface), thecate 0-16—-0-21
athecate Me ie ie se N m .. OFL7—-OF
Hydrotheca, height abcauline ne Bre Avs sf .. O0°12—0°185
height adcauline, free part .. igh on sts .. O°10-0°13
free part/abcauline height .. ae is ane .. 0°65-1-:00
diameter at margin: .. a - ai aig . 227 (O02 O=aean
Nematotheca, lateral a re Mi. 4g =e .. 0:06—0:085,
inferior median. . *: os a at 2A .. 0°07—0:09
superior By yd ag an. ots a .. 0°03-0'045
cauline .. 2 aK. se ae ~ ae .. .0*0Q—O'II
Gonotheca, female ?, length without pedicel he reaching 0-69
maximum diameter a Re a reaching 0-61
male, length without pedicel, He se reaching 0°35
maximum diameter ie oe wy reaching 0-24.
Remarks. This material was compared with a slide of Jarvis’s type in the British
Museum of Natural History. In the latter the athecate internodes normally.
bear 2 nematothecae, but the number is variable and some have only one.
In the present material 1 is the normal condition and 2 occur only rarely. In
the type material the superior median nematotheca is 1-chambered and not
2-chambered as figured by Jarvis (1922, pl. 26, fig. 23 A).
Certain minor variations sometimes occur. Thus in CPR 46G one hydro-
cladium bears a branch arising from its first thecate internode and bearing
one hydrotheca.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 303
(ee ee ee ty foddtesthalinal
Fic. 8. Plumularia wasini Jarvis.
A. A portion of stem in anterior view showing male (small) and female (large) gonothecae
and origins of hydrocladia. Gonothecae removed on right side.
B. A portion of a hydrocladium.
Both from CPR 46G.
This speciés is in some ways reminiscent of Halopteris, as instance the
large hydrothecae which are not completely adnate, the shape and arrange-
ment of the gonothecae, and the presence of nematothecae on them.
The species is so far only known from Wasin, East Africa. It is a new
record from South Africa.
Subfamily Aglaopheniinae
Aglaophenia late-carinata Allman, 1877
Aglaophenia late-carinata. Millard, 1958, p. 213, fig. 14. Vervoort, 1959, p. 309, fig. 54.
Records. South coast: CPR 46F.
304. ANNALS OF THE SOUTH AFRICAN MUSEUM
Aglaophenia pluma (Linn.) 1758, subsp. dichotoma (M. Sars), 1857
eee eee var. dichotoma. Millard, 1957, p. 239, fig. 15 B, CG. Vervoort, 1959, p. 308,
g. 52 b.
Records. West coast: LAM 16W, 35B, 51A. WCD 32G.
South coast: MB 5E, 34F, 37C, 42A, 79E, 81M, 84A. KNY 22F, 28C,
70B, 71D, 127A (recorded by Day, Millard and Harrison (1952) as A. ?dicho-
toma). ‘TRA 35Q, 36A, 38A, 42A, 56W, 99M. SAMH 149, 163, 195, 215, 322,
330, 344. SCD 20C, 26A, 29D; 690A, 94C, 103D, 104F,, 108], 120) eiesee
152X, 153B, 191P, 195J, 199U, 230B.
Aglaophenia pluma (Linn.) 1758, subsp. parvula Bale, 1882
Aglaophenia pluma, var. parvula. Millard, 1957, p. 239, fig. 15 D-F. Millard, 1958, p. 215. Ver-
voort, 1959, p. 307, figs. 52 a, 53 b.
Records. West coast: SWA 5Y. N 116. LAM 45Z. B 114A. SB 141T, 194].
LB 511D. CP 17, 328, 335, 650G. TRA 156K. WCD 34F, 81A.
South coast: E 133. L 149. SAMH 201, 304.
Cladocarpus leloupi nov. nom.
Cladocarpus flexilis Leloup, 1939, p. 14, fig. 10.
Records. South coast: TRA 92J. SCD 96B, 122V, 141K, 145A, 153M, 188R.
Description. A number of colonies and fragments without root-stock, the longest
stem reaching 22 cm. Stem fascicled but unbranched, bearing alternate hydro-
cladia. Stem internodes each bearing 3-6 median nematothecae, one in the
axil of the hydrocladium and the rest below it. Cauline nematotheca with 1,
2 or 3 openings.
Hydrocladium and hydrotheca as described by Leloup, but intrathecal
ridges tending to disappear towards the distal ends of the hydrocladia. Lateral
nematotheca usually with 3 distal openings (but occasionally with 1, 2 or 4)
and 1 lateral opening on mesial surface.
Gonophores absent.
Measurements (mm.)
Hydrocladial internode, length ne ae Br ae .. 0°70-0:87
diameter at distalend .. ae a ay ne .. 0°08-0°13
Hydrotheca, depth to median tooth .. Ss camuiiger: ys .. 0°45-0°62
diameter at margin me Ns se 5 oes .. O*19—-0°26
Remarks. In view of the fact that the name Cladocarpus flexilis is preoccupied by
an American species, C. flexilis Verrill 1883, I assign to Leloup’s species the
new name C’ leloupi.
5 ie sai
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 305
Cladocarpus lignosus (Kirchenpauer), 1872
(Fig. 9)
Cladocarpus lignosus. Stechow, 1923b, p. 243, fig. J*. Stechow, 1925, p. 505. Millard, 1961, p. 206.
Records. South coast: TRA 38H, 56S, 151E. SAMH 167, 280, 284, 297, 306.
Sie 26), 52H, 84A, 153A, 201X.
(Natal: NAD aiF.)
Fic. 9. Cladocarpus lignosus (Kirch.).
A, C. Phylactocarps bearing gonothecae, from NAD 21F and TRA 38H respectively.
B. The distal part of the hydrocladium-bearing tube of the stem to show segmentation and
cauline nematothecae, from NAD alF.
D. Two nematothecae from a phylactocarp, one with 2 apertures and the other with 3.
Description. Colonies large, reaching a maximum height of 34:5 cm., easily
recognizable by the woody nature of the stem and the branching, which is
always in one plane.
Stem dividing and subdividing in a very irregular fashion, some of the
smaller branches showing a tendency towards an alternate arrangement, but
frequently opposite or unilateral—the final result is a fan-shaped colony with
well-dispersed branches.
Hydrocladia borne on the smaller and medium-sized branches, absent
306 ANNALS OF THE SOUTH AFRICAN MUSEUM
on the larger ones. Hydrocladia arising alternately from a single tube of the
fascicled branch, this tube, where it is free from the peripheral tubes at the
extremity, being regularly segmented, each internode giving rise to a long
hydrocladium-bearing apophysis at about the middle of its length. Hydro-
cladium with first internode short and athecate, with 1 median nematotheca
and 2 internodal septa; remaining internodes longer and thecate, each with a
median and 2 lateral nematothecae, and 3 or 5 internodal septa. Three of the
internodal septa situated as described by Stechow, but frequently an additional
one at the extreme proximal and another at the extreme distal end.
Hydrotheca with depth about 14 times width at mouth, slightly compressed
laterally, with 1 anterior and 2 lateral marginal teeth, which may all be of
equal size or with the anterior tooth slightly exceeding the laterals in length.
Median nematotheca arising from the basal swelling of the internode on
which the hydrotheca is seated, very short, quite free from hydrotheca, with
2 distal, tubular apertures.
Lateral nematotheca with 3 apertures, one on the mesial surface, and 2
distal and tubular, of which the anterior one is long and reaches beyond thecal
margin, and the other is shorter and reaches to the margin or just below
te
Two cauline nematothecae on one surface of each internode of the hydro-
cladia-bearing tube, 1 near base on opposite side to apophysis, and 1 in axil
of apophysis, both with 2 distal apertures. Two rows of small, single nemato-
thecae on each peripheral tube.
Phylactocarps present in the older parts of the colony, 1 or a pair, on
each hydrocladium arising one on each side of the median nematotheca of the
first thecate internode. Phylactocarp unbranched, curving towards hydro-
cladium, bearing up to 8 pairs of nematothecae. Nematothecae decreasing in
size from base to tip, basal ones stout and with 3 apertures—2 terminal and
I on mesial surface, distal ones more slender and generally with 2 apertures—
I terminal and 1 on mesial surface.
One to four gonothecae (not previously described) borne on each phylacto-
carp, 1 between the members of each pair of nematothecae on surface facing
hydrocladium. Gonotheca elongated, with more or less rectangular aperture
near distal end facing towards phylactocarp.
Measurements (mm.)
Hydrocladium, first internode, length on posterior surface .. 013-0°18
remaining internodes, length on posterior surface .. .. 0°33-0°42
Hydrotheca, depth to lateral tooth ae au 1h _. > (O23 o ag
width at mouth ae be Be au at .. O*15—-O0°19
Median nematotheca, length of free part be ee .. 0°025-0°05
Lateral nematotheca, length ae fe wf ay .. | \0°L2=016
Gonotheca, length .. ah ans i a reaching 1°78
WAIL INE eet a PN a ay a: reaching 0:68
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 307
Remarks. This endemic South African species occurs on the Agulhas Bank
from 19°E. eastwards, extending round into Natal waters as far as 31°E.
It occurs in depths ranging from 22 to 120 metres.
The plentiful material available makes possible the amplification of
Stechow’s description.
Halicornaria arcuata (Lamouroux), 1816
(Fig. 10 E)
Halicornaria arcuata. Millard, 1958, p. 218, fig. 15 D, E.
Records. South coast: MB 24U, 47G. KNY go0T, 71F. CPR 46A. SAMH 257,
300. SCD 20B, 84C, 117P, 169R.
Description. Rich colonies of unbranched stems reaching a maximum height of
12°8 cm.
2mm.
TRAN
oe
.
site
Lipkwoerer rept eepe
SS Se Sd Oesimim:
Fic. 10. Halicornaria arcuata (Lamx.).
A-D. Subsp. epizootica Millard.
A. Part ofa colony from SCD 84D growing on the back of the stem of the normal subspecies.
Hydrocladia of host cut off short.
B-D. Examples of reduced hydrothecae, B from SAMH 255, C and D from the upper and
lower parts respectively of the same hydrocladium, from SAMH 355.
E. Subsp. arcuata, two gonothecae from MB 47G.
308 ANNALS OF THE SOUTH AFRICAN MUSEUM
Hydrothecal margin with an anterior tooth, a posterior tooth and usually
3 pairs of lateral teeth. Of the latter the posterior pair is often poorly defined
in the proximal region of the hydrocladium, and the middle pair in the distal
region.
Median nematotheca usually closed at the tip, but often open in the
proximal regions of the hydrocladia. Lateral nematothecae with 2 openings,
of which 1 may be produced into a tube or into a long closed spine near the
distal ends of the hydrocladia.
One stem bears several branching hydrocladia, possibly the result of
injury. Each branch arises from the posterior surface of the hydrocladium
and bears hydrothecae on the surface facing it.
Female gonothecae borne on the hydrocladial apophyses, 1 to each, and
forming a double row on the anterior surface of the stem. Each is rounded
below and truncated above, with the width slightly exceeding the
height.
Remarks. Female gonophores are said to have been mentioned by Krauss (1837)
as occurring in the axils of the hydrocladia, but this is the first description of
these structures.
Halicornaria arcuata epizootica Millard, 1958
(Fig. 10 A-D)
Halicornaria arcuata var. epizootica Millard, 1958, p. 218, fig. 15 F.
Records. South coast: CPR 46C. SAMH 255. SCD 75B, 84D.
(Natal: SAMH 355.)
Description. Colonies epizootic on Thecocarpus formosus (Busk) and Halicornaria
arcuata (nominate subspecies); consisting of solitary hydrocladia and upright
pinnate stems.
Pinnate stems reaching 12-5 mm. in length and bearing hydrocladia with
as many as 14 hydrothecae. Solitary hydrocladia bearing as many as 13
hydrothecae.
The hydrothecae and nematothecae are very variable in structure, those
on the distal ends of the hydrocladia of pinnate stems resembling those of the
nominate subspecies (see Millard, 1958, fig. 15 F), but those on the proximal ends
of the same hydrocladia and those on solitary hydrocladia being successively
reduced towards the base of the colony (fig. 10 B-D). This reduction affects:
(a) the paired lateral and the posterior thecal teeth which become ‘smoothed
out’ until the margin is merely sinuated; (b) the median inferior nematotheca,
which becomes shorter and no longer reaches the level of the thecal margin;
thus it may have no free portion and only one opening; and (c) the lateral
nematothecae which become reduced, with the two apertures combined into
one.
Gonophores absent.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 309
Remarks. The same modifications were present in the holotype (Millard, 1958),
but owing to the poor condition of the colony and the covering of silt it was not
possible to determine whether the condition was normal or due to damage.
Hfalicornaria exserta nov. sp.
(Fig. 11 A-E)
Types and records. Holotype: SAMH 169 (South African Museum registered
number). Additional records: SCD 36N, 153Y. All from the south coast.
Description (holotype). One upright stem 8-3 cm. in length and bearing gono-
phores. Hydrorhiza of same diameter as stem, with thick perisarc. Stem unfas-
cicled and unbranched. Proximal region (about 1:5 cm.) unsegmented and
without hydrocladia or nematothecae. Middel region (about 1 cm.) divided by
transverse nodes into rather irregular internodes which tend to become pro-
gressively shorter, and which bear 1 or 2 median nematothecae each. Distal
region divided into short hydrocladia-bearing internodes by slightly oblique
nodes which slope in opposite directions so that the side of an internode bearing
the hydrocladium is longer than the other.
Hydrocladia alternate, borne on short apophyses of the stem, one to an
internode, reaching a maximum length of 1-75 cm. Each divided into thecate
internodes by transverse nodes. No internodal septa.
Hydrotheca expanding to margin, with adcauline wall free from internode
for a short region distally, and with abcauline wall bent strongly forwards.
Margin with 1 prominent anterior tooth, 3 pairs of well-defined lateral teeth,
and a low, rounded, posterior tooth. Hydropore with margin inturned.
Median inferior nematotheca short (about 4 length of hydrotheca),
adnate to hydrotheca for most of its length, with the terminal opening extending
along all of the short free part, with no communication with thecal cavity.
Lateral nematotheca ovoid, not reaching thecal margin or even beginning of
free part, with 1 broad distal aperture. Cauline nematothecae similar to
laterals: 1 inferior, 1 axillary anterior, and 1 axillary posterior.
Gonophores borne on hydrocladial apophyses and forming a double row
down front of stem, bowl-shaped, with very short pedicels and broad, truncated,
distal ends.
Measurements. See subspecies epizootica.
Remarks. This species is related to H. sibogae Billard 1918 and H. expansa
Jaderholm 1903, but differs in the unbranched stem, shorter hydrothecae,
the presence of a posterior marginal thecal tooth, and other minor details.
Halicornaria exserta epizootica nov. subsp.
(Fig. 11 F—H)
Holotype: SAMH 320 (South African Museum registered number). From
south coast.
310 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description. Hydrorhiza epizootic on back of stem and hydrocladia of Thecocarpus
flexuosus (Lamx.) umbellatus, giving off solitary hydrocladia and upright pinnate
stems. Solitary hydrocladia arising alternately, reaching a maximum height of
o-5 cm. and bearing up to 10 hydrothecae, forming an angle of about 45°
with hydrorhiza. The two rows of hydrocladia not in the same plane but shifted
towards posterior surface of host. All hydrothecae facing same direction as
those of host. Pinnate stems reaching a maximum height of 0-9 cm. and bearing
alternate hydrocladia as in nominate subspecies.
Structural details similar to those of normal form except that the hydro-
cladial internodes tend to be longer and narrower and the hydrothecae are not
so strongly bent forwards. Median and lateral nematothecae tending to be
reduced in size towards the proximal region of solitary hydrocladia, so much
so that the median nematotheca may not even reach the base of the hydrotheca.
Gonophores absent.
Measurements (mm.)
Halicornaria subsp.
exserta eprzootica
SAMH 169 SCD36N SAMH 320
Hydrocladium, internode length 0°24-0°39 0°29-0°37 0°33-0°45
diameter at distal end O°15—0°23 O°19-0°26 =0:09-0°15
Hydrotheca, length adcauline 0°24-0°34. 0°30-0°36 0°25-0°35
length adcauline, free part .. 0:03-0:06 0:03-0:06 0:04-0:08
length abcauline 0:28-0°34 0:34-0:36 0°22—-0°33
diameter at margin . 0:22-0'27 0:22-0:25 0:18-0-26
Median nematotheca, length O'10-0°I2 O°10—-0°12 0:06—0-09
Lateral nematotheca, length 0:09-0'I12 O°I0-0'12 0:05-0:09
Gonotheca, length 0:26
maximum diameter 0°42
Remarks. This subspecies has a similar growth-form to that of H. arcuata
epizootica, and shows the same tendency for reduction of nematothecae in
solitary hydrocladia.
A-E. Subsp. arcuata.
A, B. The stem, and the gonotheca, from the holotype, SAMH 169.
C. Anterior view of stem showing the cauline nematothecae and scars for gonothecae
(striped), from SCD 36N.
D, E. Anterior and lateral views respectively of part of a hydrocladium from the holotype.
F-H. Subsp. epizootica, from the holotype SAMH 320.
F. Part of a colony on the back of a stem of Thecocarpus flexuosus umbellatus showing
solitary hydrocladia and 2 pinnate stems. Hydrocladia of host cut off short.
G, H. Anterior and lateral views respectively of part of a hydrocladium.
eet
311
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA
q{
"wu G-O
Hohe Gd =a
SS BRS.
Vee aI
Le
we 0 O—————OoO
Fic. 11. Halicornaria exserta n. sp.
312 ANNALS OF THE SOUTH AFRICAN MUSEUM
Flalicornaria ferlusi Billard, 1901
Halicornaria Ferlusi. Billard, 1907, p. 370, pl. 25 (fig. 8), fig. 14.
Records. South coast: CPR 46B.
(Additional from Natal: NAD 1R.)
Description. Stem reaching a maximum height of 3 cm. Structural details
exactly similar to those described by Billard.
Male gonothecae present (in NAD 1R only), similar to those described
by Jarvis (1922) for var. brevis, except that there is no aperture, and no ‘refrac-
tive discs’ are visible. Each containing one gonophore.
Remarks. This species is known only from Madagascar, and var. brevis Jarvis
from Wasin, East Africa. This is the first record from South Africa.
Lytocarpus filamentosus (Lamarck), 1816
Lytocarpus filamentosus. Millard, 1957, p. 241. Millard, 1958, p. 220.
Records. South coast: MB 15A, 24S, 26D, 39V, 47J, 52B, 55E, 58B, 70F, 72F,
83A. LIZ 7M, 13B, 27B. CPR 46E. TRA 42B. SAMH 150, 259: SCD SAE;
o4k; 117, 169);
Thecocarpus flexuosus (Lamx.), 1816
(Bigey 2)
Aglaophenia flexuosa Lamouroux, 1816, p. 167. Kirchenpauer, 1872, p. 25.
Aglaophenia plumifera Kirchenpauer, 1872, p. 31, pl. 1 (fig. 6), pl. 3 (fig. 6). Stechow, 1923b,
Pp- 255, fig. K1.
Thecocarpus giardi Billard, 1907, p. 381, pl. 25 (figs. 9-10), pl. 26 (figs. 11-17), fig. 21. Vervoort,
1946, p. 335. Millard, 1957, p. 240. Millard, 1958, p. 221, fig. 16 A-C.
Thecocarpus flexuosus. Billard, 1909, p. 330.
? Aglaophenia(?) bifida Stechow, 1923a, p. 117. Stechow, 1925, p. 515, fig. 53.
Diagnosis. Stem thick, woody and fascicled, subdividing and giving rise to
‘branches’ formed from sympodially branching pinnae. Each pinna giving
rise to a subsequent one from its anterior surface, the whole ‘branch’ twisted
in a spiral manner and with a geniculate main axis formed from the proximal
parts of successive pinnae. Each pinna consisting of a basal part bearing 1
large median nematotheca on each internode, and a distal part bearing 1
hydrocladium and 3 nematothecae on each internode. Hydrocladia alternately
arranged, divided into internodes by slightly oblique nodes, each internode
typically with 2 internodal septa. Hydrotheca adnate, deep, expanding to
margin, with an intrathecal septum near base, and with 9 marginal teeth (1
median and 4 pairs lateral). Median nematotheca short, reaching approximately
to level of intrathecal septum, with about + length free. Lateral nematotheca
reaching to thecal margin. Corbula replacing hydrocladium, with pedicel of
1-6 thecate internodes, elongated, closed. Each rib of corbula bearing a series
of nematothecae along outer edge, a hydrotheca near the base, and a crested
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 313
process of variable length below it. Base of corbula with a rounded projection
facing towards pedicel.
Remarks. This species is endemic to southern Africa, where it appears to be
widely distributed around the coast extending as far north as Madagascar
on the east. It is extremely variable in its growth form and in its microscopic
structure, necessitating the establishment of a number of subspecies. The charac-
ters of a subspecies are generally constant in a sample from a particular locality,
though the occurrence of occasional intergrading specimens shows that all the
material is conspecific. The species can always be recognized by the spiral
arrangement of the pinnae, by the method of branching (described in detail
by Billard (1907), and Millard (1958) under the name of T. gzardi) and by the
structure of the corbula.
It has already been shown (Millard, 1958) that the keel-like ridge along
the abcauline wall of the hydrotheca (i.e. the outer point of the median tooth)
may be hollow or solid. Material from the Agulhas Bank now shows that this
ridge may be very poorly developed (so that the outer point of the median
tooth may be shorter than the inner one), or absent altogether. Material with
the latter arrangement has been placed in a separate subspecies, but amongst it
rare hydrothecae occur showing the beginnings of a ridge, i.e. the first bifur-
cation of the median tooth. A series of hydrothecae can thus be arranged
(fig. 12, C-M) showing the progressive development of this structure culminat-
ing in the large hollow spine of subsp. flexuosus and subsp. umbellatus.
The structure of the hydrotheca in the ‘keelless’ variety (fig. 12 C) closely
resembles Stechow’s diagram (1923b, fig. Kt) of Kirchenpauer’s type material
of Aglaophenia plumifera. The preserved material of the type of A. plumifera has
unfortunately been lost, but Stechow states that the pinnae arise alternately
and Kirchenpauer that they arise irregularly. Probably both are correct, for a
series of alternate pinnae twisted in a spiral manner would give an irregular
appearance. I have seen whole mounts of A. plumifera prepared by Stechow
(loaned from the Munich Museum) including a complete pinna, and this
shows absolute agreement with the ‘keelless’ variety including the presence of
hinge-joints and the large median nematothecae on the internodes below them.
The two are thus considered to be synonymous, and are given the status of a
subspecies, under Kirchenpauer’s name plumiferus.
Thecocarpus flexuosus plumiferus (Kirch.), 1872
(Fig. 12 €, D)
Aglaophenia plumifera Kirchenpauer, 1872, p. 31, pl. 1 (fig. 6), pl. 3 (fig. 6). Stechow, 1923b,
p- 255, fig. K?.
Diagnosis. Pinnae arranged in a loose to tight spiral around a ‘main axis’ which
is seldom definitely geniculate. Intervals between pinnae regular and approxi-
mately equal. Basal and distal regions of pinnae separated by hinge-joints.
314 ANNALS OF THE SOUTH AFRICAN MUSEUM
SS S\ GH
ey \ LAF
<f qt = ZZ
‘ A ANG W]W]WXxkkfyzz
ys WW A \ A Mill Ka Zi LE
| Ny > = SS a oy —
Fic. 12. Thecocarpus flexuosus (Lamx.).
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 315
Hydrocladia short, 3-6 mm. in length. Hydrotheca with no abcauline ‘keel’,
but abcauline wall somewhat thickened near margin. First and second pairs of
marginal thecal teeth not bifid.
Records. South coast: SCD 36L, 79H, 104G. SAMH 353.
Description. One of the colonies (SAMH 353) very large, reaching 46 cm.,
and with base of stem 12 mm. thick. The spiral twisting of the ‘branches’ is
sometimes difficult to recognize in the lower, heavily fascicled regions of the
colony, where also the geniculate appearance is lost. It is, however, quite
distinct in the distal regions.
Unlike other subspecies a number of internodes below the hinge-joints
may bear hydrocladia. Hinge-joints distal to origin of next pinna.
Corbulae sometimes very long, reaching a maximum length of 20 mm.
and bearing up to 39 pairs of ribs.
pce also Pp. 313.
Measurements. See under subsp. umbellatus.
Thecocarpus flexuosus flexuosus (Lamx.) 1816
(Fig. 12 A, J-L)
Thecocarpus giardi Billard, 1907, pp. 381-385, pl. 25 (fig. 9), pl. 26 (figs. 1 1-16), fig. 21. Vervoort,
1946, p. 335. Millard, 1957, p. 240. Millard, 1958, p. 221, fig. 16 A.
? Aglaophenia(?) bifida Stechow, 1923a, p. 117. Stechow, 1925, p. 515, fig. 53.
Diagnosis. Pinnae arranged in a loose to tight spiral, around a geniculate
‘main axis’. Intervals between pinnae regular and approximately equal. No
hinge-joints. Hydrocladia short, about 6 mm. in length. Hydrotheca with
abcauline ‘keel’ extended beyond median tooth as a hollow spine. First and
second pairs of marginal thecal teeth typically bifid.
Records. South coast: KNY go0R.
ie
A. Subsp. flexuosus. Part of colony showing a spirally twisted ‘branch’ and its pinnae, from
30R.
B. Subsp. umbellatus n. subsp. Part of colony showing one ‘branch’, from the holotype SCD 36M.
C-M. Selected hydrothecae showing variations, particularly of the anterior ‘keel’.
C, D. Subsp. plumiferus (Kirch.), a typical hydrotheca with no ‘keel’ from SCD 36L, and a
rare hydrotheca with a slight indication of a ‘keel’, from SAMH 353.
E-H. Subsp. solidus Millard, a rare terminal hydrotheca with the ‘keel’ barely distinct from
the median tooth, from TRA 92C; a hydrotheca with well-developed ‘keel’ but restricted
abcauline thickening, from TRA 92C; a typical hydrotheca with abcauline thickening
reaching to base, from AFR 1028A (described by Millard 1958); and a rare hydrotheca
with a hollow ‘keel’, from AFR 1028A (described by Millard, 1958).
J-L. Subsp. flexuosus, a rare hydrotheca with a solid ‘keel’, from PF 396A (described by
Millard, 1957); a typical hydrotheca with a hollow ‘keel’ and a bifid lateral tooth,
from PF 12308B (described by Millard, 1958); and a hydrotheca with an extra long
‘keel’ pierced at distal end, from KNY 30R.
M. Subsp. umbellatus n. subsp., a typical hydrotheca, very similar to K except that the lateral
teeth are not bifid, from the holotype SCD 36M.
316 ANNALS OF THE SOUTH AFRICAN MUSEUM
Thecocarpus flexuosus solidus Millard, 1958
(Fig. 12 E-H)
Thecocarpus giardi Billard, var. solidus Millard, 1958, p. 222, fig. 16 B, C.
Diagnosis. Pinnae arranged in a loose to tight spiral around a geniculate ‘main
axis’. Intervals between pinnae regular and approximately equal. Basal and
distal regions of pinnae separated by hinge-joints. Hydrocladia short, about
6 mm. in length. Hydrotheca with abcauline ‘keel’ forming a solid spine. First
and second pairs of marginal thecal teeth not bifid.
Records. South coast: TRA 92C. SCD 29F, 36K, 84E. SAMH igo.
Remarks. One of these samples (TRA 92C) shows characters tending towards
subsp. plumiferus (fig. 12 E, F). The abcauline ‘keel’ of the hydrotheca is poorly
developed and restricted to the margin, the typical chitinous thickening which
usually extends down to the junction of the median nematotheca being
absent.
In the same sample there is a number of branching hydrocladia. A hydro-
cladium may produce as many as 7 subsidiary hydrocladia, and in rare cases
the subsidiary hydrocladia branch too. The hydrocladial internodes which
give rise to branches are without hydrothecae and similar in every respect to
those of the distal region of the pinna.
Thecocarpus flexuosus umbellatus nov. subsp.
(Fig. 12 B, M)
Diagnosis. Pinnae (usually 5) arranged in a very close spiral forming an umbel-
like cluster at the end of a long ‘stem’ formed by the proximal region of the first
pinna. Hinge-joints present on first pinna only. Hydrocladia long (reaching
15 mm.). Hydrotheca with abcauline ‘keel’ extending beyond median tooth
as a hollow spine. First and second pairs of marginal thecal teeth not bifid.
Types and records. Holotype: SGD 36M (South African Museum registered num-
ber — sa Wie o2)e
Further records, south coast: SCD 33B, 117M, 153C. SAMH 168, 258,
286, 305, 3109.
Description of holotype. Colony reaching a height of 12:3 cm. Stem fascicled and
dividing near base to give the branching pinnae or ‘branches’.
First pinna with basal part long (several cm.), usually lightly fascicled
(though the peripheral tubes do not extend on to the distal part), divided into
internodes each with 1 large median nematotheca with several (3-6) openings.
This part terminated by 2 oblique hinge-joints, the internode between them
bearing 2 large median nematothecae. Distal part bearing 2 hydrocladia on
the first internode and 1 on each of the rest, and giving origin to the next pinna
from the anterior surface of the first to fifth internode.
HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 317
Second pinna facing towards the first, seated on a short apophysis, with no
basal part and no hinge-joints, consisting only of hydrocladia-bearing inter-
nodes, and giving rise to the third pinna from its second internode.
Subsequent pinnae (usually 3) as second, and the ‘main axis’ (formed
by the origins of consecutive pinnae) twisted in a spiral manner.
Hydrocladia alternate, long, reaching 15 mm. Hydrotheca similar in
structure to subsp. flexuosus except that the first and second lateral teeth are
not bifid. Hydrothecal depth and hydrocladial internode length varying from
that found in subsp. flexuosus to that in subsp. perarmatus Billard 1907.
Corbula (not from holotype) with up to 20 ribs and reaching 12 mm. in
length. In some cases there is an enormous development of the terminal parts
of the ribs and the ‘crests’ arising from their bases, giving to the whole a spidery
appearance similar to that sometimes encountered in T. formosus.
Measurements (mm.)
subsp. subsp. Aglaophenia
umbellatus plumiferus plumifera*
_Hydrocladium, internode length .. O*19-0°32 0:28-0:37 0:28-0°30
Hydrotheca, height (to tip of inner point
of median tooth) .. si .. 0°22-0°30 0°23-0°31I 0°24-0°27
diameter at margin (inside) .. O°16-0-2I 0-14-0°205 0°13-0°18
Median nematotheca, length .. .. O°14-0°24 O°13-0°2I 0°15-0°18
length of free part .. a: .. 0°03-0:06 0:03-0:04 0:03-0:04
Corbula, length .. ay, reaching 8-50 20°00
diameter . ae reaching 1-40 ge
Remarks. Although the arrangement described above is remarkably constant,
variations do occur (e.g. longer intervals between origins of pinnae) which
show that this form is not worthy of specific rank. The top part of the colony
breaks easily at the hinge-joints and many specimens brought up by the dredge
consist of crowns only. The pinnae often terminate in long tendrils.
Thecocarpus formosus (Busk), 1851
Thecocarpus formosus. Millard, 1958, p. 221. Maillard, 1961, p. 206.
Records. South coast: MB 24Y, 47F. KNY 30S, 57K, 71A (recorded by Day,
Millard and Harrison, 1952). LIZ 7N, 27A. BMR 12K. L 55. CPR 46D.
SCD 20A, 29E, 36H, 50B, 75A, 84B, 94D, 117N, 1698S. SAMH 200, 210,
254, 285, 299.
* Measurements from Stechow’s slides borrowed from the Munich Museum.
318 ANNALS OF THE SOUTH AFRICAN MUSEUM
SUMMARY
A total of 35 species of plumularian hydroids is described from the south
and west coasts of South Africa. Among these are 4 new species and 4 new
records from South Africa.
A new subfamily, the Halopterinae, is created for genera with cauline
hydrothecae and related forms, and possible evolutionary trends within it
discussed.
The genus Antennopsis is sunk in the synonymy of Nemertesia, and a new
genus, Corhiza, created. The genus Paragattya is sunk in the synonymy of Gattya.
Various problems of nomenclature are dealt with, including that of
Thecocarpus flexuosus, and the subspecies of the latter are revised.
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Bate, W. M. 1914a. Report on the Hydroida coHected in the Great Australian Bight and other
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BALE, W. M. 1914b. Report on the Hydroida collected in the Great Australian Bight and other
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Bate, W. M. 1915. Report on the Hydroida collected in the Great Australian Bight and other
localities. Part III. Zool. Res. Fish. Exp. ‘Endeavour’ 3, 241-336.
Bepot, M. 1921. Notes systématiques sur les plumularides. 2me Partie. Rev. suisse Zool. 29,
I-40.
Bepot, M. 1923. Notes systématiques sur les plumularides. 3me Partie. Rev. suisse Zool. 30,
213-243.
BILLARD, 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), 72 335-396.
BILLARD, A. 1909. Revision des espéces types d’hydroides de la collection Lamouroux, con-
servée a l'Institut botanique da Caen. Ann. Sci. nat. zool. (9), 9, 307-336.
BILLARD, A. 1910. Revision d’une partie de la collection des hydroides du British Museum.
Ann. Sci. nat. zool. (9), 115 1-67.
BILLARD, A. 1913. Les hydroides de l’expédition du Siboga. I. Plumulariidae. Siboga Exped.
monogr. 7a, I-I15.
BILLARD, A. 1918. Notes sur quelques espéces d’hydroides de l’expédition du ‘Siboga’. Arch.
Kool. exp. gén. 5°75 21-27.
Briccs, E. A. 1914. Notes on Tasmanian Hydrozoa. 7. roy. Soc. N.S.W. 48, 302-318.
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Brocu, H. 1933. Zur Kenntnis der adriatischen Hydroidenfauna von Split. Skr. norske Vidensk-
Akad., Mat.-natur. Kl. 1933, no. 4, I-115.
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.
Day, J. H. 1959. The biology of Langebaan Lagoon: a study of the effect of shelter from wave
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HYDROZOA OF SOUTH AND WEST COASTS OF SOUTH AFRICA 319
Eyre, J., BRoEKHUYSEN, G. J. & Cricuton, M. I. 1938. The South African intertidal zone and
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STECHOW, E. 1923a. Ueber Hydroiden der deutschen Tiefsee-Expedition, nebst Bemerkungen
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STECcHOW, E. 1925. Hydroiden der deutschen Tiefsee-Expedition. Wiss. Ergebn. ‘Valdivia’ 17,
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STEPHENSON, T. A., STEPHENSON, A. & pu Torr, C. A. 1937. The South African intertidal
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Soc. S. Afr. 24, 341-382.
STEPHENSON, T. A., STEPHENSON, A. & Bricut, K. M. F. 1938. The South African intertidal
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VerRvoorT, W. 1946. Exotic hydroids in the collections of the Rijksmuseum van Natuurlijke
Historie and the Zoological Museum at Amsterdam. Zool. Meded. 26, 287-351.
VeERVooRT, W. 1959. The Hydroida of the tropical west coast of Africa. Atlantide Rep. 5, 211-325.
Warren, E. 1908. On a collection of hydroids, mostly from the Natal coast. Ann. Natal Mus. 1,
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hydroid. Ann. Natal Mus. 3, 83-102.
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Bettrdége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
HartTMAN, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
IzuxA, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
202.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.— All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
' tive citations following in chronological order after each name... .
B. Form of bibliographic references to synonymic names.—The first reference following
any name in the synonymy should be to the earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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INDEX
XXXI.
XXXIT.
XXXITI.
XXXIV.
XXXV.
XXXVI.
XXXVI.
XXXVITI.
XXXIX.
XL.
XLI.
XLII.
XLII.
XLIV.
XLV.
XLVI.
1900-1902
1903-1905
1903-1908
1906-1910
1908-1910
1908-1913
IQII—1918
IQII-I9I4
IQII—1918
1913-1924
IQI13—-1923
IQI5-1924
1914-1916
1917-1933
1917-1920
IQ2I
1924-1925
1924-1926
1925-1928
1925-1926
1929-1038
1927-1928
1928
1929
1929-1932
1929-1931
1931-1935
Zoology and Geology (excl. Parts 1-3, 5, 7,
Zoology (excl. Part I)
Palaeontology ..
Geology, Palacontology, Zooloay, Anthropology (excl.
Parts 1,2; 5,7, 8; .9) 3
Zoology (excl. Part i, Index)
Palaeontology (excl. Parts 1-4)
Botany (excl. Parts 1, 2)
Zoology (excl. Parts 1, 2) :
Zoology (excl. Parts 2, 7, Index)
Palaeontology and Geology
Archaeology and Zoology
Zoology
Zoology
Botany
Zoology
Zoology
Zoology
Zoology oe
Palaeontology ..
Zoology
Anthropology al Bie olony (eon Part De
Zoology : Be Ke
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ANNALS
OFR-TEHE
PVE APRICAN MUSEUM
VOLUME XLVI
pana XI
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA, WITH
DESCRIPTIONS OF FIVE NEW SPECIES
By
ie Ee Srock
Koblogisch Museum, Amsterdam, Holland
ISSUED MARCH 1963 PRICE 40c.
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[ CONTINUED ON INSIDE BACK COVER
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA, WITH
DESCRIPTIONS OF FIVE NEW SPECIES
By
J. H. Stock
Koblogisch Museum, Amsterdam, Holland
[Accepted April 1960]
(With to figures in the text)
CONTENTS
PAGE
Introduction ‘ ‘ : 5 Ber
Descriptions of species . : 5) OH
Summary . : ; ; a 340
Acknowledgements : , 5 Bylo)
INTRODUCTION
The interesting collection of deep-sea Pycnogonida on which the present
paper has been based was entrusted to me by Dr. F. H. Talbot of the
South African Museum. Taking into consideration the considerable depths in
which the animals were caught (ranging from 1,470 to 1,785 fathoms), and
the relative rarity of Pycnogonida in general, the present collection is pretty
rich both in species and in individuals.
The material collected belongs to 5 genera only: Nymphon (1 species),
Ascorhynchus (1 species), Colossendeis (6 species), Anoplodactylus (1 species), and
Pantopipetta (3 species). Although all of these were known to be restricted to,
or to have representatives in, the deep sea, the occurrence of not less than 3
species of Pantopipetta, a very rare genus, is most surprising.
The strong armature of body and legs with long spurs and long, stiff
setae in several of these abyssal species is a curious morphological phenomenon,
and is at least for the genus Nymphon quite unusual.
Pycnogonid material was taken in the following deep trawlings (gear: 15’
beam trawl); Station A 193 especially was very rich in species and specimens.
Station A 192. 33° 45’ S., 16° 234’ E. 1,480 fms. Aug. 27, 1959.
Station A 193. 33° 49’ S., 16° 30’ E. 1,500—1,580 fms. Aug. 27, 1959.
Station A 315. 34° 37’ S., 17° 03’ E. 1,580-1,620 fms. Dec. 8, 1959.
Station A 316. 34° 42’ S., 16° 54’ E. 1,725-1,780 fms. Dec. 8, 1959.
Station A 319. 34° 05’ S., 16° 58’ E. 1,470-1,490 fms. Dec. 9, 1959.
Station A 322. 34° 36’ S., 17° 00’ E. 1,500 fms. Dec. 10, 1959.
The holotypes of the new species are in the South African Museum,
Cape Town; duplicates are in the Zoological Museum, Amsterdam.
321
Ann. S. Afr. Mus. 46 (12), 1963, 321-340, 10 figs.
322 ANNALS OF THE SOUTH AFRICAN MUSEUM
DESCRIPTIONS OF SPECIES
Nymphon laterospinum n.sp.
(Fig. 1)
Material. 1 § ovig. Station A 193.
Description. Trunk fairly slender, completely segmented. Lateral processes
separated by nearly 1} times their own diameter; armed distally with spurs,
/
Fic. 1. Nymphon laterospinum n.sp., $ (holotype). a, dorsal view of the trunk; 5, inner margin of
the claw of a leg; c, first leg; d, distal segments of the first leg; e, chela; f; oviger; g, some special
oviger spines; h, terminal part of the oviger; 7, palp.
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA B28
which are very tall and spiniform on lateral processes 1, 2 and 3, much shorter
on lateral process 4. A protuberance of about the same size as that of the 4th
lateral process is present on the neck, at the base of the chelifore. No eye
tubercle, nor middorsal spurs. No eyes. Oviger implantation in contact with the
first lateral process. Neck short, fairly broad. Abdomen long, reaching to at
least a third of coxa 2 of leg 4. Proboscis about as long as the neck.
Fingers of chela nearly twice as long as the hand, curved; movable finger
with about 16 short, spiniform teeth; immovable finger with the same number
of stronger teeth.
Palp hairy; segment 1 short, segment 2 the longest (subequal to segments 3
and 4 combined), segment 5 slightly longer than segment 4.
Oviger segment 4 with a swelling at about a third of its length; segment 5
long, curved, distinctly clubbed distally; segment 6 very hairy; segments 7 to 10
with a small number of compound spines of a relatively simple structure,
bearing only 1 to 3 serrations, according to the formula 5 + 3 + 3+ 3= 14.
Terminal claw slender, slightly curved, longer than the roth oviger segment,
bearing 7 teeth at the inner margin. Eggs few (only two on each oviger in the
only male available); they are collapsed now, but originally were probably
large and rich in yolk.
Legs slender, setose; particularly coxa 2, femur and tibia 1, wach very
long, stiff setae; tibia 1 ane 2 subequal; tarsus about 2 of the oncwarhss propo-
dus nearly straight; propodal sole with 20 to 25 small spinules of a size; claw
about # of the propodus, slightly curved, its inner margin armed with about 8
irregular denticulations.
Remarks. The only other slender, uniunguiculate, blind Mymphon that has the
oviger implantation in contact with the first lateral process is N. hedgpethi
Stock, 1953. Instead of spurs on the neck and the lateral processes, NV. hedgpethi
possesses long hairs on these spots. The new species is, by the presence of these
spurs, and by its crenulated claw of the legs, particularly well characterized
against the 120 other named taxa in this genus.
Measurements of the holotype (3) in mm.
Length (frontal margin neck to base of abdomen) aD wy. Pe a,
Length of proboscis (in dorsal we ss se see ae Sy OROO
Length of abdomen : ue Bie fe Bi cf oa, On7
Width across 2nd lateral processes sti dhe a: : dae
First leg—ist coxa, 0°52; 2nd coxa, 1°13; Card coxa, 0°395 femur, 2°20; 1st
tibia, 2°44; 2nd tibia, 2-35; tarsus, 0°56; propodus, 0-89; claw, 0°34
Ascorhynchus inflatum n.sp.
(Figs. 2, 3) 4
Material. 4 specimens (2 of which adult females). Station A 193.
Description. Body completely segmented. Lateral processes separated by more
than 14 times their own diameter. Neck rather slender, bearing two small
324 ANNALS OF THE SOUTH AFRICAN MUSEUM
Og i
Sh ean
Fic. 2. Ascorhynchus inflatum n.sp., 9 (holotype). a, lateral view of the trunk; 6, fourth leg; ¢c, dorsal
view of the trunk; d, chelifore.
~~
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA 325
tubercles near the implantation of the chelifores. Basal protuberances for palps
and ovigers distinct. Eye tubercle arising slightly in front of the oviger implanta-
tions, very tall, pointed, spine-like; eyes absent. Body segments 1, 2 and 3
with a tall, sharp medio-dorsal tubercle. Tubercles of about the same size
and shape arise from the dorso-distal margin of the lateral processes. Abdomen
curved, slender, much longer than the 4th lateral process. The posterior part
of trunk segments 1, 2 and 3 and the anterior part of segments 2, 3 and 4 are
strongly inflated. The articulation line runs just over this inflated part of the
trunk segments. The proposed specific name refers to the curious gonflated
aspect of the trunk segments.
Fic. 3. Ascorhynchus inflatum n.sp., 2 (holotype). a, palp; b, distal segments of fourth leg; c, terminal
part of the oviger; d, compound oviger spine; e, oviger.
_ Basal third of proboscis distinctly separated from the distal two-thirds
Chelifore scape 1-segmented, rather long; chelae reduced.
Palps 10-segmented; basal two segments small; segment 3 the longest
4 distal segments gradually diminishing in length.
Oviger 10-segmented. Terminal claw slender, smooth. Compound spines,
bearing 3 to 5 strong basal denticulations and numerous fine crenulations at
themiip 4 ake present) on iSesments! 7 to. 10," according: to the fonnula
Pet hae 7 28.
Legs slender, rather setose, especially the tibiae, but without tubercles or
protuberances. Tarsus less than 4 of the propodus; it is about twice as long as
wide. Propodus slightly curved, the sole armed over its entire length with
326 ANNALS OF THE SOUTH AFRICAN MUSEUM
numerous small spinules of equal length. Claw short, distinctly less than half
the propodus, feebly curved. No auxiliary claws. Genital pores (2) on a very
slight swelling of the ventro-distal surface of the 2nd coxa of all legs.
Remarks. This species is not unlike A. abyssi G. O. Sars, 1877, a blind northern
Atlantic deep-sea species, but it differs notably from it in the much taller
dorsal spurs, the taller eye tubercle, and the different mutual position of eye
tubercle and oviger implantation.
Measurements of 9 (holotype) in mm.
Length of trunk somite 1 .. she ap ae oP JS med 7
Length of trunk somite 2 .. oe os Be Als 7 ies aa
Length of trunk somite 3... “e Se ns Be 1% RR os
Length of trunk somite 4 .. ber :3 ae a Bi *. §.0°66
Length of proboscis . . ns ni fe Se aa bid he Paes
Length of abdomen Bs 5 ne el ae Ng (Se
Width across 2nd lateral processes it am We ¢ eo eee
Length of scape oe : soe elas
Fourth leg—tst coxa, 66 gade COXAy 209F ard coxa, oe fenitt. 52 Ist
tibia, 5:2; 2nd tibia, 5-7; tarsus, 0-44; propodus, 1°54; claw, 0-66
Genus COLOSSENDEIS Jarz.
Brevitarsal group
Colossendeis macerrima Wilson
C. macerrima Wilson, Stock, 1953, pp. 308-311, fig. 17 e—h (lit., syn.). Barnard, 1954, p. 85.
C. japonica (non Hoek), Hedgpeth, 1949, pp. 299-300, fig. 46 a-d. Fage, 1956, p. 176 (new
synonymy).
Material. 7 specimens. Station A 193.
Remarks. Barnard, 1954, and Flynn, 1928, have already recorded this species
from South African waters. Concerning the proposed new synonymy, compare
the remarks under C. minuta.
Colossendeis minuta Hoek
(Fig. 4) 3
C. minuta Hoek, 1881, pp. 73-74, pl. 10 figs. 12-14. Schimkewitsch, 1893, pp. 34-35 (tabulated
only). Gordon, 1944, p. 13, fig. 2 f-g (remarks on holotype). Hedgpeth, 1948, p. 272, fig.
50 ¢.
C. japonica Hoek, 1898, pp. 295-296, figs. 11-13 (new synonymy).
[Non] ‘C. japonica Hoek’, Hedgpeth, 1949, and Fage, 1956 (= C. macerrima).
Material. 3 specimens. Station A 193.
Remarks. Three specimens, quite striking as they have preserved their crimson
colour, even in alcohol, agree in all particulars with the description of Hoek
(1881) and the notes given by Gordon (1944) on Hoek’s holotype. Their size
is somewhat larger than in Hoek’s immature material, though not yet quite
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA 327
as large (‘about 25 cm. in extent’) as the specimens recorded by Hedgpeth
(1948).
Characteristic of the species are the blunt eye tubercles, without eyes but
with distinct lateral sense organs; the ear-like lobes at the antero-lateral angles
of the cephalic segment; the (more or less distinct) groove immediately in front
of the first pair of lateral processes; the very short terminal claw of the legs;
Fic. 4. Colossendeis minuta Hoek. a, neck and proboscis, in dorsal view; 6, palp; c, terminal part
of the oviger; d, distal segments of the fourth leg; e, compound spine from oviger segment 10.
the shape of the proboscis, that is about as long as the trunk; the relative length
of the palp segments (3 as long as 5; 7 very long; 8, 9 and 10 subequal).
I consider Hoek’s C. japonica identical with minuta. Hoek stressed the great
similarity between the two forms, but kept them apart, chiefly because of the
different number of oviger spines and the relative length of the distal segments
of the legs. The number of oviger spines in Colossendeis is considered nowadays
of less value than in Hoek’s time, and as to the relative length of the distal
328 ANNALS OF THE SOUTH AFRICAN MUSEUM
segments of the legs, the differences are too slight to be considered of specific
value.
The ratios of tarsus, propodus, and claw of the 4th leg are:
in the holotype of C. minuta (cf. Gordon, 1944, p. 13, footnote) 1-5 : 1 : 0°46
in C. japonica (cf. Hoek, 1898) 1°5 fone
in C. minuta (present material) { BEM kis 2
145 pls Oren
The form recorded by Hedgpeth, 1949, and by Fage, 1956, under the
name of C’. japonica is quite distinct from Hoek’s species. It has, as distinct from
the real japonica, (1) a 5th palp segment that is at least 14 times as long as
the 3rd; (2) a proboscis that is much longer than the trunk (14-12 times as
long); (3) a very long terminal oviger claw; (4) the 6th and 7th palp segments
subequal. All these features, and also the general aspect of the animal (Hedg-
peth, 1949, fig. 46 a), indicate clearly that the animals in question belong to
the macerrima complex, instead of to the minutia group.
Measurements in mm.
Proboscis: specimen 1, 9°5; specimen 2, 10-0.
Trunk, including abdomen: specimen 1, 12°53; specimen 2, 9°5.
Length of legs: 75.
Fourth leg: femur, 23:0; Ist tibia, 23:5; 2nd tibia, 18-0; tarsus, 3:5; propodus,
2°42; claw, 0°83.
Colossendeis oculifera n.sp.
(Fig. 5)
Material. 21 specimens, including 1 juvenile. Station A 193.
Description. The trunk is unsegmented. The lateral processes are separated by
their own diameter. The conical eye tubercle varies somewhat in height (fig.
5 ¢, d); it is usually a very narrow cone with the slender apex directed forwards;
in some specimens it is shorter (due to damage ?). The 2 anterior eyes are much
larger than the 2 posterior ones; all are well pigmented (brown in alcohol).
Lateral sense organs, between the bases of the eyes, distinct. Cephalic segment
without ear-like projections on the anterior margin. Proboscis narrowly
cylindrical, straight, somewhat dilated in the middle, shorter than the rest of
the body. Ratios proboscis : trunk + abdomen of 5 specimens (in cm.):
0°60 : 1°00; 0:77 : 0:90; 0:72 : 0°83; 0-80 : 0:98; 0:63 : 0-70. The abdomen
is arched and somewhat clavate; it reaches to the middle of coxa 2 of leg 4.
Palp segment 3 slightly longer than 5; segment 7 twice as long as 6;
segments 8, 9 and 10 subequal. Palp segments not very setose.
Ovigers slender; the 6th segment is a trifle longer than the 4th; the terminal
claw is rather robust. The oviger spines on segments 7, 8 and g are longer than
those on segment 10; they are tongue-shaped, and bear minute denticulations
on the margin of their distal part.
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA 329
Legs not very slender. In a specimen in which the trunk + proboscis
measure 1°4. cm., the 3rd leg is 6:3 cm. In another specimen the femur is
1:95 cm., the Ist tibia 1-80 cm. and the 2nd tibia 1-73 cm.; the tarsus is 0-33 cm.,
the propodus 0:18 cm. and the claw 0-08 cm. The tarsus is always 14 to 12
times as long as the propodus. The ventral side of the tarsus bears some robust
spines; the sole of the propodus bears 4 or 5 spines, one or two of which are
very robust, in its distal fourth. The claw is much less than half as long as
the propodus. Female sexual pores are distinct on a slight swelling of the
ventral surface of the 2nd coxa of all legs. The juvenile in the present collection,
though still having an incompletely developed oviger, already possesses all the
characteristic features of the species: in the length of the proboscis, ocular
tubercle, relative length of palp and of leg segments, and propodal armature.
Fic. 5. Colossendeis oculifera n.sp. a, head and proboscis, in dorsal view; b, proboscis, from the left;
c, d, ocular tubercle, from the right, of two different specimens; ¢, distal segments of a leg;
J; propodus and claw; g, palp; h, distal part of the oviger; 7, compound spine from oviger seg-
ment 7; 7, abdomen in dorsal view.
330 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. The only species of the brevitarsal group that shares the presence of a
well-developed eye tubercle with a long 7th palp segment is C. hoeki Gordon,
1944. The latter has, however, the proboscis longer than the trunk, and the
claw of the legs ?—# as long as the propodus. Moreover, the eyes of C. hoeki,
though present, are unpigmented. We know nothing about the armature of
the propodus of C. hoekz, but since Gordon compared this species with C. gracilis,
C. media, C. brevipes, and C. minuta, all of which have practically unarmed propo-
dal soles, we may safely assume that the sole of C. hoeki does not show such a
curious armature as in the new species.
Longitarsal group
Colossendeis gracilis Hoek
(Fig. 6 a)
C. gracilis Hoek, 1881, pp. 69-70, pl. 9 figs. 6-8, pl. 10 figs. 6-7.
Material. 3 specimens, Station A 193; 6 specimens, Station A 315; 2 specimens,
Station A 316; 1 specimen, Station A 319; I specimen, Station A 322.
Remarks. 1 am aware that most pycnogonid specialists have synonymized
Hoek’s gracilis with C. angusta Sars, but the present material (which I have
compared with North Atlantic angusta) does not support this view. Calman,
1938, who was an advocate of the identity of gracilis and angusta, curiously
enough stressed plainly their differences.
The South African material, consisting of adults and juveniles, agrees with
C. gracilis (and consequently disagrees from C. angusta) in the presence of a
highly conical, pointed ocular tubercle (fig. 6 a), in the 3 short distal palp
segments (which are, combined, shorter than palp segment 7), in the small
size (length trunk + abdomen 5-10 mm. in gracilis, 15-20 mm. in angusta),
and in the relatively slender legs (6 to 7 times as long as the trunk + abdomen).
All specimens of C. gracilis in the present collection have neotenic chelifores,
a phenomenon also met with, though much less frequently, in C. angusta.
Of the published records, only the North Atlantic (and possibly the North
Pacific) ones belong to C. angusta. This species sometimes occurs in very shallow
waters. The deep-sea records from the tropics and from the Southern hemi-
sphere seem to belong to C. gracilis, but a revision of the material would cer-
tainly be necessary to make this statement more reliable.
Colossendeis curtirostris n.sp.
(Fig. 6 b-h)
Material. t 2 (holotype), Station A 192. 1 specimen, Station A 315. 2 specimens,
Station A 316.
Description of the holotype. Trunk elongated. Width across the 2nd lateral processes
only slightly more than half the length of the trunk. Lateral processes separated
by more than their own diameter. Intersegmental suture lines vaguely indicated
(in other specimens absent), certainly no functional articulations present.
ee ee eet ee a, tS a
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA 331
Fic. 6. (a) Colossendeis gracilis Hoek, front view of the ocular tubercle. (b—-h) Colossendeis curtirostris
n.sp. b, trunk in dorsal view; c, distal segments of the palp; d, distal segments of the first leg;
e, two special oviger spines; f, distal part of the oviger; g, ocular tubercle from the left; h, palp.
332 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ocular tubercle highly conical, apical part bent forwards; anterior pair of
eyes large, well pigmented, posterior pair very small, unpigmented (in some
other specimens also the anterior eyes are not well pigmented). No anterior
tubercles on the cephalic segment. Proboscis straight, only ? of the length of
trunk and abdomen combined, somewhat club-shaped. Abdomen ee
nearly to the distal end of coxa 1 of leg 4.
No trace of chelifores. Palp segment 3 about 14 times as long as segment 5;
of the 5 distal segments, segment 6 is the shortest, segment 7 the longest; the
latter is slightly less than 24 times as long as wide; segments 8, g and Io
diminish gradually in length. Oviger segment 4 shorter than segment 6. Distal
4 segments with very densely placed, smooth-edged spines, mostly of a very
peculiar, truncated appearance. Terminal oviger claw over half as long as the
10th segment.
Legs smooth, slender; 4th leg about 5 times as long as trunk and abdomen
combined. Femur the longer segment. Tarsus at least } longer than the propo-
dus. Claw at least equal to the propodus. Propodal sole practically unarmed.
The 2nd leg is the longest.
Remarks. The new species is apparently close to C. draket Calman and C. gracilipes
Bouvier (the latter being possibly synonymous with C. glacialis Hodgson).
It has the smooth legs, the slender, non-globular distal palp segments and the
long claw of draket, and may represent a northern subspecies of this Antarctic
and antiboreal shallow-water form.
C. curtirostris differs from C. drake: in the following respects: the much
shorter proboscis, the longer abdomen, the more widely spaced lateral pro-
cesses, the longer 2nd palp segment, the distal 4 palp segments that decrease
successively in length, the simple oviger spines, and the longer tarsus.
Another South Atlantic species, C. geoffroyi Mafie-Garzon, is considered
by its author to be close to C. drakei. This species differs from curtirostris in the
more slender 7th palp segment, in the very short 8th palp segment, and in the
lateral processes which are separated by less than their own diameter.
Measurements of the holotype (2) in mm.
Length of proboscis . , ve ssh nh ae ey Mh 7-3
Length of trunk + Abdoinen a 2) AORe
Palp—grd segment, 4:05; 4th segment, 0°51; an seomientt 2: ey 6th segment,
0-68; 7th segment, 0-91; 8th segment, 0-86; gth segment, 0-82; 10th
segment, 0°75
Diameter of 7th palp segment 0:38
Fourth leg—femur, 14:0; Ist tibia, 13-0; 2nd tibia, 11-5; tarsus, 5-5; propodus,
4:0; claw, 4:3
One of the specimens of Station A 316, a 9, has the following measurements:
proboscis, 8 mm.; trunk + abdomen, 11:5 mm.; width across the 2nd lateral
processes, 6 mm. ‘teotal length 4th leg, 82 mm. This specimen has much longer
legs (about 7 eines as long as trunk and abdomen combined) than the holotype.
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA 333
Fic. 7. Colossendeis orcadensis Hodgson, 3 (Station A 322). a, palp; 6, distal segments of the
oviger; c, compound oviger spines; d, ocular tubercle, from the right; e, proboscis in dorsal view;
f; proboscis from the right.
334 ANNALS OF THE SOUTH AFRICAN MUSEUM
Colossendeis orcadensis Hodgson
(Fig. 7)
Colossendeis orcadense Hodgson, 1908, pp. 161, 184-186, pl. 2 fig. 3.
Material. 1 flabby specimen, Station A 192; 4 small specimens, Station A 193;
1 large g, Station A 315; 1 small specimen, Station A 316; 1 large 3, Station
Avgoo.
Remarks. Since the original description in 1908, this species has not been found
again. ‘The holotype and only specimen known was taken at the South Orkneys.
The depth is not given, but in the introduction of Hodgson’s account we find
the following indication: ‘it [= C. orcadensis] is quite blind—not an unusual
character of the genus, but the more surprising as it is a shallow-water species.’
The present material, taken in the deep sea, is, as far as we can judge from
the poor illustration of the holotype, identical with C. orcadensis.
The following measurements are derived from a big male specimen
(Station A 315): Length of proboscis, 19 mm.; length trunk + abdomen,
15 mm.; length and leg, 119 mm.; femur, 34 mm.; Ist tibia, 27 @nmeomd
tibia, 22 mm.; tarsus, 12 mm.; propodus, 8-7 mm.; claw, 6-2 mm.
Another large male (from Station A 322) has distinct, although unpig-
mented, eyes. In all other specimens eyes are lacking. The specimen with eyes
comes close to C. megalonyx Hoek; as a matter of fact, I believe that, when more
material becomes available, we shall have to synonymize orcadensis and megalonyx.
Genus PANTOPIPETTA nom. nov.
It is a regrettable circumstance that the generic name Pzpetta Loman, 1904,
used universally in our pycnogonid literature, is preoccupied by Haeckel, 1887,
for a genus of Protozoa. Reluctantly, I propose the new name Pantopipetta for
the pycnogonid genus.
This curious deep-sea genus is represented in the present collection by
not less than 3 species: P. capensis (Barnard), which was hitherto known from
a single specimen only, P. brevicauda n.sp., and P. spec. (? longituberculata
Turpaeva). All are present in fair numbers of both sexes. It is remarkable that,
although several females are quite mature (bearing fully developed eggs in
their trunks and legs), no ovigerous males are known. This, together with the
structure of the oviger, in which segments 2, 4 and 6 are elongated, is a strong
indication for attributing the genus to the family Colossendeidae.
I insert here (fig. 8 f-h) some new illustrations of the genotype, P. weberi
(Loman), drawn from the holotype and only specimen known. This species is
chiefly characterized by its low ocular tubercle (which may, however, be the
result of a previous injury), but also by the very slender 2nd coxa and the
long 5th palp segment.
The species of the genus are very similar to one another; they are known
from some widely scattered localities in the deep sea: northern Pacific (Kurile-
Kamchatka trench), East Indies, Antarctic, and off South Africa.
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA 335
Fic. 8. (a—-e) Pantopipetta capensis (Barnard). a, trunk in dorsal view (2); 6, distal segments of the
palp (3); ¢, distal segments of the third leg (2); d, third leg (3); e, terminal part of the femur,
with the cement gland cone (¢). (f-h) Pantopipetta weberi (Loman), 9 (holotype). f, coxae;
g, distal part of the palp; A, distal segments of the leg.
336 ANNALS OF THE SOUTH AFRICAN MUSEUM
Key to the known species:
1 (a) Lateral processes smooth, 2.
(6) Lateral processes 2, 3 and 4 with a long dorsal spur; lateral process 1 with a dorsal
tubercle. P. capensis (Barnard, 1946).
2 (a) Propodus 5 to 6 times as long as the tarsus. P. australis (Hodgson, 1914).
(6) Propodus less than 4 times as long as the tarsus, 3.
3 (a) Abdomen reaching to the distal end of coxa 1 of leg 4. P. brevicauda n.sp.
(b) Abdomen reaching nearly or entirely to the distal end of coxa 2 of leg 4, 4.
4 (a) Ocular tubercle high conical. Coxa 2 about 4 times as long as its distal diameter. P.
longituberculata (Turpaeva, 1955).
(b) Ocular tubercle low. Coxa 2 at least 6 times as long as its distal diameter. P. weberi
(Loman, 1904).
Pantopipetta capensis (Barnard)
(Fig. 8 a—e)
Pipetia capensis Barnard, 1946, p. 60. Barnard, 1954, pp. 86-88, fig. 1.
Material. 8 9, 6 3 and 1 juvenile. Station A 193.
Remarks. Only 1 specimen, judging from Barnard’s figure a female, of this
interesting species was known. It was taken off Cape Point, N. 81° E. 32 miles,
in 460 fathoms.
The present material, from the same general region, but taken in a much
greater depth (1,500-1,580 fathoms) agrees perfectly with the description of
the type.
The sexual differences are feebly developed; no such differences are found
in the palps, ovigers and legs, with the exception of the presence in the male of
a cement gland cone on the femur. This ‘cone’ or projection is situated at a slight
distance from the distal end of the femur, somewhat variable in shape in
different individuals, but roughly triangular in outline. Nothing about this
cone was known in literature, although I suppose that the projection figured
very vaguely and described superficially by Hodgson (1927) on the femur of
P. australis is in fact the cement gland cone.
One of the specimens shows a curious anomaly: the first left leg, its corre-
sponding lateral process and the left oviger have disappeared, probably as a
result of a previous injury. Since no scar is visible, the animal in question offers
a peculiar unbalanced aspect.
Pantopipetta brevicauda n.sp.
(Figs. 9, 10 a)
Material. 3 3, 3 9, 1 juvenile. Station A 193.
Description. Very slender. Body and appendages appreciably more delicate
than in capensis. Trunk completely segmented. Lateral processes smooth,
separated by intervals that are 2 to 3 times as large as their own diameter
(slightly narrower between the lateral processes of segments 1 and 2). Eye
tubercle a very tall cone; no eyes. Abdomen separated from the trunk by a
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA 337
\
e
Cc
a
PO Ness
a
‘ Ae,
a eg
Fic. 9. Pantopipetta brevicauda n.sp. a, dorsal view of the trunk (¢); 6, ocular tubercle in front
view (3); ¢, special spine from oviger segment 8; d, palp (3); e, distal segments of palp (3);
J, third leg ($); g, distal segments of the third leg (3); A, femur (9).
338 ANNALS OF THE SOUTH AFRICAN MUSEUM
kind of suture, short, just reaching to the end of coxa 1 of leg 4. Proboscis
tubiform, longer than the trunk and the abdomen combined.
Palps practically as in capensis, the 5th segment being less than 3 times as
long as wide.
Ovigers also very similar to those of capensis (and of weberi). The formula of
the compound spines on the distal segments is in a dissected 9 3 :2:2:3::0
in a dissected ¢ 3:1:2:3::0. The shape of the compound spines is as in
capensis.
Legs very slender, except for the sexual pores and the cement gland very
similar in both sexes. Coxa 2 about 5 times as long as wide; coxa 3 with a long,
twig-shaped, flexible dorsal projection. Femur with an inconspicuous distal
spur; on the ventral surface of the femur of the 4, not far from the distal end,
a conical projection indicates the place where the cement gland discharges.
Tibia 1 subequal to the femur; tibia 2 much shorter. Tarsus about 2 of the
propodus. Claw half as long, or slightly less than half as long, as the propodus.
Inner margin of the claw armed, as in P. capensis and P. weberi, with irregular
denticulations. Ventral margin of propodus, tarsus and distal part of tibia 2
armed with a row of minute spinules.
Remarks. ‘Though occurring at the same station as P. capensis, the new species
differs clearly from it, even at first sight, by its much more delicate appearance.
The absence of projections on the lateral processes, the shorter abdomen and
the different proportions of the distal segments of the legs form useful additional
distinctions.
The only species which has about the same relative length of tarsus, propo-
dus and claw is P. webert. However, this East Indian species has the 5th palp
segment much slenderer (about 4 times as long as wide), a much longer
abdomen (reaching the end of coxa 2 of leg 4), and a short eye tubercle.
Measurements in mm.
¢ )
Length of proboscis ae on 3°07) aoaTio
Length (frontal margin of neck | to base of Abdomenyh ss 2-10 (21g
Length of abdomen Ve : - ae ee ie 0°53 0°65
Width across 2nd lateral processes a ys ae: ae 1°74 ey
Third leg—coxa 1 ae ae ee uh Re ee 0:28 = =—s_ 0:28
coxa 2 Up sh f nf Me es 0:98 1-02
COXa 3 a ur om ve fae + | 0290 on
femur ee ae Bi oe Me me I°QI ‘QI
tibia I a ee ae Bs x ae 1:86 1°86
tibia 2 us wv uy a a i 1°35 1°33
tarsus oe 40 °° 3'¢ ae a4 0°40 0°37
propodus.. big os Ne iN a 0-60 = 056
claw .. sie oe at on ep .. O26 ioe
SOUTH AFRICAN DEEP-SEA PYCNOGONIDA 339
Pantopipetta sp.
(Fig. 10 b-c)
Material. 1 3, 1 Q. Station A 316.
Remarks. The two available specimens can be distinguished from both capensis
and brevicauda by their much more robust appearance and less spaced lateral
processes. They come close to P. longztuberculata (Turpaeva), a deep-sea species
from the Kurile-Kamchatka trench, except for the relatively longer propodus
in the South African material, without attaining the enormous elongation
found in P. australis. In longituberculata the propodus is less than 3 times as long
5
sseeg |
ee
Il)
ie
Fic. 10. (a) Pantopipetia brevicauda n.sp., oviger (3). (b-c) Pantopipetta spec., 2. b, dorsal view of
the trunk; c, distal segments of the fourth leg.
340 ANNALS OF THE SOUTH AFRICAN MUSEUM
as the tarsus, in the present specimens more than 3 times as long, while in
australis it is 5 to 6 times as long. The lateral processes bear a low elevation on
their dorsal surface. The 9 has a proboscis of 4:6 mm.; the trunk is 3-6 mm. and
the abdomen 1-3 mm. in length. ‘The male is too much damaged to give exact
measurements.
Anoplodactylus pelagicus Flynn
Anoplodactylus pelagicus Flynn, 1928, p. 25, fig. 14. Barnard, 1954, p. 128, fig. 19.
Material. 1 § (damaged), Station A 316. 1 g¢ (damaged), off Cape Natal
(Durban), 24 miles, 440 fathoms (don. Dr. K. H. Barnard). Z.M.A. Pa. 1327.
Remarks. The specimen from Station A 316 differs from Barnard’s material
(described in his 1954 paper) in having a highly conical ocular tubercle. Eyes,
however, are, as in the previously recorded material, absent. Otherwise very
similar to the published descriptions.
The species has up to now not been recorded below 300 fathoms.
SUMMARY
A systematic account is given of a collection of Pycnogonida from the
abyssal zone of the sea south-west of Cape Town. The collection included 12
species, 5 of which are new.
ACKNOWLEDGEMENTS
The Trustees of the South African Museum wish to acknowledge a grant
from the Council for Scientific and Industrial Research towards the purchase
of the deep trawling equipment with which the specimens reported on were
collected, and a grant for the publication of this paper. Grateful acknowledge-
ment is also due to the Director, Division of Fisheries, Department of Com-
merce and Industries, for making the collecting possible.
REFERENCES
The works cited in this paper can be found in the bibliography of my 1956 paper; and
those published prior to 1935 also in Helfer & Schlottke. Not included in these bibliographies are:
Face, L. 1956. Les pycnogonides (excl. le genre Nymphon). Galathea Rep. 2, 167-182.
HE.LFER, H., AND SCHLOTTKE, E. 1935. Pantopoda. Bronn’s Klassen 5, Abt. 4, Buch 2, 1-314.
Stock, J. H. 1956. Tropical and subtropical Pycnogonida, chiefly from South Africa. Vidensk.
Medd. dansk naturh. Foren. Kbh. 118, 71-113.
TuRPAEVA, E. P. 1955. [New species of Pantopoda from the Kurile-Kamchatka trench.] Trud.
Inst. Okeanol. 12, 322-327. [In Russian. ]
[ CONTINUED FROM INSIDE FRONT COVER
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Hartman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.— All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name... .
B. Form of bibliographic references to synonymic names.—'The first reference following
any name in the synonymy should be to the earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalis capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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INDEX
XXXI.
XXXIT.
XXXII.
XXXIV.
XXXV.
XXXVI.
XXXVII.
XXXVIITI.
XXXIX.
XL.
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D7 .63
ANNALS
OF THE
PeUlh AFRICAN MUSEUM
VOLUME XLVI
Pano XIit
THE SOUTH AFRICAN SUBSPECIES OF THE BUFFY PIPIT,
ANTHUS VAALENSIS SHELLEY
By
J. M. WINTERBOTTOM
Percy FitzPatrick Institute of African Ornithology
and
South African Museum, Cape Town
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[ CONTINUED ON INSIDE BACK COVER
THE SOUTH AFRICAN SUBSPECIES OF THE BUFFY PIPIT,
ANTHUS VAALENSIS SHELLEY
By
J. M. WINTERBOTTOM
Percy FitzPatrick Institute of African Ornithology
and
South African Museum, Cape Town
[Accepted July 1962]
(With 1 figure in the text)
CONTENTS
PAGE
Introduction : : : : oar
Discussion . : ; : : = 342
Taxonomic List . i : : = 346
Summary . t : : : wT 350
Acknowledgements : ; : = 350
References . : ; : é 5 UXO)
Appendix: Statistical Analysis
by A. N. Rowan . : : 3 GH
INTRODUCTION
The Buffy Pipit Anthus vaalensis was originally described from Newcastle,
Natal, by Shelley in 1900. For some time it was regarded as a race of the Plain-
backed Pipit A. leucophrys and is so treated by Sclater (1930). Roberts, however,
pointed out that the two forms overlapped widely and must be regarded as
distinct species; and the problem was discussed in greater detail by White
(1948).
Three races of vaalensis have been described from within the limits of
geographical South Africa, for Roberts separated off a bird from Griqualand
West as daviesi in 1914 and others from the Chobe River as chobiensis in 1932.
All three subspecies were accepted by Vincent (1952); and Benson and White
(1957) and Smithers, Irwin and Paterson (1957) have recognized chobiensis
as distinct. Vaurie, White, Mayr and Greenway (1960) and White (1961),
however, synonymize daviesi with vaalensis, and chobiensis, together with marungen-
sis Chapin and muhingae White, with neuwmanni Meinertzhagen, originally
described (as Anthus leucophrys angolensis Neumann) from Ambaca, Angola.
Hall (1959), in her discussion of the Angola forms of A. leucophrys and A.
vaalensis, also considers it is impracticable to separate marungensis and muhingae
from neumanni, but does not discuss chobiensis.
541
Ann. S. Afr. Mus. 46 (13), 1963, 341-352, 1 fig. eee VA) Ai ist
342 ANNALS OF THE SOUTH AFRICAN MUSEUM
DiIscussIon
For the present study I was able to utilize a series of 241 skins; and I am
most grateful to the following for the loan of material: the Director, Mr. J. D.
Macdonald and Mrs. B. P. Hall of the British Museum (Natural History) ;
the Director (Mr. P. A. Clancey) of the Durban Museum and Art Gallery;
the Director (Miss M. Courtenay-Latimer) of the East London Museum; the
Director and Mr. O. P. M. Prozesky of the Transvaal Museum, Pretoria; the
Director and Mr. M. P. S. Irwin of the National Museum of Southern Rho-
desia, Bulawayo; and the Director (Dr. T. H. Barry) of the Albany Museum,
Grahamstown.
Current literature (Vincent, 1952; Vaurie et al., 1960) often includes the
whole of South Africa within the range given for Anthus vaalensis and its sub-
species; but this is misleading. The range is correctly given by McLachlan and
Liversidge (1957). There appear to be no authentic records of the occurrence
of this Pipit in the coastal strip of South Africa from Walvis Bay round to
Portuguese East Africa (records from Black Rock, east of East London, and
from the Bathurst District are due to confusion with A. leucophrys). On the
Karoo, the records are sparse and scattered in the west and south, more fre-
quent in the east. The sole record from the Transkei is the type of davies, from
Matatiele; from Natal the species is known only from Newcastle, Utrecht and
Blood River; and from the Orange Free State only from the neighbourhoods of
Bloemfontein and Vredefort Road. It becomes commoner in the Transvaal
and is widely distributed in Southern and Northern Rhodesia (Smithers,
Irwin and Paterson, 1957; Benson and White, 1957). In South West Africa
(Hoesch and Niethammer, 1940; Macdonald, 1957) it has only been recorded
from the north (Omaruru, Klein Otavi, Kaukurus, Franzfontein, Kamanjab).
It is not mentioned by Smithers (1959) or Smithers and Paterson (1959) from
Bechuanaland, but on the evidence of specimens is widely distributed there.
In Nyasaland (Benson, 1953) it is recorded from a few localities west of the
Shiré, but not from the east.
The three races described from south of the Zambezi were differentiated
solely on size. Roberts (1940) considered davies: to be the southern race and
larger than vaalensis (w., g, 109-112 mm., as against 100-108 mm.), and that
Ngamiland and Southern Rhodesian birds represented the smaller chobiensis
(w., 3, 98-99 mm.). Smithers, Irwin and Paterson (1957), however, conclude
that most Southern Rhodesian birds cannot be differentiated from typical
vaalensis, and this point is further discussed below. McLachlan and Liversidge
(1957) restrict the range of chobiensis to Ngamiland.
The present series shows that the average wing-lengths of males of various
populations vary with no very clear geographical divisions between 101-1 mm.
and 1053 mm., with two marked, and one less striking, exceptions. Birds from
north-eastern Bechuanaland Protectorate average markedly smaller (95:0 mm.)
and those from the near-by Caprivi Strip also run smaller (100-0 mm.)
S.A. SUBSPECIES OF THE BUFFY PIPIT ANTHUS VAALENSIS SHELLEY 343
‘Apricorn
~ -Trpic of Ca
490 Miles
)
2)
°
=
a
=
2
=)
a
i=}
=
bs
o
2
rr
E
2
E -]
t eonbokskeorkring.
23
5
-
2
RANGE oF Anthus vaalensis AND Irs SUBSPECIES IN SOUTHERN AFRICA
Each dot represents a locality from which at least one specimen has been examined. Type
localities of named subspecies, whether valid or not, are named. The approximate ranges
of four subspecies are indicated; the rest of the area is the range of Anthus vaalensis vaalensis
. and A. v. chobiensis (N. of about 23°S.).
344 ANNALS OF THE SOUTH AFRICAN MUSEUM
than the general population; and at the other extreme, birds from the Karoo
average larger (112-0 mm.), though near the coast (Klaarstroom, Uniondale,
Grahamstown, Zuurberg) they are smaller again (103:0 mm.). If we leave
these aberrant populations out of account, then there are two minor population
size-groups, that in Natal, the northern Cape, the Transvaal and southern
Bechuanaland Protectorate comprising bigger birds (w., g, av. c. 104-105 mm.)
and those in Southern and Northern Rhodesia and South West Africa smaller
(c. 101-103 mm.). This broad grouping follows Bergmann’s rule, but the rule
does not apply within the groups (e.g. the average of 17 gs from Selukwe,
IOI'l mm., is less than that of 17 gs from Northern Rhodesia, 102:3 mm.).
Nevertheless, the broad difference is highly significant statistically.
In females, the average wing-lengths fall into three groups: Karoo (105
mm.); the rest of the Cape, Natal, southern Bechuanaland, Orange Free State
and the Transvaal (99-101 mm.); and the rest (94-98 mm.), in which the
smallest birds are in northern Bechuanaland. Here again, Bergmann’s rule
is broadly conformed to, but not within each group (e.g. of the middle group,
the birds from the southern Cape are the smallest; Barotseland birds are bigger
than any Southern Rhodesian population; and Angola birds are bigger than
those from South West Africa). The two sexes agree in that Karoo birds are the
biggest; that birds from south of the Limpopo are larger than those from the
north; and that the smallest are those from northern Bechuanaland. ‘These
differences, too, are statistically significant.
Commenting on size-differences, White (1948) says: ‘The birds of the
high ground in the interior of the Cape Province seem to form a definable large
race. ... Those from northern Bechuanaland are smallest on the Chobe River,
but not worth separating on size alone.’
We may note that males with wings of 108 mm. or more (108 mm. being
the minimum given by White for male davies) occur sporadically as far north
as Ngoma (15° 54’ S.) in Northern Rhodesia; and that the shortest-winged
population in northern Bechuanaland occurs not on the Chobe but in the
Makarikari Pan area.
Turning now to colour, no recent worker has suggested any variation in
populations in geographical South Africa. Hall (1959), discussing the Angola
population, distinguishes two variants—a sandier form and a greyer form—_
the former predominating in the west, the latter in the east, so that, if considered
in isolation, Angola birds might be classified in two subspecies; but she points
out that ‘some from as far east as Mzimba, Nyasaland, are indistinguishable
from the co-types of newmanni’ (a sandy bird). She therefore considers it is
impracticable to recognize more than one subspecies; but she believes ‘that
A. v. neumanni can be distinguished from A. v. vaalensis by smaller size and less
sandy colour’.
In the present series, two populations stood out from the others on colour
grounds. One was from the neighbourhood of Bloemfontein, where the birds
are distinctly greyer above than surrounding populations, as well as being
S.A. SUBSPECIES OF THE BUFFY PIPIT ANTHUS VAALENSIS SHELLEY 345
bigger than any except that to the south (daviest), though the latter difference
is not statistically significant. A small series from Vredefort Road, 150 miles
farther north in the Orange Free State, probably belongs here too, but the
skins are old, faded and mostly very badly prepared, so that it is difficult to
be sure.
The second distinctive population is that from South West Africa, whence
the birds are decidedly redder above and below than the general series. It
should be noted, however, that a May bird from Darwendale, Southern
Rhodesia, fits perfectly into this series, and a number of other skins from the
Rhodesias approach the less extreme South West African examples fairly
closely. I have not seen any Angola specimens but I have been unable to
detect any general colour variation between series of birds from the Transvaal,
Natal, southern Bechuanaland, northern Cape, Southern Rhodesia and
Northern Rhodesia.
Summarizing the variation in this species, I consider that there are four
well-marked but very localized variations:
(i) A large-sized population in the higher ground of the Cape from Matatiele
to the west coast escarpment. For this the name davies? Roberts is available.
(it) A grey population in the Orange Free State. This is named, below,
clanceyt.
(iii) A small-sized population in north-east Bechuanaland. Unfortunately
the name chobiensis (Roberts) cannot be used for this population. Roberts
does not give the wing-measurement of the type of chobiensis separately ;
he only says ‘males, 98-99 mm.’; but I have measured the type twice
myself and asked Dr. G. J. Broekhuysen to measure it independently
and all three measurements were just over 101 mm. Of the three Chobe
birds available to me, two, including the type of chobiensis, are too big
to be placed in this small race, which I have accordingly named, below,
exasperatus.
(iv) A reddish population in South West Africa. In view of the evidence cited
above, I believe this to be indistinguishable from the birds named
neumannt Meinertzhagen, and propose to use this name for them, which is
the name used also by Hoesch and Niethammer (1940), though Mac-
donald (1957) calls his birds from this area chobiensis, and White (1948)
synonymizes neumanni with vaalensis but keeps chobiensis and marungensis
separate.
The rest of the population consists of birds intermediate in various degrees
between these four subspecies but divisible at the Limpopo into two populations,
on average size. If we divide the males into those with wings of 105 mm. and
over and those with wings of 104. mm. or less, than 20 out of 29 (69%) of the
birds from south of the Limpopo fall into the former class and 78 out of 101
(77%) of those from north of that river into the latter. Treating the females
346 ANNALS OF THE SOUTH AFRICAN MUSEUM
similarly but drawing the line between 98 and 99 mm., then 23 out of 27 (85%)
from south of the Limpopo fall into the large group and 55 out of 70 (78%)
from north of that river into the small one. The combined total separable in
size alone is 176 out of 227 (78%). If it is considered that this is sufficient to
warrant nomenclatorial recognition, then the birds north of the Limpopo
must be called A. v. chobiensis (Roberts). Considerable differences exist as to the
exact meaning of the ‘seventy-five per cent’? rule (Amadon, 1949), but in
view of the comments of Mr. A. N. Rowan in the Appendix, I propose to
revive chobiensis for the birds north of the Limpopo.
Two possible explanations of the curious distribution pattern exhibited
by the races of this species may be advanced: (i) A uniform population is in
the process of differentiating into geographical subspecies; (ii) the species is
in the course of extending, or has recently extended, its range, bringing into
contact discrete populations previously isolated. The latter seems to fit the
facts better, and also seems inherently the more probable. White (1948) points
out that, although A. vaalensis and A. leucophrys frequently occur on the same
ground, yet on the whole vaalensis prefers drier and leucophrys damper conditions.
During the last pluvial (or hypothermal) period, the areas suitable for vaalensis
would have been markedly more restricted than they are today; and I suggest
that the four subspecies, daviesi, clancey1, exasperatus and neumanni, were differen-
tiated at that time. With the increasing desiccation since then, the species has
expanded its range and a ‘hybrid swarm’ has come to occupy most of the area,
including, probably, some of the original ranges of the four distinct subspecies,
and has begun to ‘swamp’ their distinctiveness. The ‘newmanni’ specimen from
Darwendale has already been mentioned; and a January female from 3 m.
south of Nata, with a wing of 102 mm., 6 mm. longer than the wing of any
other female and 4 mm. longer than that of any male from within the range of
exasperatus, is another example: though in view of the nomadic habits of Anthus
spp. in the off-season, the possibility that these are stragglers cannot be excluded,
even if the date renders it unlikely in the second case.
TAXONOMIC LIST
Taxonomically, six subspecies may be recognized.
1. Anthus vaalensis vaalensis Shelley
Anthus vaalensis Shelley, 1900, 311 — Newcastle, Natal.
Plumage, size and degree of sexual dimorphism intermediate between
the other races. Greyish brown above, unstreaked; buff below, usually ss
some dark flecking on the breast, but never very ain
Range: South Africa from the Little Karoo and inland eastern Cape to the
Transvaal and southern Bechuanaland, except those areas occupied by the
next two races.
S.A. SUBSPECIES OF THE BUFFY PIPIT ANTHUS VAALENSIS SHELLEY 347
Measurements: 29 38; wing 97-110 mm., av. 104:9 mm.; tail 69-80 mm.,
av. 74°4 mm.; culmen 17-21 mm., av. 19°5 mm.; tarsus 24-30 mm., av. 26:9
mm.; hind-claw 8-12 mm., av. 9°5 mm.
22 9s; wing 94-105 mm., av. 100°1 mm.; tail 64-76 mm., av. 70:8 mm.;
culmen 17-20 mm., av. 18-7 mm.; tarsus 25-28 mm., av. 26-7 mm.; hind-claw
8-II mm., av. 9°3 mm.
Material examined :
Cape Province: South African Museum 3 (Klaarstroom, Kimberley,
Molopo R.); Durban Museum 6 (Uniondale, Riverton, Griquatown-Niekerks-
hoop); Albany Museum 4 (Grahamstown, Zuurberg, Kimberley); National
Museum, Bulawayo 3 (Riverton, Langley, Kuruman); British Museum 1
(‘Cape of Good Hope’); total, 17.
Natal: British Museum 1 (Ingangani R.): Durban Museum 2 (Blood R..,
Utrecht); total, 3.
Transvaal: British Museum 3 (Rustenburg, Lydenburg, Potchefstroom) ;
Durban Museum 8 (Piet Retief, 10 m. E. of Wakkerstroom); Transvaal
Museum 15 (Hammanskraal, Pretoria, Groot Spelonken, Bloemhof, Wol-
maransstad, Barberton, Carolina, Warmbaths); Albany Museum 1 (Pieters-
burg); South African Museum 1 (‘Transvaal’); total, 29.
Swaziland: Transvaal Museum 1 (Ndhlovudwalili) ; total, 1.
Bechuanaland: National Museum, Bulawayo 7 (23 and 25 m. S. of
Letlaking, Lephepe-Dabeete, 41 m. N.W. of Molepole, Lotlelekane Wells) ;
totale 7. <>
2. Anthus vaalensis daviest Roberts
Anthus daviesi Roberts, 1914, 172— Matatiele.
Larger than vaalensis and with less sexual dimorphism of size.
Range: ‘The higher parts of the Cape Province, from Matatiele to Nieu-
woudtville and the Cold Bokkeveld, north to Philipstown and Colesberg.
Occasional birds from as far north as Ngoma, in Northern Rhodesia, are
inseparable.
Measurements: 6 3s; wing 108-113 mm., av. 112°0 mm.; tail 70-83 mm.,
av. 75°3 mm.; culmen 18-20 mm., av. 19°5 mm.; tarsus 23-30 mm., av. 27°6
mm.; hind-claw 9-10 mm., av. 9°6 mm.
I 9; wing 105 mm.; tail 71 mm.; culmen 20 mm.; tarsus 26 mm.; hind-
claw II mm.
Material examined: South African Museum 4 (Nieuwoudtville, Cold
Bokkeveld, Hanover, Glen Grey); British Museum 1 (Deelfontein) ; Transvaal
Museum 1 (Matatiele (type, daviesi)); Albany Museum 1 (Colesberg) ; Durban
Museum 1 (Steynsburg); total, 8.
3. Anthus vaalensis clanceyt subsp. nov.
Decidedly greyer than vaalensis or daviesi; intermediate in size; virtually
no sexual dimorphism of size.
348 ANNALS OF THE SOUTH AFRICAN MUSEUM
Type: Ad 4, in Durban Museum: ‘Glen Lyon’, Glen, Modder River,
Bloemfontein; collected by Durban Museum personnel, 8 June, 1960.
I name this subspecies after Mr. P. A. Clancey, who first drew my attention
to it.
Range: The Central Orange Free State, from Bloemfontein to (?) Vrede-
fort Road.*
Measurements: 8 gs; wing 103-109 mm., av. 105°7 mm.; tail 72-79 mm.,
av. 74°7 mm.; culmen 18-19 mm., av. 18:5 mm.; tarsus 25-30 mm., av. 27:2
mm.; hind-claw 8-10 mm., av. 9:4 mm.
5 Qs; wing 102-111 mm., av. 106-8 mm.; tail 70-83 mm., av. 74:4 mm.;
culmen 18-19 mm., av. 18-4 mm.; tarsus 25-28 mm., av. 26:4 mm.; hind-claw
g-10 mm., av. 9°4 mm.
Material examined: Durban Museum 6 (Glen (type, clanceyz)); British
Museum 3 (Vredefort Road); Albany Museum 2 (Vredefort Road, Bloemfon-
tein); East London Museum 2 (Glen); total, 13.
4. Anthus vaalensis exasperatus subsp. nov.
Smaller than vaalensis or any other race (w. normally less than 100 mm.);
little sexual dimorphism in size (w. 9, 99% of 3).
Type: Ad 3, in National Museum of Southern Rhodesia, Bulawayo, No.
29147, 3 m. S. of Nata, Bechuanaland Protectorate; collected 28 January,
1957, collector’s number M.118. |
Range: North-eastern Bechuanaland, from the Nata River to Francistown.
Occasional males from Matabeleland and Northern Rhodesia are inseparable;
and females are only distinguishable in series.
Measurements: 4 gs; wing 91-98 mm., av. 95:0 mm.; tail 64-69 mm., av.
67-0 mm.; culmen 18 mm., av. 18-0 mm.; tarsus 25-29 mm., av. 26-7 mm.;
hind-claw 10 mm., av. 10:0 mm.
7 Qs; wing QI-102 mm., av. 94:4 mm.; tail 60-71 mm., av. 67-9 mm.;
culmen 17-19 mm., av. 18-1 mm.; tarsus 24-28 mm., av. 26-7 mm.; hind-claw
G-1T im AV. Ora mate
Note that females of this race are less obviously smaller than those of
neighbouring populations (S.W.A., w. av. 94:7 mm.; Caprivi Strip, w. av.
95 mm.) than are the males, in correlation with the lesser sexual dimorphism of
size shown by this race.
Material examined: National Museum, Bulawayo 12 (Mache, Nata (type,
exasperatus), 4m. 8S. of Francistown, Makarikari); total, 12.
5. Anthus vaalensis chobiensis (Roberts)
Meganthus vaalensis chobiensts Roberts, 1932, 29— Kabulabula.
Anthus leucophrys marungensis Chapin, 1937, 342—Kaschi, Marungu, Belgian Congo.
Anthus vaalensis muhingae White, 1944, 6— Muhinga plain, Kasendoge, Katanga.
* Since the above was written, Mr. Clancey has found specimens of this race in Griqualand
West.
S.A. SUBSPECIES OF THE BUFFY PIPIT ANTHUS VAALENSIS SHELLEY 349
Similar to A. v. vaalensis but smaller; larger than A. v. exasperatus, especially
in the males.
Range: From the Limpopo to western Nyasaland, Northern Province of
Northern Rhodesia, Katanga and eastern Angola, except the north-eastern
corner of Bechuanaland.
Measurements: 99 3s; wing 90-109 mm., av. 101-9 mm.; tail 60-79 mm.,
av. 72°0 mm.; culmen 15-22 mm., av. 18-9 mm.; tarsus 21-29 mm., av. 24:2
mm.; hind-claw 7-12 mm., av. 9:1 mm.
78 Qs; wing 90-105 mm., av. 96-4 mm.; tail 60-78 mm., av. 69:4 mm.;
culmen 16-21 mm., av. 18-4 mm.; tarsus 2I-30 mm., av. 25:7 mm.; hind-claw
7-1) mm, av. Ol mim.
Material examined
Southern Rhodesia: Durban Museum 1 (Tjolotjo); National Museum,
Bulawayo 104 (Gazuma Pan, Wankie Game Reserve, Kana R., Dorset Siding,
Hopefountain, Nyamandhlovu, Copleston, Tjolotjo, Lupane, Bulawayo,
Shangani R., Matopos, Bembezi, Inseze, Limpopo R., Khami, Sebungwe,
Umvuma, Darwendale, Headlands, Salisbury, Rusape, Inyanga, Sabi-Lundi
confluence, Hunyani R., Selukwe); Albany Museum 2 (Matopos); total, 107.
Northern Rhodesia: South African Museum 2 (Balovale, Lusaka) ; National
Museum, Bulawayo 33 (Ngoma, Mazabuka, Choma, Lochinvar, Chuunga,
Chilanga, Lusaka, Lukanga Swamp, Luanshya, Ntemwa-Lufupa confluence,
Moshi-Lufupa confluence, Mweru Marsh, Mpika, Mwinilunga, Kasanga,
Mporokoso, Kasusu, 8 m. N. of Nangweshi, Matabele Plain, Chiolola, Chunga
Pools, Luete, Senenga) ; total, 35.
Caprivi Strip: Transvaal Museum 3 (Kabulabula (type, chobdiensis),
Kabuta); total, 3.
6. Anthus vaalensis neumanni Meinertzhagen
Anthus leucophrys neumanni Meinertzhagen, 1920, 23—Ambaca, Angola.
Decidedly redder than any other race.
Range: South West Africa and western Angola. Some specimens from
eastern Angola, the Rhodesias and Nyasaland are indistinguishable from this
form.
Measurements (S.W.A. birds only); 3 gs; wing 101-103 mm., av. 102:0 mm. ;
tail 73-76 mm., av. 74:7 mm.; culmen 17-18 mm., av. 17°7 mm.; tarsus
24-26 mm., av. 25:0 mm.; hind-claw g-12 mm., av. 10-3 mm. (Hall, 1959,
gives w. 93-105 for Angola birds, and White, 1948, 95-106 mm. for S.W.A.
birds.)
3 9s; wing 92-98 mm., av. 94:7 mm.; tail 63-73 mm., av. 67:7 mm.;
culmen 17-18 mm., av. 17:7 mm.; tarsus 27-28 mm., av. 27:7 mm.; hind-claw
g-I2 mm., av. 10°3 mm. (Hall gives w. 89-102 mm. for Angola birds and
White 94-99 mm. for S.W.A. birds.)
350 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material examined: Durban Museum 4 (Omaruru); British Museum 2
(Kamanjab, Franzfontein) ; total, 6.
SUMMARY
The forms of Anthus vaalensis are discussed, and six subspecies: vaalensis
Shelley, daviest Roberts, clanceyt subsp. nov., exasperatus subsp. nov., chobiensis
Roberts, and newmanni Meinertzhagen, are recognized. A statistical analysis is
made of the measurements of the specimens.
ACKNOWLEDGEMENTS
I am deeply indebted to Mr. A. N. Rowan for statistical analysis of the
measurements; to Mr. P. A. Clancey for drawing my attention to the peculiar
series from the Orange Free State; and to Miss A. Schweitzer for the map.
REFERENCES
Amapon, D. 1949. The Seventy-five per cent Rule for Subspecies. The Condor 51, 250-8.
Benson, C. W. 1953. A Check List of the Birds of Nyasaland. Blantyre.
Benson, C. W., AND Wuirte, C. M. N. 1957. A Check List of the Birds of Northern Rhodesia. Lusaka:
Government Printer.
Cuapin, J. P. 1937. The Pipits of the Belgian Congo. Rev. Zool. Bot. afr. 29, 336-45.
Hatt, B. P. 1959. The Plain-backed Pipits of Angola. Bull. Brit. Orn. Cl. 79, 113-16.
Hoescu, W., AND NIETHAMMER, G. 1940. Die Vogelwelt Deutsch-Siidwestafrikas. 7. Orn., Lpz.
88, Sonderh.
Macpona.Lp, J. D. 1957. A Contribution to the Ornithology of Western South Africa. London: British
Museum (Natural History).
McLacuian, G. R., AND LIVERSIDGE, R. 1957. Roberts’ Birds of South Africa. Cape Town: South
African Bird Book Fund.
MEINERTZHAGEN, R. 1920. New subspecies of birds. Bull. Brit. Orn. Cl. 41, 19-25.
RosBeErts, A. 1914. Notes on birds in the collection of the Transvaal Museum with descriptions
of several new sub-species. Ann. Transv. Mus. 4, 169-79.
Roserts, A. 1932. Preliminary descriptions of sixty-six new forms of South African birds.
Ann. Transv. Mus. 15, 21-34.
Roserts, A. 1940. The Birds of South Africa. London: Witherby.
ScLATER, W. L. 1930. Systema Avium A&thiopicarum. London: British Ornithologists Union.
SHELLEY, G. E. 1900. The Birds of Africa, vol. 2. London: R. H. Porter.
SMITHERS, R. H. N. 1959. Notes on the Avifauna of the South-West Kalahari Area of Bechuana-
land Protectorate. Proc. Pan. Afr. Orn. Cong. Ostrich, Suppl. 3, 126-32.
SMITHERS, R. H. N., Irnwin, M. P. S., anp PATERsON, M. 1957. A Check List of the Birds of Southern
Rhodesia. Bulawayo: Rhodesian Ornithological Society.
SmiTHERs, R. H. N., AND Paterson, M. 1959. Notes on the Avifauna of the Eastern Sector of the
Great Makarikari Lake, Bechuanaland Protectorate. Proc. Pan. Afr. Orn. Cong. Ostrich,
Suppl. 3, 133-43.
VauriE, C. H., Wuits, C. M. N., Mayr, E., AND GREENWAY, J. C. 1960. Family Motacillidae.
In Check List of Birds of the World. g (ed. Mayr and Greenway). Cambridge, Mass.: Museum
of Comparative Zoology.
VINCENT, J. 1952. A Check List of the Birds of South Africa. Cape Town: South African Bird Book
Fund.
Wuirte, C. M. N. 1944. New Races of Lark, Pipit and Cordon Bleu from Northern Rhodesia,
Angola, and the Belgian Congo. Bull. Brit. Orn. Cl. 65.
Wuire, C. M. N. 1948. The African Plain-backed Pipits—a Case of Sibling Species. The Ibis go,
547-53:
Wurite, C. M. N. 1961. A Revised Check List of African Broadbills, Pitias, Larks, Swallows, Wagtatls
and Pipits. Lusaka: Government Printer.
S.A. SUBSPECIES OF THE BUFFY PIPIT ANTHUS VAALENSIS SHELLEY 351
APPENDIX
STATISTICAL ANALYSIS OF THE NUMERICAL DATA
By
A. N. Rowan
In his study of Anthus vaalensis, Winterbottom measured the lengths of
wing, tail, culmen, tarsus and hind claw in 258 museum specimens, of which
150 were males. These data, arranged geographically, were submitted for
statistical analysis. As a first step, male wing lengths were tested for homo-
geneity using the F test, and were found to be very significantly heterogeneous,
thus indicating that there were statistically valid differences between various
populations.
A preliminary study of the figures suggested that male wing lengths tended
to decrease with decreasing latitude, and the possible existence of a cline was
examined by plotting these two measurements against each other. However,
the correlation coefficient was not significantly different from zero, showing
that a cline did not exist.
From a further inspection of the data it appeared (1) that males from
northern Bechuanaland had smaller wings and tails than other groups; and
(2) that the entire series could be separated into two groups, one with larger
dimensions south of the Limpopo River, and the other comprising smaller
birds north of that river. Accordingly, the ‘t’ test was employed to test for
significant differences between the postulated northern and southern groups
and between them and the birds of northern Bechuanaland. Tables A and B
give the results for wing, tail, tarsus and culmen lengths. The dimensions of the
hind claw were omitted from this analysis since they showed no clear relation
to other measurements and the over-all differences in length were too small in
relation to the units of measurement employed.
From Table A it is clear that southern males are very significantly larger
than their northern counterparts in all measurements and that southern females
are significantly larger in all measurements except culmen length.
TABLE A: COMPARISONS BETWEEN NORTHERN AND SOUTHERN BIRDS
AVERAGES
Southern Northern
Character Sex group CB: group t Probability
mm. mm.
Wing length Male 105-80 0-60, 101-76 2°58 0:01-0:02
Female 101 °33 0:70 96°53 >6 Far less than 0-001
Tail length Male 74°50 0°45 71°64 3°40 Less than 0-:01%
Female 71°60 0°25 69°78 2°24 0:*02-0°05
Length of tarsus Male PG 5 0°33 26-06 3:26 Less than o-o1
Female 26°63 0°34 25°59 2°53 O°OI-0'02
Length of culmen Male 19°23 0:27 18-72 2°47 O°OI-0°02
Female 18-63 a-12 18-39 1-11 Not significant
352 ANNALS OF THE SOUTH AFRICAN MUSEUM
These results were further examined using the Coefficient of Difference,
as recommended by Mayr et al. (Methods and Principles of Systematic Xoology,
New York, 1953: 146). Table A shows the C.D. for various combinations.
The conventional level of subspecific difference is 1:28. None of the coefficients
of difference approaches this figure. Thus, using this criterion, it would appear
that no single character is sufficient to indicate subspecific difference, but if all
of them are considered together a good case might be made for subspecific
difference between the populations north and south of the Limpopo.
Table B gives similar figures for northern Bechuanaland birds and those
of the northern groups. It can be seen that while the northern males are larger
than the northern Bechuanaland birds in wing, tail and culmen length the
difference is highly significant only in wing length and moderately significant
in tail length. The females of the two groups are not significantly different. In
no case does the coefficient of difference reach the conventional level for sub-
specific difference. In view of the few specimens of northern Bechuanaland
birds examined and because only male wing length is very significantly larger,
it is considered very doubtful whether they should be separated on the mensural
data alone.
TABLE B: COMPARISONS BETWEEN NORTHERN BIRDS AND NORTHERN
BECHUANALAND BIRDS
AVERAGES
Northern
Character Sex Northern Bechuanaland t Probability
birds birds
mm. mm.
Wing length Male 101+ 76 95°00 3°51 Less than 0-01
Female 96-53 94°23 1:76 Not significant
Tail length Male 71°64 67:00 2°58 0-'OI-0:02
Female 69:78 67-86 1:50 Not significant
Length of tarsus Male 26-06 26°75 0:82 Not significant
Female 25°59 26°71 1°41 Not significant
Length of culmen Male 19°72 18-00 1:24 Not significant
Female 18-39 18-14 0°65 Not significant
[ CONTINUED FROM INSIDE FRONT COVER
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrdége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
HartMan, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.—All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name... .
B. Form of bibliographic references to synonymic names.—'The first reference following
any name in the synonymy should be to the earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order. . . .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalis capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulaha viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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INDEX
XXXII.
XXXII.
XXXII.
XXXIV.
XXXV.
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XXXVII.
XXXVITI.
XXXIX.
XL.
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XLII.
XLII.
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1900-1902
1903-1905
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1906-1910
1908-1910
1908-1913
IQII—-1918
IQII-I1914
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1914-1916
1917-1933
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1921
1924-1925
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ANNALS
OF THE
Pol ih sank AN MUSEUM
VOLUME XLVI
PART XIV
POLYCHAETE FAUNA OF SOUTH AFRICA: PART 7
SPECIES FROM DEPTHS BETWEEN 1,000 AND 3,330 METRES
WEST OF CAPE TOWN
By
: J. H. Day
Koology Department, University of Cape Town
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[ CONTINUED ON INSIDE BACK COVER
POLYCHAETE FAUNA OF SOUTH AFRICA: PART 7
SPECIES FROM DEPTHS BETWEEN 1,000 AND 3,330 METRES
WEST OF CAPE TOWN
By
Alt DAY,
[Accepted July 1962]
Koology Department, University of Cape Town
(With 3 figures in the text)
CONTENTS
PAGE
Introduction 353
Systematic section 355
Summary . ; 271
Acknowledgements 371
References . 371
INTRODUCTION
At the request of Dr. F. H. Talbot of the South African Museum, the
Director of the Division of Fisheries kindly allowed R.V. Africana IJ to make a
series of deep-water trawls on the continental slope west of Cape Town during
August 1959, and a further series during December. The 15-foot Agassiz trawl
brought up rich hauls of fish and invertebrates, and I am grateful to Dr. Talbot
for allowing me to examine the Polychaeta. I have taken the opportunity of
describing in this paper not only Dr. Talbot’s collection, but also material
from the same area and depth range obtained by means of a 0-1m.? van Veen
grab operated from the University of Cape Town’s research vessel, the John D.
Gilchrist. As might be expected, the grab sample contained several small species
which were absent from the trawl samples.
Details of the stations which contained Polychaeta are as follows:
Vessel Station Date Position Depth (m.) Bottom
Africana II
JENS TUBIO) RISO) (0) he 1g] 21k de 1098 Globigerina ooze
ASI QO!) 25/0/59 93926) 5-/ 10733). 2269 e a
ano N2O)(3)/ 501682290) 9) TOukny Eau lin 27/45 5 93
A.193 27/8/59 + 33°49'S./16°30"E. = 2745 39 25
A.315 8/12/59 34°37'S./17°05'E. —- 2930 ») :
A.316 8/12/59 34°42'S./16°54’E. = 3203 29 »
LA BNG OVNAix0) PS0) iS yiilOeeio) 10, 2850 55 3
A.318 9/12/59 33°52’'S./16°51’E. 2520-2780 A; Ep
A.319 = 9/12/59 = 3.4.°05'S./16°58’E. 2700 ss ;
A.321 10/12/59 34°33'S./16°42’E. 3330 os =
A.322 10/12/59 34°36'S./17°00"E. = 2745 >» se
J. D. Gilchrist
WCD 73-75 ~=8/10/60 =34°25’S./17°36’E. 1240 on. S.M.
393
Ann. S. Afr. Mus. 46 (14), 1963, 353—371, 3 figs.
eo ||
354 ANNALS OF THE SOUTH AFRICAN MUSEUM
These are not the first deep-water collections made in this area. In November
1921 the late Professor Gilchrist occupied several abyssal stations off Cape
Town. The Polychaeta from these stations were described by McIntosh (1925).
The total list of species recorded from depths exceeding 1,000 metres in this
area is shown below.
Species McIntosh 1925 Present records
Aphrodita alta Kbg. x
Hermonia hystrix (Sav.) x
Laetmonice benthaliana Mcl. x
= L, filicornis (Mcl. non Kbg.) x
Eunoe assimilis Mcl.
Eunoe macrophthalma Mcl. x
Harmothoe profunda n. sp. x
Harmothoe sp.
Euphione elizabethae Mcl. x
Macellicephala mirabilis Mcl.
Leanira hystricis Ehl. x
Leanira tetragona (Oersted)
Nereis (Neanthes) papillosa n. sp.
Nephthys (Aglaophamus) macroura Schm.
Nephthys hystricis Mcl.
Nephthys paradoxa Malmgren
Hyalinoecia sp.
Lumbrineris brevicirra (Schm.)
Lumbrinerts magalhaensis Kbg.
Paraonis gracilis oculata Hartman
Aricidea suecica simplex nov.
Prionospio steenstrupi. Malmgren
Spiophanes soderstromt Hartman
Poecilochaetus serpens Allen
Notomastus latericeus Sars. x
Maldanella fibrillata Chamberlin
Lumbriclymene minor Arwidsson
Phalacrostemma elegans Fauvel
Neosabellides elongatus (Ehl.)
Amphitrite cirrata Miller
Streblosoma abranchiata n. sp.
Streblosoma chilensis (MclI.)
Terebellides stroemi Sars
Pista cristata (Miiller)
x
x
x
x
Wee Xe Oe DS Pe ee He OG Ow
eS eK OG XK
x
The list includes 31 identified species of which 4 are new. A further 9
species are new records for South Africa; in fact only 4 of the 31 species listed
are known to occur at depths less than 100 metres in South African seas. Many
POLYCHAETE FAUNA OF SOUTH AFRICA 355
of the others such as Laetmonice benthaliana, Euphione elizabethae, Macellicephala
mirabilis, Maldanella fibrillata and Phalacrostemma elegans are abyssal species
from such widely separated regions as the sub-Antarctic, the North Pacific,
New Zealand, the Gulf of Panama and Madeira. This does not indicate any.
affinity between the faunas of these areas, merely that abyssal species are very
widely distributed and poorly known. It also emphasizes the importance of
indicating the depth when listing distribution records.
The study of this collection was facilitated by side-by-side comparisons
with the rich collections housed in the British Museum. I wish to thank my
many friends at the British Museum for their help while I worked there and
the Oppenheimer Memorial Trust which financed my research.
The material described in this report has been deposited in the South
African Museum, Cape Town, and the registration number of each species
is given in the text.
SYSTEMATIC SECTION
Family Aphroditidae
Sub-family Hermoninae
Genus Laetmonice Kinberg 1855
Both L. filicornis Kbg. and L. producta var. benthaliana MclI. have been
recorded from South Africa but the published descriptions do not indicate
clearly the differences between them. Before the new material was identified
specimens of Laetmonice in the British Museum were examined. The results are
set out below.
Laetmonice filicornis Kinberg 1855 (type locality Sweden)
Material examined: British Museum specimen 1865 :9 : 23:13 from
Bohuslan, Sweden, and 1921 : 5 : 1 : 94-100 from the Shetland Islands.
The two samples are conspecific and the following description is based
on the Bohuslan specimen. Length 30 mm. for 32 segments. Ommatophores
not pigmented. Tentacular segment without purse-shaped lobes at the sides
of the prostomium. 15 pairs of elytra. A felt present. Harpoon-setae numerous
and much larger than the notopodial spines of cirrigerous feet. Ventrum covered
with rounded papillae. Ventral cirri well developed, their tips reaching the
bases of the inferior neurosetae.
Laetmonice producta Grube 1877 (type locality Kerguelen)
Material examined: British Museum specimens 1941 : 3 : 3 : 439-50 from
Kerguelen.
Length 40-100 mm. for 42-46 setigers. Each ommatophore with a small
eye-spot. Tentacular segment with a pair of purse-shaped lobes at the sides
of the prostomium. 18 pairs of elytra. No felt. Harpoon-setae rare and the
notopodial spines of cirrigerous feet as strong as those of elytrigerous feet.
Ventrum covered with rounded papillae. Ventral cirri are well developed
and reach the bases of the inferior neurosetae.
356 ANNALS OF THE SOUTH AFRICAN MUSEUM
Laetmonice producta var. wyville: McIntosh 1885
Material examined: British Museum type 1885: 12:1 : 34 dredged off
Prince Edward Island, and type 1885 : 12:1 : 35 from Challenger station
157 in the Antarctic.
The first is a specimen of L. producta differing from Kerguelen specimens
only in the possession of smaller lobes on either side of the prostomium and
indistinct eyes. All other characters agree. The second, chosen as the lectotype
of L. producta var. wyvillei, may be described as follows:
Length 40 mm. for 42 segments. Ommatophores not pigmented. Tenta-
cular segment without lobes at the sides of the prostomium. 18 pairs of elytra.
Felt poorly developed. Harpoon-setae well developed and stronger than the
notopodial spines of cirrigerous feet. Ventrum with a few papillae. Ventral
cirri are small and do not reach the bases of the inferior neurosetae.
Laetmonice benthaliana McIntosh 1885
Material examined: ‘Types of L. producta var. benthaliana in the British —
Museum numbered 1885 :12:1:1 from Challenger station 147 off Prince
Edward Island (lectotype). 1885 : 12:1 :36 from Challenger station 244
in the Pacific. 1885 : 12: 1:37 from Challenger station 241 in the North
Pacific. Types of L. producta var. willemoest in the British Museum numbered
1885 : 12:1 : 31 from Challenger station 169 off New Zealand. 1885 : 12 : 1 : 32
from Challenger station 146 off Prince Edward Island.
The lectotype measures 35 mm. for 32 segments. Ommatophores not
pigmented. Tentacular segment without lobes at the sides of the prostomium.
15 pairs of elytra. Felt present but scanty (? lost). Harpoon-setae common
and markedly larger than the notopodial spines of cirrigerous feet. Ventrum
smooth. Ventral cirri are very small and do not reach the bases of the inferior
neurosetae.
Key To Laetmonice producta, producta wyvillet, filicornis AND benthaliana
1. Body with 42 or more segments; 18 pairs of elytra Li s) 3 ae oe hee
Body with about 32 segments; 15 pairs of elytra
2. Purse-shaped lobes at the sides of the prostomium. No felt. Notopodial Seiiaes ae cirri-
gerous feet as strong as those of elytrigerous feet ie ie .. L. producta
No purse-shaped lobes. A scanty felt. Notopodial spines of sinieecous feet weaker
than those of elytrigerous feet .. : Ae L. producta wyvillei
3. Ventrum covered with rounded papillae. Tree of ventral cirri reach bases of inferior
neurosetae : si .. L. filicornis
Ventrum smooth or ih a aur aerate at the pee ot the panned. Ventral cirri small
and not reaching the base of the neurosetae .. a A a L. benthaliana
ReEcorps oF Laetmonice FROM SoUTH AFRICA
Laetmonice benthaliana McIntosh 1885
Laetmonice producta var. benthaliana McIntosh 1885, p. 45, pl. 8 figs. 4-5, pl. 4 fig. 12.
? Laetmonice filicornis McIntosh 1925, p. 20.
Records: Station A.1g1 (2) and A.193 (9)—S.A. Museum Register No.
A.19778.
POLYCHAETE FAUNA OF SOUTH AFRICA 357
Station A.316 (15), A.317 (2), A.318 (2), A.gig (4) and A.322 (1)—
Register No. A.19959.
Notes: This. species seems to be one of the commonest polychaetes at
abyssal depths west of Cape Town, and these South African specimens agree
closely with the type described above. McIntosh (1925) recorded L. filicornis
from five stations in this area and from 4-10 fathoms in Saldanha Bay. The only
two specimens from this collection, which are now in the British Museum
(numbers 1924: 7 : 21 : 21-22) are Hermonia hystrix; possibly the missing
specimens were L. benthaliana. At any rate the record of L. filicornis must be
eliminated.
Laetmonice producta wyville: McIntosh 1885
Laetmonice producta var. wyvillei McIntosh 1885, p. 44.
Laetmonice producta (non Grube) Day 1934, p. 18.
Notes: The two specimens recorded by me (Day, 1934) from deep dredgings
off Portuguese East Africa have been re-examined. They are L. producta wyviller
with 18 pairs of elytra, a smooth ventral surface and very small ventral cirri.
Aphrodita alta Kinberg 1855
Aphrodita alta Kinberg 1857, p. 2, pl. 1 fig. 1 a-g. Day, 1960, p. 274.
Records: Station A.319 (1 juvenile) —S.A. Museum Register No. A.19964.
Sub-family Polynoinae
Harmothoe profunda n. sp.
(Fig. 1 a-e)
Records: Station A.193 (2)—S.A. Museum Register No. A.19780. Stations
A.316 (2) and A.318 (1) —Register No. A.1g9960.
Description: The holotype from station A.193 is the only complete specimen.
It is 16 mm. long for 33 segments. The dorsum is black or possibly purple
when fresh but the elytra and parapodia are pale. The prostomium (fig. 1a)
is bilobed with obvious frontal peaks and the anterior pair of eyes is large and
half-way back on the sides of the head. The median antenna is missing and the
laterals are small, about half as long as the prostomium, markedly tapered
and obviously ventral in origin. The dorsal cirri are about as long as theneuro- |
setae and like the antennae they are sparsely beset with long papillae.
The whole length of the body is covered with 15 pairs of elytra. Individual
elytra (fig. 1b, 6’) vary from rounded to broadly oval and each is densely
covered with chitinous tubercles which increase gradually in size from the
anterior to the posterior margin. The small anterior ones are bluntly conical,
those near the centre are larger and stouter, and a few near the posterior margin
are almost spherical. Some are smooth, others rugose and some even have small
blunt projections. A few small soft papillae are to be found near the external
margin and it might be said that the elytra are minutely fringed.
358 ANNALS OF THE SOUTH AFRICAN MUSEUM
The notopodium is well developed (fig. 1c) and bears numerous sabre-
shaped notosetae much stouter than the neurosetae. Each notoseta (fig. 1d, d’)
bears numerous rows of strong serrations and has a short, flanged and bluntly
pointed tip. The neuropodium is rather short with a triangular presetal lobe
and rounded postsetal lip. The neurosetae (fig. 1e, ce!) have rather long blades
with 15-20 rows of well-marked spinules. The tips are of normal length and
bidentate, but the relative size of the two teeth varies. In superior setae the
secondary tooth is not much smaller than the terminal one, but in the middle
of the series the secondary tooth decreases and inferior setae have a strong
terminal tooth and a weak secondary one, only half the length of the terminal.
There are many resemblances between this species and H. exanthema var.
bergstromt Monro 1936 from the Magellan area. An examination of the type in
the British Museum proved that the main difference is in the character of the
tubercles on the elytra. In Monro’s species there is a scattering of uniformly
small, slightly curved conical tubercles and a few very large, soft, pedunculate
vesicles but no intermediate forms. Moreover the antennae and cirri are more
densely papillose, the notosetae have finer serrations and the neurosetae have
longer naked tips and a smaller secondary tooth.
Ss SSS
= =
PG EG
Fic. 1. Harmothoe profunda.
(a) Head. (6, 5’) Elytron and details of marginal tubercles. (c) Parapodium. (d, d’) Notoseta and
details of tip. (e, e’) Neuroseta and details of tip.
POLYCHAETE FAUNA OF SOUTH AFRICA 359
Harmothoe sp.
Records: Station A.190 (1)—S.A. Museum Register No. A.19771.
Notes: Although the specimen is too poorly preserved to give it a specific
name it is obviously different from any other recorded from South Africa.
The characters may be briefly summarized as follows. Antennae and cirri
smooth. Elytra with minute conical tubercles anteriorly and 4-5 large cylindro-
conical soft papillae on the external margin. Notosetae with small, close-set
serrations and strongly flanged tips. Neurosetae with long, weakly spinulose
blades and minutely bidentate or unidentate tips.
Eunoe assimilis McIntosh 1925
Eunoe assimilis McIntosh 1925, p. 21, pl. 2 figs. 1-2, pl. 3 fig. 3.
Records: Station A.193 (2)—S.A. Museum Register No. A.19782. Station
A.316 (1) —Register No. A.19961. Station A.318 (1) —(doubtful identification)
Register No. A.19962.
Description: MclIntosh’s original description is incomplete and a
re-examination of the type in the British Museum and the discovery of a
complete specimen with elytra allow me to give a summary of the diagnostic
characters.
Body 20-30 mm. long with 36-37 segments, rather broad but tapered
posteriorly. Body completely covered with 15 pairs of elytra. Colour generally
pale, but the ventral cirri and outer margins of the elytra are tinged with
purple.
Prostomium with small frontal peaks but without eyes. Median antenna
three times the prostomial length; laterals ventral in origin and about 1°5
times the prostomial length. Antennae and cirri appear smooth but are really
clad with short papillae. Dorsal cirri are tapered and reach the ends of the
neurosetae. Elytra large and delicate with smooth margins; the surface is
glabrous apart from a small patch of tiny rounded tubercles near the anterior
margin.
Notosetae fairly numerous, each stout with close-set rows of weak serra-
tions and a long, narrowly flanged and bluntly pointed tip. Neurosetae few
and unidentate, the swollen blades having about 20 rows of short spinules and
a long hooked and sharply pointed tip. There is no trace of a secondary tooth.
E. assimilis is close to E. abyssorum McIntosh 1885 dredged in 2,600 fathoms
south of Australia. The type was examined in the British Museum. Like E.
assimilis it lacks eyes and has papillose antennae and dorsal cirri. The elytra
are missing. The setae are not markedly different, but the notosetae are more
strongly serrated and the tips are quite smooth, not flanged. On the other hand
each neuroseta does have a flange preceding the strong terminal tooth. £.
assimilis may prove to be a synonym of E. abyssorum, but until the elytra of
E. abyssorum are known the two should be kept separate.
360 ANNALS OF THE SOUTH AFRICAN MUSEUM
Sub-family Sigalioninae
Leanira tetragona (Oersted, 1845)
Leanira tetragona (Oersted), Fauvel 1923, p. 117, fig. 43 a-g.
Records: Station A.189 (2)—S.A. Museum Register No. A.19768.
Notes: This is a new record for South Africa but although the specimens
are in poor condition and lack elytra their other characters agree with speci-
mens from northern Europe (the type locality is Norway). In particular the
prostomium lacks eyes, the median antenna is long and unjointed with a pair
of flaps at its base, there is a dorsal cirrus on setiger 3 and there are several
slender stylodes on the parapodia; the notopodium has 5-7 and the neuro-
podium a series of 10-12. Cirriform branchiae appear on setiger 5 or 6, and 3
ciliated swellings or ‘cupuliform branchiae’ are present between the noto- |
podium and the elytrophore. There are no simple setae in the neuropodia—
all are compound with smooth or weakly serrated shaft-heads and laddered
blades. It may be added that European specimens have a few delicate papillae
on the external margins of the elytra, a character which distinguishes this
species from L. hystricis and L. incisa which have both been recorded from
southern Africa. Both of them also lack a dorsal cirrus on setiger 3.
Fauvel (1923) was doubtful of the presence of a dorsal cirrus on setiger 3
of any species of Leanira, but an examination of the types of L. magellanica
McIntosh 1885, L. areolata McIntosh 1885 and L. japonica McIntosh 1885
shows that all three have a dorsal cirrus on setiger 3. A revision of the genus is
necessary.
Leanira hystricis Ehlers 1874
Leanira hystricis Ehlers, Fauvel 1923, p. 118, fig. 43 h-m.
L. hystrix McIntosh 1925, p. 38.
Records: Stations A.315 (1), A.317 (4) and A.gi9 (1)—S.A. Museum
Register No. A.19963. WCD 73 (1).
Notes: McIntosh’s specimen in the British Museum agrees with the present
material. There are no eyes, the median antenna lacks ctenidial flaps, there is
no dorsal cirrus on setiger 3, and cirriform gills appear between the 24th and
30th foot. The stylodes are all sausage-shaped; there is one on the notopodium
and 2—5 on the neuropodium. There are no simple neuropodial setae.
Family Nereidae
Nereis (Neanthes) papillosa n. sp.
(Fig. 2 a—g)
Records: Station A.1g1t (1)—S.A. Museum Register No. A.19779.
Description: ‘The holotype is a complete specimen 25 mm. long with 58
segments. It is pale in alcohol without any colour pattern. It is an incipient
heteronereid female with enlarged eyes and swimming setae developing in
setigers 16-35.
POLYCHAETE FAUNA OF SOUTH AFRICA 361
The prostomium (fig. 2a) is broadly oval with short broad palps but the
antennae and tentacular cirri do not present any features of special interest.
The anterior pair of eyes is greatly enlarged but the posterior pair is normal.
The proboscis (fig. 2b and c) when dissected proved to have very pale, poorly
chitinized paragnaths difficult to see; in particular some doubt must remain
as to the presence of paragnaths on groups VI, VII and VIII. Group I = 1,
mie lp — 4 IN — 5) im a wedge,, Vi — 0, Vi = 20r' 3 (very faint),
VII + VIII = a single row of about 4 very pale points.
Fic. 2. Nereis (Neanthes) papillosa.
(a) Head. (4, c) Dorsal and ventral views of proboscis. (d) 10th foot. (e) 25th foot. (f) 45th foot.
(g) Posterior neuropodial falciger.
Anterior feet (fig. 2d) have conical lobes and a rather short dorsal cirrus.
There are only two notopodial lobes but the notosetae arise from a dorsal bulge
on the inferior lobe so that the notopodium might be said to have 24 lobes.
The neuropodium is normal. In the middle of the body (fig. 2e) there is a
marked gap between the notopodium and the neuropodium, and all the lobes
bear numerous club-shaped papillae. It is unlikely that these papillae are
modifications due to the heteronereid phase for this does not occur in other
species and in any case the papillae continue on the posterior feet where
swimming setae are not developed. The posterior feet (fig. 2f) have long dorsal
362 ANNALS OF THE SOUTH AFRICAN MUSEUM
cirri without any basal flaps, the parapodial lobes are slender and the papillae
are less numerous.
The notosetae are homogomph spinigers throughout, there being no
notopodial falcigers. ‘The neurosetae include the usual homogomph and hetero-
gomph spinigers and heterogomph falcigers (fig. 2g). These have rather long
straight blades. In middle segments the tips of the developing swimming setae
project from both setigerous lobes.
There is a general similarity between WN. kerguelensis, N. unifasciata and
N. papillosa. All have two notopodial lobes in anterior feet and a single row of
paragnaths on groups VII + VIII. WN. papillosa is unique in having papillae
on the parapodial lobes.
Family Nephthydidae
Nephthys (Aglaophamus) macroura Schmarda 1861
Aglaophamus macroura (Schmarda), Hartman 1950, p. 118.
Nephthys (Aglaophamus) macroura Schmarda, Day 1960, p. 327.
Records: WCD 732 (1).
Notes: The single specimen is a juvenile 25 mm. long and its identification
is therefore doubtful. The ventral cirrus on setiger 1 is large but the dorsal
cirrus is rudimentary. Cylindrical, involute branchiae appear on setiger 3.
An average parapodium from the middle of the body has conical setigerous
lobes but lacks presetal lamellae on both rami. The postsetal lamella of the
notopodium is divided into a large rounded superior part and a small inferior
part. The notopodial cirrus is well developed, tapered and rounded in section,
not flattened and blade-like. The postsetal lamella of the neuropodium is
auricular and just exceeds the setigerous lobe. Anterior setae are normal
laddered capillaries and posterior setae are long, fine and minutely denticulate
on one side.
The absence of a dorsal cirrus on setiger 1 and the early appearance of
the branchiae suggest V. macroura, but the structure of the feet, particularly
the notopodial cirrus, is closer to WN. malmgreni, which is also a deep-water
species. Possibly the early appearance of the branchiae is a juvenile character.
Nephthys hystricis McIntosh 1900
Nephthys hystricis McIntosh, Fauvel 1923, p. 373, fig. 146 a-e.
Records: WCD 73 (1). |
Notes: The single specimen is a juvenile and is referred to WN. hystricis with
hesitation.
Nephihys (Nephthys) ? paradoxa Malmeren 1874
Nephthys (Nephthys) ? paradoxa Malmgren, Day 1960, p. 327.
Records: Station A.315 (1)—S.A. Museum Register No. A.19965.
Notes: The specimen is broken and badly preserved. While it is similar to
the one reported by Day, 1960, the identification must remain uncertain.
A large parasitic nematode lies in the body cavity.
POLYCHAETE FAUNA OF SOUTH AFRICA 363
Family Eunicidae
Sub-family Onuphidinae
Ayalinoecia sp.
Records: WCD 73—15 specimens and 3 juveniles.
Notes: These specimens approach H. bilineata var. rigida as described by
Fauvel, 1923, p. 424, but differ in several respects. They are encased in fragile
tubes covered with detritus quite unlike the tough quill tubes of H. tubicola,
the only species of the genus previously recorded from South Africa. The two
frontal antennae are rather broad and the five occipital antennae are mounted
on 4-ringed ceratophores, each ring having a projecting lateral lobe. There are
no eyes and no tentacular cirri. Conical ventral cirri are present on the first 3
setigers. Gills start on setiger 9 and continue for about 12 segments. Each gill
is a small inconspicuous filament no larger than the dorsal cirrus. The hooded
hooks of setigers 1 and 2 are unidentate and very faintly pseudo-compound.
I know of no species of Hyalinoecia with branched ceratophores to the
occipital antennae, but similar structures do occur in the genus Epzdiopatra
and it is just possible that all these specimens may develop spiral gills at a later
stage. For this reason they have not been named as a new species.
Sub-family Lumbrinerinae
Lumbrineris magalhaensis Kinberg 1864
Lumbrineris magalhaensis Kinberg, Day 1960, p. 362, fig. 12 hj.
Records: Station A.189 (1)—S.A. Museum Register No. A.19769.
Lumbrineris brevicirra (Schmarda 1861)
Lumbrineris brevicirra (Schmarda), Day 1961, p. 361, fig. 12 e-g.
Records: Station A.193 (1)—S.A. Museum Register No. A.19781.
Family Paraonidae
Paraonis gracilis oculata Hartman 1957
Paraonis gracilis oculata Hartman 1957, p. 331, pl. 44, figs. 1-3.
Records: WCD 74 (3 juveniles).
Notes: All three specimens are broken but probably measured less than
8 mm. when complete. The prostomium bears a pair of subdermal eyes and
there are only 3 pairs of gills on setigers 6, 7 and 8. Posterior neurosetae include
1-2 capillaries and 2-3 short stout acicular hooks with curved, sigmoid shafts
and blunt unidentate tips. There is no trace of an arista. Ehlers (1913) recorded
Aonides gracilis (now recognized as a synonym of Paraonis gracilis) from False
Bay but did not mention whether his specimen had eyes or not.
364. ANNALS OF THE SOUTH AFRICAN MUSEUM
Aricidea suecica simplex var. nov.
(Fig. 3 a—b)
Records: WCD 73 (3).
Description: All three specimens are broken and only one is sufficiently
long to identify the posterior neurosetae. It has been chosen as the holotype.
It is 8 mm. long with 65 segments and probably represents about half of the
original worm.
The prostomium (fig. 3a) is short and broad with well-marked nuchal
slits, a very short median antenna, but no eyes. The anterior region is flattened
dorsally and the posterior region rounded. Branchiae start on setiger 4 and
continue to setiger 15 so that there are 12 pairs in all. Each gill is stout with a
pointed tip and just meets its fellow in the mid-dorsal line. All gills are subequal
except for the last 2-3 which are small. ‘The postsetal notopodial lobe of setiger 1
is a small papilla but later ones soon increase in size and in the branchial
region they are 2/3 the length of the gills. The neuropodia are merely low
CIAAIE SH
Cd ZA 1 ; 2 j i ! Pye
v [es \ \ \ 3 : \ i
c epee IN : eee ies
Fic. 3. Aricidea suecica var. simplex.
(a) Head and anterior segments. (b) Posterior neuropodial hook. Streblosoma abranchata:
(c) Lateral view of head and anterior segments. (d, ¢) Lateral and edge-on views of thoracic
uncinus. (f) Thoracic notoseta.
POLYCHAETE FAUNA OF SOUTH AFRICA 365
lateral ridges. Anterior notosetae and neurosetae are similar. Each is a curved
capillary with a slender blade.
The posterior region as defined by the appearance of the neuropodial
hooks starts on setiger 24. The segments are rounded in section, each with a
slender postsetal notopodial lobe. The posterior notosetae are 2-3 fine capil-
laries but the neurosetae are more numerous. Each neuropodium bears about
5 long slender capillaries and an increasing number of hooks; at first there are
only 1-2 but later as many as 10. The hooks (fig. 3) are all similar from segment
24. onwards. Each is sigmoid in shape with a shaft which is constricted where it
leaves the surface and a curved unidentate tip which never has any prolonga-
tion or sign of an arista.
The stem form A. suecica Eliason 1920 from northern Europe is described
as having an arista or slender blade projecting beyond the apex of the neuro-
podial hook in the middle of the body, but this is lost later and posterior neuro-
podial hooks are sigmoid and unidentate. There is also a pair of eyes. This is a
new record for South Africa.
Family Spionidae
Prionospio steenstrupi Malmgren 1867
Prionospio steenstrupi Fauvel 1927, p. 60, fig. 21 f-1.
Prionospio malmgreni var. dubia Day 1961, p. 489, fig. 3 j—n.
Records: WCD 73 (1).
Spiophanes soderstromt Hartman 1953
Spiophanes soderstromi Hartman 1953, p. 41, fig. 14 a-c. Day 1961, p. 484.
Records: WCD 74 (1 juvenile).
Family Disomidae
Poecilochaetus serpens Allen 1904
Poecilochaetus serpens Allen, Fauvel 1927, p. 67, fig. 23 a-m. Day 1961, p. 497.
Records: WCD 73 (1 juvenile).
Family Maldanidae
Maldanella fibrillata Chamberlin 1919
Maldanella fibrillata Chamberlin 1919, p. 413, pl. 72 figs. 1-6, pl. 73 figs. 1-2.
Records: Station A.190 (5)—S.A. Museum Register No. A.19773.
Description: The 5 specimens were encased in fragile tubes covered with
grey silt. Their bodies were broken and the number of segments is unknown.
Head with an oval cephalic plate inclined at 45° to the axis of the body and
surrounded by a high, smooth rim which is continuous apart from a small
anterior gap occupied by the prostomium. No eye spots. Nuchal grooves
straight and half the length of the cephalic plate but their anterior ends are
366 ANNALS OF THE SOUTH AFRICAN MUSEUM
continuous with a streak which curves sharply back around the inside of the
rim. Anterior segments short, hardly longer than broad, and without collars
but with glandular rings on the first 7. Middle segments long but the posterior
ones shorter and sausage-shaped. 3 achaetous preanals which decrease in
length. The last is not very distinct and is situated at the base of the pygidial
funnel which is rimmed with 30 equal anal cirri. Anus on a pleated cone with-
out a marked ventral valve.
Setiger 1 with notosetae only. Setiger 2 with notosetae and 15-20 neuro-
podial hooks. Later segments similar but with more hooks. The notosetae are
all narrow-winged capillaries, some long and some short. The hooks on setiger
2 have a vertical series of 2-3 teeth above the main fang while those of later
segments have 3—4 teeth.
These characters agree with Chamberlin’s description and it may be
added that his original specimen was complete and measured 70 mm. by 5°5
mm. and had 109 setigerous segments.
M. fibrillata is a rare abyssal species from the Pacific off the coast of Panama.
The only species of Maldanella known from South Africa is M. capensis Day
which has 4 achaetous preanals (not 3) and only 2-3 neuropodial hooks on
setiger 2 (not 12 or more).
Lumbriclymene minor Arwidsson 1906
Lumbriclymene minor Arwidsson: Fauvel 1927, p. 196, fig. 68 k—q.
Records: Station A.315 (1)—S.A. Museum Register No. A.19966.
Notes: The tube is free and encrusted with foraminiferan shells. The body
is 20 mm. long with 19 setigers and 3 indistinct preanals in front of the pygidium.
The anterior end is bluntly rounded without a well-defined prostomium,
cepalic plate or crest. Even the curved nuchal grooves are poorly marked. The
19 setigers do not differ greatly in length, each being about three times as long
as broad. The anterior margins of anterior setigers are slightly glandular but
the intersegmental constrictions are not deep and there is no sign of collars.
The three achaetous preanal segments are short and poorly marked with
indistinct lateral tori. The pygidium is blunt and slightly swollen with a dorsal
anus. The ventral valve is very large and is continuous with the bluntly conical
posterior end. The ventral surface of the pygidium is somewhat flattened and
slopes upwards. There are no anal cirri. The notosetae are all winged capil-
laries, some smooth-edged and some striated towards the distal end. Each of
the first 4 setigers bears a single stout, smoothly pointed, acicular seta. Subse-
quent neurosetae are rows of about 6 hooks, each with a vertical series of 4
teeth above the rostrum and a few lateral denticles. Below the rostrum is a
short ‘neck’ and then a marked swelling in the shaft.
This is a new record for South Africa. Its pygidium differs from L. cylindri-
cauda Sars which has recently been found in shallow water in this area (unpub-
lished record). In L. cylindricauda as the name implies the posterior end is
POLYCHAETE FAUNA OF SOUTH AFRICA 367
cylindrical with a terminal anus; in L. minor the anus is dorsal and the ventral
surface of the pygidium is flattened and slanting.
Family Sabellariidae
Phalacrostemma elegans Fauvel 1911
Phalacrostemma elegans Fauvel 1914, p. 270, pl. 24, figs. 1-16.
Records: Station A.315 (1)—S.A. Museum Register No. A.19967.
Notes: The single specimen is poorly preserved and lacks a tube, but the
available characters, in particular the setae, agree completely with Fauvel’s
description.
The opercular lobes are separate and each bears a single ring of long,
tapered and spirally serrated ‘paleae’ with about 3 stout acicular setae at the
base. At the junction of the opercular lobes there is one pair of hooks dorsally
and a single median tentacle ventrally. The buccal tentacles and palps have
disintegrated. There are 4 parathoracic segments with oar-shaped setae. There
are at least 12 abdominal segments bearing uncini and capillaries, but the gills
have been lost and the posterior end has disintegrated so that the exact Tae?
of segments and details of the caudal region are unknown.
This is a new record for South Africa. Fauvel’s original specimen came
from 1,968 metres off Madeira.
Family Ampharetidae
Neosabellides cf. elongaius (Ehlers) 1913
[?] Sabellides elongatus Ehlers 1913, p. 551, pl. 42, figs. 1-6.
[?] Neosabellides elongatus (Ehlers), Hessle 1917, p. 104. Monro 1936, p. 175.
Records: Station A.319 (3)—S.A. Museum Register No. A.19968.
Description: ‘The 3 specimens are encased in characteristically slender and
closely ringed brownish mud tubes. The body is slender, slightly swollen
anteriorly, about 30 mm. long and 1-2 mm. wide at the broadest part of the
thorax.
The prostomium is bluntly spade-shaped without lateral grooves but has a
single pair of eyes. The tentacles are largely retracted and lateral pinnules are
poorly marked. Segment II is narrow but unusually distinct. Segment III
lacks paleae and is fused to segment IV which bears a small bundle of noto-
podial capillaries. Segments V and VI also bear notopodial capillaries but no
uncini. Segment VII and the next 10 segments bear both notopodial capil-
laries and neuropodial uncini so that there is a total of 14 segments with noto-
setae of which the first three lack uncini. Segment III bears two groups of 3
smooth, tapered gills which project well beyond the prostomium. The two
groups of gills are well separated in the dorsal median line and each group is
arranged in a transverse row, but there is no obvious branchial ridge nor is
there any sign of nephridial papillae between the two groups of gills. The first
368 ANNALS OF THE SOUTH AFRICAN MUSEUM
5 thoracic segments are short, each being about three times as broad as long,
but subsequent segments become much longer until in the middle of the thorax
each segment is three times as long as broad. The length of the segments affects
the glandular ventral pads which are recognizable on all except the last thoracic
segment. Anterior ones are well marked and contiguous, but posterior pads are
poorly developed and well separated from one another. The abdomen consists
of 32 segments. Each uncigerous pinnule is a small, roughly square lateral lobe
with a small papilla above the row of uncini. The pygidium bears a circle of 6-8
tapered anal cirri and a larger pair of ventral lobes.
The notosetae are smooth-winged capillaries. Thoracic uncini bear 8
teeth in two vertical rows of 4.
These South African specimens agree with Ehlers’s original description
of Sabellides elongatus from Antarctica in regard to most characters, but the first
three setigers lack uncini whereas Ehlers states that only two anterior setigers
lack uncini. Moreover Ehlers found only 19 abdominal segments or less, whereas
these South African specimens have over 30. It may be that Ehlers (1913),
Hessle (1917) and Monro (1936), who all describe material from the Antarctic
or sub-Antarctic, missed the first bundle of notosetae, which is small and very
close to the second, but for the present the identity of the South African speci-
mens must remain doubtful.
Family Terebellidae
Amphitrite cirrata Miller 1771
Amphitrite cirrata Miller, Fauvel 1927, p. 251, fig. 86 7-0.
Records: Station A.319 (6)—S.A. Museum Register No. A.19971.
Notes: The present material agrees perfectly with Fauvel’s description.
There is a general resemblance to a species of Thelepus since the three pairs of
gills are filamentous, with the right and left tufts well separated. Each tuft
arises from a basal stump. There are 17 thoracic segments starting on segment 4
(3rd branchiferous), each bearing a bundle of notopodial capillaries with
minutely denticulate tips. There are small lateral lobes on segments 2, 3 and 4,
and 12 ventral pads. Uncini start on segment 5 (setiger 2) and are arranged in
double rows after the first few. Each uncinus is avicular with a close-set cap
of denticles above the main fang. The denticles are irregularly arranged but
approximate to the formula: MF : 4-5 : 5-6 : 8-10 : 10-15.
The abdomen is broken in every specimen but has at least 20 segments
with square uncigerous pinnules.
This species is well known from high latitudes in the North Atlantic.
Fauvel (1914) recorded Amphitrite cirrata profunda from abyssal depths off the
Azores, but as he says himself it is very doubtfully distinct from the stem form.
This is the first record of the genus Amphitrite from South Africa.
POLYCHAETE FAUNA OF SOUTH AFRICA 369
Pista cristata (Miller 1776)
Pista cristata (Miller), Fauvel 1927, p. 266, fig. 93 a-g.
Records: Station A.317 (1), A.319 (6)—S.A. Museum Register No. A.19970.
Notes: These specimens agree well with Fauvel’s description. There are 2
pairs of gills, each with a long stem and a terminal pompon of spirally arranged
filaments. There are 3 pairs of lateral lobes; the pair on segment 2 is small and
continuous across the ventrum, the pair on segment 3 is large and lateral in
position, and the pair on segment 4 is small. 17 thoracic segments bear bundles
of smooth-winged notopodial capillaries. The first four rows of uncini have a
close-set crest of denticles, short ‘necks’ below the main fang, rounded bases
and well-developed shafts; later uncini have no shafts.
P. cristata var. capensis, reported by McIntosh (1925) from Portuguese
East Africa, was said to have a single tooth above the main fang. The type has
been lost. P. brevibrancha Caullery, reported by me (Day, 1951 and 1957)
from Portuguese East Africa, may be distinguished from P. cristata by the short-
ness of the branchial trunks and the uncini. Only the first row of uncini has
well-developed shafts, the neck of the uncinus is longer and the base larger and
more triangular.
Terebellid— ? gen. et sp.
Records: Station A.316 (12)—S.A. Museum Register No. A.19972.
Notes: The specimens are very soft and poorly preserved, and some of the
characters are doubtful. They are obviously different from the other species
recorded here but cannot be identified with certainty.
An average specimen is about 50 mm. long and 4 mm. wide across the
anterior thorax. The collar-shaped tentacular lobe bears numerous orange
tentacles. No gills were found; it is possible that they have been lost, but there
was no clear sign of scars. No lateral lobes were visible. ‘There are 17 bundles
of notosetae starting on segment 3 or possibly 4. The notosetae are of two
lengths, both with smooth, broad wings and very long attenuated tips which
appear to be quite smooth. Uncini start on setiger 2 (i.e. segment 4 or 5),
and on the posterior thorax they are arranged in double rows. The thoracic
uncini are all similar, each being avicular with a crest of numerous irregularly
arranged denticles above the main fang. There are at least 12 ventral pads.
The abdomen consists of 40 or more segments bearing short uncigerous pinnules.
These specimens belong to the sub-family Amphitritinae as shown by the
arrangement and structure of the uncini. I know of no species which has
smooth-bladed notosetae and lacks gills. However, a better preserved specimen
is necessary to confirm that gills are really absent.
Streblosoma abranchiata n. sp.
(Fig. 3 cf)
Records: Station A.190 (numerous tubes and specimens) —S.A. Museum '
Register No. A.19770.
370 ANNALS OF THE SOUTH AFRICAN MUSEUM
Stations A.3g15 (2), A.g19 (Common) and A.321 (1)—Register No.
A.19969.
Description: ‘The type material is No. A.19770 from Station A.igo. The
tubes are long, fragile and heavily encrusted with foraminiferan shells. The
worms themselves are poorly preserved, but judged by the size of the larger
fragments the body was cylindrical, rather slender and about 30 mm. long by
I°5 mm. wide with over 60 segments.
The tentacular lobe (fig. 3c) is short and collar-like, with about a dozen
long, grooved tentacles. There are no eye-spots. An oral hood overhangs the
ventral mouth but the lower lip is small. There are no lateral lobes on anterior
segments and no sign of gills although all specimens were carefully examined.
The first bundle of notosetae is on segment 2 and there are at least 19 segments
with notosetae. ‘The longest anterior fragment had disintegrated at this point
but posterior fragments of 20 or more uncigerous segments are without noto-
setae. The notosetae (fig. 3f) are smooth-winged capillaries. Uncini appear on
setiger 4 (segment 5). They are arranged in single rows throughout and borne
on poorly marked uncigerous ridges even in abdominal segments. Each
uncinus (fig. 3d and e) has a close-set cap of denticles above the main fang.
These are irregularly arranged but approximate to the formula MF:
4-5 :ca. 8: ca. 12. The base of the uncinus has the characteristic clog-shape
of the sub-family Thelepinae with a forwardly produced rounded prow and a
well-marked dorsal button.
All other species of Streblosoma have at least 2 pairs of filamentous gills,
and the present species was assigned to the genus with some hesitation; but it
obviously belongs to the Thelepinae, and the segmental arrangement of the
setae and the lack of lateral lobes on anterior segments agree with Streblosoma.
Streblosoma chilensis (McIntosh 1885)
Euthelepus chilensis McIntosh 1885, p. 467, pl. 51 figs. 4-5, pl. 28A figs. 14-15.
Records: WCD 73 (1 juvenile).
Notes: The genus Euthelepus was erected by McIntosh 1885 for two species,
E. setabulensis and E. chilensis. The types of both have been re-examined in the
British Museum. FE. setabulensis, which has page priority, has been accepted as
the type species by both Fauvel (1927) and Hartman (1959). It has 3 pairs of
filamentous gills on segments 2-4 and notosetae from segment 3 (not segment 2
as suggested by McIntosh). Uncini start on segment 5 (setiger 3). Lateral lobes
are present on segments 2-4. F. chilensis has 2 pairs of simple filamentous gills on
segments 2 and 3. Notosetae start on segment 2 (the first branchiferous) and
uncini on segment 5 (setiger 4). There are no lateral lobes on anterior segments.
It thus differs from EF. setabulensis in the possession of notosetae on segment 2
and in the absence of lateral lobes. It should thus be transferred to the genus
Streblosoma.
The single South African specimen is a juvenile but agrees in all essential
respects with McIntosh’s type from abyssal depths off Chile.
POLYCHAETE FAUNA OF SOUTH AFRICA 371
Terebellides stroemt Sars 1835
Terebellides stroemi Sars, Fauvel 1927, p. 291, fig. 100 7-@.
Records: WCD 73—2 juveniles.
SUMMARY
Twenty-eight species of Polychaeta are described from abyssal dredgings
west of Cape Town. The collection includes 4 new species and 11 new records
for this area. Only 4 of these species are known from depths less than 100
metres; the rest are widespread at abyssal depths. There is no obvious Antarctic
component.
ACKNOWLEDGEMENTS
The Trustees of the South African Museum wish to acknowledge a grant
from the Council for Scientific and Industrial Research towards the purchase
of the deep-trawling equipment with which the specimens reported on were
collected, and a grant for the publication of this paper. Grateful acknowledge-
ment is also due to the Director, Division of Fisheries, Department of Commerce
-and Industries, for making the collecting possible.
REFERENCES
CHAMBERLIN, R. V. 1919. The Annelida Polychaeta. Mem. Mus. Comp. Zool. Harvard, 48, 1-514.
Day, J. H. 1951. The polychaete fauna of South Africa. Part 1: The intertidal and estuarine
Polychaeta of Natal and Mocambique. Ann. Natal Mus. 12, 397-441.
Day, J. H. 1957. The polychaete fauna of South Africa. Part 4: New species and records from
Natal and Mocambique. Ann. Natal Mus. 14, 59-1209.
Day, J. H. 1960. The polychaete fauna of South Africa. Part 5: Errant species dredged off
Cape coasts. Ann. S. Afr. Mus. 45, 261-373.
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. 1913. Die Polychaeten-Sammelungen der deutschen Siidpolar-Expedition 1g01—
1903. Deutsch. Siidpol.-Exped. 13, 397-598.
FAuvEL, P. 1914. Annélides polychétes non-pélagiques provenant des compagnes de l’ Hirondelle
et de la Princesse-Alice (1885-1910). Result. Comp. Sci. Monaco. 46, 1-432.
FAUVEL, P. 1923. Polycheétes errantes. Faune Fr. 5, 1-488.
FAUVEL, P. 1927. Polycheétes sedentaires. Faune Fr. 16, 1-494.
HARTMAN, O. 1950. Goniadidae, Glyceridae and Nephtyidae. Allan Hancock Pacif. Exped. 15
(1), 1-181.
Hartan, O. 1953. Non-pelagic Polychaeta of the Swedish Antarctic Expedition 1901-1903.
Further Zool. Res. Swed. Antarct. Exped. 4 (2), 1-185.
Hartman, O. 1957. Orbiniidae, Aspidobranchidae, Paraonidae and Longosomatidae. Allan
Hancock Pacif. Exped. 15 (3), 211-392.
Hartman, O. 1959. Catalogue of the Polychaetous Annelids of the world. Part II. Allan Hancock
Publ. occ. paper 23, 355-628.
Hesste, C. 1917. Zur Kenntnis der terebellomophen Polychaeten. Zool. Bidr. Uppsala 5, 39-258.
MclIntosu, W. C. 1885. Report on the Annelida Polychaeta collected by H.M.S. ‘Challenger’
during the years 1873-1876. Challenger Rep. Zool. 12, 1-554.
MclIntosu, W. C. 1925. A second contribution to the marine Polychaeta of South Africa.
Rep. Fish. Mar. biol. Surv. S. Afr. 4, spec. rep. 4, 1-93.
Monro, C. C. A. 1936. Polychaete worms. II. Discovery Rep. 12, 59-198.
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References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrdége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
HartmMan, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuka, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. and ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.—All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name. . .
B. Form of bibliographic references to synonymic names.—'The first reference following
any name in the synonymy should be to the earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalis capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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XXIII.
XXIV.
XXV.
XXVI.
XXVII.
XXVIII.
XXIX.
XXX.
INDEX
XXXI.
XXXII.
XXXII.
XXXIV.
XXXV.
XXXVI.
XXXVII.
XXXVITI.
XXXIX.
XL.
XLI.
XLII.
XLII.
XLIV.
XLV.
XLVI.
1900-1902
1903-1905
1903-1908
1906-1910
1908-1910
1908-1913
IQII—-1919
IQII-I9I4
1911-1918
1913-1924
1913-1923
IQI5-1924
1914-1916
1917-1933
1QI'7—1920
1921
1924-1925
1924-1926
1925-1928
1925-1926
1929-1933
1927-1928
1928
1929
1929-1932
1929-1931
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ANNALS
OF THE
SOUTH APRICAN MUSEUM
VOLUME XLVI
Pas fy XV
THE PSEUDODIAPTOMIDAE (GOPEPODA; CALANOIDA)
OF SOUTHERN AFRICAN WATERS, INCLUDING A NEW SPECIES,
PSEUDODIAPTOMUS CHARTERI
By
Joun R. GRINDLEY
South African Museum, Cape Town
ys a
eee
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[ CONTINUED ON INSIDE BACK COVER
THE PSEUDODIAPTOMIDAE (COPEPODA; CALANOIDA)
OF SOUTHERN AFRICAN WATERS, INCLUDING A NEW SPECIES,
PSEUDODIAPTOMUS CHARTERI
By
JouN R. GRINDLEY
South African Museum, Cape Town
[Accepted July, 1962]
(With 5 figures in the text)
CONTENTS
PAGE
Introduction < ie rae 379
The genus Pseudodiaptomus . . ja OU
Pseudodiaptomus hesset (Mrazek) .. 374
Pseudodiaptomus charteri sp. n. SOL
Pseudodiaptomus nudus ‘Tanaka -. 364
Discussion .. sf os Hi) 380
Key to southern African species .. 389
Summary .. ne See S57) BOD
INeferencesi a: 5a Bi ee QO
INTRODUCTION
The Pseudodiaptomidae are a very interesting group of calanoid copepods
which occur in marine, brackish and fresh water in many parts of the world.
They are frequently the dominant organisms within their particular habitat,
while their interesting distribution makes them of zoogeographical importance.
While working on the plankton of South African estuaries it became clear to
me that a revision of the South African species of Pseudodiaptomidae was
required. The only Pseudodiaptomid previously recorded from southern
African waters was Pseudodiaptomus serricaudatus (Cleve, 1904; Marques, 1958;
etc.). I was, however, doubtful whether our specimens should be referred to
this species for they differed slightly from the form originally described by
T. Scott (1894). Tanaka (1960) confirmed my opinion by describing specimens
of the South African form, obtained from the Agulhas bank by the Japanese
Antarctic Research Expedition, as a new species, Pseudodiaptomus nudus. This
species occurs chiefly in neritic waters. Two species of Pseudodiaptomus are the
dominant planktonic animals in South African estuaries and lagoons. One
species from the west and south coasts was found to be Pseudodiaptomus hesset
which was recorded (in a preliminary report only) from the mouth of the Congo
(Mrazek, 1894). After this it does not appear to have been reported again,
and it was never fully described. The species found on the Natal coast was
apparently previously unknown and is described here. A complete description
2k
Ann. S. Afr. Mus. 46 (15), 1963, 373-391, 5 figs. Mert
il MAY 21 63
374 ANNALS OF THE SOUTH AFRICAN MUSEUM
is given of P. hessei and the other species are described largely by comparison
with it. The terminology I have used to describe the regions of the copepod
body follows the recommendations of Gooding (1957).
Family Pseudodiaptomidae
Genus PSEUDODIAPTOMUS
Pseudodiaptomus Herrick 1884, 180.
Schmackeria (non Poppe & Richard 1890) Mrazek 1894, 1.
Heterocalanus 'T. Scott 1894, 39.
Weismanella Dahl 1894, 10.
Head smoothly rounded, separated from or fused with first pedigerous
prosome segment. Fourth and fifth pedigerous segments usually fused, with
rounded or pointed posterior angles, frequently bearing spines or hairs, par-
ticularly in the female. Genital segment of the female usually swollen, bearing
setae and spines arranged asymmetrically. Caudal rami at least twice as long
as wide, caudal setae often jointed or thickened. First four pairs of legs biramous,
rami 3-segmented. Female fifth legs uniramous and 4-segmented. Male fifth
legs 4-segmented, uniramous or bearing 1-segmented endopods on either or
both legs. The right endopod, when present, is usually rudimentary. Three
species represented in southern African waters. Pseudodiaptomus pelagicus Herrick
(1884), which is the type of the genus, has not been seen since that time and the
original description is brief and the figures unfortunately poor.
Pseudodiaptomus hesser (Mrazek)
(Figs. 1 a-k; 2 a-f; 3 af)
Schmackeria hessei Mrazek 1894, 1, figs. 1, 2, 3.
Pseudodiaptomus hessei (Mrazek) Giesbrecht & Schmeil 1898, 65. Marsh 1933, 36, pl. 19, figs. 1, 2.
Occurrence
Abundant in estuaries and lagoons on the west and south coasts of South
Africa as far east as the Kleinmond estuary (Bathurst division). Material from
the latter locality was collected for me by Professor J. Omer-Cooper. Specimens
have been found in water with salinity ranging from about 10°/5) to over 4.0/9.
Known distribution
Banana Creek (brackish water) at the mouth of the Congo River (Mrazek,
1894). |
Description
FEMALE: Length 1°55-1:80 mm. (1:20 mm., Mrazek). Prosome slender‘
viewed dorsally approximately two and a half times as long as wide, rounded
anteriorly (fig. 1 a, b). Fourth and fifth pedigerous segments fused. Posterior
angles of metasome bearing a sharp spine on each side on the postero-dorsal
margin. Rostrum well developed with two strong filaments which reach the
middle of the proximal segment of the first antennae.
PSEUDODIAPTOMIDAE OF SOUTHERN AFRICAN WATERS 375
Urosome 4-segmented and about two-thirds as long as prosome. Genital
segment (fig. 1 f; h) asymmetrical, with irregular swellings laterally and a
prominent genital boss ventrally. A number of setae are present mainly on the
lateral swellings, and there is a patch of fine hairs in the middle of the dorsal
surface. The genital boss is of characteristic form. It bears a row of spinules
anteriorly and the genital flaps each have a distal seta. The first three urosome
segments are furnished with rows of coarse teeth on their postero-dorsal margins.
The caudal rami (fig. 1 g) are divergent, more than three times as long as wide
and furnished with fine hairs on their inner margins. The third or middle
caudal seta (not counting the sensory bristle between the fourth and fifth setae)
is broad and flattened in a characteristic blade-like form. It is about three times
as wide as the other setae and is easily visible under low magnification so that
it is a useful feature for distinguishing the females of this species. Similar
broad setae occur in a number of species of the Schmackeria group from Asia.
Antenna I (fig. 1 ¢) of 21 segments, reaching the second urosome segment
when extended. Segments 1 + 2, 3 + 4,8 + 9 and 24 + 25 apparently fused.
Proportional lengths and setation of segments as shown in figure 1 ¢c. Giesbrecht
and Schmeil (1898) and Marsh (1933) reported 22 segments, but Mrazek
(1894) does not mention the number of segments.
There is one aesthetask on each segment up to and including no. 16,
one on no. 19 and one on the terminal segment. (Aesthetasks are not shown in
figure 1 c.) A specialized comb-like seta (fig. 1 7) is present on the third last
segment. It lies parallel to the antenna so that it is not readily noticed. Its distal
end is curved and slightly expanded and bears about three distinct recurved
hooks and a number of much smaller ones which grade into a series of minute
comb-like teeth (3 long and tp apart) along the shaft of the seta. There is a
bulbous swelling near the base of this seta. There is also an unspecialized
plumose seta on this segment. The comb-like seta is an interesting feature,
for modified setae (apparently of varying form) have been described on this
segment of the first antenna of the female and the unmodified antenna of the
male from a number of widely scattered species within the genus. Some form
of comb-like seta has been noted in Pseudodiaptomus stuhlmanm, P. serricaudatus
and P. hesser (Poppe and Mrazek, 1895); P. richardi (Mrazek, 1901); P. hickmani,
P. mason and P. salinus (Seymour Sewell, 1932) ; P. ardjuna (Ummerkutty, 1961),
and P. charteri and P. nudus in this present work. ‘This feature thus occurs in
species from America to the orient and is not confined to any particular sub-
group of this genus.
Antenna 2 (fig. 2 a) with basipod partially divided by an incomplete
suture, bearing 1 lateral and 2 terminal setae. Exopod 3-segmented, third
segment only partially separated, bearing 2 setae on the first segment, 7 setae
on the second, and 6 terminal setae and a lateral fringe of fine hairs on the
distal segment. Endopod apparently 4-segmented, third segment small and
indistinct, bearing 1 seta on segment 1, 2 lateral and 3 distal on segment 2,
I on segment 3 and 2 lateral and 3 terminal on segment 4.
376 ANNALS OF THE SOUTH AFRICAN MUSEUM
yy.
wey
)
N
Fic. 1. Pseudodiaptomus hesset (Mrazek). a, 9 dorsal; b, 9 lateral; c, antenna 1 9; d, fifth legs 9;
e, distal part of latter enlarged; f, 2 genital segment ventral; g, caudal ramus 9; h, 2 genital
segment lateral; 7, comb-like seta enlarged; j, fourth swimming leg; &, first swimming leg.
PSEUDODIAPTOMIDAE OF SOUTHERN AFRICAN WATERS Ba
Mandible (fig. 2 c) with gnathal lobe heavily chitinized and bearing about
10 fine teeth and a plumose spinule. Basipod of palp with 4 inner marginal
setae. Exopod indistinctly 3-segmented, bearing 1 lateral and 5 terminal setae.
Endopod 2-segmented, bearing 4 setae on segment 1 (1 separate) and 8 setae
distally on segment 2 (1 jointed and crooked). This last segment is narrowly
triangular and bears a row of tiny bristles.
Maxilla 1 (fig. 2 d) with first inner lobe or gnathobase bearing about 10
strong spines, small second inner lobe bearing ?3 setae and third inner lobe
bearing 3 terminal setae. Outer lobe or coxal epipod bearing 8 long setae.
Exopod with 9 marginal setae. Endopod 3-segmented bearing 4 setae medially
Fic. 2. Pseudodiaptomus hessei (Mrazek). a, antenna 2; b, divided seta from maxilliped; c, mandible;
d, maxilla 1; e, maxilliped; f, maxilla 2.
on the basal portion (second basal segment) and groups of 4 setae medially on
segments 1 and 2 and approximately 7 terminal setae on segment 3.
Maxilla 2 (fig. 2 f) with 5 large medial lobes or endites and 2 smaller
terminal lobes each bearing 3 or 4 setae. Distal portion indistinctly segmented.
Maxilliped (fig. 2 ¢) 6-segmented, 2 basal segments large, 4 distal segments
small and decreasing in size distally. First segment with 3 medial lobes bearing
2 setae, 3 setae and 2 setae and a spine respectively. There are short fine hairs
medially and tiny hooked spinules on the third lobe. Second segment expanded
medially with 3 setae and a fringe of short fine hairs on the medial margin.
Third segment bearing 2 + 3 setae, fourth segment 2 setae, fifth segment 2
setae, sixth segment 2 setae medially and 6 on an outer lobe. Among the setae
on segments 3, 4 and 5 are four with a peculiar branched structure. These
378 ANNALS OF THE SOUTH AFRICAN MUSEUM
branched setae (fig. 2 6) were noticed by Mrazek (1894) on what he called
the ‘hinteren Maxillipeden’. He said: “Bei unserer Form sind es gewéhnliche
Borsten die aber von der Mitte ihrer Lange an gespalten sind.’ It is clear that
Mrazek must have examined this species very carefully although he never
described it fully. These branched setae have a thickened basal portion and
divide near their middle, one branch continuing as a thinner seta forming the
extension of the proximal part while the other is short, broad and spatulate,
fringed with short curved spinules. (. . . l6ffelformig erweitert ist und einige feine
steife Harchen tragt.— Mrazek, 1894.)
Similar branched setae were noted in Pseudodiaptomus stuhlmann by Poppe
and Mrazek (1895). They were described as being on the second maxilla, but
Seymour Sewell (1932, 243) has shown that it is clear from their figure that the
appendage on which these modified setae were present is that which is now
termed the maxilliped. Branched setae have also been described on the maxilli-
ped of P. dauglisht (Seymour Sewell, 1932) and P. euryhalinus (Johnson, 1939)
and they are here recorded in P. chartert and P. nudus.
Swimming legs 1-4 biramous, with 2-segmented basipodite and
3-segmented exopodite and endopodite. They are similar in both sexes.
The first pair (fig. 1 k) are somewhat different to the following three pairs.
The outer marginal spines of the exopod are not serrate and blade-like. The
first segment of the exopod has one outer edge spine. the second no spine, and
the terminal segment has two short lateral spines and a long, slender terminal
spine. On the endopodite the numbers of setae on the successive segments are
1 on the first, 1 on the second and 5 on the third. On the exopodite the setation
is 1 on the first, 1 on the second and 4 on the third segment.
The remaining swimming legs (2-4) (fig. 1 7) bear serrated blade-like
spines on the outer distal corners of each exopod segment, an extra spine on
the distal part of the outer margin of the terminal segment and a long blade-like
terminal spine serrate only on its outer margin. On the exopodite the setation of
the segments is: 1 on the first, 1 on the second and 5 on the third. Endopodite:
I on the first, 2 on the second and 8 on the third. All the setae on the swimming
legs are jointed and slightly thicker proximal to the joint. The first basal segment
of all the legs has 2 groups of short stiff hairs on the outer margin and a long
seta on the inner distal margin. The second basal segment is completely naked
in the first leg and bears only a medial fringe of fine hairs in the remainder.
Fine short hairs occur on the outer margins of exopod segments 2 and 3.
The ornamentation of the swimming legs may be represented by the table
on the next page.
The fifth pair of legs in the female (fig. 1 d, e) is uniramous, 4-segmented
and asymmetrical, particularly as regards the hyaline projections of the inner
margin of the second basal segment. There seems to be some individual varia-
tion in the forms of these projections. Marsh (1933) suggests that these might
be regarded as rudimentary indications of an endopod. Such rudiments are
present also in P. stuhlmanni and P. charteri, but otherwise endopods of the female
PSEUDODIAPTOMIDAE OF SOUTHERN AFRICAN WATERS 379
LEG PROTOPOD ENDOPOD EXOPOD
or OF Ot +
pe ee Pel
—i—i— a=
Si, Se, St represent internal, external and terminal spines or setae respectively. The number of
setae is shown in arabic numerals and spines in roman numerals.
fifth legs are completely lacking in the Pseudodiaptomidae. The outer distal
angles of the first and second basal segments are produced into very small
spines. The first segment of the ramus bears a spine near the distal end of the
outer margin. The second segment bears a spine on the outer distal angle and
is produced distally on the medial side into a large curved spiniform process
bearing short bristles on the anterior and posterior margins and reaching about
the midpoint of the terminal spine. The terminal spine has a small distinct basal
portion bearing a slender plumose seta on its posterior face, while the long
terminal blade-like portion is furnished with minute bristles on its anterior
and posterior margins. ‘Terminal spine, including the basal portion, slightly
longer than the third segment. Marsh (1933, 28) points out that the distinct
basal portion of the terminal spine found in some species is not a real segment
but a modification of the terminal spine or ‘hook’.
The egg sac is single, containing about 25 eggs arranged in a single layer
curved to fit the urosome.
Mate: Length 1:35-1:45 mm. (1:05 mm., Mrazek). Habitus as in the female
(fig. 3 a, b). Posterior angles of the metasome without spines. Urosome 5-
segmented, uniformly slender, first and last segments shorter than the remainder.
Posterior margins of the second, third and fourth segments fringed partially (2nd
segment) or completely (3rd and 4th segments) with rows of coarse teeth. The
caudal setae (fig. 3 e) are distinctly different from those of the female, for the
middle seta is not expanded and all the setae are jointed about one-third of
their length from the proximal end.
Left antenna 1 as in female, including setation and arrangement of aesthe-
tasks. Right antenna 1 geniculate (fig. 3 c), with 21 segments, reaching the
second urosome segment when extended. The last four long segments are
apparently each formed by the fusion of two segments. Porportional lengths
and setation as shown in figure 3 c. There is a hooked spine on segment 10.
There is one aesthetask on segments 2, 5, 7, 9-16, and the second last and last
segments.
Male fifth legs (fig. 3 d) biramous and 4-segmented. In the right leg the
first basal segment is naked except for some rugosity on the medial surface,
ANNALS OF THE SOUTH AFRICAN MUSEUM
380
BAMUAVA A A pS
Bay
Fic. 3. Pseudodiaptomus hessei (Mrazek). a, 3 dorsal; 6, g lateral; c, right antenna 1 ¢; d, fifth
legs 3; e, caudal ramus 3; f, spermatophore.
PSEUDODIAPTOMIDAE OF SOUTHERN AFRICAN WATERS 381
The second basal segment is larger with some small spinose processes on the
inner margin, and bears a small rod-like endopodite with a short terminal seta.
On the outer margin there are a row of minute spinules and a fine seta near
the distal angle. The exopodite is 2-segmented, the first segment produced
into a curved spine at its outer distal angle, which just reaches the base of the
spine on the following segment, and bears some spinules near the inner distal
angle. The second segment bears a long, partly plumose spine on the distal
part of the outer margin and a rugose swelling on the proximal part of the inner
margin. The terminal hook has a triangular basal portion, broadest distally
where it bears a row of small spinules and is produced at the outer corner
into a slightly curved hook as long as the two exopod segments. The tip is
recurved and minutely serrate. In the left leg the first basal segment is naked and
the second basal segment bears a few spinules near the outer margin, some
rugosity on the inner margin and a long hyaline endopodite. This has an
expansion basally, is apically acute, and bears a fringe of about 8 fine hairs
near the apex. The exopodite is 2-segmented, the first segment short, bearing a
short spine at its outer distal angle. The second segment is expanded and
flattened, somewhat convex posteriorly and emarginate medially. The outer
distal margin is strongly convex and the inner distal margin nearly straight.
There is a spine near the middle of the outer margin, and a number of small
setae and spinules on the posterior surface and margins and around the apex.
The left leg reaches the base of the terminal hook of the right leg.
The spermatophore (fig. 3 f) is fusiform, stalked, and 250-400p long.
CoLour: Unpigmented except for some orange coloration around the mouth-
parts.
Types
Hypotypes, S.A.M. A1og58, in the South African Museum.
Remarks
This species was named after P. Hesse who first collected it. It was
originally described as a Schmackeria but was correctly transferred to the genus
Pseudodiaptomus by Giesbrecht and Schmeil (1898). The species is easily dis-
tinguished by the broad middle caudal seta in the female and by the charac-
teristic fifth legs in the male. Our specimens are considerably larger than those
originally described.
Pseudodiaptomus charteri sp. n.
(Figs. 4 4-7)
Occurrence |
Abundant in the estuaries of St. Lucia and Richards Bay on the Natal
coast. Specimens have been found in water with salinity ranging from about
109/59 to about 389/50.
382 ANNALS OF THE SOUTH AFRICAN MUSEUM
aN
=a?
ae
QO
\
i
\
Fic. 4. Pseudodiaptomus charteri sp. n. a, 2 lateral; b, 2 genital segment lateral; c, g dorsal; d, fifth
leg 2; e, posterior angle of g metasome lateral; f, 2 urosome dorsal; g, ¢ fifth legs; h, first
swimming leg; 7, fourth swimming leg.
PSEUDODIAPTOMIDAE OF SOUTHERN AFRICAN WATERS 383
Description
FEMALE: Length 1-40-1-55 mm. Prosome slender, viewed dorsally more than
two and a half times as long as wide. Head rounded anteriorly. Fourth and
fifth pedigerous segments fused. Posterior angles of metasome bearing a sharp
spine on each side near the middle of the posterior margin. Rostrum well
developed with two strong filaments which reach the middle of the proximal
segment of the first antenna.
Urosome (fig. 1 f) 4-segmented and less than two-thirds as long as the
prosome. Genital segment (fig. 4 6) asymmetrical with irregular lateral swellings
in the posterior part and a prominent genital boss ventrally. A few coarse
teeth are present on the postero-dorsal margin, some setae dorsally and laterally
and a patch of fine hairs on the mid-dorsal surface. The genital boss is similar
to that of P. hesser with genital flaps, setae and an anterior fringe of spinules.
The second and third urosome segments are furnished with rows of coarse
teeth on their postero-dorsal margins. The caudal rami are divergent, more
than 3 times as long as wide and furnished with fine hairs on their inner margins.
The caudal setae are all similar, unjointed but tapering sharply to the thinner
distal part.
Antenna 1 of 21 segments, reaching the second urosome segment when
extended. Proportional lengths, setation and arrangement of aesthetasks as in
P. hesset.
Antenna 2, mandible, maxilla 1 and 2 and maxillipeds all apparently
identical with those of P. hessei. The same peculiar branched setae are present
on the maxillipeds.
‘Swimming legs 1-4 as in P. hesser except in the following respects. In the
first leg (fig. 4 h) the terminal exopod spine is shorter and there are apparently
no spinules on the first basal segment. In legs 2-4 (fig. 1 7) the second basal
segment is longer and narrow proximally and there are fine short hairs on the
outer margin of the exopod and between it and the endopod on the second
basal segment. ‘The setal formula is identical with that of P. hessez.
The fifth pair of legs in the female (fig. 4 d) is similar to those of P. hessev.
The inner processes of the second basal segment are furnished with minute
spinules medially. The first outer spine is minutely serrate. A single hair is
present on the medial margin of segments 3 and 4. The spiniform process of
the fourth segment reaches beyond the midpoint of the terminal spine. Terminal
spine, including the basal portion, slightly shorter than the third segment.
Ege sac as in P. hesset.
Mate: Length 1:25-1:30 mm. Habitus similar to the female (fig. 4 c). Posterior
angles of the metasome bearing very small spines on the posterior margin
(fig. 4 e). Urosome 5-segmented, as in P. hesset except for some postero-lateral
spinules on the first segment and a mid-ventral patch of hairs on the second
segment. Caudal setae jointed about one-third of their length from the proximal
end as in the male of P. hessei, Left antenna 1 as in female. Right antenna
384. ANNALS OF THE SOUTH AFRICAN MUSEUM
1 geniculate with 21 segments, reaching the second urosome segment when
extended. Proportions and setation as in P. hesset.
The male fifth legs (fig. 4 g) are similar to those of P. hessei in general form
but show the following differences. In the right leg the first basal segment is
naked and smooth, second basal segment without inner spinose processes or
outer row of spinules. Outer spine of first exopod segment straight and not
reaching the base of the second outer spine. The latter non-plumose and not
reaching beyond the thickened basal portion of the terminal hook. There are
some rugose patches on the medial and distal margins of the second segment
but no inner swelling. Terminal hook not as long as the two exopod segments.
In the left leg the second basal segment is without medial or outer spinules.
The endopodite is apically rounded and bears only 2 fine hairs near the apex.
The second exopodite segment is a different shape to that of P. hessez, with the
outer distal margin only slightly convex and the inner distal margin strongly
convex. The outer marginal spine is longer and more proximal.
The spermatophore is similar to that of P. hesse1, 400-600 long. Frequently
more than one are attached to a female.
CoLour: Unpigmented except for some orange coloration in the anterior part
of the metasome.
Types
Types are in the South African Museum, Cape Town. Holotype, male,
S.A.M. A10g59; allotype, female, S.A.M. A1og60; paratypes, S.A.M. A1og61;
from St. Lucia Lagoon, Natal, plankton sample STL-—Ga, collected by
R. 8S. Crass on 21/9/54. !
Remarks
This species is named after Mr. R. R. Charter in recognition of his extensive
unpublished work on the Copepoda of South Africa. P. charter: differs from
P. hessei in a large number of characters which are mentioned in the above
description. It may be distinguished most easily in the female by the caudal
setae which are all similar and in the male by the fifth legs, particularly the
shorter marginal spines on the right leg.
Pseudodiaptomus nudus ‘Tanaka
(Figs. 5 a7)
Pseudodiaptomus serricaudatus (non 'T. Scott 1894). Cleve 1904, 196.
Pseudodiaptomus nudus Tanaka 1960, 47, pl. 21, figs. 1-9.
Occurrence
Present in neritic waters off the Cape and abundant over the Agulhas bank.
Found mostly in sea water with salinity near 35°/ 9) but does penetrate into
water of reduced salinity.
PSEUDODIAPTOMIDAE OF SOUTHERN AFRICAN WATERS 385
Fic. 5. Pseudodiaptomus nudus Tanaka. a, ¢ dorsal; b, ¢ urosome lateral; c, 9 lateral; d, 2 urosome
dorsal; e, 2 genital segment ventral; f, fourth swimming leg; g, first swimming leg; /, fifth leg 9;
J, fifth legs 3.
386 ANNALS OF THE SOUTH AFRICAN MUSEUM
Known distribution
Station 8 of the Japanese Antarctic Research Expedition off Cape Agulhas |
(Tanaka, 1960).
Description
FEMALE: Length 1:30-1:40 mm. Prosome slender, viewed dorsally approxi-
mately two and a half times as long as wide (fig. 5 c). Head rounded anteriorly.
Posterior angles of the metasome rounded and naked. Rostrum well developed
with 2 strong filaments which reach the proximal segment of the first antenna.
Urosome (fig. 5 d) 4-segmented, slightly over half as long as metasome.
Genital segment (fig. 5 ¢) asymmetrical, the right side inflated and the left side
depressed, and a prominent genital boss ventrally. The right distal angle is
furnished with a group of small spines. The left lateral swelling bears 2 setae
distally and some minute spinules. The genital boss is similar to that of P. hessez
but also bears spinules on the periphery. There are rows of coarse teeth on the
postero-dorsal margins of the second and third urosome segments and a few
on the genital segment. Proportional lengths of urosome segments are given by
Tanaka (1960). The caudal rami are divergent, approximately 3 times as long
as wide, and the setae are jointed and not expanded.
Antenna 1 of 21 segments, reaching the posterior margin of the second
urosome segment when extended. Proportional lengths of segments are given
by Tanaka (1960). Setation and arrangement of aesthetasks as in P. hessev.
Antenna 2, mandible, maxilla 1 and 2 and maxillipeds all apparently
identical with those of P. hessez. The same peculiar branched setae are present
on the maxillipeds.
Swimming legs 1-4 resemble those of P. hessez except in the following
respects. In the first leg (fig. 5 g) the terminal exopod spine is serrate and about
as long as the last 2 exopod segments. On the first exopod segment there are 3
small spinules proximal to the outer marginal spine which is shorter than in
P. hessei. The second basipod segment bears a small inner distal spinule. The
distal endopod segment bears 6 setae. In swimming legs 2-4 (fig. 5 f) the distri-
bution of fine short hairs is similar to that in P. chartert but even more extensive.
The ornamentation of the swimming legs may be represented as follows:
LEG PROTOPOD ENDOPOD ExXoPop
Si Se | Si’ Se’'| Si Se | St Se | Si St Se] Si Se} Si Se SipeSteesc
Jee LO i © |e £6) TS gicOid ly hoya Aue ie k 1 IV |.1... © | {35a hae
P, La (O) Or nO Tit On ale) ee Oa aeA emo aD eee | 1. . 1o| 255) Seer
ee gC) oO, 0 100 22 KOpa\ Aue eo ienten | I BUS ae
Ie, i 1@) Oo Oo i 0 2 © Ae aA DUNT yi Rea ie | t Di spa ieee
The female fifth legs (fig. 5 h) are uniramous and 4-segmented, lack
medial processes on the second basal segment, and are nearly but not quite
PSEUDODIAPTOMIDAE OF SOUTHERN AFRICAN WATERS 387
symmetrical. The terminal spine is shorter than the spiniform process of the
fourth segment, and bears a slender plumose seta in place of the short, stout,
serrate spine of P. serricaudatus.
The egg sac is single and similar to that of P. hessez.
Mate: Length 1-15-1-20 mm. Habitus similar to the female (fig. 5 a). Urosome
(fig. 5 b) 5-segmented and similar to that of P. hessez. Proportional lengths of the
segments are given by Tanaka (1960). The second urosome segment bears a
mid-ventral patch of hairs. Caudal setae as in the female. Antenna 1, 21-
segmented, right geniculate, as in P. hesset. Proportional lengths of the segments
are given by Tanaka (1960). The male fifth legs (fig. 5 7) are in the main
identical with those of P. serricaudatus (T. Scott) (1896), but in the right leg
the second marginal spine reaches as far as the terminal hook does. The endo-
pod is more complex than in serricaudatus. ‘There are also minor differences in
setation and proportions. The spermatophore is similar to that of P. hessez.
CoLour: Scattered orange pigment is present throughout the metasome and
the female genital segment. The urosome is colourless apart from a distinctive
red patch in the anal segment.
Types
3 99,8 3d and 1 juv. deposited in the National Science Museum of Japan.
There are many hypotypes and topotypes in the South African Museum
(S.A.M. Arog62).
Remarks
Pseudodiaptomus nudus Tanaka differs from P. serricaudatus (T. Scott) (1896),
to which it is closely allied, in the following characters. In both sexes the pos-
terior thoracic margin is spineless, from which (presumably) the name nudus
is derived. This species is somewhat larger. In the female the genital segment is
asymmetrical and the terminal spine and seta of the fifth legs are different.
In the male fifth legs the right endopod is more complex and the terminal
spine of the right leg is longer. P. nudus is easily distinguished from the other
South African species by the naked posterior angles of the metasome in the
female and the characteristic fifth legs of the male. The above account of this
species includes some small additions and corrections to Tanaka’s (1960)
description. I am somewhat doubtful as to whether this species is really dis-
tinct from serricaudatus but a re-examination of T. Scott’s types will be necessary
to determine this, for his original description is inadequate. I have examined
Cleve’s (1904) specimens of serricaudatus in the South African Museum (S.A.M.
A2067, A2068) from off Cape Infanta, and they are of the nudus form.
Both nudus and serricaudatus should be maintained in the genus Pseudodiapto-
mus rather than being placed in the genus Schmackeria as suggested by Marsh
(1933). They do not possess the characteristic long, curved projection on the
inner border of the second segment of the left fifth leg of the male, but have a
normal endopod similar to that of the African and American species.
388 ANNALS OF THE SOUTH AFRICAN MUSEUM
DIscussION
It is perhaps appropriate now to discuss the relationships of this group of
species, for the description of these three species clarifies the picture of the
distribution of this group round the coast of Africa. It is now clear that the
African species have a number of important features in common and tend to
constitute a distinct morphological and zoogeographical unit. Early authors
tended to overlook this. Mrazek (1894), for example, in his note on P. hessez
compares it to gracilis Dahl (1890) from South America and forbes: Poppe &
Richard (1890) from China. It is now apparent that both of these are really
extremely divergent forms and the only resemblances are those common to the
genus or family. Brehm (1951), when discussing the relationships of P. pauliani
from Madagascar, persists in comparing African and American species, on the
basis of such characters as the possession of a left endopod. He also says that
all attempts to arrange the Pseudodiaptomidae into groups that are mor-
phologically and also geographically common are in vain, but it would seem
that this is now possible. All the species found round the coast of Africa are
characterized by the possession of both left and right endopods in the fifth
legs of the male. Certain oriental species also possess both endopods but they
may be separated on the basis of other features such as the possession of a
“Y’-shaped right endopod in the fifth leg of the male.
A series of species related to P. hessex and P. chartert are found on the east
coast of Africa. P. stuhlmanni (Poppe & Mrazek) (1895) was described from the
estuary of the Quilimane river on the coast of Mozambique north of Beira.
The male fifth legs of this species are almost identical with those of P. chartera
except for the strikingly different right endopod which is spatulate, furnished
with bristles terminally and with a large stout claw at its base. P. salinus (Gies-
brecht) (1896) is a marine species known from the north-west Indian Ocean
and Red Sea (Sewell, 1947). The male fifth legs of this species although still
basically similar in structure to the South African species show further differences
in details and proportions. The two species described from Madagascar,
P. paulian Brehm (1951) and P. batillipes Brehm (1954), are also related but
reveal some distinctive modifications such as the inner projection on the second
basal segment of the left male fifth leg. P. ardjuna Brehm (1953) from Salsette
Island near Bombay on the west coast of India falls into the African group also,
on the basis of the structure of the male fifth legs as described by Brehm (1953).
Ummerkutty (1961), however, has shown that specimens of this species
from off Mandapam in the Gulf of Mannar region of south-east India have a
“Y’-shaped spinous right endopod in the fifth leg of the male such as is found
in the oriental ‘hickmani group’. It would be interesting to re-examine material
of this species from Bombay to see whether this difference really exists.
On the west coast of Africa P. hessez has been recorded from the mouth of
the Congo, while a related species is figured by Marques (1951) from Portu-
suese Guinea.
PSEUDODIAPTOMIDAE OF SOUTHERN AFRICAN WATERS 389
The status of the remaining South African species, P. nudus Tanaka 1960,
is somewhat doubtful. It is obviously very closely allied to P. serricaudatus.
(T. Scott) (1894) described from the Gulf of Guinea, but differs slightly from the
original description and figures. Marques (1947), however, gives figures of
specimens of P. serricaudatus from Portuguese Guinea that resemble the nudus
form. P. serricaudatus has also been recorded from Sierra Leone (Bainbridge,
1959) and Angola (Marques, 1958) and various other localities on the West
African coast. In the Indian Ocean P. serricaudatus has been recorded from the
east and west coasts of India, Ceylon, the Arabian Sea, south Arabian coast,
Gulf of Aden and Red Sea (Sewell, 1947). Sewell suggested that it appeared to
be an Indian Ocean form that had managed to get round the Cape of Good
Hope into the Gulf of Guinea, where it was originally taken. If the nudus form
from the Cape is really distinct it is likely that the Indian Ocean form will also
prove to be different from the originally described West African form, which is
completely geographically isolated.
The left exopod of the fifth leg of P. serricaudatus and P. nudus with two long
curved spines is quite different from that of any other pseudodiaptomid, but the
presence and nature of the left and right endopods indicates that their affinities
are with the other African species.
P. nudus, like P. serricaudatus, appears to be essentially a neritic marine
species not found far from land except over the wide, shallow Agulhas bank.
It penetrates into the mouths of estuaries, however, and can live in water of
reduced salinity. P. hessei and P. charteri, like P. stuhlmanni, are essentially
estuarine species found in water of reduced salinity and in hypersaline lagoons,
and occur only rarely together with P. nudus. There is thus an ecological separa-
tion between these species despite the geographical overlap.
Specimens of all three species occurring in South Africa are considerably
larger than the same or related species from elsewhere on the African coast.
The question of the grouping of the species of Pseudodiaptomus, sensu-lato
and the relationships and zoogeographical significance of these groups is being
considered further in a forthcoming paper.
KEY TO THE SOUTHERN AFRICAN SPECIES OF PSEUDODIAPTOMUS
Key to the females (urosome 4-segmented )
1. Posterior corners of metasome without spines as os HE Sd He nudus
2. Posterior corners of metasome bearing spines me os as Ae oe act eet ag
3. Middle caudal seta broad and blade-like .. Ae ay Ag 7 is hessei
4. Middle caudal seta identical to other setae a x ss i ie charteri
Key to the males (urosome 5-segmented )
In the right fifth leg:
1. Second marginal spine reaching as far as tip of terminal hook .. es ys nudus
2. Second marginal spine much shorter than terminal hook f . ) 48
3. Second marginal spine not reaching beyond thickened basal part af terminal hook chartert
4. Second marginal spine reaching well beyond thickened basal part of terminal hook hesset
390 ANNALS OF THE SOUTH AFRICAN MUSEUM
SUMMARY
Descriptions and figures of the three species of Pseudodiaptomidae occur-
ring in southern African waters are given. A full description is given of P. hessei
(Mrazek, 1894), which was never fully described. A new species, P. charteri,
is described. A further description of P. nudus (Tanaka, 1960) is given. The
affinities of these species with other members of this interesting genus are
discussed. The existence of a distinct African group of pseudodiaptomids
characterized primarily by the possession of both left and right endopods in
the fifth legs of the male is noted. It is considered that nudus and serricaudatus
should be maintained in the genus Pseudodiaptomus and not placed in the genus
Schmackeria as had been suggested. It is further noted that the affinities of the
latter two species lie with the African group of species.
ACKNOWLEDGEMENTS
For going through the manuscript and making useful suggestions, I am
indebted to Col. R. B. Seymour Sewell (Cambridge), Professor J. E. G. Ray-
mont (Southampton) and Professor J. H. Day (Cape Town). Many people
helped collect the material on which this paper is based, but in particular I
should like to thank Professor J. Omer Cooper (Grahamstown), Professor
J. H. Day (Cape Town) and Mr. R. S. Crass (Natal Parks Board).
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.
REFERENCES
BAINBRIDGE, V. 1959. The Plankton of inshore waters off Sierra Leone. London: H.M. Stationery
Office.
BreHM, V. 1951. Pseudodiaptomus pauliani (Crustacea, Copepoda). Der erste Vertreter der
Pseudodiaptomiden in der madagassischen Fauna. Mem. Inst. Sct. Madagascar (A), 6, 4.19-425.
BreHM, V. 1953. Indische Diaptomiden, Pseudodiaptomiden und Cladoceren. Ost. zool. Z. 6,
241-345.
BREHM, V. 1954. Pseudodiaptomus batillipes spec. nov., ein zweiter Pseudodiaptomus aus Mada-
gaskar. S.B. dst. Akad. Wiss. Math.-nat. K1., Abt. I, 163, 603-607.
CievE, P. T. 1904. The Plankton of the South African Seas. I. Copepoda. Mar. Invest. in. S Afr.
3, 177-210.
Dau, F. 1894. Die Copepodenfauna des unteren Amazonas. Ber. Naturf. Ges. Freiburg 1.B. 8,
10-23.
GIESBRECHT, W. 1896. Uber pelagische Copepoden des Rothen Meeres, gesammelt vom
Marinestabsarzt Dr. Augustin Kramer. Zool. Jahrb. (Abt. Syst.) 9, 315-328.
GrEsBRECHT, W., & SCHMEIL, O. 1898. Copepoda. I, Gymnoplea. Tierreich. Lief 6, 1-169.
Goopine, R. U. 1957. On some copepoda from Plymouth mainly associated with invertebrates,
including three new species. 7. mar. biol. Ass. U.K. 36, 195-221.
Herrick, C. L. 1884. A final report on the crustacea of Minnesota included in the orders
Cladocera and Copepoda. Rep. Minn. geol. nat. Hist. Surv. 12, 1-191.
Jounson, M. W. 1939. Pseudodiaptomus (Pseudodiaptallous) euryhalinus. A new sub-genus and species
of copepoda with preliminary notes on its ecology. Trans. Amer. micros. Soc. 58, 349-355-
Margues, E. 1947. Copépodes da Guiné Porguguesa. Ann. Junta. Invest. Ultramar., Lisboa 2,
Tom. 3, 31-46.
PSEUDODIAPTOMIDAE OF SOUTHERN AFRICAN WATERS 391
Marguges, E. 1951. Copépodes encontrados no conteudo gastrico de alguns clupeideos da
Guiné Portuguesa. Ann. Junta. Invest. Ultramar., Lisboa 6, Tom. 4, Fasc. 1, 11-18.
Margugs, E. 1958. Copédodes marinhos de Angola. 2 campanha (1952-1953). Mem. Junta.
Invest. Ultramar., Lisboa (2) 4, 199-222.
Marsh, C. D. 1933. Synopsis of the calanoid crustaceans, exclusive of the Diaptomidae, found
in fresh and brackish waters, chiefly of North America. Proc. U.S. Nat. Mus. 82, 1-58.
MrAzexk, A. 1894. Uber eine neue Schmackeria (Schm. hessei n. sp.) aus der Kongo-Miindung.
S.B. bohm. Ges. Wiss. Math-nat. KI]. 1894, art. 24, 1-3.
MrAzexk, A. 1901. Siisswasser-Copepoden. Ergeb. Hamburg Magalhaens. Sammelr., Lief 6, no. 2,
29 pp. 4 pl.
Poppe, S. A., & MrAzex, A. 1895. Die von Herrn Dr. F. Stuhlman auf Zanzibar und dem
gegentiberliegenden Festlande gesammelten Siisswasser—Copepoden. Mitt. nat. Mus.
Hamburg 12 (1894), 123-134.
Poppe, S. A., & RICHARD, J. 1890. Description du Schmackeria forbesi n.g. et sp. calanide nouveau
recueilli par M. Schumacker dans les eaux douces des environs de Shanghai. Mem. Soc.
zool. France 3, 396-400.
SEWELL, R. B. S—EymMour. 1932. The Copepoda of Indian Seas. Calanoida. Mem. Ind. Mus.
Calcutta 10, 223-407.
SEWELL, R. B. SEymMour. 1947. The free swimming planktonic Copepoda. Systematic account.
Sci. Rep. Murray Exped. 8 (1), 1-303.
Scott, T. 1894. Report on the Entomostraca from the Gulf of Guinea, collected by John
Rattray, B.Sc. Trans. Linn. Soc. Kool. 6, 1-162.
TANAKA, O. 1960. Pelagic Copepoda. Biol. Results Jap. Antarct. Res. Exped. 10, 1-191.
Ummerkutty, A. N. P. 1961. Studies on Indian copepods 4. Description of the female and a
redescription of the male of Pseudodiaptomus ardjuna Brehm (Copepoda, Calanoida) with
notes on the distribution and affinities of the species. 7. mar. biol. Ass. India 2, 179-185.
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Bettrdége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
HartMAN, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
HartMan, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.—All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name. ...
B. Form of bibliographic references to synonymic names.—The first reference following
any name in the synonymy should be to the earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalis capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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REFERENCES. Harvard system—authors’ names and dates of publication given
in the body of the text; references arranged at the end of the paper in alphabetical
order of authors’ names.
Bibliographical arrangement of references at the end of the paper must give:
1. Name of author, followed by his initials. Names of joint authors connected by
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suffixes a, b, etc., are used.
3. Full title of the paper. Initial capital letters only for the first word and for proper
names, except in German, where the usage of the language is followed.
4. ‘Title of the journal, abbreviated according to the World list of scientific periodicals,
and underlined to indicate italics.
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[CONTINUED ON INSIDE BACK COVER
SUPPLEMENTARY CONTRIBUTIONS TO THE REVISION OF THE
BOMBYLIIDAE (DIPTERA) OF SOUTHERN AFRICA: THE GENUS
SYSTROPUS
By
AS Esse
South African Museum, Cape Town
[Accepted June, 1962]
CONTENTS
PAGE
Descriptions of new species . F308
Key to southern African species of
Systropus , : ; = 400
Summary . ‘ : : . 404
Acknowledgement : : . 404
References . : ‘ : - 405
DESCRIPTIONS OF NEw SPECIES
Subfamily Systropinae
Genus sysTRopus Wied.
Systropus Wiedemann 1820, p. 18. Wiedemann 1828, p. 359. Hesse 1938, p. 991.
Since my revision of the South African representatives of this genus in
1938 the South African Museum has acquired some new species from the
Cape Province, Southern Rhodesia and Portuguese East Africa. Descriptions
of these are given below.
Systropus bicoloripennis n. sp.
This peculiar smallish species which differs from all the other known
South African forms in certain respects is characterized as follows:
Body mainly black; frons, sides of face and genae yellowish in 3, dark or black
in 9; buccal rim black in both sexes; antennae entirely black in 9, but segment
1 ivory yellowish and silvery-haired above in J; propleural callosity or a spot
on it ivory yellowish in both sexes; base of sternite 2 and sometimes to a smaller
and lesser extent apex of sternite 2 and base and apices of 3 and even 4 very
pale yellowish; a large transverse humeral spot passing into a narrow noto-
pleural streak on each side, broadened on false tegulae in front of wing-bases,
continued in a curve above wings to include postalar calli and narrowly across
hind margin of thorax, pteropleuron or greater part of it, down middle of
pleurae (posterior part of sternopleuron or middle epimeral part), more or
less anterior half of hypopleural (or posterior episternal) part, the broad anterior
393
Ann. S. Afr. Mus. 46 (16), 1963, 393-405, mSTITi
394 ANNALS OF THE SOUTH AFRICAN MUSEUM
border (much broadened above) of metasternal part, postnotal process and
area around it, a large basal spot on each side of tergite 1, narrowly continuous
with its fellow along anterior margin, broadish sides (narrowed in middle)
and to an obscure extent narrowish dorsal hind margins of tergites 2-5, hind
margins of rest of tergites in g and of 6 and 7 in Q, anterior faces of front coxae,
sides in basal halves and hinder part of middle coxae, a ring-like fascia round
posterior coxae (broader on front face), femora, tibiae and basal segments
of tarsi, excepting only the dark callus-area on front femora and dark apical
parts of hind femora and tibiae, yellowish red, reddish or ferruginous red;
scutellar callosities also ferruginous reddish or brownish. Vestzture with the hairs
on antennae black, excepting silvery white ones on segment 1 above in dg;
those on face also black; pruinescence on head in front silvery; scaling on thorax,
pleurae and especially hypopleural and metasternal parts and on sides of
tergite 1 glittering iridescent and silvery, definitely scale-like, the individual
scales broad, leaf-shaped or ovate, not fine and hair-like as in all the other
known South African forms, excepting namaquensis Hesse; fine scaling on
abdomen dark or black in a broad dorsal streak, in a broadish streak on each
side of venter and on club, the dorsal band being broadened at about middle of
tergites 2-5, suggesting a node-like or spindle-shaped structure; scales on
pale coloured sides of tergites 2-5 gleaming sericeous yellowish, becoming
silvery white in apical patches, especially on 2-4, and those across hind margins
of these segments also silvery; fine hair-like scaling on legs greyish or whitish
on coxae, mainly dark on femora and tibiae, but yellowish on inner faces of
hind femora and to a much lesser extent on the others and outer faces of front
and middle tibiae. Head with the eyes above in ¢ separated by only a narrow
line, about or a little more than 3 times length of ocellar tubercle, in 2 by a
narrow space about as wide as front ocellus and a little shorter than ocellar
tubercle; antennal segment 1 in ¢ longer than in 9 and also more distinctly
longer than segments 1 and 2 combined; segment 2 markedly short, markedly
broadened apically, more so in 3; segment 3 flattened, narrowed and pointed
apically, apparently slightly longer in 9, a little more than 2, to nearly 3,
times as long as 2 in ¢ and about 2 to 3 times as long as 2 in 9; proboscis
about 2-5-3 mm. Thorax, excluding scutellum, subquadrate, only a little
broader than long, both it and scutellum above fairly coarsely rugulosely
punctured; scutellum transverse, markedly broader, or much broader, than
long; hypopleural and metasternal parts rugosely punctured, the latter trans-
versely grooved behind; postnotum on each side at base of tergite 1 produced
into a distinct or prominent, somewhat triangular, dentate process or spine
which in most other species is blunt or only rounded. Wings with a very charac-
teristic pattern of infuscation, the basal part up to level of end of axillary lobe
and the anterior costal half to discoidal cell and cubital fork brown to dark
brown in both sexes, but evidently slightly darker anteriorly, with the spot-like
infuscations at base of first submarginal cell, on apical cross veins of second basal
cell, on middle cross vein and at base of upper cubital branch darker, con-
REVISION OF THE BOMBYLIIDAE (DIPTERA) OF SOUTHERN AFRICA 395
spicuous; entire discoidal cell, almost entire first posterior cell (excepting its
extreme base only), second submarginal cell and more or less apical half of
third posterior cell uninfuscated and clear, sharply demarcated and con-
trasting with especially the anterior infuscated half; only two submarginal
cells present; alula entirely wanting; vein between submarginal cells less
sinuous than in most other species; first posterior cell markedly narrowed and
acuminate apically, often closed and subtending on hind margin or even
shortly stalked; middle cross vein much beyond middle of discoidal cell; knobs
of halteres brownish to dark brown above. Abdomen rather thick and stout, its
petiole composed of 3 segments; last sternite of 2 elongate, shining, spout-like,
sharply pointed and slightly produced apically where it is also laterally com-
pressed, and apex slightly bent downwards; last tergite of 3 produced apically
below on each side into an inwardly directed and upwardly curving process,
the apices of which are slightly dilated and rounded; beaked apical joints of
fused basal parts (taking up the position of a last sternite) with their sharp
spine-like or hook-like apices curved inwards and slightly downwards; exposed
and projecting part of medial aedeagal process stoutish, rod-like, rounded and
slightly dilated apically; accessory processes on each side sharply pointed
(similar to those depicted for macilentus Wied., cf. Hesse 1938, p. 1004, text-fig.
310); fused apical part of lateral rami (cf. text-fig. 310, R.) tongue-shaped,
curved down towards aedeagal process and rounded apically; callus-area on
terminal plates in last tergite roughly bean-shaped. Legs with the hind femora
unarmed and rather abruptly clavate apically.
From 17 §¢ and 9 99 (types in the South African Museum).
Length of body: about 9-25-13 mm.
Length of wing: about 5-8 mm.
Localities. Western Cape: Leipoldtville-Elands Bay (Mus. Exp., Oct. 1947)
(types, 14. ¢ and 6 @ paratypes); Ysterfontein (S.A.M., Sept. 1960) (1 @
paratype). Namaqualand: Bowesdorp (S.A. Mus. Exp., Sept. 1941) (2 ¢ and
1 Q paratypes).
The smallish size, striking bicolorous wings, much narrowed or even
stalked first posterior cell, flattened iridescent and silvery scales on thorax and
metasternum, transverse scutellum, the dentate postnotal processes and very
short second antennal segments as well as the distribution of the red on body,
distinguish this species from all other known South African forms. It is apparently
restricted to the west coastal belt of the Cape. In general appearance and method
of flight it simulates species of the Conopid genera Conops and Physocephala.
Systropus namaquensis Hesse
Systropus namaquensis Hesse 1938, p. 1006.
Since the publication of the specific description of a damaged 3 specimen
of this species which I compared with macilentus Wied. at the time, the Museum
has acquired numerous other 33 and also quite a number of 2 specimens. ‘To
396 ANNALS OF THE SOUTH AFRICAN MUSEUM
include the as yet undescribed 9 the original description may therefore be
supplemented as follows:
Body mainly black; frons, face and genal parts mainly dark, with silvery
pruinescence, except on middle part of face; buccal rim ivory yellowish;
antennae entirely black and black-haired; following parts yellowish red to red:
a humeral spot on each side continued as a narrow notopleural line to the broad
false tegulae and to include the postalar calli, propleural callosity and sclerite
above it, middle sutural (or posterior sternopleural and anterior hypopleural)
part, lower part of posterior episternal (or hypopleural) part, metapleural
part to a certain extent, sides of tergites 2-5, greater part of hind femora
(excepting their black apical parts), at least or more than basal halves of hind
tibiae, at least outer faces of front and middle tibiae, and sometimes to a
much lesser and variable extent outer faces of anterior and middle femora;
tergite I sometimes obscurely dark reddish at its basal corners like the exposed
postnotal parts below; scutellar callosities flattened, ivory yellowish. Vestzture
peculiar in that, as in bicoloripennis n. sp., the fine silvery whitish hair-like scales
or hairs of most other species are in this case replaced by distinct, narrow,
flattened scales (or scale-like hairs) on thorax, especially posteriorly, on scutel-
lum, hypopleural and metasternal parts and on sides of tergite 1. Head with
the eyes above contiguous in ¢ for a distance a little more than 3 to about 4
times length of ocellar tubercle, narrowly separated for a short distance in Q,
narrowest part being subequal in length to that of tubercle and about as wide
as front ocellus; frons in 9 thus much longer; antennal segment 1 in ¢ longer
than in 9; segment 2 relatively short, equally long in both sexes; segment 3
flattened, more sharply pointed apically, slightly shorter in g, about 3 or a
little more (34 or nearly 4) times length of 2 in ¢ and from 44 to nearly 5 times
length of 2 in 9, subequal or more usually shorter than segment 1 in 4, but
longer than 1 in 9. Wings with only 2 submarginal cells, smoky greyish to dusky
in 4, darker in costal part and marginal cell, distinctly darker and very much
more infuscated in 9, smoky brown, more so basally and in anterior half; first
posterior cell only slightly narrowed apically; knobs of halteres very dark
brown to almost black above. Abdomen rather thick and long; petiole composed
of 4 segments; last sternite in 9 elongate, scoop-like; last tergite in ¢ like that of
macilentus Wied., with inwardly and upwardly curved, apically thickened
apical prongs, which are however distinctly more U-curved; exposed part of
hypopygium in g with the beaked apical joints similar to those of macilentus,
but their inwardly tumid beaks apparently blunter, the aedeagal process more
broadened and bifid apically and the accessory processes similar, but the other
on each side dorsally which I labelled as ‘ramus’ in macilentus (cf. text fig. 310,
p. 1004, 1938) is fused apically with its companion and is produced into a
flattened pointed process.
From 19 @¢ (including the original holotype) and 12 99 in the South
African Museum.
REVISION OF THE BOMBYLIIDAE (DIPTERA) OF SOUTHERN AFRICA 2070
Length of body: about 13—18-5 mm.
Length of wing: about 8-10 mm.
Localities. Namaqualand: Giftsberg (van Rhynsdorp) (Sept. 1911) (original
holotype §); Wallekraal (Mus. Exp., Oct. 1950) (1 ¢); Graafwater (Mus.
Exp., Oct. 1947) (Q allotype, 4 9 paratypes and 4 ¢¢). Koup Karoo: Lammers-
kraal in Prince Albert Dist. (Mus. Exp., Sept. 1947) (2 9 paratypes and 8 33);
Koup Siding—Laingsburg (Mus. Exp., Oct. 1952) (1 9 paratype). Moordenaars
Karoo: Lammerfontein (Mus. Exp., Oct. 1952) (1 2 paratype and 1 4).
North Western Karoo: Augusfontein near Calvinia (Mus. Exp., Sept. 1947)
(3 2 paratypes and 4 3).
Systropus sheppardi n. sp.
A large species which superficially resembles the wasp Belonogaster junceus
(Vespidae) and which is characterized as follows:
Body mainly black; frons, face, genal parts, interior of buccal cavity, first
antennal segments, palps, and base of proboscis yellowish or pale yellowish
reddish; following parts ferruginous reddish: a broad humeral quadrate patch
on each side anteriorly on thorax, continued as broad sides of thorax, including
the false tegulae, to postalar calli, sides basally of scutellum, metanotal part,
area around wing-bases, posterior part of pteropleuron, middle episternal
and epimeral (or sternopleural) parts, metapleural part, anterior margin and
lower lappet of posterior episternal (or hypopleural) part, the extreme upper
margin of metasternal part, the area in upper half dividing the metasternal
parts, the narrowish area surrounding the base of postnotal process on each
side, the coxae and legs; a variable spot on propleural callosity, and scutellar
callosities yellow; sides of tergites 2-4, sides of base of 5 and entire club of
abdomen black; exposed parts of hypopygial structures also ferruginous.
Vestiture with the hairs on antennae and on face black; those on declivous
anterior part of thorax in humeral region and a characteristic brush of dense
hairs in groove on thorax above wing-base on each side also black; frons,
face and genae with silvery pruinescence; fine hairs or hair-like scaling on
thorax above composed of greyish and dark ones, mostly in broadish streaks,
those posteriorly and on scutellum longer, more silvery; hair-like scaling on
pleurae relatively sparse, denser posteriorly and on metasternal part, silvery
white, but relatively shorter than in most species; fine scaling on abdomen
dark or black on dark parts, yellowish on pale parts, but those on last two or
three segments of club greyish white or silvery; scaling on legs mainly dark,
even on middle and hind femora below, pale basally below on front ones;
fine hairs or hair-like scaling on callosities on front femora (in certain lights),
on outer faces of front and middle tibiae silvery whitish, the dark ones on front
or inner faces gleaming golden in certain lights, especially in 9. Head with the
eyes in both sexes in contact above for a little more than 3 times length of
ocellar tubercle; frons more depressed centrally in 2; antennal segment |
398 ANNALS OF THE SOUTH AFRICAN MUSEUM
slender, elongate, very slightly longer in ¢ than in 9, about or almost as long
as last four tarsal segments together, a little more than 3, or nearly 4 times
length of segment 2 (segment 3 missing in specimens); palps quite or nearly
half length of antennal segment 1; proboscis long, slender, about 6:52-6:72 mm.
Thorax dull above, leathery in appearance, more finely punctured in hinder
half, slightly more coarsely on scutellum; metasternal part transversely grooved.
Wings dusky, tinged smoky brownish, very much darker in 9, darker in costal
part in both sexes, but more so in 9; veins dark brown; two submarginal cells
present; first posterior cell narrowed apically; alula and outer squamal lobe
wanting; knobs of halteres very dark or black above. Abdomen long, with a
petiole composed of three segments; last sternite in 9 elongate, produced
apically into two sharp, spine-like processes; last tergite in g only deeply
emarginate apically, not armed with a spine or process on each side; exposed
hypopygium of ¢ with the fused basal parts (ventral in position) broad, scoop-
like, truncate apically, armed apically on its inner side on each side with an
inwardly directed, curved, somewhat flattened, spine-like apical segment
(clasper) which is also curved downwards apically; exposed accessory struc-
tures or aedeagal processes in form of a medial, dorsally directed, broadish,.
hook-like structure, flanked on each side by a broad, flattened, apically rounded,
lobe-like structure. Legs without spines on hind femora below, the latter not
clavate apically; hind legs relatively long and with rather well-developed
spicules on hind tibiae.
From a ¢ and 2 99 (¢ holotype and @ allotype in the South African
Museum and 1 @ paratype in Transvaal Museum).
Length of body: about 25-25:5 mm.
Length of wing: about 14-14°5 mm.
Localities. Southern Rhodesia: Vumba in Umtali Dist. (P. A. Sheppard, March
1931) (¢ holotype); Umtali Dist. (P. A. Sheppard, April 1931) (Q allotype
and 2 paratype).
Distinguished from the other large Belonogaster-like Rhodesian species
marshalli Bezz. by its different colour-pattern in which the base of the thorax,
scutellum, propleural callosities and base of tergite 1 are not reddish, and the
wings are not yellowish, but dusky or smoky brownish. From snow: Adams, a
slightly smaller species from Rhodesia with a similar colour-pattern, it may,
apart from the larger size, be at once distinguished by the much smaller
propleural yellow spot, the non-pallid outer faces of front and middle tibiae,
less red on lower part of hypopleuron, presence of black hairs on face, anterior
humeral part and densely in groove above wing-bases, and the much darker
wings.
Systropus gracilis n. sp.
A slender-bodied species superficially resembling Jleptogaster Lw., but
agreeing and differing from the latter as follows:
REVISION OF THE BOMBYLIIDAE (DIPTERA) OF SOUTHERN AFRICA 399
Body mainly black; frons, face, genal parts, humeral angle and a fascia from
it down on each side to include propleural callosity (but not transversely on
thorax above as in leptogaster), front coxae, apical halves of front femora, part
of underside of middle femora and outer faces of front and middle tibiae and
their basal tarsal segments, and scutellar callosities ivory yellowish; hind
femora below even more extensively pallid or yellowish and white-scaled than
in leptogaster; thorax above on false tegulae without any oblique yellow fascia or
spot; pleurae entirely black, the middle sutural part between sterno- and
hypopleural parts dull velvety black, not reddish; tergites 2-5, especially 4
and 5, laterally also ochreous yellowish, but even bases of 6 laterally also
yellowish; apical halves of front and middle tarsi black; apices of hind tibiae
and tarsi also black or dark. Vestiture with the fine tomentum on frons, face
and genae silvery; fine hairs on antennae black; fine and sparse hairs on disc
of thorax appearing greyish yellowish to brownish in certain lights; hairs or
hair-like scales on pleurae, metasternum, scutellum, and sides of tergite 1
silvery white, those on metasternum longer, but sparser than in leptogaster;
fine hair-like scaling on disc of tergite 1 blackish brown; that on dorsum of rest
of tergites and also on sides ventrally below black like most of those on abdomi-
nal club; that on sides of tergites 2-5, but less so on sides of 2, yellowish; fine
hair-fringes across hind margins of same tergites more silvery white, more
conspicuous than yellowish or dark ones of leptogaster; fine scaling on upper
sides of femora and tibiae dark, contrasting with pale ones on lower surfaces
and with the silvery white fine hairs and scales on outer sides of front and
middle tibiae and basal segments of their tarsi. Head with the eyes above in g
in contact for a very much shorter distance, only a very little more than 3
times length of ocellar tubercle, not actually about 5 times as in leptogaster;
antennal segment 3 longer, elongate, a little longer than segment 1, more or
less spindle-shaped and bluntly pointed and about or nearly 3 times length of
segment 2 in ¢ (proportions of segments 7 : 2-6 : 8) whereas in ¢ of leptogaster
segment 3 is about or only a little more than twice length of segment 2; pro-
boscis about 3-6 mm. long. Thorax relatively much shorter (excluding scutel-
lum), distinctly much shorter than width of head, its upper surface shining,
more coarsely rugosely punctured (dull and more finely rugulose in leptogaster) ;
scutellum shorter, more rounded and not subemarginate posteriorly; meta-
sternal part distinctly much narrower, more finely transversely rugose, without
the normal transverse groove-like sculpture of most other species, in profile
sloping more sharply forwards, the hind coxae appearing shifted far forwards
to below level of neck (in leptogaster to about below middle of thorax). Wings
tinted only a little less smoky greyish than in ¢ of leptogaster, the costal cell
also darker and similarly yellowish brownish, veins brownish; spot-like infusca-
tions on middle cross vein and base of discoidal cell, apart from one at common
base of second and third veins, slightly less conspicuous than in leptogaster;
a delicate, shiny, hyaline, lobe-like outer squama (or part of alula) present
which in leptogaster is entirely wanting as in other known South African species ;
400 ANNALS OF THE SOUTH AFRICAN MUSEUM
three submarginal cells present, but base of normal second submarginal cell
distinctly very much shorter; middle cross vein apparently nearer middle of
discoidal cell; first posterior cell less narrowed apically; knobs of halteres
paler, more yellowish brown above. Abdomen more slender, but also with a
three-segmented petiole, though segment 4 is relatively less broadened (side
view) and may almost be considered as a fourth segment of the petiole. Legs as
in leptogaster; hind femora not clavate apically, only gradually thickened.
Hypopygium of 3 with the callus-area on terminal plates in last tergite very
much broader and more oval than in leptogaster; aedeagal process flattened,
ending into two slightly diverging flattened lobes, not cowl-like or hood-like
as in latter.
From a 4 in the South African Museum.
Length of body: about 11-5 mm.
Length of wing: about 6-5 mm.
Locality. Portuguese East Africa: Nyaka (R. Lawrence, Feb. 1924).
Superficially this species also resembles the species zchneumoniformis which I
described from the Congo (1958, p. 64), and which also has similar hyaline,
lobe-like, outer squamae at base of wings, dusky wings, shiny thorax, elongated
third antennal segments, etc. From the latter, gracilis differs, however, in its
ivory yellowish face, genal parts, ivory yellowish fascia from humeral angle to
coxae, ivory yellowish front coxae, more yellowish legs, shorter, more convex
and more coarsely sculptured thorax, much longer line of contact between
eyes above, etc.
KEY TO THE SOUTHERN AFRICAN SPECIES OF SYSTROPUS
A revised and descriptive key to all the known southern African species.
1(a) Wings with only two submarginal cells. . , : : : : i =»
(b) Wings with three submarginal cells. 3 5 : ‘ 5 nae Ta
2(a) Antennal segment 2 distinctly much shorter, much less than half length of segment 3,
or shorter than, subequal in length to, or only a little longer than anterior tarsal segment 3;
club of abdomen without or with scarcely any, or with only poorly developed, silvery
hairs in apical part, these hairs if pale usually not silvery white; pteropleuron without or
with only very short silvery hairs or hair-like scales posteriorly, and sides of tergite 1 and
metasternal part with sparser and usually shorter silvery elements; outer faces of front
and middle tibiae usually without conspicuous, resplendent, silvery hairs or pubescence,
and if yellowish, with feebler and less conspicuous silvery hairs, and if resplendently
silvery, antennal segment 2 short; hind femora more abruptly and ne clavate
apically. . ; ®
(6) Antennal segment 2 distnedly ee jensen ais or alae? half lenedh of 3, or arnt or
very much longer than anterior tarsal segment 3; club of abdomen usually with con-
spicuous or dense silvery hairs or with conspicuous silvery-gleaming pubescence in
apical part; pteropleuron with more conspicuous and distinctly longer (or with a tuft
of distinctly longer) silvery hairs posteriorly, and sides of tergite 1 and metasternal part
usually also with longer and much denser silvery elements; outer faces of front and
middle tibiae, even if not yellowish, usually with conspicuous, Pare silvery hairs;
hind femora not or scarcely abruptly clavate apically. . : : 5 Ca
3(a) Wings markedly short, anterior half and basal part up to end of ee lobe darkly
infuscated, the cross veins more darkly so, but rest or lower apical half clear; first pos-
(0)
(0)
5(a)
(0)
6(a)
REVISION OF THE BOMBYLIIDAE (DIPTERA) OF SOUTHERN AFRICA 401
terior cell very much narrowed apically, acute, closed or sessile on hind margin; antennae
relatively short, segment 2 markedly broadened apically and 1 whitened and white-
haired above in 43; postnotum on each side at base of tergite 1 produced spine-like or
dentately prominent; abdomen relatively thick or stoutish, with 3-segmented petiole,
the apical parts of which have a large quadrate patch of fine silvery hair-like scaling
on each side; base of sternite 2 conspicuously bone yellowish; mesopleural, posterior
episternal and epimeral (metasternal) parts and sides of tergite 1 with small broadish
iridescent or pearly scales; hind femora without spines; smallish form, not more than
13 mm. : - 6 & bicoloripennis n. sp. (W. Cape, Namaqualand)
Wings relatively loneen ant SO , (lagers infuscated, either tinged uniformly through-
out or with only costal part darker and with the spots on cross veins less conspicuous;
first posterior cell either not or only slightly narrowed apically, usually broadly open
apically; antennae distinctly much longer, segment 2 not or less broadened apically,
and 1 not whitened above, dark-haired in both sexes; postnotal angle on each side at
base of tergite 1 less sharply prominent, bluntly angular or rounded; abdomen distinctly
more slender, with 4-segmented petiole, the apical parts of which with more yellowish
or pallid fine scales or hairs or with darker hairs; base of sternite 2, if pale, more normally
yellowish; pleural, metasternal parts and sides of tergite 1 with silvery hair-like scales or
with longer, less broadish silvery ones; hind femora usually with some spines below at
about middle; larger forms, usually more than 13 mm. . ; : ees)
Wings distinctly more darkly infuscated, either in both sexes, or in 99 more darkly
than in $4; propleural tubercle, front coxae and outer surfaces of front and middle
tibiae castaneous brownish, reddish brown or yellowish brown, both pairs without even
faint silvery hairs; apices of hind femora and tibiae conspicuously dark or blackened,
greater part of hind legs being pale reddish or yellowish brown; hyposternal and meta-
sternal parts or at least hind part of metasternum black
‘Wings less dark, more cinereous or smoky greyish, darker only at (ase and aastall part,
especially in 2; propleural tubercle, front coxae, a subapical spot on front femora below
and outer faces of front and middle tibiae ivory yellowish or whitish, both pairs with
faint, but distinct, silvery whitish hairs on ivory faces; legs dark or black, hind ones
entirely so; pteropleuron, middle epimeral, metapleural, posterior episternal and epi-
metal (or metasternal) parts entirely pale reddish or yellowish brownish.
3d & barnardi Hesse (1938, p. 1006) (S.W. Cape)
Wings in both sexes more uniformly reddish brownish, only very slightly darker in 9;
halteres slightly paler, the knobs more yellowish brownish to castaneous brown above;
entire or greater part of antennae, face, sides of thorax broadly, base of thorax, apical
spot on or hind margin of scutellum, pteropleuron, middle epimeral, metapleural,
posterior episternal parts and front part of posterior epimeral (or metasternal) part,
base of tergite 1, tergites 2-5 above and medially below, and legs including basal seg-
ments of tarsi pale reddish brown, yellowish brown to ochreous brownish; silvery vesti-
ture on thorax, pleurae, metasternum and sides of tergite 1 in form of finer hair-like
scales or hairs. . & & macilentus Wied. (1938, p. 1001) (W. Cape, Karoo)
Wings 1 in 9 very dark sina brownish, darker in anterior half and basally, less infuscated
in g, more cinereous or smoky greyish, darker only in costal part and base; halteres
darker, more dark castaneous to blackish brown, the knobs dark to almost Black above;
antennae, face or middle part of face, upper half or even greater part of genae, greater
part of thorax above, entire scutellum, greater part or entire pteropleuron, hypopleuron
or posterior episternal part (excepting only its reddish lower part), entire posterior
epimeral part, entire tergite 1, dorsum and sides below of tergites 2-5 and abdominal
club black; coxae, front and middle femora, more or less apical parts of their reddish
brown tibiae, apical parts of hind femora, almost apical halves of hind tibiae, and all
the tarsi very dark to black; silvery vestiture on thorax, pleurae, metasternum and sides
of tergite 1 tending to be more in form of scale-like hairs or even scales in part.
3 2 namaquensis Hesse (W. Karoo, Namaqualand)
Wings entirely glassy hyaline in ¢ at least; antennal segment 1 entirely dark or black,
apical part of 3 more distinctly and more sharply produced outwards; humeral angle
and across anterior spiracle, including propleural tubercle, a large rounded spot on each
402
(0)
7(a)
ANNALS OF THE SOUTH AFRICAN MUSEUM
side of thorax above wings, postalar calli, front coxae and to a certain extent outer face
of front tibiae ivory whitish; vestiture on posterior episternal (hypopleural) and epimeral
(or metasternal) parts slightly longer, denser, that towards apex of abdomen not con-
spicuously silvery; black callus-area on terminal plates of last sternite (tergite) of gd
very narrow, linear. : - & munroi Hesse (1938, p. 1010) (Transvaal)
Wings even in ¢¢ not ids skis hyaline, either tinged smoky brownish or with at
least costal and basal parts brownish; antennal segment 1 not entirely dark, either basal
part, basal half or entire segment ‘acral, reddish brown or reddish, ain apical part
of 3 even if acute not distinctly produced or prolonged; humeral angle, anterior spiracular
part, spot or macula above wing-base and postalar calli not ivory whitish, usually
reddish or ferruginous red; vestiture on hypopleural and metasternal parts on the whole
shorter, less dense and shaggy (if dense and longish other characters do not differ), that
towards apex of abdomen usually more conspicuously silvery; callus-area on terminal
plates of last sternite (tergite) of ¢¢ distinctly broader. ; : ‘ : Lie Sie
Wings glassy hyaline, but base, costal cell and marginal cell to a variable extent in g,
and in @ also greater part of first basal cell, to a variable extent base of second basal cell
and also basal half of first submarginal cell dark brown or reddish brownish; knobs of
halteres pale above; frons, entire antennal segment 1 at least above, face, genae and
base of tergite 1 in addition to other red on thorax pale reddish or pale reddish brown;
propleural tubercle ivory yellowish only in upper part and sclerite above it, black; outer
faces of front and middle tibiae and tarsi, though silvery-haired, only slightly yellowish;
scutellar callosities very dark, blackish brown or black; eyes above contiguous for only a
short distance in both sexes; thorax above distinctly more coarsely rugulose-punctate;
vestiture on body, especially on thorax above, distinctly and markedly denser, longer,
that on antennal segment 1 above and to a certain extent also on 2 denser, longer,
entirely silvery white, especially in 3.
3 2 crudelis Westwd. (1938, p. 1012) (N.W. Cape, Bechuanaland, Transvaal, Natal)
Wings not glassy hyaline, distinctly tinged more greyish or cinereous, smoky or even
dark smoky or reddish brownish, the darker basal and costal parts not contrastingly
demarcated from less dark parts; knobs of halteres dark or black to a variable extent;
first antennal segments darkened in apical part or half or entirely dark or black, but
apical part of frons, face or sides of face, and genae yellow; tergite 1 entirely dark;
propleural tubercle and sclerite above it usually entirely ivory yellowish and if darker
other characters do not differ; outer faces of front and middle tibiae and tarsi usually
much paler or yellowish, contrasting with inner faces; scutellar callosities very pale
yellowish, almost white; eyes above in actual contact for a much longer distance; thorax
above only very finely rugulose; vestiture on body above distinctly very much sparser,
shorter and finer, and that on antennal segment 1 finer, less shaggy, entirely dark or with
fewer intermixed pale ones. . : : : , : é : : sO
Wings distinctly less darkly infuscated in both sexes, only tinted greyish, cinereous or
faintly smoky or yellowish brownish, the basal and costal parts slightly darker, more so
in 99 than in ¢ 4; sides of thorax above continuously pale reddish from broad humeral
spot; legs on the whole very much paler, pale reddish or yellowish reddish, the femora
not or scarcely darkened above, their fine dark scaling present only tan ; vestiture
on thorax above entirely or predominantly whitish. . : 9
Wings distinctly more dusky or darkly infuscated smoky beouralslh, even in 38, but ee
more so in 99, the basal and costal parts being darker still; sides of thorax above darker
red or more ferruginous reddish or the red is smuo anced just behind humeral spot;
legs darker, darker brownish or reddish brown, the femora also distinctly more darkened
above, to a large extent due to denser or more extensive fine black scaling; vestiture
on thorax above not entirely silvery or pale, but with at least some fine dark or black
hair-like scales or hairs either in streaks or on anterior declivous part or above wing-
bases. . : 5 2 : i : é ‘ : : : ; pee 0)
Pleurae predominantly black, the greater part or at least upper half of hypopleuron (or
posterior episternal part), entire metasternal part and metanotal part also black; front
coxae dark or black in front or on sides; tergites 2-5 darker, more blackish above and
8
(0)
10(a)
(9)
I1(a)
(5)
12(a)
REVISION OF THE BOMBYLIIDAE (DIPTERA) OF SOUTHERN AFRICA 403
5 with dorsum and apical half of sides also very dark or black; vestiture on antennal
segment I sometimes entirely black, even in d.
3 2 snowi Adams (1938, p. 1015) (Rhodesia)
Pleurae with greater part a preropicurar: entire hypopleural (or posterior episternal)
part and metanotal part as well as middle part of metasternum behind red; front coxae
paler, tending to be largely pallid or ivory yellowish in front; tergites 2-5 less dark above,
more brownish, and 5 entirely or predominantly pale reddish yellowish or ochreous;
vestiture on antennal segment 1 with more intermixed pale or yellowish hairs or even
pale-haired in basal half in ¢ at least.
3 zuluensis Hesse (1938, p. 1018) (Zululand)
Smaller form, about 14:5 mm.; first posterior cell in wings markedly narrowed apically;
frons, face, entire propleural tubercle and sclerite above it, anterior faces of front coxae
and outer faces of front and middle tibiae and apical half of halteral knobs ivory whitish;
red on thorax above not continuous on sides, the humeral spot being isolated; sterno-
pleural part mainly black and at least lower half of hypopleuron (or posterior episternum)
reddish; hairs in groove above wing-bases sparse and pale, and those on anterior declivity
in front of humerus and on face also pale; fine hairs on hind coxae and hair-like scaling
on femora below pale, sericeous or silvery.
3 fumosus Hesse (1938, p. 1019) (Portuguese East Africa)
Much larger foe About 25-25°5 mm.; first posterior cell only slightly narrowed; frons
and face reddish or reddish brownish, ole a smallish spot on propleural Alvevelo: ivory
yellowish or whitish, the front coxae reddish and outer faces of front and middle tibiae
more pale yellowish reddish, not ivory whitish; knobs of halteres entirely very dark
blackish brown or black above; ferruginous red on sides of thorax continuous and con-
fluent with humeral spot; sternopleural part mainly ferruginous and only extreme sternal
part of hypopleural part ferruginous; hairs in grooves above wing-bases dense, con-
spicuous and black like those on anterior declivous part of humerus and on face; fine
hairs on hind coxae and hair-like scaling on entire femora black.
3S 2 sheppard n. sp. (S. Rhodesia)
Wings dusky, darkly tinged smoky brownish or reddish brownish, darker in basal and
costal parts, more so in 29; veins darker, brownish or blackish brown; eyes in actual con-
tact in both sexes for a long distance; antennae darker, segment 1 dark reddish brown
to almost black, segment 2 black and usually longer; scutellum entirely or predominantly
black, and scutellar callosities yellow; hind legs with the tibiae darkened or blackened
apically and hind tarsi entirely very dark or black; callus on front femora larger, nearly
half length of femora; knobs of halteres very dark or blackish above; club of abdomen
shorter; smaller forms, usually less than 20 mm. J 2
Wings tinged predominantly yellowish, the costal sal basal apts praeally yellowish,
the posterior clearer part more greyish, being slightly more mauvish at apex; veins
mainly yellowish; eyes subcontiguous or very nearly separated in both sexes; antennal
segments 1 and 2 very pale yellowish red and 2 much shorter; scutellum predominantly
reddish and its callosities dark velvety brown; hind legs mainly yellowish red, even basal
half of tarsi reddish, only bases of femora and apical parts of tarsi dark; callus on front
femora much shorter, narrower, less than half length of femora; knobs of halteres paler,
more yellowish brown above; club of abdomen more elongate; larger form, about
20-22 mm. . : : ‘ 3 2 marshalli Bezz. (1938, p. 1026) (Rhodesia)
Wings more uniformly dusky or tinged smoky brownish, only a narrow costal part
slightly darker in both sexes; apical or third submarginal cell normally broadened apically,
vein separating it from second submarginal not markedly curved hindwards; antennal
segment 2 much longer, usually more than a third length of segment 3, sometimes at
least half in ¢¢; sides of thorax not reddened all round, but humeral angle or sometimes
a transverse fascia continuous with it, a continuous fascia from humerus including pro-
pleural tubercle and sometimes a transverse stripe in front of wing-bases ivory yellow;
greater part of pleurae, excepting sometimes only the reddish sutural part between
sterno- and hypopleuron, black; tergite 1 entirely black and rest also more darkened or
even black above; femora more darkened or dark-scaled above, and front coxae and
outer faces of front and middle tibiae and tarsi ivory yellowish, silver-haired; hind
404
(4)
(b)
ANNALS OF THE SOUTH AFRICAN MUSEUM
femora without spines below; thorax and scutellum above and mesopleuron usually
with finer sculpture; hairs on face pale and with the two longitudinal bands of darkish
hairs on thorax scarcely discernible. : ne
Wings with a more delimited, darker, more devik yeaa, ivthgentlia ¢ in aia half
up to level of upper vein of discoidal cell, including basal part of second basal cell,
more so in 9; apical or third submarginal cell characteristically narrowed apically, its
lower vein characteristically curved hindwards at about middle; antennal segment 2
much shorter, only about a third or less length of 3; sides of thorax and across base,
propleural tubercle, pteropleuron, middle parts of pleurae and entire hypopleural part
and even upper and lower parts of metasternum reddish; base of tergite 1 also broadly
reddish and rest of tergites scarcely or not darkened above; legs on the whole paler
reddish, front coxae dark, and outer sides of front and middle tibiae not ivory yellowish,
nor strikingly silver-haired; hind femora with some spines below at about middle;
thorax, scutellum and mesopleuron more coarsely sculptured; hairs on face black and
with more distinct dark ones on thorax above.
3 Y sanguineus Bezz. (1938, p. 1021) (S.W. Cape)
A lobe-like outer squama (part of alula) absent; knobs of halteres much darker above or
even blackish; thorax (excl. scutellum) longer, subequal to width of head, duller above,
with very much finer rugulose sculpture; metasternal part normally broader, more
coarsely sculptured and grooved, not sloping or markedly shifted forwards, hind coxae
not below level of neck; transverse ivory yellowish fascia on humeral part and propleural
part on each side extending transversely across front part of thorax for some distance
towards middle; a variable oblique transverse ivory yellowish fascia or mark present in
front of wing-bases or at least a yellowish spot on false tegulae; middle pleural part
between sterno- and hypopleural parts as well as extreme lower part of hypopleuron
reddish; silvery vestiture on metasternal part distinctly denser, though shorter, and fine
pale hairs across hind margins of tergites 2-5 not in form of conspicuous silvery rings;
larger form, about 15°5-18 mm.
: 3 2 leptogaster Lw. (1938, p. 1024) (Natal, E. Transvaal, Portuguese E. Africa)
A distinct delicate, transparent, lobe-like outer squama (part of alula) present; knobs
of halteres much paler, more yellowish brownish above; thorax (excl. scutellum) much
shorter, distinctly much shorter than width of head, more shiny above, distinctly more
coarsely rugulosely sculptured; metasternal part relatively much narrower, much finer
transversely rugose, distinctly more sloping or more markedly shifted forwards, the hind
coxae being at level of neck; transverse ivory yellowish fascia on humeral angle and
propleural part not continued dorsally for some distance across anterior part of thorax;
no spot on fascia present above wing-bases; pleurae black and middle pleural part velvety
black; silvery vestiture on metasternal part, though less dense, distinctly longer, and fine
hairs across hind margins of tergites 2-5 in more conspicuous silvery rings; smaller and
more slender form, about 11°5 mm. ; . 6 gracilis n. sp. (Portuguese E. Africa)
SUMMARY
Descriptions are given of three new species of Systropus (Systropinae): S.
bicoloripennis, S. sheppard: and S. gracilis. A further description of S. namaquensis
is given, including the previously undescribed female. A revised and descriptive
key to all the known southern African species of Systropus is included.
for
ACKNOWLEDGEMENT
The Trustees of the South African Museum are grateful to the Council
* Scientific and Industrial Research of South Africa for the award of a
grant towards the cost of publishing this paper.
2)
REVISION OF THE BOMBYLIIDAE (DIPTERA) OF SOUTHERN AFRICA 405
REFERENCES
Bezz1, M. 1921. On the Bombyliid Fauna of South Africa (Diptera) as represented in the
South African Museum. Ann. S. Afr. Mus. 18, 102-5.
Bezzi, M. 1924. The Bombyliidae of the Ethiopian Region, 116-28. London: Brit. Mus. (Nat.
Hist.).
Hesse, A. J. 1938. A revision of the Bombyliidae (Diptera) of Southern Africa. Ann. S. Afr. Mus.
34, 991-1028.
Hesse, A. J. 1958. Bombyliidae (Diptera Brachycera). Explor. Parc nat. de ’ Upemba, Miss. de Witte,
50 (7), 69-78.
Loew, H. 1860. Die Dipteren-Fauna Siidafrika’s 1, 173-245. Berlin: Bosselmann.
WIEDEMANN, C. R. W. 1820. Nova Dipterorum genera, 18. Kiliae: Mohr.
WIEDEMANN, C. R. W. 1828. Aussereuropaische zweifliigelige Insekten 1, 359-360. Hamm: Schulz.
‘, i i
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[ CONTINUED FROM INSIDE FRONT COVER
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Bettrdége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Hartman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sct. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
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by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name... .
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should be followed by references to all subsequent citations of the same name,
arranged in chronological order. .. .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalis capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 250.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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se «6ANNALS
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POUT APRICAN MUSEUM
VOLUME XLVI
PART XVII
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT,
SOUTH AFRICA
By
K. H. Barnarp
South African Museum, Cape Town
psy
ee
ah OM
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DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH
AFRICA
By
K. H. BARNARD
South African Museum, Cape Town
[Accepted July 1962]
(With 11 figures in the text)
CONTENTS
PAGE
Introduction : : : 407
List of species : : . 408
Species found at each station AS
Descriptions and notes ; 5 AILS)
Summary . ‘ : : SN 451
Acknowledgments : 3 nee sy
References : ; ; S451
INTRODUCTION
The material described here was obtained by Dr. F. H. Talbot of the
South African Museum, on board the r.s. Africana IJ of the Division of Sea
Fisheries in August and December 1959, at twelve stations off Cape Point
and west of the Cape Peninsula. It comprises approximately 590 specimens, of
which more than half are Prosobranch Gastropods. The number of species
- in the groups is as follows: Gastropods 43; Heteropods 2; Pteropods 5; Tecti-
branchs 2; Nudibranch 1; Solenogastres 2-3; Scaphopods 6; Cephalopods 3-4;
Lamellibranchs 11; Brachipod 1. Total: about 78.
One Cephalopod is here recorded. Two or three species of Octopods were
also obtained, but these have not yet been identified.
An attempt has been made to identify the Solenogastres, but owing to
technical difficulties in section-cutting, the attempt has been temporarily
postponed. There appear to be two, possibly three, species.
The material contains several species obtained many years ago by the
Cape Government trawler s.s. Pieter Faure, under the direction of the late Dr.
J. D. F. Gilchrist. The Africana II, however, has sampled depths greater than
those worked by the Pieter Faure. Therefore, as might have been expected,
several new records and new species have been obtained.
Most of the new forms belong to the family Turritidae (Gastropoda) ; the
Pieter Faure also collected more representatives of this family than of any other;
and similar results have been obtained in other regions by other expeditions.
407
Ann. S. Afr. Mus. 46 (17), 1963: 407-452, 11 figs.
408 ANNALS OF THE SOUTH AFRICAN MUSEUM
The most interesting discovery has been the Volute Guwivillea alabastrina
(Watson), originally taken by the Challenger between Marion Island and the
Crozets; and later by the Scotza near the South Orkneys.
Some of the species have already been recorded in Part III of the author’s
‘Contributions to. . . fauna of South African Marine Mollusca’, and in the
Journal of Conchology (see list of titles).
The whole collection made by Dr. Talbot and submitted to me for report
is now deposited in the South African Museum and is catalogued with South
African Museum registration numbers.
The Lamont Geological Observatory research vessel Vema obtained abyssal
molluscs from stations off the west coast of South Africa and south-west of
Cape Town (St. 14, 16, 18, 51, 52, 53), which have been reported on by Clarke
(1961). This author seems to have been misinformed as to the true position of
the Agulhas Basin, and has erroneously localized Stations 51 and 52 (1,000
and 800 miles respectively south-west of Cape Town) in the ‘Agulhas’ Basin.
The naming of two new species from Station 51 as ‘agulhasae’ is also very
misleading.
LisT OF SPECIES
GASTROPODS
Station
No.
Terebra sp. oe ye - .. A322 1 dead; more material wanted
Surcula scalaria Brnrd. 1958 A 7) Atego. dead
A1go 2 dead
Atg2 I living
A193 1 living, 1 dead (fresh)
Clavatula lobatopsis n. sp. St We me ; ce |
A3i7 11 living
A318 2 living
A322 6 living, 3 dead
Moniliopsis psilarosis n. sp. AST Oa eae
A317 2 living
i 93 2 living
Argi 2 living
A319 1 dead
A322 3 dead
Typhlomangelia(?) polythele n. sp.
A315 1 dead (ex anemone)
A318 1 living, 2 dead (ex anemone)
Allg! 5 living
A192 3 dead
A315 1 dead
A317 5 dead
A322 1 living, 1 juv. dead
Cythara(?) glaucocreas n. sp.
Ciythara(?) dagama n. sp.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 409
Typhlosyrinx pyrropelex n. sp.
Typhlosyrinx chrysopelex n. sp.
Typhlosyrinx subrosea n. sp.
Philbertia cala (Watson) . .
Mangilia sp.
Daphnella(?) verecunda n. sp.
Daphnella(?) bitrudis n. sp.
Gymnobela sp.
? Gen. ( Turritidae)
Cancellaria euthymer Brnrd. 1960..
Admete decapensis Brnrd. 1960
Guivillea alabastrina (Watson)
? Gen. (?Fasciolariidae) ..
Charitodoron pasithea Tomlin
Charitodoron thalia ‘Tomlin
Nux alabaster Brnrd. 1960
Neptunea bonae-spei n. sp.
Prosipho torquatus n. sp. ..
A193 1 living
..< A318 2 living, 2 dead
Agi 1 living, 1 dead
A317 6 dead
A318 2 dead
A319 1 dead
fe 3 living
A322 1 living
A318 1 living, 1 dead
A316 3 living
A189 1 dead
A315 1 dead
Ao. 2 dead
A18g 2 living, 1 dead
A193 2 dead
f A189 1 dead; more material wanted
VAST 1 dead
a A317 1 dead; more material wanted
"Agig 2 dead
A322 1 living
A3I5 1 living
A315 1 dead (apex only)
asec I protoconch
A317 1 fragment (columella)
Agig 1 dead (half grown)
A322 1 dead (half grown) and frag-
L ments
A315 2 dead (ex anemone); more
material wanted
f A189 2 living, 1 dead
"| Aga2 1 dead
Aigo 1 living
[Aso 1 living
Arg2 2 living
A1g3 2 living, 1 dead
A318 1 living
ae 5 dead
Argo 1 living
A318 2 living
A322 1 living
A315 1 living
A317 1 dead
A322 1 dead
410 ANNALS OF THE SOUTH AFRICAN MUSEUM
Pyrene cf. profundi Dall
Trophon acceptans Brnrd. 1959
Trophon cf. droueti Dautzenberg
Columbarium rotundum Brnrd. 1959
Columbarium angulare Brnrd. 1959
Thallassocyon bonus Brnrd. 1960. .
Odcorys watsont Locard
Polynices cleistopsila Brnrd. 1963
Falsilunatia pseudopsila Brnrd. 1963
Turbonilla sp. (cf. krausst) A
Cerithiella taylori Brnrd. 1963
Lamellaria capensis Bergh
Scala bonae-sper Brnrd. 1963
Abyssochrysos melanioides Tomlin
Calliotropis metallica (W.-M. & A.)
eee 1 dead
A322 8 living, 10 dead
A189 4 living, 7 dead
Atgi 9 living, 2 dead
Atg2 2 living
A193 8 living, 1 dead
A315 4 living, 1 dead
A316 2 living, 2 dead
A317 10 living
A319 2 living
A322 3 dead
A322 3 dead
A189 6 living, 16 dead
A318 2 living, 4 dead
( Argo 1 living
A193 1 living
A315 1 dead
A317 1 living
Atgo 2 dead
Atgo2 2 living, 1 dead
A1g3 I and 2 juv. living
A316 1 dead (large)
A317 2 dead
A318 1 living, 2 dead
A31Qg 1 living, 18 dead
ta 2 living, 3 dead
A18g 1 living
Aigo 1 dead
Agi 3 living
A1g2 2 dead
A317 1 living, 2 dead
A319 6 dead
A322 3 dead
A315 1 living
A18g 1 dead; more material wanted
Argo 1 dead
A316 2 living
“ie 93 2 living
A316 1 living
Aigo 1 dead
A319 2 living, 5 dead
Argo 1 living, 1 dead
A322 1 living
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA All
Calliotropis pompe n. sp. ..
Basilissa gelida n. sp.
Calliostoma glaucophaos n. sp.
? Solariella
HETEROPODS
Ailanta sp. $2 ae
Cardiapoda richardi Vayss
PTEROPODS
Cavolinia tridentata (Forskal)
Cavolinia limbata D’Orb...
Cavolinia ? globulosa
Diacria trispinosa (Lesueur)
FHlerse (Cuvieria) columnella (Rang)
TECTIBRANCHS
_ Scaphander puncto-striatis Mighels
Gastropteron sp.
NUDIBRANCHS
Doridoxa benthalis n. sp.
SOLENOGASTRES
In course of study
SCAPHOPODS
Dentalium capense Tomlin
ue 16 1 dead
A317 1 living, 1 dead
Aigo 1 living
A318 2 living
A1go 1 dead; more material wanted
A193 1 dead
Aigo 1 living
Aigo 18 dead
A193 14 dead
A315 4 dead
A317 1 dead
A318 11 dead
A319 24 dead
A322 3 dead
Aigo 1 dead
A315 2 dead
[Ase 2 dead
ae 1 dead
A322 3 dead
A322 2 dead
( A1g2 1 juv. living
| Agis 9 living
A317 1 living
A318 1 living
A3ig 3 dead
A322 1 juv. living
oe 1 living
A319 1 living
A316 1
AIgl
A193
he (large)
A316 (small)
A18Q 1 living, 2 dead
412 ANNALS OF THE SOUTH AFRICAN MUSEUM
Dentalium eualdes n. sp. ..
Denialium lardum n. sp. ..
Denialium sp. (9 ribs)
Dentalium sp. (18-22 ribs)
Cadulus promontortt Brnrd. MS. ..
CEPHALOPODS
Eggs
Octopus sp. 3
Octopus sp. juv.
Octopus sp.
Octopus sp. large ..
Octopus sp. small
Leachia cyclura Lesueur ..
LAMELLIBRANCHS
Nucula (Pronucula) benguelana Clarke
Malletia estheriopsis n. sp.
Leda parsimonia n. sp.
Leda macella n. sp.
? Sarepta sp.
Argo 1 living
Atog1 8 living
Atg2 1 and 1 juv. living
A1o3 14 living, 3 dead
A315 1 adult living, 1 juv. dead
A317 18 living, 2 dead
A318 1 living, 1 dead
A3ig 12 living, 1 dead
A322 4 living, 1 juv. dead
A193 5 dead
A318 2 dead
A322 1 living, 1 juv. dead
A1go 2 dead; more material wanted
A315 1 dead
A322 2 dead
A189 17 dead; more material wanted
A189 3 living
Jf Argo
Neen
A189 I
A1g2 I
A318 1
A319 I
EPTUG) it
Alg2 I
A322 1 living
(A317 2 living
A319 3 living
; : Ag2i 1 living
| A322 2 living
A317 1 living
f Aigo t living
Atg2 1 living
A317 1 living
A319 a lot living
A322 1 valve
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 413
( Argo 3 living
| Argr 17 living
| Aig2 12 living
Limopsis cf. straminea Smith... aut a 1 ane
315 3 living
A317 6 living, 5 valves
A3ig 34 living, 4 valves
A322 2 living, 1 valve
Bs A189 1 valve
A18q 1 living, 2 dead
is aa 1 living
a A18qg 1 living
a A18g 1 living
A315 I valve, 1 broken valve
r ae 1 broken valve
Thyasira investigatoris (Smith)
Abra longicallus (Scacchi)
Halicardia flexuosa (V. & S.)
Cuspidaria sp. (cf. meridionalts)
Cuspidaria sp. (cf. maxima)
BRACHIOPODS
Terebratula sp... Sy Hh Te Ago) 2) livin:
SPECIES FOUND AT EACH STATION
33° 50’ S., 17° 21’ E., 600 fathoms. (Station number A189) *
Surcula scalaria Brnrd. 1958
Mangilia sp. me bs Sys ag .. new to fauna-list
Daphnella verecunda . ee ee eS pHs alive
Gymnobela sp. pa mY, 3. ee .. new to fauna-list
Charitodoron pasithea Tomli
Trophon acceptans Brnrd. 1959
Columbarium rotundum Brnrd. 1959
~ Polynices cleistopsila Brnrd. 1963 .. ae er aS 0
Turbonilla sp. ae Si Ai se .. more material wanted
Cavolinia tridentata (Forskal)
Dentalium capense Tomlin
Dentalium sp. (18-22 ribs) oe Sa .. new to fauna-list
Cadulus promontorti Brnrd. MS. .. ee 55) 10g Sons Aaeres
Octopus sp. 3
Halicardia flexuosa (V. & S.)
Cryptodon investigatoris Smith
Abra longicallus (Scacchi) . . Pe ae .. new to fauna-list
Cuspidaria sp. (cf. meridionalis) .. ee .. new to fauna-list
33° 26’ S., 16° 33’ E., 1,240-1,300 fathoms (A190)
Clavatula lobatopsis ae he axe ve patlespe
Charitodoron thalia ‘Tomlin
*Not to be confused with South African Museum registration numbers which have the
prefix A.... and four numerals. For this reason, in the descriptions and notes only the localities
are given, not the station number.
414 ANNALS OF THE SOUTH AFRICAN MUSEUM
Nux alabaster Brnrd. 1960 Ne ui 5.) Shea. Spas laine
Thalassocyon bonus Brnrd. 1960 .. ue ses MSA INL Se
Odcorys watsont Locard
Polynices cleistopsila Brnrd. 1963 .. ah Paty ESD
Cerithiella taylori Brnrd. 1963s yj a 1c DESDE
Abyssochrysos melanioides ‘Tomlin
Calliotropis metallica (W-M. & A.)
Basilissa gelida .. ae Ae we 0) Ml. Spas Iiyoe
? Solariella a be - be .. more material wanted
Cavolinia tridentata (Forskal)
Cavolinia limbata D’Orb.
Diacria trispinosa (Lesueur)
Cardiapoda richardi Vayss. oe ae .. new to fauna-list
Dentalium eualdes .. we He se Dito) sey SIs
Dentalium sp. (9 ribs) i cH Si ... new to fauna-list
Cephalopod eggs
Limopsis sp. cf. straminea Smith .. EY .. new to fauna-list
Leda macella ae eT 01S) oF
33° 36'S., 16° 15’ E., 1,520-1,570 fathoms (A191)
Clavatula lobatopsis be ae me n. sp.; [Types
Cythara(?) dagama we a ie 2 4'ne sp.) Eyes
Typhlosyrinx pyrropelex ... Us we: .. n.sp.; Types (of juvenile)
Charitodoron thalia Tomlin
Trophon acceptans Brnrd. 1959
Polynices cleistopsila Brnrd. 1963 .. ne Soe WOM SOK
Dentalium eualdes .. i iS a fo LM. SP. bypes
Solenogastres oe ss at As .. new to fauna-list
Limopsis sp. cf. straminea Smith .. .. new to fauna-list
33° 454’ S., 16° 2334’ E., 1,480 fathoms (A192)
Clavatula lobatopsis Ane ae ie Gee ten OSES 0}.
Cythara(?) dagama ae Be ne oe eT SDs
Charitodoron thalia ‘Tomlin
Trophon acceptans Brnrd. 1959
Odcorys watsont Locard
Polynices cleistopsila Brnrd. 1963 .. Me it 2 le |S:
Scaphander puncto-striatus Mighels !
Dentalium eualdes .. Be ie Bi ae ORSON
Octopus sp. juv.
Leachia cyclura Lesueur .. ue oe .. new to fauna-list
Limopsis sp. cf. straminea Smith .. ve .. new to fauna-list
Leda macella a ane is sts Wei IY SO.
33° 49’ S., 16° 30’ E., 1,500 fathoms (A193)
Clavatula lobatopsis ae ae Le Cie Molt SE
Cythara(?) glaucocreas a Me ee ate STD.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA
Daphnella(?) bitrudis
Charitodoron thalia Tomlin
Neptunea bonae-spet. .
Pyrene cf. profundi Dall
Trophon acceptans Brnrd. 1959
Thalassocyon bonus Brnrd. 1960
Oécorys watson Locard
Scala bonae-sper
Atlanta sp.
Cavolinia tridentata (Forskal)
Solenogastres
Dentalium eualdes
Dentalium lardum
Peps sp. cf. straminea Smith
R437 9., 17 03 E., 1,580—1,620 Gathers (Agus
Clavatula ee
Cythara(?) glaucocreas
Cythara(?) dagama
Mangilia sp.
? Fasciolarudae ? gen.
Admete decapensis Brnrd. 1960
Guivillea alabastrina Sak
Prostpho torquatus
Trophon acceptans Brnrd. 1959
Thalassocyon bonus Brnrd. 1960
Falsilunatia pseudopsila
Cavolinia tridentata (Forskal)
Cavolina ? globulosa
Scaphander puncto-striatus Mighels
Dentalium eualdes
Dentalium sp. (9 ribs)
Limopsis sp. cf. straminea Smith
Cuspidaria sp. cf. maxima ..
[Fragment of Argonauta|
34° 42’ S., 16° 54’ E., 1,725-1,780 fathoms (A316)
Clavatula lobatopsis
Moniliopsis psilarosis
Philbertia cala (Watson)
Guivillea alabastrina (Watson)
Trophon acceptans Brnrd. 1959
Odcorys watson Locard
Lamellaria capensis Bergh
Scala bonae-sper
Calliotropis pompe
n. sp.; Types
ie Spt
new to fauna-list
n. sp.
TM. SP-5 2 lunes
new to fauna-list
n. sp.
NSp.; Ly pes
new to fauna-list
Tes Sz
n. sp.
n.sp.
new to fauna-list
new to fauna-list
n. sp.;, Lype
new to fauna-list
NaspsaLype
n. sp.
lM. SPsjel ype
ne sp:
new to fauna-list
new to fauna-list
new to fauna-list
n. sp.
n. sp.; Types
new to fauna-list
new to fauna-list
n. sp.
n. sp.
A15
A416 ANNALS OF THE SOUTH AFRICAN MUSEUM
Doridoxa benthalis ..
Solenogastres
Brachiopod
33° 50’ S., 16° 30’ E., 1,480—1,660 fathoms (A317)
Clavatula lobatopsis
Typhlomangelia polythele
Cythara(?) dagama
Typhlosyrinx pyrropelex
Mangilia sp.
Gymnobela sp.
Turritid. Gen. ?
Guivillea alabastrina (Watson
Prosipho torquatus
Trophon acceptans Brnrd. 1959
Thalassocyon bonus Brnrd. 1960
Odécorys watsont Locard
Polynices cleistopsila Brnrd. 1963 ..
Calliotropis pompe a
Cavolinia tridentata (Forskal)
Scaphander puncto-striatus Mighels
Dentalium eualdes
Limopsis sp. cf. straminea Smith
Leda macella
Leda parsimonia
Malletia estheriopsis
[Fragment of Fanthina]
n. sp.; Type
new to fauna-list
. Spe
. sp.; Types
sp.
Sp.
new to fauna-list
new to fauna-list
new to fauna-list
Sie 6
Ne Sp:
n. sp.
Ne Sp:
n. sp.; lypes
DE S]OS
new to fauna-list
in, SO
Spa slay
n. sp.; Types
33° 52’ S., 16° 51’ E., 1,380-1,520 fathoms oe
Clavatula lobatopsis
Cythara(?) glaucocreas
Typhlosyrinx pyrropelex
Typhlosyrinx subrosea
Charitodoron thalia ‘Tomlin
Nux alabaster Brnrd. 1960
Neptunea bonae-spet. .
Columbarium angulare Bawidl 1959
Odcorys watsont Locard
Calliostoma glaucophaos
Cavolinia tridentata (Forskal)
Scaphander puncto-striatus Mighels
Gastropteron sp.
Dentalium eualdes
Dentalium lardum
Cephalopod eggs
Octopus sp.
i, SDs
n. sp.; Type
Me spe
n. sp.; Types
ile BJO
n. sp.; Types
n. sp.; Types
new to fauna-list
Tey Spe
Ne SO
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA
34° 05’ S., 16° 58’ E., 1,470-1,490 fathoms (A319)
Clavatula lobatopsis Te SOF
Typhlosyrinx pyrropelex .... Ae a Ee easoe
turritid. (Gen. ? 1.2 us ay .. new to fauna-list
Guivillea alabastrina (Watson)
Trophon acceptans Brnrd. 1959
Odcorys watson Locard
Polynices cleistopsila Brnrd. 1963 .. a ee so tS) 0)
Abyssochrysos melanioides ‘Tomlin
Cavolinia tridentata (¥orskal)
Scaphander puncto-striatus Mighels
Gastropteron sp... Ede Ns oe .. new to fauna-list
Dentalium eualdes
Octopus sp. (large)
Octopus sp. (small)
Limopsis sp. cf. straminea Smith .. es .. new to fauna-list
Leda macella we aN we a 5 Wy SOR Iqores
Malletia estheriopsis oy. es n. sp.
Bameeor ton Or 42) Ho 1.770-1,600 fathoms (Agar)
Malletia estheriopsis : Ae ae So: 1) Toy SOE
34° 36’ S., 17° 00’ E., 1,500-1,760 fathoms (A322)
Terebra sp. . new to fauna-list
Clavatula lobatopsis n. Sp.
Moniliopsis psilarosis n. sp.
Cythara(?) dagama ily G0.
Typhlosyrinx pyrropelex Ms SO
Typhlosyrinx chrysopelex se ily S08. JE\qore
Cancellaria euthymet Brnrd. 1960 .. ial, Sous Ivor
~ Guwwillea alabastrina (Watson)... as .. new to fauna-list
Charitodoron pasithea Tomlin
Charitodoron thalia Tomlin
Neptunea bonae-spet. . ins bu ee £5 10s BOF
Prosipho torquatus .. ae cr aM so 1 Oe SOs
Pyrene cf. profund: Dall... 1g Le .. new to fauna-list
Trophon acceptans Brnrd. 1959
Trophon cf. droueti Dautzenberg .. o .. new to fauna-list
Odcorys watsoni Locard
Calhotropis metallica (W.-M. & A.)
Cavolinia tridentata (Forskal)
Diacria trispinosa (Lesueur)
Herse (Cuvierta) columnella (Rang)
Scaphander puncto-striatus Mighels
Dentalium eualdes .. Re . i 3.) ERS s
Dentalium lardum .. ui ac ae se fa easy OF
417
418 ANNALS OF THE SOUTH AFRICAN MUSEUM
Dentalium sp. (9 ribs) i ce ve .. new to fauna-list
Abra longicallus (Scacchi) sb re .. new to fauna-list
Limopsis sp. cf. straminea Smith .. Bie .. new to fauna-list
Cuspidaria sp. cf. maxima .. os .. new to fauna-list
Nucula (Pronucula) benguelana Glarke in .. new to fauna-list
? Sarepta sp. Ne Ae Ae ay. .. new to fauna-list
Malletia estheriopsis De es i JS ne Spe
DESCRIPTIONS AND NOTES
GASTROPODA
Terebridae
Terebra sp.
(Fig. 1 a)
Protoconch 2 (24) whorls, last whorl bulbous, smooth. Postnatal whorls 7.
Axial plicae 14-15 on 2nd whorl, increasing to 18 on last whorl, straight, slightly
protractive; intervening grooves shallow. Slight indication of a subsutural
spiral groove, and 4—5 very faint spiral lirae in the axial grooves. Base with
growthlines and very faint spiral striae. 6-5 x 2 mm. Very pale corneous,
protoconch opaque white.
34° 36’ S., 17° 00’ E., 1,500—-1,760 fathoms, 1 dead (S. Afr. Mus. Ag854,
F. H. Yalbot coll.).
Remarks. As further and better specimens may be obtained later, no specific
name is attached to this specimen.
Turritidae
Subfam. Turrinae
[Turris lobata]
In 1958 I united Pieter Faure specimens from Cape Point with specimens
from Cape Natal (Durban)-East London under the specific name lobata Sow.,
and transferred it to the genus Turris. The new material raises doubts as to the
conspecificity of the shells, and the generic position.
Comparison of the 7 additional specimens with the previous material
shows that the Cape Point shells can be separated on conchological characters
from those of the Cape Natal-East London area, though the differences are
subtle (slightly exaggerated in the figures herewith).
From the Pieter Faure material two radulae were obtained, one from an
East London shell and one from a Cape Point shell, both of them extracted
from poorly preserved animals. They seemed to show a slight difference in the
shape of the lateral plate. On re-examination, and comparison with 4 radulae
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 419
from the new material, the difference appears to be due merely to the slightly
different position in which the plates are lying in the mounted preparation.
A more important point, however, is that the 4 additional radulae show
distinctly the presence of a central plate and of an accessory wing-like appen-
dage, albeit both are very delicate.
S
Ss
.
&
Fic. 1. a, Terebra sp. Apex and base, with
cross-section of whorl. 6, Daphnella (?)
bitrudis n. sp.
These Cape Point shells must therefore be placed in Clavatula. Possibly
when more material is obtained from the Natal-East London area, it will
show that lobata has been incorrectly transferred to Turris; but for the present
I retain it in Turris.
Turris lobata (Sow.)
(Fig. 2 5)
Turris lobata (Sow.), [partim] Barnard, 1958, p. 107, figs. 3 7, 6 profile.
To the description should be added: upper margin of whorl straight,
suture visible; the sharp keel continuous, without any trace of nodules; the
mid-whorl nodules always rounded, though they may be divided by a slight
sulcus.
In my description the number of midwhorl tubercles was not given.
Sowerby’s original description gave 13 on the penultimate whorl, and his
figure seems to confirm this. There is, however, a possibility that °13’ was a
misprint for 18, because a lobate specimen, labelled by Sowerby, has 13 on the
3rd whorl, 18 on the gth and 22 on the roth; other specimens agree, none having
less than 16 tubercles on the 8th whorl.
Natal and East London area, 440 and 310 fathoms (S. Afr. Mus. A1673,
ATO7A, PE. coll.)
420 ANNALS OF THE SOUTH AFRICAN MUSEUM
The remarks in the above reference on the formation of the lobe on the
outer lip apply to lobata (Sow.).
There is one dead shell from the Cape Point area, 380-475 fathoms (S. Afr.
Mus. A1675, P.F. coll.), however, which seems referable to lobata. Although
damaged several times and repaired by the animal, and corroded, nevertheless
it shows the diagnostic features of lobata, not those of the other Cape Point
shells from much greater depths.
Clavatula lobatopsis n. sp.
(Fig. 2 a)
Turris lobata (Sow.), [ partim] Barnard, 1958, p. 107, fig. 3).
Extremely like Turris lobata but upper margin of whorl undulate, and
slightly raised so that the actual suture is scarcely or only partly visible in
lateral view; instead of the sharp keel in lobata there is a blunt lira with small
nodules, corresponding in number with those in the mid-whorl series, often
divided by a slight sulcus; between this lira and the mid-whorl nodules there
may be 2—3 feeble lirae, or none at all; the mid-whorl nodules are sharper than
in lobata, more tubercular than nodular, and they may be divided by a faint
sulcus; on the last whorl (or last half thereof) in the larger shells the tubercles
Fic. 2. a, Clavatula lobatopsis n. sp. Sculpture for comparison with b. b, Turris lobaia (Sow.). Sculpture.
c, Typhlosyrinx pyrropelex n. sp. Protoconch and base; radula tooth. d, Cythara (?) glaucocreas n. sp.
e, Daphnella (?) verecunda n. sp. With radula tooth. Apex of latter further enlarged. f, Mangilia (?) sp.
g, Cythara (?) dagama n. sp. h, two radula teeth.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 421
tend to disappear, leaving only a continuous well-marked lira (or costa) at
the lip sinus.
Lirae below the mid-whorl girdle as in lobata, but with no tendency to
become costate and form a lobe on the outer lip; the lira next below the mid-
whorl girdle may have small nodules.
Number of midwhorl tubercles 13-14 on 3rd whorl, increasing to 21-24
on gth whorl. Towards the end of the last whorl the tubercles often tend to be
feeble and irregular.
Among the specimens from Station A317 (S. Afr. Mus. Ag800) was one
with 12 midwhorl tubercles on the 3rd whorl, increasing to 14 on the last (8th)
whorl. This can be regarded only as a casual variation.
Up to 40 X 14 mm. and 39 X 16 mm. (apices corroded).
Operculum oval, nucleus apical.
Animal pale. Eyes absent, or sometimes indicated by a minute brown or
black speck. Radula with c. 70 rows, central plate narrow, acicular, extremely
delicate, lateral plate with accessory appendage.
Cape Point NE. x E.4E. 46 miles, 900 fathoms, 1 living; N. 70° E. 40
miles, 800 fathoms, 2 dead; NE. x E.{E. 40 miles, 800—g00 fathoms, 1 living,
3 dead (S. Afr. Mus. A1676—A1681, P.F. coll.).
Come Ouno a 10n 3oe be ai soOOmathomsy) 2adeads. 39° 96, S., 16: 15 EA,
1,520-1,570 fathoms, 3 living (Types; 33° 454’ S., 16° 234’ E., 1,480 fathoms,
macad: 93-49 S., 160° 30 E., 1,500 fathoms, 1 living, 1 dead (fresh) ; 34°37 S.,
17° 03’ E., 1,580-1,620 fathoms, 2 living; 33° 50’ S., 16° 30’ E., 1,480-1,660
fathoms, 11 living; 33° 52’ S., 16° 51 E., 1,380-1,520 fathoms; 2 living;
PaO os On 56) Ee. 1.470 -1,400) fathoms, 1 dead; 94° 26° S., 17° 00 EL,
1,500 fathoms, 6 living, 3 dead (S. Afr. Mus. Ag712, Ag730 (Types), Ag740,
Ag752, Ag771, Ag800, Ag820, Ag838, Ag855; F. H. Talbot coll.).
Surcula scalaria Brnrd.
_ Surcula scalaria, Barnard, 1958, p. 146, fig. 22 d.
Like the Preter Faure specimens, the present shells are dead; the generic
position therefore remains uncertain.
BoMnTOu os, 721 ak. OOO) fathoms, 9 dead (S:. Atry Mus. Ag6o5, F. Ho.
Talbot coll.).
Subfam. Brachytominae
Moniliopsis psilarosis n. sp.
(Fig. 4 a)
Protoconch and ? 2 whorls missing. Remaining postnatal whorls 6. First
3 whorls (probably the 3rd—5th) distinctly but not strongly shouldered, profile
of following whorls evenly convex. Oblique, protractive axial riblets 17-18
on first 2 whorls, 18-19 on 3rd whorl, forming small knobs at the shoulder,
petering out below and scarcely reaching suture; becoming evanescent and
obsolete on following whorls; crossed by impressed spiral striae 4 on first 2
422 ANNALS OF THE SOUTH AFRICAN MUSEUM
whorls, 4—5 on 3rd, increasing to 8-9 on 4th, and c. 13 on last 2 whorls; some-
times 2-3 fine striae above the shoulder on the sulcus. Base with ¢c. 24 (main
and interpolated) spiral striae. Sulcus feebly concave, lip sinus moderately
deep. Canal rather short and narrow. 47 X 16 mm. Operculum narrow oval,
13 X 5mm. Drab or brownish, columella and interior of aperture dull pinkish;
operculum amber.
Animal pale; eyes at base of short tentacles. Radula with 15 pairs of rather
elongate, unbarbed teeth.
34° 42’ S., 16° 54’ E., 1,725-1,780 fathoms, 1 living, 1 dead (fresh) ;
34° 36’ S., 17° 00’ E., 1,500-1,760 fathoms, 3 dead (worn and corroded)
(S. Afr. Mus. Ag789 (Types) and Ag856; F. H. Talbot coll.). .
Remarks. Seems to fit best into the genus Monzliopsis. The sculpture on the
later whorls resembles a bare ploughed field.
The smallest worn specimen has lost the protoconch, but retains the first
2 postnatal whorls (corroded); the full complement of postnatal whorls would
appear to be 8. The two largest specimens, corroded and comprising 4th—8th
whorls, measure 51:5 X 19 mm. I have seen a larger one, comprising 3th—8th
whorls, measuring 56 X 20 mm. (in coll. Fisheries Survey).
Typhlomangelia (?) polythele n. sp.
(Fig. 3 ¢, f)
Protoconch and ? 2 whorls corroded. Postnatal whorls 44; profile angularly
shouldered a little above middle of whorl. Small peripheral knobs on the
shoulder, c. 20 on 2nd whorl, c. 23 on gard, c. 26 on last whorl, evanescent
towards outer lip, not continued below shoulder (or only very slightly); low
flat spiral lirae 3-4 on Ist whorl, 4-5 on and, 5-6 on 3rd, 7-8 on last whorl;
8-9 additional lirae on base, plus about the same number of finer lirae on
rostrum. Sulcus scarcely concave, with a keel forming a distinct cingulum
below the suture. Growth-lines distinct, especially on sulcus where they are
subpliculose. Lip sinus deep, semicircular. 13 X 5:5 mm. Operculum oval,
nucleus apical. White, operculum amber.
Animal pale. No eyes. Radula with 22 pairs of dagger-like, unbarbed teeth.
33° 50’ S., 16° 30’ E., 1,480-1,660 fathoms, 2 living (S. Afr. Mus. Ag8oa2,
FF. Ho Lalbot coll):
Remarks. Placed provisionally in Typhlomangeia although the radula teeth
are not elongate as in nivalis (see Sars, 1878, pl. ix, fig. 10).
Subfam. Cytharinae
Cythara (?) glaucocreas n. sp.
(Fig. 2 d)
Protoconch corroded. Postnatal whorls 6, apical whorls more or less
corroded; profile of whorls moderately convex, shoulder distinct, base rather
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 423
ventricose. Oblique axial ribs on penultimate and ultimate whorls 26-30,
from shoulder to suture below, more or less traceable on base; spiral lirae 7-8
or g on sulcus; ribs crossed by 8-10 spiral lirae below shoulder, c. 20-24 on
base (main and intermediaries). Growth-lines distinct across sulcus, often
forming pliculae, nearly straight on upper half, curved when nearing the
shoulder. Columella curved, canal wide, very short. 25:5 X 11°5 mm. and
21 X 11 mm. No operculum. White.
Animal greenish, no eyes, Radula with 21 pairs of short dagger-like,
unbarbed teeth (similar to those of D. verecunda, see fig. 2e ).
Boe Ono selOresOneh 01 500 pfathoms,., 2. living; 94° 97) S.717° 03’ E.,
1,580—1,620 fathoms, 1 dead (extracted from an anemone) ; 33° 52’ S., 16° 51’
E., 1,380-1,520 fathoms, 1 living (Type) and 2 dead (S. Afr. Mus. Ag753,
Ag773, Ag821 (Type), Ag824; F. H. Talbot coll.).
Remarks. Belongs to one of the Cytharine genera and is provisionally placed
in Cythara.
Cythara (?) dagama n. sp.
(Fig. 2 g, h)
Protoconch corroded. Profile of whorls convex, shoulder not prominent
owing to corrosion, except in the smallest (6-whorled) shell. Postnatal whorls 8.
Oblique axial ribs 14 on 3rd whorl, 16 on 4th, 16-17 on 5th, 19 on 6th, 20-22
on 7th, but becoming obscure towards end of whorl, ribs on 8th whorl (only
one shell) uncountable owing to corrosion, from shoulder to suture below,
evanescent on base; 5-7 spiral lirae on sulcus (chiefly on lower part), obscure
on later whorls; ribs crossed by 6—7 lirae between shoulder and suture on 4th
and 5th whorls, 7-8 on 6th, 8-g (10) on 7th whorl (? ro-11 on 8th whorl,
corroded), 12-15 on base, lirae regular, without intermediaries except one or
two on base. Growth-lines forming a nearly even curve on the sulcus, slightly
- pliculose on earlier whorls. Columella curved, canal short, moderately wide.
Bom a HO-hy ama. 9495 mth; 390 < 19) mm.; 29 x 11° mm. White. No
operculum.
Animal pale. Tentacles short, no eyes. Radula with 20 pairs of dagger-like
teeth, proximally not divided, a short process on inner margin slightly nearer
to base than to apex, distally expanded with short lateral tangs, but not barbed.
33° 36’ S., 16° 15’ E., 1,520-1,570 fathoms, 5 living (Types); 33° 453’ S.,
16° 23%’ E., 1,480 fathoms, 3 dead; 34° 37’ S., 17° 03’ E., 1,580—1,620 fathoms,
WG@ecias 92 950) 9... 10, 90) |B 1,480-1,060 fathoms, 5 dead, corroded; 34°
36’ S., 17° 00’ E., 1,500-1,760 fathoms, 1 living, 1 juv. dead (S. Afr. Mus.
Ag731 (Types), Ag741, Ag772, Ag806, Ag860; F. H. Talbot coll.).
Typhlosyrinx pyrropelex n. sp.
(Fig. 2 c)
Pleurotoma (Surcula) dissimilis (non Watson). Barnard, 1958, p. 147, fig. 23 a (protoconch).
Shell smooth, polished. Protoconch 34—4 whorls, last 2 or 3 whorls with
regular oblique (protractive) pliculae, becoming slightly sigmoid near junction
A424 ANNALS OF THE SOUTH AFRICAN MUSEUM
with 1st postnatal whorl. Postnatal whorls 64 (7); profile convex, with slight
shoulder. Growth-lines strongly sigmoid, irregularly pliculose near the suture,
becoming strongly protractive (nearly horizontal) on the shoulder, and forming
in some specimens obseure rounded axial ribs below shoulder on 3rd and 4th
whorls, c. 14 on 4th whorl, best seen as marginal undulations in apical view.
Fine indistinct spiral striae below shoulder, c. 5-6 on 2nd whorl, 7-8 on 3rd,
8-9 on 4th, increasing to c. 25 on 7th whorl, on base 25-30 on 4th whorl, 50-66
on 7th. Juveniles: up to 22°5 x 9 mm. (4 whorls). Creamy-white, glossy,
protoconch fulvous brown. No operculum.
Animal pale; eyes represented by a minute pigment speck or absent.
Radula (juveniles) with 25-30 pairs of slender doubly-barbed teeth, with a
projecting knob proximally.
Cape Point N. 77° E. 650-700 fathoms, 2 dead; NE. x E.4E. 43 miles,
goo fathoms, 2 dead; NE. x E.2E. 38 miles, 750-800 fathoms, 1 dead (S. Afr.
Mus. (Types) A1643, A1644, A1645; P.F. coll.). :
33° 36’ S., 16° 15 E., 1,520-1,570 fathoms, 1 living, 1 deadgiagsijamae
16° 30’ E., 1,480—-1,660 fathoms, 6 dead; 33° 52’ S., 16° 51’ E., 1,380-1,520
fathoms, 2 dead; 34° 05’S., 16°58’ E., 1,470-1,490 fathoms, 1 dead; 34° 36’S.,
17° 00’ E., 1,500-1,760 fathoms, 3 living (S. Afr. Mus. Ag732, Ag805, Ag823,
Ag9839, Ag858; F. H. Talbot coll.).
Remarks. The radula corresponds with that of 7. vepallida von Martens
(see: Thiele, 1903, pl. 9, fig. 74; and 1929, fig. 450) and the species may be
provisionally included in Typhlosyrinx.
The shells obtained by Dr. Talbot are evidently juveniles of the same
species as was obtained farther to the south-east by the Preter Faure. In 1958
I was in two minds whether to refer the Cape shells to the Philippine dissimilis
or the Cape Verde alberti. I now consider that slight differences in shape are
unimportant, but that, on the other hand, the strong protractive bend in the
erowth-lines is sufficient to distinguish the Cape shells from both the other
Fic. 3. a, Typhlosyrinx subrosea n. sp. b, protoconch. c, radula as arranged in radula sac. d, radula
tooth, with apex further enlarged. e, Typhlomangelia polythele n. sp. f, radula tooth, with apex
further enlarged. g, Typhlosyrinx chrysopelex n. sp. h, radula tooth, with apex further enlarged.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 425
species. The protoconch was missing in disszmilis, and though present in the
living example of alberti, was not stated to be coloured.
Several species with coloured (brown or yellow) protoconchs on a white
shell have been described (see: Dautzenberg, 1927), mostly assigned to
‘Pleurotoma’.
The Pieter Faure shells are regarded as Types of the adult. One of the shells
of S. Afr. Mus. A1643 was sent to Tomlin, and presumably remains in his
collection. The specimens 8. Afr. Mus. Ag858, from the largest of which the
radula was extracted, may be regarded as Types of the juvenile and radula.
In the 1958 description (p. 147), for protoconch ‘lip’, read ‘tip’.
Typhlosyrinx chrysopelex n. sp.
(Fig. 3 g)
Protoconch 3 whorls, somewhat worn, whorls pliculose, cancellate on
lower half. Postnatal whorls 44, profile shouldered slightly above middle of
whorl. Slightly oblique axial ribs from shoulder to suture, petering out on base,
13 on Ist whorl, 15 on 2nd, 16 on grd, and 19 on last whorl. No spiral sculp-
turing, except 12-15 feeble lirae on rostrum. Sulcus slightly concave, lip sinus
shallow. Growth-lines distinct on sulcus, some of them pliculose below the
suture. 19 X 9 mm. No operculum. White, glossy, protoconch yellowish-
brown (faded).
Animal pale, eyes present. Radula with 25 pairs of dagger-like, barbed and
flanged teeth, base broad and concave.
34° 36’ S., 17° 00’ E., 1,500—-1,760 fathoms, 1 living (S. Afr. Mus. Ag857,
eye Walbot coll.).
Remarks. Also placed provisionally in Typhlosyrinx.
Typhlosyrinx subrosea n. sp.
(Fig. 3 ad)
Thin-shelled. Protoconch 34-4 whorls, with (except 1st) oblique (protrac-
tive) pliculae, crossed below the periphery by retractive pliculae, producing a
micro-clathrate sculpture. Postnatal whorls 44, profile evenly convex, the
sulcus not concave, scarcely distinguishable from rest of profile. No axial
sculpture except the growth-lines, which are sigmoid but not very concave
across the sulcus. Impressed spiral striae scarcely indicated on ist and 2nd
whorls, but becoming distinct near end of 2nd whorl, c. 16 on 2nd—3rd, ¢. 20
on 3rd—4th, c. 24 on 4th whorl, with 2-4 additional finer ones on the sulcus
above the ‘shoulder’. On base at least 36 striae, extending to end of rostrum.
Columella curved, canal rather short and narrow. In the larger living shell no
columellar callus concealing the spiral striae; in the smaller dead shell a weak
callus partly concealing the striae, especially on the rostrum. 36 x 16 mm.
No operculum. Very pale translucent pink, protoconch golden-brown.
4.26 ANNALS OF THE SOUTH AFRICAN MUSEUM
Animal pale, no eyes. Radula with 10 pairs of rather short, dagger-like
teeth, apically barbed and flanged. |
33° 52’ S., 16° 51’ E., 1,380—1,520 fathoms, 1 living, 1 dead (S. Afr. Mus.
Aog822, F. H. Talbot coll.).
Remarks. Somewhat similar to the shells described as pyrropelex, but the
sulcus not so distinct and the columella more curved.
Philbertia cala (Watson)
(Fig. 4 b-e)
Clathurella cala Watson, 1886, p. 361, pl. 26, fig. 11.
Protoconch 34 whorls, last 24 with fine oblique protractive pliculae, and
on the lower half of the whorl oblique retractive pliculae between the pro-
tractive ones, giving a faint cancellate or granulate sculpture. Postnatal
whorls 54, profile strongly convex, shoulder well marked but rounded (Watson:
‘hunchy’). Oblique protractive axial riblets 12 on 1st whorl, increasing to 18
Fic. 4. a, Moniliopsis psilaropsis n. sp. with radula tooth. 6, Philbertia cala (Watson). ¢, proto-
conch. d, two views of radula tooth. e, aperture with animal, as preserved, showing grooved
foot.
on last, on the early whorls traceable across the sulcus, but not on the last 2
(or 3) whorls, continued across base; crossed by fine spiral lirae 4 on 1st whorl,
5 on 2nd, 5-6 on 3rd, 8 on 4th and 10 on last whorl (6-7 main lirae plus
intermediaries). On base 6-7 main lirae plus intermediaries, but on rostrum
lirae subequal. Sulcus concave, lip sinus rather deep. 19 X 10 mm. No oper-
culum. White, protoconch chestnut-brown.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA NPI)
Animal pale, no eyes. Radula with 20-25 pairs of short, dagger-like,
unbarbed teeth.
34° 42’ S., 16° 54’ E., 1,725-1,780 fathoms, 3 living (S. Afr. Mus. Ag790,
EEL. Walbot coll.)
Distribution. South Atlantic: 32° 24’ 8., 13° 5’ W., 1,425 fathoms (Watson).
Remarks. These specimens agree so well with Watson’s description that the
identification seems certain. There are only two points to note: the largest of
the present specimens is 19 mm. long with 54 whorls, the Challenger shell only
0°55 inches with the same number of whorls; and the Cape shells have 2-3
ribs more than the Challenger shell. These differences seem insignificant as
against the essential similarities.
One of the present specimens with 4 postnatal whorls has all the lirae
on the 3rd and 4th whorls subequal, not divided into main and intermediaries.
In all three specimens the elongate grooved tongue-like foot has not been
withdrawn within the aperture. This reaction to the preservative seems to be
peculiar to this species, because it has not occurred in any of the other Turritid
specimens in the present collection, all of which were preserved in the same
manner. In the latter the foot is contracted into a compact mass and withdrawn
within the aperture.
Comparable (conchologically) species in deep water off the New England
coast seem to be Pleurotomella saffordi V. & S. and benedicti V. & S., 1884.
Mangilia (?) sp.
(Fig. 2 f)
Protoconch corroded. Profile of whorls convex, with a slight shoulder.
Postnatal whorls 5. Oblique axial ribs 14 on 3rd whorl, increasing to 16 on 5th
whorl, crossing the sulcus, and also base; c. 15 spiral lirae between shoulder
and suture on 3rd whorl, crossing the ribs, increasing to at least 20 on 5th whorl;
on base at least 30 lirae. Growth-lines forming an even curve on sulcus, those
which are continued as axial ribs stronger than the others. Lip sinus moderately
deep. Columella slightly curved, canal short, rather wide. 15 x 6:5 mm.
White.
33° 50’ S., 17. 21° E., 600 fathoms, 1 dead (S. Afr. Mus. Ag696, F. H.
Talbot coll.). |
Three specimens resemble the above described shell, but have 20 axial
ribs on the last whorl. |
gAe 9775.41 7 03" H., 1.500-1,020 fathoms, 1 dead; 33° 50’ S.,\16° 30 E.,
1,480—1,660 fathoms, 2 dead (S. Afr. Mus. Ag775, Ag803; F. H. Talbot coll.).
Daphnella (?) verecunda n. sp.
(Fig. 2 e)
Protoconch corroded. Profile of whorls angular. Postnatal whorls 7.
Oblique axial ribs 12-13 on 4th and 5th whorls, 14-15 on 6th and 7th, from
4.28 ANNALS OF THE SOUTH AFRICAN MUSEUM
shoulder to suture, evanescent on base; crossed by 6—7 spiral lirae on 4th and
5th whorls, 7-8 on 6th and 7th, c. 15 on base, including on the latter some
intermediaries; no spiral lirae on sulcus, or only extremely faint ones visible
in places. Growth-lines forming a nearly even curve on sulcus, without pliculae.
Columella curved, canal short, moderately wide. 22 x 10°5 mm. White,
middle portion of columella with faint salmon flush. No operculum.
Animal pale. Tentacles short, no eyes. Radula with c. 25 pairs of dagger-
like teeth, proximally bifid, enclosing the poison gland, a short process on
inner side proximally, apex sharply pointed, not barbed.
33° 50’ S., 17° 21’ E., 600 fathoms, 2 living, 1 dead (S. Afr. Mus. Ag697,
IF. H. Talbot coll.). | :
Remarks. Differs from Surcula sulcicancellata Brnrd. 1958 in having fewer
ribs and no sculpturing on the sulcus.
The radula teeth have some similarity with those figured by Thiele (1929,
fig. 456) for a species of Daphnella.
Daphnella (?) bitrudis n. sp.
(Fig. 1 5)
Very narrow fusiform. Point of protoconch broken, and apical whorls
corroded; 7 postnatal whorls remaining. Profile of whorls evenly convex, no
shoulder. Growth-lines for the most part distinct, somewhat variable but not
forming axial ribs, strongly protractive on sulcus before passing over on to
whorl. Fine spiral lirae 4 on 4th whorl, 6 on 5th, 8-9 on 6th and 10 on 7th,
on base c. 15 additional lirae but not well defined on rostrum. Columella
sinuous, canal long, narrow. 15 X 3:5 mm. White, glossy except where corroded.
39° 4g’ S., 16° 30’ E., 1,500 fathoms, 2 dead, but fresh (6) AicseMans:
Nome, 1, 1BL, Wealleoe Cally,
Remarks. May be compared with Mangilia scipio Dall (1889, p. 117, pl. 10,
fig. 12) from the West Indies, 124 and 982 fathoms; and Clathurella (Daphnella)
monoceros Watson (1886, p. 365, pl. 20, fig. 1) from off Sierra Leone, 2,500
fathoms.
The genus is quite provisional; perhaps the species might fit into Spergo,
but the suggestion is made without much confidence.
One of the most slender of the Turritids, being slightly more slender than
Pleurotoma torta Dautzenberg (1912, p. 11, pl. 1. figs. 3, 4).
Gymnobela sp.
Two dead specimens, 9°5 X 6:5 mm. and 13 X 8 mm., closely resemble
the figures of G. blakeana Dall 1889 and G. rhomboidea Thiele 1925, but the axial
ribs are obsolete while the growth-line pliculae across the sulcus are distinct.
Although in fair condition, it is preferable to wait for more material, with
the animal, before describing these shells.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 429
33° 50'S.; 17° 21’ E.,, 600 fathoms, 1 dead; 33° 50’ S., 16° 30’ E., 1,480—
1,660 fathoms, 1 dead (S. Afr. Mus. Ag698 and Ag804, F. H. Talbot coll.).
Gen. ?
Somewhat resembling Pleurotomella lotiae Verrill 1885, from the New
England coast, 1,525 fathoms, but narrower and less ventricose. Five to six
whorls, profile convex. Sulcus ?, not clearly marked. No axial sculpture.
Spiral lirae over greater part of whorl, c. 10, finer above and encroaching on
the ‘sulcus’. Growth-lines sigmoid, more or less pliculose below suture, especially
on early whorls. 11-5 X 7, 13 X 7°5 and 14 X 8 mm. White.
33° 50’ S., 16° 30’ E., 1,480—1,660 fathoms, 1 dead; 34° 05’ S., 16° 58’ E.,
1,470-1,490 fathoms, 2 dead (S. Afr. Mus. Ag807, Ag840, F. H. Talbot coll.).
Cancellariidae
Cancellaria euthymer Brnrd.
Cancellaria euthymei Barnard, 1960c, p. 438, fig. 1 0b.
34° 36’ S., 17° 00’ E., 1,500-1,760 fathoms, 1 living (S. Afr. Mus. Ag888,
F. H. Talbot coll.).
Admete decapensis Brnrd.
Admete decapensis Barnard, 1960c, p. 439, fig. 1 a.
34° 37’ S., 17° 03’ E., 1,580—1,620 fathoms, 1 living (S. Afr. Mus. Ag777,
F. H. Talbot coll.).
Since the above description was published, a second specimen has been
found among the Fisheries Survey collections. Presumably it is from the same
locality as the Type. It is of the same size as the Type. The columellar pleats
are slightly more prominent; and posterior to the upper one is a pair of small
narrow pleats close together.
Type and second specimen in the South African Museum.
Volutidae
Guivillea alabastrina (Watson)
(Fig. 5)
Wyvillea alabastrina Watson, 1882, p. 332.
Guivillea alabastrina (Watson), Watson, 1886, p. 262, pl. 15, fig. 2. Pelseneer, 1888, p. 3, pl. 1,
figs. 1,2 (animal). Melvill & Standen, 1907, p. 140. Barnard, 1960a, p. 398. South Afri-
can Museum Report, 1961, pl. 4, fig. C.
Four worn and broken specimens were obtained. An apex consisting of
protoconch plus 2 whorls; a portion of a very worn columella, identifiable by
comparison with the following specimen; a protoconch plus 24 whorls, length
77 mm.; and a protoconch plus 2? whorls, length go mm., together with
fragments.
430 ANNALS OF THE SOUTH AFRICAN MUSEUM
The protoconch agrees with Watson’s description. Its extent is uncertain
owing to corrosion of the surface, but 14 (possibly 2) whorls would seem a
reasonable estimate.
The columella (pillar), however. has no kink as has the Challenger shell,
and it has a very slight groove, visible on the 77 and 90 mm. apices, but dis-
appearing on the basal part of the columella as seen in the fragments.
The canal is not so markedly truncate as the figure of the Challenger shell
would seem to suggest, even when seen in approximately the same position.
Perhaps the edge of the canal was broken, but drawn by the artist as if unbroken.
The fragments from the same haul as the 90 mm. apex include: a portion
of the outer wall of the shell with sutural inflexion, which does not fit on to the
.
4
t]
ie
=
4
Fic. 5. Guivillea alabastrina (Watson). a, b, two views of columella, with
(slightly enlarged) sections. c, aperture of last whorl of specimen (proto-
conch + 2? whorls), showing grooved columella. d, specimen Ag841
with, e, view of broken end of columella at a point opposite x, at right
angles to frontal view. (All figures about + natural size.)
apex as far as the latter is preserved; the canal with adjacent columella (pillar)
to the upper end of which another fragment of columella appears to join (the
opposed surfaces are not large enough to form an undeniable ‘fit’).
There is some doubt whether the 90 mm. apex and the columella fragments
belong to one or two shells.
Dr. Talbot tells me that the contents of the dredge when it came aboard
formed such a compact mass of globigerina ooze that the extraction of the
animals was difficult. Nevertheless he thinks that if a second shell (apex) had
been present it would not have been overlooked.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 431
If the apex and the columella are placed end to end in their correct relative
position, and without allowance for the probable loss of small intervening
pieces, the shell would be at least 7} inches long; the Challenger shell was 64
inches long, and the species may well grow an inch larger.
34° 37’ S., 17° 03’ E., 1,580-1,620 fathoms, 1 protoconch plus 2 whorls;
33° 50’ S., 16° 30’ E., 1,480—-1,660 fathoms, 1 columella (worn); 34° 05’ S.,
16° 58’ E., 1,470-1,490 fathoms, 1 protoconch plus 24 whorls; 34° 36’ S.,
17° 00’ E., 1,500-1,760 fathoms, 1 protoconch plus 2? whorls, and fragments
(S. Afr. Mus. Ag776, Ag809, Ag841, and Ag870; F. H. Talbot coll.).
Distribution. Between Marion Island and the Crozets, 46° 16’ S., 48° 27’ E.,
1,600 fathoms (Watson: Challenger); South Orkneys (Melvill & Standen:
Scotia).
Remarks. This is the most interesting of the results of Dr. Talbot’s deep-sea
dredging.
Watson gave only a general description of the external appearance of the
Challenger animal. In 1882 he said that Prof. Huxley had undertaken the detailed
description of the anatomy, and in 1886 he said the description would appear
elsewhere (i.e. not in his Challenger Report). I have not been able to trace any
description by Huxley.
The animal, however, was submitted to Pelseneer, and a brief account
appeared in a later volume of the Challenger Reports (Pelseneer, 1888, p. 3,
pl. 1, figs. 1, 2). Pelseneer figured the foot and cephalic region from the right
side, but undertook no dissection or anatomical investigation except to remove
and section one of the rudimentary unpigmented eyes (Thiele, 1929, repeated
Watson’s statement that eyes were absent).
Possibly, therefore, the radula is still within the remains of the animal.
Mr. Dance (in litt. 2 Febr. 1960) told me that the animal was intact in the
British Museum. Can no one be found to extract the radula and confirm, or
otherwise, the animal’s position in the Volutidae?
Fasciolariidae?
Gen. ?
Two broken and corroded shells, one 30 mm. long, the other 23 X 10mm.,
extracted from anemones. Whorls preserved: 5 and 4 respectively. Aperture
(incl. canal) about 14 times the spire. Profile evenly convex, but possibly with
a slight midwhorl shoulder. No sulcus. Columella slightly curved, no pleats;
canal well marked. No axial sculpture; spiral lirae on last whorl (4th) of
smaller shell 12-13, regular, subequal; on 5th whorl of larger shell 12 on upper
half, 6 on lower half of whorl. Although the numbers of lirae on the two shells
differ in number and strength, they cover the whorl completely between upper
and lower sutures. On base (of smaller shell) 12 lirae plus c. 8 on rostrum.
34° 37’ S., 17° 03’ E., 1,580—1,620 fathoms, 2 dead (S. Afr. Mus. Ag774,
F. H. Talbot coll.).
432 ANNALS OF THE SOUTH AFRICAN MUSEUM
Mitridae
Gen. CHARITODORON Tomlin
Barnard, 1960b, p. 402.
Examination of the living material brought up by Dr. Talbot’s dredging
has resulted in transferring this genus from the Buccinidae to the Mitridae.
Fam. ?
Nux alabaster Brnrd.
Nux alabaster Barnard, 1960c, p. 440, fig. 2.
The radula of this curious species indicates one of the Rhachiglossate
families, but its exact systematic position remains doubtful.
Mr. A. E. Salisbury (in litt. 20 June 1961) has drawn my attention to the
previous use of the generic name Nux, viz.: Humphrey, Mus. Callonianum, 1797,
p- 59. This work was arbitrarily rejected by the International Committee
(Opinion 51). But a future International Committee may, also arbitrarily,
reverse this opinion. The name is in Sherborn’s Index Animalium 1758-1800,
but not in Neave’s Nomenclator. For the time being I maintain the name.
Buccinidae
Neptunea bonae-sper n. sp.
(Fig. 6 a, b)
Protoconch 24 whorls, smooth, but corroded and junction with Ist post-
natal whorl indistinct. Postnatal whorls 6; profile of whorls evenly convex.
Axial ribs c. 15 on 1st whorl (but slightly corroded), 16 on 2nd, 18 on 3rd, 20
on 4th, 22 on 5th, and 26 on 6th whorl, straight or slightly retractive, from
suture to suture, obsolete on base; crossed by spiral lirae 5 or 6 on Ist, 7-8 on
2nd, 8 on 3rd and 4th, 10 plus intermediaries on last two whorls, c. 24 on base.
Canal short, rather wide. 55 < 26 mm. and 51 X 27 mm. Operculum ovate,
nucleus apical, 14 X 9 mm. Creamy-white with pale buff, thin, somewhat
scabrous periostracum; operculum amber-brown.
Animal pale. Eyes well developed. Radula with 80-85 rows, central plate
quadrangular, with median cusp, sometimes a minute denticle on one side or
on both sides; lateral plate much stronger than central plate, unequally bicus-
pid, with 2-5 tiny denticles between the two cusps, the denticles not always
symmetrical.
33° 49'.S., 16° 30’ E., 1,500 fathoms, 1 living; 33° 52; (S.5; 107 jee
1,380-1,520 fathoms, 2 living (Types), 2 dead; 34° 36’ S., 17° 00’ E., 1,500—
1,760 fathoms, 1 living (S. Afr. Mus. Ag757, Ag826 (Types), and Ag887,
HE Walboticolls):
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 433
Remarks. The assignation of this Cape species to the old boreal genus
Neptunea may seem strange; it is admittedly somewhat unsatisfactory, but it is
an alternative to instituting a new genus.
The shell is an ordinary-looking Buccinid, but the radula has unusual
features.
The central plate resembles that of Mohnia (see Thiele, 1929, fig. 342),
Chauvetia (Lachesis) (see Thiele, 1929, fig. 357), some species of Sipho (e.g.
tslandicus, gracilis, glaber) (see Sars, 1878, pl. x, figs. 19, 20, 21), and Lachesis
australis von Martens ( = albozonata Watson) (see Thiele, 1903, pl. 9, fig. 55;
also Powell, 1951, fig. K 59).* ©
On the other hand the lateral plate agrees with that of none of these genera,
but closely resembles that of Chrysodomus turtoni (see Sars, 1878, pl. x, fig. 16).
Thiele (1929) puts Chrysodomus as a synonym of Neptunea, but does not mention
a particular species as its representative. The length of the outer cusp and
shortness of the inner cusp give the lateral plate of C. turtont and Neptunea
bonae-sper a distinctive shape. (The central plate of C. turtoni has no cusp.)
The specimen from 34° 36’ S., 17° E. (Ag887), is more slender than the
other specimens; the axial ribs are evanescent on the last whorl, and obsolete
on the back of the outer lip.
I have been shown a 7-whorled specimen (in coll. Fisheries Survey)
62 X 30 mm. On the 7th whorl the axial ribs are obsolete.
Prosipho torquatus n. sp.
(Figs 6c) d, ¢)
Protoconch 14 whorls, alt. and diam. 1 mm. Postnatal whorls 4; profile of
whorls angularly shouldered, but not sharply, a little above middle of whorl.
Axial ribs on 1st whorl (partly corroded) 16, on each of the following whorls
17-18, from shoulder to suture, and extending across base; crossed by spiral
lirae 3-4 on 2nd whorl, 5-7 on 3rd, 8-9 on 4th whorl; small granules on the
intersections, those on the shoulder slightly larger than the others; 15 additional
lirae on base. Below the suture a circlet of granules, about twice as many as the
axial ribs. 15°5 X 7 mm. Operculum 4 x 2 mm., ovate, nucleus apical.
Dirty white, operculum pale amber.
Radula with 75 rows, central plate excised in front, with 3 cusps, lateral
plate strong, twice as long as the central plate, with 2 apical cusps.
34° 37’ S., 17° 03’ E., 1,580-1,620 fathoms, 1 living (Type); 33° 50’ S.,
16° 30’ E., 1,480-1,660 fathoms, 1 dead; 34° 36’ S., 17° 00’ E., 1,500-1,760
fathoms, 1 dead (S. Afr. Mus. Ag884, Ag801 and Ag886 respectively; F. H.
Talbot coll.).
Remarks. P. astrolabiensis (Strebel) seems to be the only other species with a
bicuspid lateral radula plate, the others having more than two (3-6). The shape
*For the last-mentioned species, from Kerguelen, Powell (1951) proposed the generic name
Falsimohnia.
A434 ANNALS OF THE SOUTH AFRICAN MUSEUM
of the lateral plate, however, in the present species is different trom that of
astrolabiensis as figured by Powell (1951, fig. K 56). When mounting the radula
some of the lateral plates were purposely displaced into various positions, but
none of them assumed the shape shown in Powell’s figure.
The shell of astrolabiensis is quite different from the present shell.
This record forms a noteworthy extension of the known distribution of this
Antarctic and sub-Antarctic genus.
—<
——
G
Uy
SN
Fic. 6. a, Neptunea bonae-spei n. sp. b, central and lateral plates of radula. c, Prosipho
torquatus n. sp. d, two views of protoconch. e, central and lateral plates of radula.
Pyrenidae
Pyrene cf. profundi (Dall)
Astyris profundi Dall, 1889, p. 192, pl. 35, fig. 3.
Columella (Astyris) profundi Dautzenberg, 1927, p. 89.
Protoconch nucleus diam. 0:25 mm., plus 7 whorls. Profile of whorls
slightly convex. Spire longer than aperture. Surface smooth, without any
sculpture, the growth-lines for the most part very indistinct; but a few spiral
lirae on rostrum. Outer lip sometimes with feeble varicoid thickening; no
plicae on inner surface. No periostracum. 11 X 5 mm. Operculum subtriangu-
larly ovoid, thickened on inner surface in basal half (i.e. from nucleus onwards),
the thickening extending along both lateral margins and also forming a midrib,
leaving a semi-oval thinner area between the latter and the margins; midrib
not (or scarcely) visible on external surface. White, operculum amber.
Radula normal, proximal cusp on lateral plate well separated from the
bifalcate apex.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 435
BA 36) (9.,17: 00; E.,)1,500-1,760 fathoms, & living, 9 dead; 33° 40’'S.,
16° 30’ E., 1,500 fathoms, 1 dead (S. Afr. Mus. Ag864 and Ag758 respectively ;
F. H. Talbot coll.).
Remarks. A perfectly plain, smooth and slightly glossy species, comparable
with both Astyris diaphana Verrill from off east coast of North America and with
A. profundi Dall from the same region and also the Azores and Cape Verde;
but intermediate between the two in proportions.
The appearance of a trident on the internal surface of the operculum is not
distinctive, because it occurs in P. filmerae and in Columbella fulgurans.
Muricidae
Trophon acceptans Brnrd.
(Fig. 7 a)
Trophon acceptans [ partim] Barnard, 1959, p. 202, figs. 40 d (radula), 43 6 (only the fig. of adult)
(only the adults A3449, A3473 and A348o). .
39° 50'S.,17 21 E., 600 fathoms, 4 living, 7 dead; 33° 36’ S., 16° 15’ E.,
1,520-1,570 fathoms, 9g living, 2 dead; 33° 454’ S., 16° 232’ E., 1,480 fathoms,
2 living; 33° 49’ S., 16° 30’ E., 1,500 fathoms, 8 living, 1 dead; 34° 37’ S.,
17° 03’ E., 1,580—1,620 fathoms, 4 living, 1 dead; 34° 42’ S., 16° 54’ E., 1,725-
1,780 fathoms, 2 living, 2 dead; 33° 50’ S., 16° 30’ E., 1,480-1,660 fathoms,
1o living; 34° 05’ S., 16° 58’ E., 1,470-1,490 fathoms, 2 living; 34° 36’ S.,
17° 00’ E., 1,500-1,760 fathoms, 3 dead (S. Afr. Mus. Ag7o1, Ag734, Ag743,
Ag759, A9778, Ag791, Ag811, Ag842, Ag865; F. H. Talbot coll.).
The new material comprises 56 specimens ranging from 10 to 46 mm. in
length, most of them living.
Fic. 7. a, Trophon acceptans Brnrd., multicostate variations. b, Trophon cf.
droueti Dautzenberg.
436 ANNALS OF THE SOUTH AFRICAN MUSEUM
In the original description were included some juveniles, 3:5 to 8 mm.,
which appeared to be conspecific with the adults, the smallest of the latter
being 18 mm. in length. I am now inclined to think this was an error, and that
not enough importance was attached to the position of the shoulder. There is,
in fact, in these juveniles no angular shoulder, and the highest part of the profile
of the ribs is at, or nearly at, a level with the suture (see figs. of juv.). The
recently obtained 10 mm. specimen shows, although the apex is corroded, that
there is a definite angular shoulder from at least the 2nd (postnatal) whorl
onwards. Therefore the juveniles from the Agulhas Bank, Algoa Bay and East
London are now excluded from acceptans, and considered as belonging to a
separate species. This is best left without a name pending the discovery of
further and better material.
The original description, omitting those characters which apply to the
juveniles, may be emended and added to as follows: Postnatal whorls 7 (1st
whorl corroded in all specimens); 2nd and following whorls angularly shoul-
dered; axial ribs on 2nd and grd whorls 11-12, on 4th and 5th 12-13, on 6th
and 7th 13-14, sharply keeled and slightly squamosely lamellate at the shoulder
(when not worn or corroded); on the later whorls the intervals between the
ribs become U-shaped, and when the ribs are far apart the intervals are very
open and flat. The rostrum and canal may be slightly curved in the largest
shells. A thin, pale brown periostracum, which usually comes away when the
investing Epizoanthus is removed. Radula of large specimens with 125-135
rows.
Remarks. All the material collected by the Pieter Faure and Dr. Talbot
came from the same area (see original description and the localities given
above). The bathymetrical distribution is as follows:
At 630-800 fathoms 7 specimens 18-21 mm., somewhat corroded but clean,
1 of them living (Preter Faure).
At 600 fathoms 11 specimens 10-22 mm., somewhat corroded but clean,
4 of them living.
At 1,480 fathoms 2 living specimens 31 and 35 mm., somewhat corroded,
part of each shell covered with the beginning of a colony of purple Epizoanthus
(Coelenterate), with one or two polyps.
At 1,470—-1,490 fathoms 2 living, 22 and 25 mm., clean.
At 1,480-1,660 fathoms to living, with Epizoanthus colonies.
At 1,500 fathoms 9 specimens, 8 of them living, 21-45 mm., somewhat
corroded, covered with Epizoanthus colonies, that on the largest shell with
7 polyps.
At 1,500—1,760 fathoms 2 dead.
At 1,520-1,570 fathoms 11 specimens, 9 of them living, 30-46 mm.,
somewhat corroded, covered with Epizoanthus colonies, with up to 10 polyps
on a shell.
At 1,580-1,620 fathoms 5 specimens, 4 of them living, with Epizoanthus
colonies, and 1 dead extracted from an anemone (Actinian).
1
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 437
At 1,725-1,780 fathoms 4 specimens, 2 of them living, 30 and 32 mm.,
clean.
No examples have been found in less than 600 fathoms, and all those
obtained at this depth, and down to 800 fathoms, were not more than half-
grown. The largest shells, and also half-grown (23 mm.) and three-quarter-
grown shells were obtained at 1,480 fathoms and greater depths. The animals
from lesser depths, though smaller, may nevertheless be sexually mature and
represent a dwarf form. There is as yet no evidence on this point.
The purple Epizoanthus is found only at the greater depths, 1,480 fathoms
onwards. It settles on half- or three-quarter-grown shells, and completely
envelops the largest shells, including the whole ventral surface, though of
course the polyps arise only dorsally and laterally. When the mollusc is with-
drawn into its shell there is nothing to indicate that the object is other than a
clump of polyps, distasteful to fishes (as many Coelenterates are known to be)
and possibly also to predaceous molluscs or Echinoderms.
The original figure of the ‘adult’ will serve also for the larger shells, and
represents the typical form; two figures are here given showing multicostate
variations.
In the original description the one living specimen (S. Afr. Mus. A3473)
was designated the Type. The new material contains specimens which, because
they show the size to which it grows, are really more typical of the species (?
hypertypes).
A resemblance to tenuirostratus Smith 1899 and 1901 was noted in the ori-
ginal description ; but there is a considerably stronger resemblance to obtuseliratus
Schepman 1911. These are resemblances between specimens from the Cape
and from localities in the Indian Ocean and the East Indies.
A more serious question is the possible identity of the Cape shells with
guineensis Thiele (1925, p. 169, pl. 30(18), fig. 11) from 2,278 metres in the Gulf
of Guinea. Comparison of Thiele’s figure and mine leaves little choice, and I
fully expect that acceptans will not be accepted when further material is obtained
from the Atlantic trough along the west coast of Africa. For the present the
Cape shells are retained as a separate species.
Variation. The following examples I consider as no more than individual
multicostate variations.
One (30 mm.) of the two examples from 1,480 fathoms has 16 ribs on the
5th, and 18 on the 6th whorl.
One (32 mm.) of the specimens from 1,500 fathoms has 15 ribs on the 3rd
whorl, and 16 on the 4th, 5th and 6th whorls.
One (27 mm.) of the specimens from 1,500 fathoms has 16 ribs on the 3rd
whorl, 18 on the 4th and 5th whorls, and 14 on the first three-quarters of the
6th whorl followed by 2 ribs widely separated.
One (46 mm.) of the specimens from 1,520-1,570 fathoms has 15 ribs on
the 4th whorl, 17 on the 5th and 6th whorls, and 15 on the 7th whorl.
In the last-mentioned shell (46 mm.) the shoulder disappears on the last
438 ANNALS OF THE SOUTH AFRICAN MUSEUM
(7th) whorl, and consequently the shell approximates in shape to the figure of
declinans Watson, though the latter has no shoulder on any of the whorls.
Faint indications of 2 spiral lirae below the shoulder on the 6th whorl
were noted in the original description. In these multicostate variations there are
indications of 3 or even 4 such lirae.
Trophon cf. droueti Dautzbeg.
(Fig. 7 4)
Trophon droueti Dautzenberg, 1889, p. 37, pl. 2, figs. 1 a, b, c (hand-drawn).
1927, p. 92, pl. 7, figs. 26-28 (photo).
Protoconch 14 whorls, alt. and diam. c. 1 mm. (slightly corroded). Post-
natal whorls 34—4, profile angularly shouldered, but shoulder becoming rounded
on last whorl. Axial ribs 16-17 on 1st whorl, increasing to 24 on last, retractive
from suture to shoulder, straight below, sharp, becoming distinctly lamellate
on back of outer lip. On 2nd and 3rd whorls a feeble lira at the shoulder and
another below it produce small nodules on the ribs; on last whorl 2 more lirae
below the subperipheral one. 10-11°5 X 5 mm. White.
34° 36’ S., 17° 00’ E., 1,500-1,760 fathoms, 3 dead (S. Afr. Mus. Ag866,
F. H. Talbot coll.).
Distribution. Azores, 1,287 metres.
Remarks. These shells are remarkably like droueti, and I deem it advisable
not to institute a separate species for them, at least not until further material is
available. They are slightly more slender (droueti: 8 * 4 mm.), thus possibly
representing var. elongata Locard, 1897. The spiral lirae are very feeble, but
sufficiently in relief to cast slight shadows, comparable with the grey bands in
Dautzenberg’s hand-drawn figures (1889).
Dautzenberg estimated from fragments that the species reached a size of
16 mm. The present specimens have a protoconch as large as that of acceptans,
and the species may possibly reach a greater size than 16 mm. in the Cape area.
In Dautzenberg’s fig. 1 ) the number of riblets seems to be greater than
would be expected.
Columbariidae
Columbarium rotundum Brnrd. and angulare Brnrd.
The localities from which specimens of these two species were obtained
confirm the results obtained by the Preter Faure (Barnard, 1959, pp. 235, 236).
C. rotundum occurs in depths of 250-760 fathoms; but angulare, which the Pieter
Faure obtained in depths of 720-900 fathoms, has now been shown to extend
down to 1,520 fathoms.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 439
Cymatiidae ©
Thalassocyon bonus Brnrd.
Thalassocyon bonus Barnard, 1960c, p. 440, fig. 3.
Excepting Guwivillea, this is the most interesting Mollusc obtained by Dr.
Talbot. The shell resembles in shape a Semifusus, but the animal was found to
have a taenioglossate radula similar to that of Cymatium.
Odcorythidae
Odcorys watson Locard
Oécorys sulcata (non Fischer) Watson, 1886, p. 412, pl. 17, fig. 11.
Oécorys watsoni Locard, 1897, p. 288. Tomlin, 1927, p. 80. Barnard, 1963, p. 9.
Largest specimen 44 X 31 mm. Dead specimens were previously taken by
the Pieter Faure off Cape Point in 720-1,000 fathoms.
Naticidae
Polynices cleistopsila Brnrd.
Polynices cleistopsila Barnard, 1963, p. 64.
Falsilunatia pseudopsila Brnrd.
Falsilunatia pseudopsila Barnard, 1963, p. 64.
Pyramidellidae
Turbonilla cf. kraussit Clessin
Turbonilla cf. kraussi Clessin, Barnard, 1963, p. 85.
Although closely similar to the littoral and shallow-water kraussi, the single
dead specimen will probably prove to be a distinct species when more material
is obtained.
Cerithiopsidae
Cerithiella taylor: Brnrd.
Cerithiella taylori Barnard, 1963, p. 126.
Lamellariidae
Lamellaria capensis (Bergh)
Lamellaria capensis Bergh, Barnard, 1963, p. 58.
Scalidae
Scala bonae-sper Brnrd.
Scala bonae-spei Barnard, 1963, p. 104.
Abyssochrysidae
Abyssochrysos melantoides ‘Tomlin
Abyssochrysos melanioides Tomlin, 1927, p. 78, figs. 1-3. Barnard, 1963, p. 141.
Previously taken by the Pieter Faure off Cape Point in 800-1,000 fathoms;
now shown to occur down to 1,490 fathoms.
440 ANNALS OF THE SOUTH AFRICAN MUSEUM
Trochidae
Calliotropis metallica (W.-M. & A.)
(Fig. 8 a)
Solariella metallica Wood—Mason & Alcock, 1891, p. 444, fig. 12 a, b.
Previously taken by the Pieter Faure off Cape Point. A notable extension of
the hitherto known distribution: Gulf of Manaar, East Indies, East Africa.
An account of the species will be given in Part IV of Barnard, Contributions...
South African Marine Mollusca. :
Calliotropis pompe n. sp.
(Fig. 8 b)
Shell thin-walled. Protoconch nucleus plus 7 whorls. First to 3rd whorls with
c. 23-24 slightly retractive axial pliculae; on 3rd and following whorls crossed
by a peripheral spiral lira at lower third of whorl, and at end of 3rd whorl
and on 4th and following whorls by a second lira at upper third; the upper lira
forms conical tubercles at the intersections with the pliculae, c. 18-20 increas-
ing to c. 25 on 6th whorl, but becoming feeble and eventually evanescent on
7th whorl; on 5th-7th whorls the pliculae are distinct from suture to upper lira,
Cc
Fic. 8. a, b, sculpture of penultimate whorl of Calliotropis metallica (W.-M. & A.)
and C. pompe n. sp. c, Calliostoma glaucophaos n. sp., with two views of protoconch,
central and 1st marginal plates of radula.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 441
extending less distinctly to the lower lira; from end of 5th whorl onwards
accessory pliculae develop at the suture, 2-3 between each pair of main pli-
culae; similar accessory pliculae develop on the lower lira, so that the latter
becomes finely granulate; on 7th whorl all the pliculae become less distinct
and more or less indistinguishable from the growth-lines; the lower lira becomes
almost smooth. On base growth-lines and pliculae continued, the latter becom-
ing stronger towards the umbilicus; 5 spiral lirae, the outer 3 nearly smooth,
the next one granulate, and the one bordering the umbilicus strongly granulate;
umbilicus plicate within. 19 X 17 mm. (6 whorls); 22 x 20 mm. (7 whorls).
White, with a faint greenish tinge due to the nacreous interior. Operculum
pale corneous.
Jaws and radula as in granolirata.
Off Cape Point: 34° 42’ S., 16° 54’ E., 1,725-1,780 fathoms, 1 dead;
33° 50’ S., 16° 30’ E., 1,480-1,660 fathoms, 1 living, 1 dead (Types) (S. Afr.
Mus. Ag795 and Ag883 (Types); F. H. Talbot coll.).
Remarks. The procession (pompe) of close-set axial pliculae, and the granu-
late, instead of tuberculate (as in metallica and other species), lower spiral lira,
seem distinctive.
Gen. BasiutissA Watson
Wearson 1879, p. 593; 1886, p. 96. Schepman, 1908, p.61. Thiele, 1925, pp. 43, 44; 1929, p. 48.
Dall (1881) instituted the genus Fluxina, and considered that it should
probably be placed in the Solaridae, occupying in this family an analogous
position to that of Basilissa among the Trochidae.
Fluxina discula Dall (1889, p. 273, pl. 23, figs. 5, 6), F. marginata Schepm.
1908, F. trochiformis Schepm. 1908 and the present species are very much alike
in shape, differing from Basilissa by being strongly depressed. Admittedly the
difference is one of degree only, because brunnea Dall 1881 (genotype of Fluxina),
lampra Watson 1879 (genotype of Basilissa) and alta Watson var. delicatula Dall
(see 1889, pl. 22, fig. 2) form a series transitional to the higher species simplex
Watson 1879 and superba Watson 1879.
The radulae of only a few species are known, e.g. lampra (see Schepman,
1908), sibogae Schepman 1908, and trochiformis (see Thiele, 1925). Thiele gave
no figure of the latter. The present species has a radula somewhat resembling
that of szbogae, but not at all like that of lampra.
Provisionally this n. sp. is included in Baszilissa.
Basilissa gelida n. sp.
(Fig. 9)
Protoconch nucleus plus 5 whorls. Smooth, polished, periphery very sharply
keeled. No spiral sculpture; fine close growth-lines, sigmoid both above the
keel and on base. Umbilical wall smooth, vertical; umbilical margin rect-
angular, not keeled, no impressed line outside the margin. 8, alt. 3:5 mm.
442 ANNALS OF THE SOUTH AFRICAN MUSEUM
White, transparent, slightly iridescent when wet.
Operculum not observed.
Jaws present, reticulate. Radula with c. 45 rows, central plate quadrangu-
lar, with slightly overturned cutting-edge, with feeble median cusp (? other
serrations), lateral plate wide, with slightly overturned serrulate cutting-edge,
Ist marginal plate, hastate, distally obscurely serrulate, and 3 slender hamate
marginals.
33° 26’ S., 16° 33’ E., 1,240—-1,300 fathoms 1 living (S. Afr. Mus. Ag720,
F. H. Talbot coll.).
Fic. 9. Basilissa gelida n. sp., with radula plates.
aaa
oe
"Ah iy
ma
SPT TH FH
== BH f
Remarks. Differs from Fluxina discula Dall 1889 from the West Indies, 982
fathoms, only in having a non-carinate umbilical margin, without impressed
line; and in being slightly nacreous.
There are 3, possibly 4, slender outer marginal plates in the radula, in
addition to the stouter 1st marginal plate.
Calliostoma glaucophaos n. sp.
(Fig. 8 c)
Shell like Solariella in shape, slightly wider than high. Protoconch nucleus
plus 34 whorls. Protoconch alt. 0-8, diam. 1 mm., smooth. Profile of whorls
rounded, but with tabulate shoulder at upper third. One spiral lira forming the
shoulder and one at middle of whorl, both beginning on rst whorl; a third,
peripheral lira concealed in the suture until the last half-whorl. On the tabulate
shoulder 1 lira near the suture followed by 1 (2nd whorl), 3-4 (3rd whorl), 5
(last half-whorl) very fine lirae, also between inner lira and suture 2—3 very fine
lirae visible on last half-whorl. Beginning on 2nd whorl 1 lira between the
shoulder and mid-whorl lirae, and 1 between the latter and the peripheral
lirae. Base with 1 lira almost as strong as the peripheral lira, starting at junction
of outer lip and body-whorl, followed by c. 18 feeble lirae ; umbilicus bordered by
a strong lira; 1 feebler lira within the umbilicus, which is pervious but narrows
rapidly. Growth-lines mostly faint, not pliculose. Aperture subcircular, slightly
angular where outer lip meets the narrow lira-like columella. 11 (long) x 12°5
(diam.) mm.
White, iridescent, umbilical and columellar lirae opaque white. Operculum
amber.
Jaws with intercalated platelets. Radula with c. 50 rows, resembling that of
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 443
perfragile, but the central plate is broader, with a broadly triangular cusp,
minutely serrulate distally; 5 lateral plates, 1st marginal plate strong, hooked
and serrate distally. The lateral plates and especially the central plate are so
very delicate that the shape of their bases could not be determined.
33° 52’ S., 16° 51’ E., 1,380—-1,520 fathoms, 2 living (S. Afr. Mus. Ag830,
F. H. Talbot coll.).
Remarks. The specimen with the strong outer basal lira is figured. In the
other shell this lira is much weaker, but both it and the next one are more
distinct than the other basal lirae. The columella appears to have been injured
and repaired, and consequently is broader and somewhat concavely angular.
Although clearly distinct, these shells are not unlike Trochus (Margarita)
charopus Watson 1879 and 1886 from Kerguelen, 105 fathoms, and, though less
so, T(M) brychius Watson 1879 and 1886, also from Kerguelen, 1,260 fathoms.
There is a general resemblance to a Solariella; and unless the animal had
been present these shells might perhaps have been assigned to Solariella.
NUDIBRANCHIATA
Doridoxidae
Doridoxa benthalts n. sp.
(Fig. 10)
Length of animal as preserved c. 32 mm. Dull brown, the retracted
rhinophores orange.
Frontal veil with lateral processes, on the underside of each process a
wrinkled fold of skin connecting with the wrinkled margin of the mouth.
Fic. 10. Doridoxa benthalis n. sp. Lateral and dorsal views of animal; ventral view of
anterior end; front and inner views of left mandible; two central plates of radula in
side view; central and lateral plates of radula.
444 ANNALS OF THE SOUTH AFRICAN MUSEUM
Foot tapering posteriorly. Dorsal surface smooth, but with faint indications of
4 pairs of small warts. No gills. Genital opening, nephroproct, and anus on
right side, the nephroproct a short distance in front of anus.
Jaws large, 4-5 mm. long, cutting-edge entire. Radula with 30 rows,
central plate very strong, lateral plates 11-12, dagger-like, graduated, the
middle ones slightly larger than the others.
Internal organs not well enough preserved to determine the presence of a
blood-gland, or whether there were 2 sphermathecae. There were, however,
no liver diverticula. )
34° 42’ S., 16° 54’ E., 1,725-1,780 fathoms, 1 (S. Afr. Mus. Ag796,
F. H. Talbot coll.).
Remarks. In spite of uncertainty about the internal anatomy, this specimen
seems certainly to be a species of Doridoxa Bergh 1900, which was instituted for
ingolfiana Bergh taken in the North Atlantic. There seem to be no later records.
I have not seen the original description, but Bergh (1906, pl. 31, figs. 7-12)
gave 6 figures (presumably reproduced from his Ingolf Report, 1900), and
Thiele (1929, fig. 521, after Bergh) figured the radula. The resemblance is
close, but the present specimen appears to have no tentacles, larger lateral
processes, and small differences in the radula plates. A n. sp. seems warranted
and desirable.
SCAPHOPODA
Dentaliidae
Dentalium eualdes n. sp.
Thick-walled, moderately curved, ribs very numerous. Ribs 16-18 on the
smallest shells (30 mm), increasing to 65-85 in the largest shells; ribs subequal
to the grooves in larger specimens, but in juveniles usually narrower; the
interpolation of intermediaries tends to narrow the grooves, and in large shells
the ribs may be at least as wide as the grooves.
Ribs extending to aperture, but in the two largest shells there is an unribbed
(or with only faint traces of ribs) ‘collar’ 3-4 mm. long around the aperture.
Apical slit usually present, may be 5—7 mm. long; juveniles may show 2-4
elongate perforations.
86, diam. aperture 13, apex 3 mm.; 90 X 12°5 X 2°75; 98 X 14°5 X 2°35
99 X 13 X 1°75 mm.
Dull grey, the unribbed collar, when present, white.
Radula as in salpinx.
33° 36’ S., 16° 15’ E., 1,520-1,570 fathoms, 8 living (Types) ; 33° 454’ S.,
16° 232’ E., 1,480 fathoms, 1 living and 1 juv.; 33° 49’ S., 16° 30’ E., 1,500
fathoms, 11 living and 3 juv.; 33° 26’ S., 16° 33’ E., 1,300 fathoms, 1 living;
34° 37’ S., 17° 03’ E., 1,580-1,620 fathoms, 1 adult living, 1 juv. dead; 33°
50’ S., 16° 30’ E., 1,480—-1,660 fathoms, 18 living, 2 dead; 33° 52’ S., 16° 51’ E.,
1,380-1,520 fathoms, 1 living, 1 dead; 34° 05’ S., 16° 58’ E., 1,470—-1,490
fathoms, 12 living, 1 dead; 34° 36’ S., 17° 00’ E., 1,500-1,760 fathoms, 1 juv.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 445
dead (S. Afr. Mus. Ag736 (Types), A9747, Ag765, Ag767, Ag784, Ag815,
Ag9834, Ag849, A9875; F. H. Talbot coll.).
Remarks. Grows to a larger size than salpinx Tomlin, and has many more
ribs.
It is not capillosum: it is more strongly curved during early growth, and the
diameter increases more rapidly. On one of the smaller examples (52 mm.)
the diameter increases from 1-3 to 5:5 mm. in a length of 30 mm., whereas
in a specimen of capillosum (identified by Tomlin) the diameter reaches only
3 mm. in the same length. These two differential characters can also be
observed by superimposing the 52 mm. shell on Watson’s figure of the Challenger
example of capillosum (1886 pl. 1, fig. 1 a).
The present species is stouter than the figure of magnificum Smith 1898
pl. 7, figs. 5, 5a (= vernedei Hanley).
Dentalium lardum n. sp.
Moderately curved. Smooth, glossy, with fine growth-lines. Apical portion
ribbed, the ribs usually extending farther on the concave side, c. 20, increasing
to 27-30 (but somewhat obscure), obsolete on later growth. No slit or perfora-
tions. Details of the specimens as follows.
40, diam. aperture 5:5, apex 0-75 mm. Glossy white; ribbed for the apical
23 mm., faintly indicated for another 7-8 mm. on concave side, thereafter only
growth-lines; ribs 20, increasing to 27.
49 X 7 X 1°3 mm. Glossy, first two-thirds grey, thereafter white; ribs
faintly visible only in the apical 10 mm.
50 X 7 X 1-5 mm. Glossy, ivory-white; partly corroded apically, but
no trace of ribs.
65 x 85 x 2 mm. Glossy, grey or yellowish-grey; ribs faintly visible in
apical 15-18 mm., more so on concave side than on convex, ribs c. 30 (but not
easy to trace).
33° 49’ S., 16° 30’ E., 1,500 fathoms, 5 dead (Types) ; 33° 52’ S., 16° 51’ E.,
1,380—1,520 fathoms, 2 dead; 34° 36’ S., 17° 00’ E., 1,500—-1,760 fathoms, I juv.
dead (S. Afr. Mus. Ag768 (Types), Ag835, A9876; F. H. Talbot coll.).
The specific name from the smooth, somewhat greasy appearance.
Dentalium sp.
One shell 7:5, one 8, one 11, one 13, and one 17 mm. long. Slightly
curved. Ribs g on all specimens from apex onwards; at 13 mm. one inter-
mediary begins between each pair of main ribs, and on the last 4 mm. of the
17 mm. shell there are 18 ribs, the intermediaries almost as strong as the main
ribs. Apical diam. 0:4-0:5, basal diam. of 13 mm. shell 1-3, of 17 mm. shell
1-5 mm. Growth-lines but no other sculpture between the ribs.
33° 26’ S., 16° 33’ E., 1,240-1,300 fathoms, the 2 largest; 34° 37’ S.,
17° 03’ E., 1,580-1,620 fathoms, 1; 34° 36’ S., 17° 00’ E., 1,500—1,760 fathoms,
1; all dead (S. Afr. Mus. Ag885, Ag877, Ag785 resp.; F. H. Talbot coll.).
446 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. Further material is desirable before a name is given to these
specimens. But attention is drawn to the rather sudden doubling of the number
of ribs; a larger specimen with the apical 13 or 14 mm. broken off would be
regarded as an 18-ribbed species.
Dentalium sp.
93° 50° 'S., 17° 21’ E., 600 fathoms, 17 dead (S. Afr. Mus. Ao76o ieee
Talbot coll.). |
A species with 18-22 ribs, comparable with but distinct from plurifissuratum.
Up to 35 mm.
In this case also, more and better material seems desirable before attaching
a specific name to these specimens.
Dentalium capense ‘Tomlin
Dentalium capense Tomlin, 1931, p. 340.
93° 50’ S., 17° 21’ E., 600 fathoms, 1 living, 2 dead (S. Afr. Mus. Ag77o,
F. H. Talbot coll.).
Previously taken by the Preter Faure off Cape Point in goo fathoms; and
also off Durban in 440 fathoms.
Cadulidae
Cadulus promontori n. sp.
Previously taken by the Preter Faure off Cape Point in 700 fathoms.
For description and figure see: Barnard, Contributions . . . South African
Marine Mollusca, Part IV [in press].
CEPHALOPODA
Cranchiidae
Leachia cyclura Lesueur
Leachia eschscholtzu Rathke, Chun, 1910, p. 347, pl. 52, figs. 4-7.
33° 45 ©. 16.29 E., 1,480 fathoms, 1 specimen) (S: Afi MusieAe@zaias
F, H. Talbot coll.).
LAMELLIBRANCHIATA
Nuculidae
Nucula (Pronucula) benguelana Clarke
(Fig. 11 a)
Pronucula benguelana Clarke, 1961, p. 368, pl. 3, figs. 9, 11.
Shell thin, subtriangular, not very oblique, length only slightly greater
than altitude. Young shell, alt. 1 mm., large, prominent and sharply demar-
cated from rest of shell. Whole surface with numerous fine radiating striae,
except in the positions of the lunule and escutcheon, which are otherwise
undefined. Teeth anterior 7-8, posterior 6—7; ligament pit vertical to hinge-
2
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 44.7
line; margin internally crenulate. Length 3°75, alt. 3-5 mm. Corneous, young
shell paler.
34° 26'S., 17° 00’ E., 1,500-1,760 fathoms, 1 living (S. Afr. Mus. Ag882,
F. H. Talbot coll.). |
30° 14’ S., 13° 3’ E., 1,703 fathoms (approx. 400 miles north-west of
Cape Town) (Clarke).
Remarks. Somewhat similar in shape to the North Pacific profundorum
Smith (1885, p. 229, pl. 18, fig. 13), and the striae are, in Smith’s words:
‘hair-like whitish lines’.
|
I
YU 4,7 f
Lich
rll
=
=
==
Fic. 11. a, Nucula (Pronucula) benguelana Clarke. b, Malletia estheriopsis n. sp. c, Leda parsimonia
n. sp. d, Leda macella n. sp. e, Sarepta sp.
Malletiidae
Malletia estheriopsis n. sp.
(Fis! 11 5)
Thin, oblong-oval, rounded at both ends, the anterior end a little less
broadly rounded, posterior dorsal margin straight; umbones at anterior third,
not prominent. Concentric growth-lines only. Teeth anterior 12-13, posterior
28-30, 2-3 inconspicuous teeth on each side of the interruption below the
umbo; ligament entirely external, conspicuous, about 3? length of straight
dorsal margin. Pallial sinus deep. Periostracum thin, pale yellowish. Length 15,
alt. 8 mm. Siphons completely fused.
448 ANNALS OF THE SOUTH AFRICAN MUSEUM
Cape Point N. 70° E. 40 miles, 800 fathoms, 1 (S. Afr. Mus. P.F. coll.).
33° 50’ S., 16°.30’ E., 1,480-1,660 fathoms, 2 living (Types); 34° 36’ S.;
17° 00’ E., 1,500-1,760 fathoms, 2 living; 34° 33’ S., 16° 43’ E., 1,770—-1,880
fathoms, 1 living; 34° 05’ S., 16° 58’ E., 1,470-1,490 fathoms, 3 living (S. Afr.
Mus. Ag817 (Types), Ag880, Ag890, and Ag8oq1 respectively; F. H. Talbot
coll.).
Remarks. These specimens seem to differ from other species in their bean-like
shape, rounded at both ends. Jeffreys (1879, p. 573) said his Szlicula fragilis
was like an Estheria (Crustacea, Conchostraca), but these specimens qualify
even better for the epithet.
Conchologically they closely resemble the Arctic obtusa Sars (1878, p. 41,
pl. 19, figs. 3a—c) but are not so obtuse and truncate posteriorly. Geographically,
the nearest species is pallida Smith 1885 (p. 246, pl. 20, figs. 8, 8 a) from 2,250
fathoms between Tristan da Cunha and the Cape.
Ledidae
Leda parsimonia n. sp.
(Fig. 11 ¢)
Triangular, almost equilateral, umbones only slightly nearer to the anterior
end; rounded at both ends. Middle of later part of shell with concentric pliculae,
closer together near the margin, rest of shell with growth-lines only. Teeth 12
anterior, 15 posterior, with a few minute ones on each side of ligament pit.
Pallial sinus very shallow. Length 13:25, alt. 8-25 mm. Yellowish-brown,
glossy. Animal decomposed; siphons ?
33° 50’ S., 16° 30’ E., 1,480—-1,660 fathoms, 1 living (S. Afr. Mus. Ag818,
F. H. Talbot coll.). |
Remarks. Resembles Yoldia semisculpta Thiele in being only partially plicate,
but differs in shape, and does not gape.
Leda macella n. sp.
(Fig. 11 d)
Rostrate, anterior end broadly rounded, dorsal posterior margin straight
(or very slightly concave), rostrum angular above, obliquely truncate, umbones
at anterior ? of length. Border of lunule from umbo to upper corner of rostrum.
curved. Concentric growth-lines only, some on the later part of shell coarser
than the others. Teeth at shell length 12 mm. 12-13 anterior, 14 posterior,
increasing to 15-16 and 16-17 respectively, with a few minute ones on each
side of the ligament pit. Pallial sinus moderately deep. No ridge on inner side
of rostrum below, and parallel to the posterior series of teeth. Periostracum
thin, yellowish or olivaceous brown. Length 20, alt. 10:5, thickness (valves
together) 8 mm.
33° 96’ S., 16° 33” E., 1,240-1,300 fathoms, 1 living; 33°45) “Stee
231’ E., 1,480 fathoms, 1 living; 33° 50’ S., 16° 30’ E., 1,480—1,660 fathoms,
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 449
1 living; 34° 05’ S., 16° 58’ E., 1,470-1,490 fathoms, a lot living (Types)
(S. Afr. Mus. Ag729, Ag751, Ag819, and Ag851 (Types); F. H. Talbot coll.).
Remarks. Similar in shape to prostrata Thiele 1931 and silicula Thiele 1931,
but not so strongly nor so narrowly rostrate, and the rostrum is obliquely
truncate; no longitudinal ridge on inside of rostrum; and the number of pos-
terior teeth is less, in conformity with the shorter rostrum.
L. prostrata came from 981 metres in the middle of the South Atlantic
(25° 25’ S., 6° 12’ E.), and szlzcula from 400-463 metres off the East African
coast.
Sarepta sp.
(Fig. 11 e)
Interior not nacreous (but only one dead valve present). Oval, not very
oblique, length greater than altitude; posterior margin convex. Concentric
ridges over whole surface. No lunule. Teeth anterior 12, posterior 10. Ligament
pit minute. Margin internally smooth. Length 5, alt. 4 mm.
34° 26’ S., 17° 00’ E., 1,500-1,760 fathoms, 1 valve (S. Afr. Mus. Ag88r,
F. H. Talbot coll.).
Limopsidae
Limopsis cf. straminea Smith
Limopsis straminea Smith 1885, p. 255, pl. 18, figs. 5, 5 a.
Shell oblique, length greater than altitude; anterior margin convex,
posterior margin nearly straight in upper half. Umbo slightly nearer to anterior
end of hinge-line. Concentric lirae, and fine radial striae marking the insertion
of the bristles. Internally smooth, no radial ridges, and no marginal crenula-
tions; no thickened pad at posterior adductor scar. Lower margin of hinge
slightly concave, very narrowly separated from ligament pit. Teeth interrupted,
in juv. up to 10 mm. 4-5, later 5-7 on each side (occasionally 8 on posterior
side), the posterior ones slightly more oblique than the anterior ones.
Length 27-28, alt. 24-25, width (valves together) 10-11 mm.
Foot with posterior process; byssus rudimentary.
Bo 20) 194100 93") Ey 1-240-1,900) fathoms, 9 living; 33° 26 35.,, 16”
15 E., 1,520-1,570 fathoms, 17 living; 33° 454’ S., 16° 234’ E., 1,480 fathoms
12 living; 33° 49’S., 16° 30’ E., 1,500 fathoms, ro living; 34° 37’ 8., 17° 03’ E.,
1,580-1,620 fathoms, 3 living; 33° 50’ S., 16° 30’ E., 1,480-1,660 fathoms, 6
living, 5 valves; 34° 05’ S., 16° 58’ E., 1,470-1,490 fathoms, 34 living, 4 valves;
34° 36’ S., 17° 00’ E., 1,500-1,760 fathoms, 2 living, one valve (S. Afr. Mus.
Ag727, Ag9739, Ag750, Ag766, Ag786, Ag816, Ag850, Ag879; F. H. Talbot
coll.).
Remarks. In juveniles up to about 10 mm. long the obliquity of the shell
is not so noticeable as in larger shells; and the lower margin of the hinge is not
sO narrowly separated from the ligament pit.
450 ANNALS OF THE SOUTH AFRICAN MUSEUM
Assigning a specific name to these shells is difficult. A valve superimposed
on Smith’s figure of straminea agrees exactly in shape. The number of teeth is
‘about 12’. Smith did not mention whether the margin was internally crenu-
late, but presumably it was not (he was dubious about this as a specific charac-
ter: p. 257): he gave no internal view of the shell.
L. straminea was taken between Kerguelen and Heard Islands at only 150
fathoms; nevertheless the present shells may be conspecific.
Ungulinidae
Thyasira investigatoris (Smith)
Cryptodon investigatoris Smith, 1895, p. 13, pl. 2, figs. 6, 6 a; 1897, Moll. pl. 3, figs. 2-2 b.
Thyasira investigatoris (Smith) Thiele & Jaeckel, 1931, p. 218.
33° 50’ S., 17° 21’ E., 600 fathoms, 1 right valve (S. Afr. Mus. Ag708,
F. H. Talbot coll.).
Previously taken (one specimen) by the Pieter Faure off Cape Point in
720-800 fathoms.
Semelidae
Abra longicallus (Scacchi)
Abra longicallis [sic] Sars, 1878, p. 74, pl. 6, figs. 3 a—c; pl. 20, fig. 4.
Syndesmya longicallus Scacchi, Dautzenberg, 1927, p. 333.
Agreeing with Sars’s description and figures. Length 13, alt. 9 mm. Up
to 25 mm. (Sars).
99° 50 8., 17° 21 E., Goo fathoms, 1 living, 2 dead; 94° 26 (Seige eam
1,500—1,760 fathoms, 1 living (S. Afr. Mus. Ag709, Ag88qQ; coll. F. H. Talbot,
Aug. and Dec. 1959).
Distribution. Arctic and North Atlantic, Mediterranean, Canaries, Azores;
Gulf of Mexico; 20-2,435 fathoms (Jeffreys).
Verticordiidae
Halicardia flexuosa (Verrill & Smith)
Halicordia [sic] flexuosa (Verrill & Smith), Tomlin, 1937, p. 23, fig. 1 (references).
33° 50' S., 17° 21 E., 600 fathoms, 1 living (S. Afr. Mus. Agyo7 hem
Talbot coll.).
The Pieter Faure took 2 dead specimens off Cape Point in 460-650 fathoms.
Cuspidariidae
Cuspidaria spp.
Until better material is obtained it is not advisable to attach definite
names to these specimens.
39° 60’ §., 17° 21 E., 600 fachoms, 1 living (S: Afr. Mus Ag7mo yi gee
Talbot coll.).
13 X 7°5 X 6 mm. Similar in shape to the Atlantic claviculata Dall and
congenita Smith, but has no ‘clavicle’; and to the South Australian meridionalis
Smith.
DEEP SEA MOLLUSCA FROM WEST OF CAPE POINT, SOUTH AFRICA 451
34° 37’ S., 17° 03’ E., 1,580—-1,620 fathoms, 2 right valves (one of them
broken); 34° 36" S., 17° 00’ E., 1,500-1,760 fathoms, one broken left valve
(S. Afr. Mus. Ag787 and Ag878, F. H. Talbot coll.).
The complete right valve is 40 X 25 mm., the broken left valve is at least
30 mm. alt. Compare: maxima Dautzenberg & Fischer from the Azores, 1,850
metres. Surface smooth, growth-lines only. Posterior lateral tooth in right
valve forms a thickened, sausage-like ridge; the ligament pit is scarcely visible
as it lies in a plane almost perpendicular to the sagittal plane of the valve.
SUMMARY
A collection of deep-sea mollusca from west of Cape Point, South Africa,
in depths between 600 and 1,880 fathoms is described. The collection comprises
approximately 590 specimens of approximately 78 species including 25 new
species.
ACKNOWLEDGMENTS
This paper is part of my research work on South African Marine Mollusca
carried out with the aid of a research grant from the South African Council
for Scientific and Industrial Research, to whom thanks and acknowledgment
are herewith made.
The deep-trawling was done by courtesy of the Director, Division of Sea
Fisheries, Cape Town, to whom we are very grateful.
The Trustees of the South African Museum are grateful to the Council
for Scientific and Industrial Research for the award of a grant for the publica-
tion of this paper.
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452 ANNALS OF THE SOUTH AFRICAN MUSEUM
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DAUTZENBERG, P. 1927. Mollusques provenant des campagnes scientifiques du Prince Albert ter
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1-401. 9 pls.
JEFFREYS, J. G. 1879. On the Mollusca procured during the ‘Lightning’ and ‘Porcupine’ Expedi-
tions, 1868—70. Part 2. Proc. zool. Soc. Lond., vol. pp. 553-588. pls. 45, 46.
Locarp, A. 1897. Mollusques Testaces. I. Expéditions scientifiques du Travailleur et du Talisman
pendant les années 1880-1883. Paris. pp. vi, 516. 22 pls.
MELvILL, J. C., & STANDEN, R. 1907 (issued separately). The Marine Mollusca of the Scottish
National Antarctic Expedition. Trans. Roy. Soc. Edinb. 46, 1908, 119-157. 1 pl.
PELSENEER, P. 1888. Report on the Anatomy of the Deep-Sea Mollusca collected by H.M.S.
Challenger in the years 1873-76. Challenger Rep. 27, 1-42, pls. 1-4.
PowE LL, A. W. B. 1951. Antarctic and Subantarctic Mollusca: Pelecypoda and Gastropoda.
Discovery Rep. 26, 47-196. pls. 5-10, text-figs. A-N.
Sars, G. O. 1878. Mollusca Regionis Arcticae Norvegiae. pp. 1-466. 32 pls. Christiana.
ScHEPMAN, M. M. 1908. The Prosobranchia of the Siboga Expedition. Part I. Rhipidoglossa and
Docoglossa. Siboga Exp. 49, 1-107. pls. 1-9.
ScHEPMAN, M. M. to11. Idem. Part IV. Rachiglossa. Siboga Exp. 49, 247-363. pls. 18-24.
SmitH, E. A. 1885. Report on the Lamellibranchiata collected by H.M.S. Challenger during the
years 1873-76. Rep. Sci. Res. Challenger 13, 1-341. 25 pls.
SmiTH, E. A. 1895. Report upon the Mollusca dredged in the Bay of Bengal and the Arabian
Sea during 1893-94. Ann. Mag. Nat. Hist. (6) 16, 1-19. 2 pls.
SmiTH, E. A. 1899. On Mollusca from the Bay of Bengal and the Arabian Sea. Ann. Mag. Nat.
Hist. (7) 4) 237-251.
SmitH, E. A. 1897-1909. Illustrations of the Zoology of the R.I.M.S. Investigator. Mollusca. pls. 1-23.
Calcutta.
SoutH AFRicAN Museum. 1961. Report for the year ended 31 March 1960. Cape Town.
Sow_ErRsy, G. B. 1903. Mollusca of South Africa. Mar. Invest. S. Afr. 2, 213-232. pls. 3-5.
THIELE, J. 1903. Die beschalten Gastropoden der deutschen Tiefsee-Expedition 1898-1899.
B. Anatomisch-systematischen Untersuchungen einiger Gastropoden. Wiss. Ergebn. Valdivia
7, 148-180. pls. 6-9 (1-4).
THIELE J. 1925. Gastropoden der Deutschen Tiefsee-Expedition. II Teil. Wiss. Ergebn. Valdivia
17, 37-382. Text-figs., pls. 13-46 (1-34).
THIELE, J. 1929. Handbuch der systematischen Weichtierkunde. 1. Part I. Loricata: Prosobranchia.
i-vi, 1-376. Text-figs. Jena: Fischer.
THIELE, J., & JAECKEL, S. 1931. Muscheln der Deutschen Tiefsee-Expedition. Wiss. Ergebn.
‘Valdivia’ 21, 161-268. pls. 1-5 (6-10).
Tomtin, J. R. LE B. 1927. Reports on the marine Mollusca in the collections of the South African
Museum. II. Abyssochrysidae, Odcorythidae. Ann. S. Afr. Mus. 25, 77-83. Text-figs.
TomuLin, J. R. LE B. 1931. Idem. V. Scaphopoda. Ann. S. Afr. Mus. 29, 337-340.
Tomuin, J. R. LE B. 1937. Idem. X. Verticordiidae. Ann. S. Afr. Mus. 32, 23, 24. 1 text-fig.
VERRILL, A. E. 1885. Third Catalogue of Mollusca recently added to the fauna of the New Eng-
land coast and the adjacent parts of the Atlantic, consisting mostly of deep-sea species,
with notes on others previously recorded. Trans. Connect. Ac. Arts. Sci. 6, 396-452. pls. 42-44.
Watson, R. B. 1879. Mollusca of H.M.S. Challenger Expedition. 2. Solenoconchia. 3. Trochidae.
Journ. Linn. Soc. Lond. 14, 586-606.
Watson, R. B. 1882. Idem. Part XII. Journ. Lin. Soc. Lond. 16, 324-343.
Watson, R. B. 1886. Report on the Scaphopoda and Gasteropoda collected by H.M.S. Challenger
during the years 1873-1876. Challenger Rep. 15, i-v, 1-756. pls. 50 + 3.
Woop-Mason, J., & Atcock, A. 1891. Natural History notes from H.M. Indian Marine Survey
Steamer. Mollusca. Ann. Mag. Nat. Hist. 8, 443-448.
[CONTINUED FROM INSIDE FRONT COVER
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrdége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Hartman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
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Eulalia (Steggoa) capensis Schmarda
Eulalis capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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PINNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XLVI
PART XVIII
SOME ECNOMINAE FROM THE TRANSVAAL AND SOUTH WEST
AFRICA
(TRICHOPTERA: PSYCHOMYIDAE)
By
Keo: Es Sscorr
National Institute for Water Research
South African Council for Scientific and Industrial Research
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[ CONTINUED ON INSIDE BACK COVER
SOME ECNOMINAE FROM THE TRANSVAAL AND SOUTH WEST
AFRICA
(TRICHOPTERA: PSYCHOMYIDAE)
By
K. M. F. Scotrr
National Institute for Water Research
South African Council for Scientific and Industrial Research
[Accepted November 1962]
(With 5 plates and 1 map)
CONTENTS
PAGE
Introduction : : ; 5 AER
Descriptions of species . : 7 9455
Summary . ; : : 2 Ab7,
References . ; ; : - 467
INTRODUCTION
During the past few years extensive collections of aquatic insects have been
made in the Transvaal during river surveys carried out by Dr. A. D. Harrison*
ef al. (1960 and in press), Dr. B. R. Allanson} (1961), Mr. F. M. Chutter (in
press), Mr. J. D. Agnew and other members of the staff of the National Institute
for Water Research. The parts of the rivers concerned are situated in the
Highveld region of the Transvaal. A small collection was also made in the
Transvaal Lowveld by Dr. G. H. Frank of the Bilharzia Research Unit of the
C.S.I.R. The caddis collected during these surveys were sent to the author
for identification, and the present paper is the first of a series based on this
material.
Grateful thanks are due to Dr. A. D. Harrison, Dr. B. R. Allanson,
Dr. G. H. Frank and Messrs. F. M. Chutter and J. D. Agnew for permission
to study and describe their collections of caddis, and for the trouble they have
taken in the provision of extra material, maps and information; also to the
South African Museum for the loan of type material, to Mr. D. E. Kimmins of
the British Museum (Natural History) for his very kind assistance, and to
Professor J. H. Day of the Zoology Department of the University of Cape
Town in whose Department the work was carried out.
*Then with the National Institute for Water Research, now Senior Research Fellow at the
University College of Rhodesia and Nyasaland.
+Then Transvaal River Research Fellow, now Senior Hydrobiologist in the National
Institute for Water Research.
453
Ann. S. Afr. Mus. 46 (18), 1963, 453-468, 5 pls., 1 map.
454 ANNALS OF THE SOUTH AFRICAN MUSEUM
In the present paper an account is given of the Ecnomus species found in
the Transvaal collections. These include one new species, E. kimminsi sp. n.,
fresh material of a little-known species, EF. oppidanus Barnard, and records of
several other species. Some of the species have been correlated with their
larvae, descriptions of which are given. Descriptions of females are also given
where possible, because the female genitalia show well-marked differences
which render identification from drawings and descriptions feasible. A map is
given, on which the localities mentioned in this paper can be approximately
located.
: Z
(SKE TCE MAP. Ur
24° ys
24 OF j-
TRANSVAAL ~~
Seyi
HART E BE ESTPOORT- prin ZAPRETORIA
DAM
Zp) OHANNE SBURG
LESLIE y
@®LOTHAIR
\e=-) = ERMELO
ROODE BAN a i
&,
ORANGE FREE STAT
100 MILES
South African Ecnomus larvae fall into two groups, one plain yellow, the
other patterned with brown, and it is difficult to distinguish the species from
others in the same group as they are very similar. The species considered in this
paper all fall into the patterned group, in which the head pattern (clypeus and
genae particularly) has been found most useful for identification purposes,
since patterns on the thoracic nota vary considerably in intensity with age and
other factors, being darker in later instars than in earlier, and also darker and
clearer in entire larvae than in larval sclerites taken from pupal cases. The
mandibles are also helpful in identification; they are usually much alike in
shape but show differences in the development of internal spines. ‘Two types of
yellow larva also occur in the area considered, but have not been described as
they have not yet been correlated; this also applies to two of the patterned
species.
ECNOMINAE FROM TRANSVAAL AND SOUTH WEST AFRICA 455
DESCRIPTIONS OF SPECIES
Ecnomus oppidanus Barnard
(Plate XVI, A—-H; plate XVII, A—c)
One of the new species described by Dr. K. H. Barnard in 1934 was
Ecnomus oppidanus. The material then available consisted of two very old and
mildewed specimens in the South African Museum collection, labelled Cape
Town 1885, and Dr. Barnard suggested that their place of origin be accepted
with caution until more specimens were discovered. None appears to have
been found, however, until the collections of caddis from the Transvaal were
studied, when, among those from the Vaal River area and the Lowveld, many
specimens were found which apparently belonged to Dr. Barnard’s species.
Through the kindness of Dr. A. J. Hesse of the South African Museum,
the author was able to compare these new imagos with Barnard’s type material,
and found that these males from the Transvaal do correspond very closely to
Barnard’s specimens. They show, however, a range of variation from the type in
minor details, as indicated in the drawings and descriptions. Ecnomus oppidanus
Barnard may therefore be regarded either as a somewhat variable species, or
as a species embracing a number of subspecies or varieties, and whose habitat
includes the Transvaal. As the original description was not very full, a new
description and drawings of the type material are given here for comparison
with those of the Transvaal specimens.
Imagos: Barnard 1934, p. 378, fig. 45, a-d.
Locality “Cape T. 1-1-85’. 2 3.
Neither of the two specimens in the South African Museum was designated
as type by Dr. Barnard, so the better one has been selected by the author and
labelled holotype g. Both are pinned specimens, old, fragile and in poor
condition; the paratype lacks a head and its anterior portion is enveloped in
cobweb. In both cases balsam preparations have been made of the genitalia
after clearing and drawing them, and in the case of the paratype one forewing
has also been mounted; both specimens lack hind wings.
Tibial spurs 3, 4, 43; length of forewing 5-0 mm.; general colouring yellow-
ish, forewings yellowish chequered with slightly darker markings. Wings as
figured by Barnard (1934, fig. 45 a).
Genitalia of holotype 3 (pl. XVI, A—F): Lobes of tenth segment stout, shorter
than the claspers, bluntly rounded, slightly upturned and dorsally keeled;
internal apical teeth large, numerous, yellowish, some projecting apically.
Between the lobes the median part of the tenth segment is membranous and
difficult to distinguish, apparently forming a raised area, posterior to which
are paired transparent projections with a depressed emargination between
them (according to Nielsen—1957—these are the bilobed lips which lie just
below the anus). Beneath these projections lie the paired internal processes of
the tenth segment (‘spiniform processes’; ‘paraprocts’ of Nielsen), oval in
456 ANNALS OF THE SOUTH AFRICAN MUSEUM
lateral view and armed with stout spines which curve upwards and outwards
round them; the inner and lower parts of the processes are pubescent, and each
ends posteriorly in a short apical projection tipped with three bristles. The
spines on the processes show slight differences in number and position between
holotype and paratype and between left and right sides (pl. XVI, B, c, c). Ninth
tergite strongly rounded dorso-laterally; side-pieces deeply incised; sternite
long, slightly wider than tergite. Claspers almost oblong in lateral view, bluntly
triangular and turned slightly outwards in dorsal or ventral view, the dorso-
lateral margins inturned and bearing long, recurved setae. Aedeagus (phallus of
Nielsen) with narrow apodeme, then widely expanded, narrowing again to
form a trough filled by membranous folds; above this lies a median sclerotized
process, bifid for much of its length and ending apically in a pair of heavily
sclerotized tips. The aedeagus then tapers to a long, slender, downcurved point.
This bifid sclerotized process presumably represents a pair of spines or
titillators, as is indicated by its origin from the dorsal side of the aedeagus.
This must be remembered when using Mr. Kimmins’s key to the genus Ecnomus
(Kimmins, 1957), as it may be mistaken for an upper part of the aedeagus, as
was suggested in Barnard’s original description, which reads ‘penis divided into
an upper portion which is apically bifid, and a lower portion which ends in a fine
point; no curved titillators’ (Barnard, 1934, p. 378). The bifid process corre-
sponds to the ‘upper beak’ of Nielsen (1957, p. 60), which in E. tenellus is
undivided.
New material (in spirit) :
Transvaal (Highveld) males (numbers cited throughout the paper are catalogue
numbers from the collections of the National Institute for Water Research):
Olifantsvlei outlet, 1 g (OLF 73 B, A.D.H., 17/1/55); Sandspruit near Volks-
rust, 1 ¢ (VAL 456 G, F.M.C., 20/11/58); headwater stream of Vaal River
between Breyton and Lake Chrissie, 1 g (VAL 516 C, F.M.C., 15/1/59); Vaal
River where crossed by Ermelo-Lothair road, 1 ¢ (VAL 851 B, F.M.C.,
8/11/59); Lake Chrissie, 4 3g (VAL 906 X (1-4), F.M.C., 21/1/60); Waterval
River (tributary of the Vaal), below confluence of streams from Leslie and
Trichardt, 1 g (VAL 1253 A, F.M.C., 6/12/60).
Other Highveld material: 1 9 (VAL 988 P) was taken at Roodebank on the
Waterval River, and larvae from the same place and also from Lake Chrissie
and Olifantsvlei.
These males closely resemble the type material described above and appear
to be the same species (see pl. XVI, j—M, of the male from Sandspruit). The
flies have prettily mottled brownish-yellow forewings (length 5:0 mm.); palps
clearly annulate, antennae faintly so. The Lake Chrissie specimens have very
widely expanded genitalia which show some of the structures more clearly, so
drawings of one of them have been included (pl. XVII, a, 8B). These Lake’
Chrissie males have slightly differently proportioned genitalia and the claspers
are turned inwards instead of outwards. It is possible to distinguish the median
ECNOMINAE FROM TRANSVAAL AND SOUTH WEST AFRICA 457
part of X more clearly; it consists of a raised, transverse portion fused to IX,
below which is a pair of thin, diagonally placed plates, projecting visibly in both
dorsal and lateral view. These plates appear to have hinge-like sclerotized
thickenings on their outer corners, partly dividing each into two portions and
enabling them to be tucked downwards. Beneath them lie the internal pro-
cesses, which in expanded specimens are definitely upcurved; this may also
cause difficulty in the use of Kimmins’s key to Ecnomus males (Kimmins, 1957).
The difficulties can be resolved by amending couplets 12 and 193 of the key as
follows:
12. Internal processes of tenth segment plain, upcurved, tipped with about 3 setae; titillators
expanded apically, toothed, and with a basal branch.
E. aequatorialis Marlier
— Internal processes with teeth or spines; titillators without basal branch. .. ae (Dg
13. Internal processes slender, terminating in 3 teeth; titillators in form of sharp spines arising
dorso-laterally from the aedeagus.
E. ulmeri Mosely
— Internal processes oval, spiny, basally pubescent, apex truncate and tipped with about three
long setae; titillators broad with bluntly pointed tips, arising from a wide common base
dorsal to the aedeagus.
E. oppidanus Barnard
The paired median processes of X are reminiscent of those figured for
E. tridens (Marlier, 1958, fig. 5). EL. tridens appears to be related to the natalensis
group of Ecnomus with spurs, 2, 4, 4, however, and could not be confused with
E. oppidanus. In any case, as the processes represent flaps beneath the anus
(Nielsen, 1957), it is probable that they occur in many species of Ecnomus.
Transvaal (Lowveld) males (pl. XVII, c, D): A number of specimens collected by
Dr. G. H. Frank in the environs of Nelspruit also appear to belong to Barnard’s
species. The data concerning them can be summarized as follows:
Nelspruit district, taken from January to March 1957, 35 dd and 48 99.
All but one of the collections made were small, and in them FE. oppidanus was
the only species of Ecnomus collected. The large batch (LOW 179 B) included,
however, 14 gd and 5 99 of E, natalensis Ulmer in addition to 22 gg and 33 99
of E. oppidanus; also 1 9 of a new species of Ecnomus from Komatipoort (which
will be described when more material is available). The E. natalensis 92 were
recognizable from the drawings given by Barnard (1934, fig. 45, h, 7), and
are the same as two @ paratypes of EF. natalensis in the British Museum (Natural
History); personal communication from Mr. Kimmins, who very kindly
compared them with Ulmer’s paratypes there. Besides the differences in the
genitalia, the oppidanus 9° can be distinguished from the other two species by
the tibial spurs (3, 4, 4 in oppidanus and 2, 4, 4 in natalensis and the Komatipoort
species).
The above-mentioned specimens were all collected from a small ground dam
(Friedenheim Farm dam) on the Nelspruit-White River road, about 6 miles
from Nelspruit. The dam is fed by a tributary of the Crocodile River, and most
4.58 ANNALS OF THE SOUTH AFRICAN MUSEUM
of the specimens were collected either in a floating trap designed by Dr. Frank
or in a Brundin Cone trap, set either over the shallow fringe of the dam or over
deeper water (about 3 feet in depth); bottom flocculent mud. A few imagos
were also taken during random sampling with a hand-net. Recently Dr. Frank
collected some Ecnomus larvae from a shady pool feeding the dam; conditions
had changed in the dam itself which was silting up. Some of the adult oppzdanus
(LOW 159 A) were taken at the same shady pool in 1957.
In addition to the above specimens, a male and three female EF. oppidanus
were taken from the upper Krokodilpoort dam in the Kaapmuiden ‘district
(LOW 30 D, 25/10/55), and many Ecnomus larvae were collected from the
Rietspruit, a small stream with grassy banks, clear, slow flow and muddy
bottom, situated on the Krokodilpoort range about 10 miles south of Nelspruit.
Both dam and stream connect with tributaries of the Crocodile River. All the
larvae except one are similar to those collected in the shady pool, the exception
being a larva of the new species described in this paper (E. kimminsz sp. n.).
It cannot be definitely stated that the larvae collected are those of EL. oppidanus,
as the larvae of natalensis and the Komatipoort species are also unknown, and
no mature ¢ pupae were found. Identification is, however, reasonably certain
as similar larvae were also collected in Olifantsvlei, an acid vle1 where the pH
range was from 4:2 to 5:4 at the stations and seasons when the Ecnomus larvae
were collected (Harrison ef al., 1960); Olifantsvlei lies on the Klip River some
14. miles SW. of Johannesburg. Very few caddis were able to tolerate the acid
conditions, and oppidanus 33 were the only adult Ecnomus found there. Similar
larvae were also found in Lake Chrissie, a shallow eutrophic lake, where
oppidanus was again the only species of Ecnomus found. A description of the larva
is therefore given below. |
The Lowveld males are on the whole slightly smaller than the rest
(w. 4°5 mm.), and have genitalia very similar to the Lake Chrissie specimens,
with short, rather humped, upper lobes. The small inner lobes are clear and
easily visible; the spiniform processes are somewhat sinuous, with fewer teeth,
and are hairier and more granular in appearance underneath (pl. XVII, D) ; the
narrowed tips of the titillators are longer than in the type, and the lobes of
X are frequently tucked inwards, with the central processes of X turned down,
and sometimes with the ends of the spiniform processes showing (pl. XVII, c).
The yellow teeth arming the upper lobes of X vary in number from about
18 in the Lowveld and Olifantsvlei specimens, to about 28 in many Highveld
specimens. The general appearance of the genitalia is however always much
the same, and, while it might be possible to separate the material into two
varieties or subspecies, it does not seem necessary, at least until the species is
better known and more specimens have been collected.
E. oppidanus 99 (pl. XVII, £, F, G):
Seventh segment normal; eighth and ninth segments with tergites normal,
the ninth thickened posteriorly and fringed with setae. The eighth sternite
ECNOMINAE FROM TRANSVAAL AND SOUTH WEST AFRICA 459
forms a subgenital plate almost completely divided into two large leaf-like
lobes connected proximally by a bridge; the lobes are setose with about three
long marginal bristles and are thickened along their inner margins. Dorsal
to the subgenital plate lie thinner flaps, possibly representing IX; proximal
to these, and connecting with the subgenital plate, is a sclerotized bridge, seen
as triangular in ventral view, in which the genital opening lies. The thin flaps
are much folded posteriorly, curling round and connecting apically by way ofa
thickened collar with a pair of rounded, seta-studded projections, each tipped
with three soft, finger-like papillae. The central papilla on each side stands on a
small boss and has a narrowed tip. These projections and papillae possibly
represent the tenth sternite; between them on the dorsal side the tenth tergite
appears to be represented by a pair of small, rounded humps fringed with
strong setae.
Females with their genitalia much expanded present a somewhat different
appearance, so a figure of one has been included for comparison (pl. XVII, F).
The wide separation of the subgenital lobes is reminiscent of the female
designated E. kunenensis by Barnard (1934, p. 380 and fig. 45,7;see also pl. XIX, x,
of this paper). In E£. natalensis Ulmer the subgenital plate is deeply cleft, with
divergent lobes separated by a V-shaped incision (Barnard, 1934, p. 379 and
figs. 45, h, 2); a drawing of one is given here for comparison (pl. XX, pb, d).
Probable larva of E. oppidanus (pl. XIX, B—p): larvae collected from the
Nelspruit district (2/8/55, 7/3/62, G.H.F.) ; from Olifantsvlei (23/6/54, 21/7/54,
18/8/54, 6/10/55, A.D.H.); from Lake Chrissie (16/9/58, F.M.C.). (Ecnomus
larvae have also been collected from many other localities all over South
Africa, but many are too small to identify further, and many have not yet
been identified to species as the differences between them are only now being
recognized.)
Larva of the usual Ecnomus type described by Ulmer (1957) ; campodeiform,
8-10 mm. in length, all three thoracic tergites sclerotized. Head yellowish,
patterned with brown (pl. XIX, B); the patterning on the clypeus consists of
lateral strips on the anterior part only, the rest being plain yellow; the patterning
on the genae, while darkest along the posterior part of the clypeus, continues
back to the occipital foramen; there is also a small brown mark alongside the
clypeus above each eye. Anteclypeus divided into four parts; gular sclerite
short, widely triangular, with curved sides. Antennae minute, close to anterior
margin of head. Eyes large, black, placed well forward in white areas beneath
lens-like thickenings of the cuticle. Labrum (pl. XIX, c) rounded, yellowish, with
median indentation flanked by a pair of slender bristles; six pairs of dorsal
setae. Mandibles (pl. XIX, p) strong, dark, unequal, the right mandible with
several blunt teeth and a number of short, stout, internal spines, the left
mandible larger, with about five blunt teeth and a number of long, slender,
internal spines. Maxillae and labium whitish, prominent; maxillary palp well
developed, longer than lobe; labium broad basally, narrowing to a slender
shaft, labial palps present, long, two-jointed.
460 ANNALS OF THE SOUTH AFRICAN MUSEUM
Thoracic tergites (pl. XIX, B) sclerotized, with posterior margins dark;
pronotum yellowish, unpatterned, with long posterolateral points which curl
round ventrally ; meso- and metanota yellowish with more or less clear brownish
pattern and diagonal brown stripes in the anterolateral corners; lateral
margins of mesonotum also darkened.
Legs strong, subequal; claws long, each with basal bristle. Anal appendages
long, slender; anal claws large, strongly curved to form a right angle, a comb
of small teeth along the central third of the inner edge, no dorsal hooks.
Abdomen somewhat flattened, no lateral gills, no lateral line, a few hairs
along each side (there is a broad lateral] tract of fuzzy hairs there in young
larvae) ; five anal gills present.
Ecnomus kimminsi sp. n.
(Plate XVIII, a—m; plate XIX, a)
Imagos (in spirit): Vaal River shortly above confluence with Klein Vaal,
holotype 3 (VAL 860 F (2), 9/11/59, F.M.C.); Zwartkoppies on the Blaauw-
bank River, an unpolluted tributary of the Crocodile River, near Krugersdorp,
2 d paratypes (ALL 13 G (1) and (2), 7/4/57, B.R.A.) (this is not the Lowveld
Crocodile, but flows northwards through the Hartebeestpoort Dam to join the
Limpopo). Klein Vaal near confluence with Vaal River, 1 g pupa with pupal
case containing larval sclerites (VAL 1022 L, 9/2/60, F.M.C.); Klein Vaal
near confluence with Vaal River, 2 larvae (VAL 1022 M, 9/2/60, F.M.C.).
Larvae were also found at several stations on the Vaal River, and higher up
the Klein Vaal.
The above specimens all belong to the same species (larvae correlated via
larval sclerites in pupal case together with mature ¢ pupa), which does not
appear to be any of the species hitherto described from Africa. It belongs to the
natalensis group of Ecnomus, and is nearest to E. ugandanus Kimmins. None of
the females found could definitely be assigned to this species.
Tibial spurs 2, 4, 4; general colouring of head and thorax chestnut brown;
antennae and legs yellowish, anterior face of forelegs browner, fore-tarsi and
palps obscurely annulate. Forewings plain golden-brown, traces of irrorations
present but wings largely denuded, membrane brownish, a white line along the
anastomosis and the anterior edge of the median and thyridial cells. Wing
length 5:0 mm., wings of the usual Ecnomus type (pl. XIX, a). Abdominal
tergites purplish brown.
6 genitalia (pl. XVIII, a—F): lobes of tenth segment long, narrowly oval in
dorsal view, broadly so in lateral view, a little shorter than the claspers;
internal apical teeth long, slender, dark, forming a band along the apical
margin; proximal to them the inner surface of the lobe is set with long setae
arising from tall papillae (as is the case in a number of Ecnomus species). The
median part of the tenth segment forms a raised, semicircular membranous
area, partly bounded anteriorly by narrow sclerotized strips. The paired
ECNOMINAE FROM TRANSVAAL AND SOUTH WEST AFRICA 401
internal processes of the tenth segment are almost as long as the lobes, stout,
slightly smuous and rodlike, set with small spines and tipped with about three
bristles; the processes slope downwards for three-quarters of their length, then
turn upwards near the apices. The ninth tergite has a T-shaped median suture-
like sclerotization, the ninth sternite is long, with a median point. Claspers of the
same type as in the natalensis group of species, with a curved apical finger which
in dorsal view has an inturned beak-like point; the incision beneath it is
rounded and bounded ventrally by a blunt incurved process; base of clasper
about four-fifths of the depth of the part to which it is attached. In the Zwart-
koppies specimens the apex of the clasper is more rounded than in the type.
In lateral view the aedeagus has a triangular apodeme; posterior to this it is
constricted, then expanded again to form a bulb which has a ventral mem-
branous portion and is apically bifid to form upcurved, flattened plates with
rounded apices, slightly toothed on the dorsal edge. The paired titillators are
longer than the aedeagus, stout, spinelike, somewhat sinuous and terminally
upcurved.
Holotype g (VAL 860 F (2)) in spirit (genitalia cleared), in the South
African Museum; one paratype (ALL 13 G (1)) in the British Museum (Natural
History) ; rest of material in the National Institute for Water Research collec-
tion. I have much pleasure in naming the species after Mr. D. E. Kimmins.
Mr. Kimmins has informed me (personal communication) that the internal
processes of X in FE. ugandanus Kimmins are not entirely glabrous as they appear
in the figures, but that under a 4” objective they show a sparse clothing of
minute spines and a variable number of apical setae; the titillators are stouter
and. a little shorter than in kimminsi, and curved but not sinuous. E. kimminsi
therefore differs from EH. ugandanus in the more spiny internal processes, the
longer, sinuous titillators, and the shape of the apex of the aedeagus and of the
claspers. It runs down to the first part of couplet 6 on Kimmins’s key (1957,
p-. 262), and can be keyed out from that point as 6A as follows:
6A. Internal processes set with small spines; titillators slightly longer than aedeagus, sinuous;
apex of aedeagus in side view rounded, slightly toothed dorsally.
E. kimminsi sp. n.
— Internal processes glabrous or sparsely set with minute spinules; titillators slightly shorter
than aedeagus, upcurved but not sinuous; apex of aedeagus in side view triangularly
pointed. ue He ee a se seh oe oy: iA oie el
7. Apical finger of clasper elongate, the excision beneath it in side view narrow. Apex of
aedeagus in side view forming an acute spine (fig. 2, N); internal processes glabrous.
E. natalensis Ulmer
— Apical finger shorter, slightly more downcurved, the excision beneath it widely rounded.
Apex of aedeagus in side view forming a broad, acute triangle (fig. 2 U); internal processes
sparsely set with minute spinules. Bs 5 ric .. . ugandanus Kimmins
Larva of E. kimminsi sp. n. (VAL 1022 M, pl. XVIII, c—m):
Larva very similar to that of E. oppidanus. Length 9-11 mm. Head yellowish,
patterned with brown (pl. XVIII, G); the lateral brown patches on the clypeus
462 ANNALS OF THE SOUTH AFRICAN MUSEUM
join or almost join centrally, and there is a patterned brown band crossing the
middle of the head, interrupted by the plain yellow posterior part of the cly-
peus; back of head plain yellow. Anteclypeus, gular sclerite (pl. XVIII, m), eyes
and antennae as in oppidanus. Mandibles strong, dark, each with about 5 blunt
teeth and 2 dorsal bristles, the right mandible smaller than the left, with some
very short inner spines; the left without inner spines. Labrum brownish,
maxillae and labium whitish, very prominent, all very like those of oppidanus
(pl. XVIII, j, m).
Thoracic tergites all sclerotized (pl. X VIII, c), yellow with brown pattern,
posterior borders dark. Pronotum with a large triangular brown mark, meso-
and metanota with well-marked pattern and dark diagonal stripes.
Legs, abdomen and claws (pl. XVIII, H) as in oppidanus.
Ecnomus thomassett Mosely
(Plate XIX, E-x)
Ecnomus thomasseti Mosely 1932.
Imagos and larvae (in spirit): Standerton, 1 3, 1 2 (VAL 985, E 23/3/60),
caught on wing; Vaal River near Standerton, 1 ¢ (VAL 1251 E, 6/12/60);
Kafferspruit, tributary of Vaal River, near Ermelo, larvae (VAL 1107 L,
12/8/60), some of these larvae being bred out in the laboratory as follows:
2 66;.1 2 (VAL 1159 E, 12/10/60),,1 g¢ (VAL 1159 F, 12/10/60), 1g QV Ak
1160 G, 12/10/60). Crocodile River, shortly before entry into the Hartebeest-
poort Dam, 3 3d (ALL 1 a-c, 5/4/51), 3 dd, 1 2 (ALL 5, 10/3/57). The VAL
specimens were all collected by Mr. F. M. Chutter, the ALL specimens by
Dr. B. R. Allanson. The larvae collected from the Kafferspruit and reared in
the laboratory by Mr. Chutter made possible the correlation of the larva with
both male and female imagos. The males are all typical FE. thomasseti; the
female and larva are described below. Larvae were also collected at several
stations on the Vaal and Waterval rivers.
The 9, as will be seen from the drawing (pl. XIX, Kk), greatly resembles
Barnard’s ‘E. kunenensis 92 from South West Africa. Owing to Kimmins’s
description of part of Barnard’s S.W.A. material as E. barnard: (Kimmins, 1957),
it was of course possible that the ‘kunenensis’ 2 might in reality be the Q of
E. barnardi, or even of E. thomasseti, since Barnard did suggest (1934, p. 380)
that one of his ¢ Ecnomus might be E. thomasseti. The latter possibility was
supported by the resemblance of Barnard’s @ to the E. thomasseti 99 from the
Transvaal.
Dr. Hesse kindly made Barnard’s type material from $.W.A. available for
comparison; it consisted of 9 pinned specimens, whose genitalia had in most
cases been cleared and then mounted in gum arabic on card. The genitalia
were removed from the cards and studied in clove oil before mounting in
balsam. The specimens proved to be as follows (all localities given as Otjim-
ECNOMINAE FROM TRANSVAAL AND SOUTH WEST AFRICA 463
bumbe, Kunene River, except in the case of No. 6, where the locality was
Erikson’s Drift, Kunene River; date in each case March 1923):
(1) E. kunenensis Barnard, 1 g. W. 4:0 mm. Specimen rather broken up, but
parts present. This has been labelled holotype because it is the only ¢ of
this species in the collection. A figure of it is given (p]. XX, A), to show the
narrowing of the internal processes of X. Otherwise it agrees with Kim-
mins’s drawings and description (Kimmins, 1957, p. 265 and fig. 2, x).
(2) E. thomasseti Mosely, 1 3. W. 3:5 mm. Also damaged, but genitalia typical.
(3) £. barnardi Kimmins, 1 g. W. 4:0 mm. Much damaged, only forewings
and head remaining apart from genitalia, which however agree exactly
with Kimmins’s drawings and description (1957, p. 265, and fig. 2 B).
(4) . barnard: Kimmins, 1 3. As (3); the hind-legs and 2 wings which were
loose have also been mounted in balsam.
(5) £. ‘kunenensis Barnard, 1 9. W. 4:0 mm.
(6) E. ‘kunenensis’ Barnard, 1 9. W. 4:0 mm. Genitalia of both (5) and (6)
as illustrated in plate XIX, x (which is actually an E. thomasseti 9).
(7) Ecnomus sp. A, 1 9. W. 4:5 mm. (pl. XX, B). Head and right forewing which
were loose have been mounted in balsam together with the abdomen.
(8) Ecnomus sp. A, 1 9. W. 4:5 mm. (as no. 7). A brief description of this species
is given elsewhere in this paper.
(9) Ecnomus sp. Specimen with wings spread, rather broken, genitalia missing.
The new material of E. thomasseti from the Transvaal has provided correla-
tion of the ‘kunenensis’ 9 with its larva and thence also with the ¢ of thomassetz,
besides which the other Transvaal 99 of this type were in each case caught with
63 of E. thomasseti only. Thus, since E. thomasseti was also present in Barnard’s
S.W.A. collection, these 99 may be regarded as almost undoubtedly being the
29 of E. thomassett Mosely. There is as yet no final proof of this, however, as
none appears to have been taken zn copula.
Description of 2 genitalia (pl. XIX, x):
Seventh segment, and eighth and ninth tergites, normal. Eighth sternite
forms a subgenital plate divided to form a pair of leaf-like lobes, each of which is
laterally rounded, apically bilobed, tipped with three long bristles and studded
with small setae; the lobes are widely separated by a squarish indentation
with a triangular median point, which is actually the bridge bearing the genital
opening, seen end-on. Dorsal to the subgenital plate are the flaps formed by the
ninth sternite; these have rugose bands near the midline in their proximal
portions, the rugose bands vary somewhat in extent in different specimens and
appear to connect basally with a pair of oblong ‘pockets’, visible through the
subgenital plate in a cleared specimen. The genitalia end apically in a pair of
soft, rounded projections, each bearing three papillae.
464. ANNALS OF THE SOUTH AFRICAN MUSEUM
In general appearance the 99° are reminiscent of the 99 of oppidanus, but
the subgenital plate has definitely bifid lobes, and the space between them is
squarer and less deep than in that species; the ninth sternite is less folded distally,
and the ‘pockets’ visible through the plates are quite different from the small
triangular slits of oppidanus. The two species are in fact distinguishable without
clearing, even without considering the tibial spur count (3, 4, 4 in oppidanus,
2, 4, 4. in thomassett). ee
Larva (pl. XIX, E, H):
Larva of the usual Ecnomus type, length 7-8 mm. Head yellowish, with
brown patterning as indicated in the figure (pl. XIX, E) ; note that the patterning
is less extensive than in kimminsi or oppidanus, the lateral bands on the clypeus
being separated by a wide yellow area, and the patterned patches on the genae
small, not extending as far as the sides of the head as seen in dorsal view.
Anteclypeus, eyes, labrum and gular sclerite as in kimminsi, mandibles very
similar to those of oppzdanus in shape, the right smaller than the left; the inner
spines on the right mandible are well marked, and two or three long spines arise
from the deeply indented inner face of the left mandible. Condyles small, as is
usual in Ecnomus larvae. Maxillae and labium of the same type as in other
species. Patterning of the thoracic nota is much fainter, the pronotum being
scarcely darkened, and the meso- and metanota having faint brownish marks
and pale brownish diagonal stripes; all have a dark posterior band. Legs and
anal appendages as in the other species. Abdomen rather strongly segmentally
constricted, lateral tracts of fuzzy hairs well developed.
Pupal case (pl. XIX, j):
The pupal case of FE. thomasseti has been figured as being typical of these
Ecnomus species; it is composed of comparatively large sand grains with a few
minute ones filling interstices, and is lined with thick, soft, semitransparent
felt-like material. There are sieve plates at each end, allowing the passage of
water currents, and the larval sclerites are found within the case, caught up in
the soft lining. Length of case about 8 mm. One case, where the substratum
had been of very fine sand, was composed entirely of minute sand grains, so
the larva can evidently make use of whatever size is available, preferring
however, to use larger sand grains when possible.
Ecnomus sp. A 2
(Plate XX, B)
Ecnomus kunenensis Barnard, 1934: 380 (partim, 9 paratypes in the South African Museum).
Otjimbumbe, Kunene River, South West Africa, March 1923, 2 99
(K. H. Barnard and R. F. Lawrence).
A brief description of the second species of 2 associated with Barnard’s
S.W.A. material is given here; it is easily distinguished from the other species
described in this paper by the genitalia.
ECNOMINAE FROM TRANSVAAL AND SOUTH WEST AFRICA 465
Specimens old and fragile, body colour reddish. brown with pale gold
setae; eyes black; antennae slightly ringed; legs yellowish, tibial spurs 2, 4, 4.
Wings pale gold, showing traces of a few darker flecks (the lack of patterning
may be due to fading), length of forewings 4-5 and 5:0 mm.
Genitalia (pl. XX, B):
The subgenital plate is completely divided into a pair of subtriangular
lobes, contiguous or slightly overlapping in the mid-ventral line; each bears 3-4
long, strong bristles and sparsely scattered small setae. Dorsal to the lobes are
the flaps of the ninth sternite, each bearing a crescent-shaped tract which at
first sight appears to be hairy; the ‘hairs’, however, seem actually to be small
sclerotized ridges. ‘These tracts are concealed by the subgenital plate but are
visible by transparency in cleared genitalia. The papillae etc. of the tenth
segment are much crushed and distorted in both specimens.
Ecnomus natalensis Ulmer
(Plate XX, pv, d)
Ecnomus natalensis Ulmer 1931.
Friedenheim Farm dam, on tributary of Crocodile River, 6 miles from
Nelspruit (LOW 179 B (2), LOW 179 B (4), 25/3/57, G.H.F.), 14 36,
Sees
od specimens typical of EF. natalensis. The ° of this species has been briefly
described already (Barnard, 1934, p. 378 and fig. 45, A). A figure of it is, how-
ever, given here (pl. XX, pb) for comparison with those of the other species
described. The genitalia differ considerably in appearance from those of the 9°
of oppidanus and kunenensis, showing more affinity with the 99 of species A and C
described in this paper. The lobes of the subgenital plate are separated by a
deep V-shaped incision, almost the full depth of the plate. Each lobe is traversed
by a longitudinal dent or furrow, and bears a number of long setae and three
strong marginal bristles. Between the lobes the fold bearing the genital opening
can be seen, and dorsal to it the thin flaps of IX. Behind the lobes of the sub-
genital plate, and only visible in a cleared specimen, these flaps are thrown into
raised, sclerotized folds which lie in the angles made by the dents on the sub-
genital plate. It is difficult to make out the structure of these folds; each appears
to have strong, ridged walls enclosing a pocket or space (pl. XX, d). The vaginal
apparatus is lightly sclerotized, and there is a sclerotized collar round each of
the projections that bear the usual three papillae.
Ecnomus ugandanus Kimmins var.
Ecnomus ugandanus Kimmins 1957.
Volksrust (VAL 541 G (2), 11/2/59, F.M.C.), 1 3. Mr. Kimmins kindly
identified this specimen as being a variety of his species.
466 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ecnomus sp., near E. complex Mosely
Ecnomus complex Mosely 1932.
Vaal River (VAL 899 D, 20/1/60, F.M.C.). Two ¢ pupae were found in
the Vaal River between Morgenzon and Amersfoort. The pupae have well-
developed 3 genitalia with straight, out-turned upper lobes like E. similis
Mosely (1932), but have the inner branches characteristic of E. complex. The
pupal cases contain larval sclerites with almost plain clypeus, genae with a
broad band of patterning, and thoracic nota faintly patterned, without dark
diagonal bars. Definite identification must await further material.
Ecnomus sp. B 9
(Plate XX, E-H)
Two 9 spurs 3, 4, 4 were bred out from larvae taken from the upper part
of the Klein Vaal River (VAL 1158 AK, larvae; VAL 1158 AL, 9; VAL 1158
AM, 9; all 20/7/60, F.M.C.). These 99 appear to have lateral slits on abdominal
segment V, rather like those of some Hydropsychidae though smaller and less
obvious.
Genitalia (pl. XX, E):
The subgenital plate is reminiscent of EF. natalensis, but the V-shaped
indentation between the lobes is short, and the lobes are broader and turned
farther outwards apically, with four long marginal bristles, and the ninth
sternite is soft, there being no heavily sclerotized folds as in natalensis. There is a
median point visible between the lobes, which appears to pertain to the vaginal
apparatus and to lie dorsal to the subgenital plate. The usual apical papillae
are present.
Larva (pl. XX, F-H):
Length 9-10 mm. Head capsule strongly patterned, particularly on the
genae; ‘muscle spots’ very strongly marked; pattern on clypeus paler than that
on the epicranial sclerites, consisting of pale greyish-brown lateral strips which
extend back onto the triangular posterior part of the clypeus. The strong
patterning on the epicranial sclerites is reminiscent of that in the larva of
oppidanus, but is even darker except on the clypeus, which is much paler and
differently shaped. The mandibles greatly resemble those of thomassett and
oppidanus, but are particularly large and strong (pl. XX, G, H); the spines in
the right mandible are well developed, but the left mandible appears to lack
spines entirely. Pronotum with faint brownish pattern, meso- and metanota with
definite brown and yellow pattern, but lacking darker diagonal bars (pl. XX, F).
Similar larvae were also collected below the creamery and sewage outflows
at Standerton on the Vaal River; also higher up the Vaal, and in a southern
tributary, the Wilge River.
ECNOMINAE FROM TRANSVAAL AND SOUTH WEST AFRICA 407
Ecnomus sp. G &
QBlateOCrc)
Three 99, spurs 2, 4, 4, were bred out from larvae collected from the
Waterval River at Roodebank, between Standerton and Leslie (VAL 1161 A,
©; VAL 1161 B, 9; VAL 1161 QG, 9; July 1960, F.M.C.). Four 99 of the same
species were collected in Amersfoort at light (VAL 977 C (2), 22/3/60, F.M.C.).
Genitalia (pl. XX, c):
Subgenital plate bluntly triangular, nearest to that of natalensis in shape,
but with the apical points blunt and slightly bifid, each with four long bristles.
The indentation between the lobes is also deeper and is basally squared; through
it the small, rounded apex of the vaginal apparatus is visible. Dorsal to the
subgenital plate a pair of sinuous, rugose tracts is visible by transparency in
cleared specimens; these appear hairy at first sight, somewhat resembling those
of Ecnomus sp. A. The subgenital plates of the two species are, however, quite
different in shape. The rugose tracts appear to be connected with the ninth
sternite and the vaginal apparatus, as in sp. A and natalensis 99; these species all
belong to the natalensis group of Ecnomus with spurs 2, 4, 4. The 99 of thomasseti,
- belonging to the same group, also have rugose tracts, though somewhat
different in appearance. .
Only larval sclerites are available in this species, so a description of the
larva will be postponed until entire specimens have been found. The sclerites
resemble those of thomasseti, but are rather more strongly patterned.
The Ecnomus larvae described in this paper show resemblances to the larva
of EL. relictus Vaillant (1953), and to one of the larvae described by Corbet
(1958, fig. 4, Cc). Several other patterned species of Ecnomus have been dis-
covered in South Africa (two of them in the Transvaal) ; they will be described
as they are correlated.
SUMMARY
An account is given of the Ecnomus species found in the Transvaal collec-
tions of the National Institute for Water Research. These include £. kimminsi
sp. n. and new material of E. oppidanus Barnard. Females and larvae of several
of the species are also described, and an account given of Barnard’s Ecnomus
material from South West Africa, now in the South African Museum.
REFERENCES
ALLANSON, B. R. 1961. Investigations into the ecology of polluted inland waters in the Transvaal.
Hydrobiologia 18, 1-76.
BARNARD, K. H. 1934. South African caddis-flies (Trichoptera). Trans. roy. Soc. S. Afr. 21,
291-394-
CuutTter, F. M. (in press). Hydrobiological Studies on the Vaal River in the Vereeniging area
(Part 1). Introduction, water chemistry and biological studies on the fauna of habitats
other than muddy bottom sediments. Hydrobiologia.
468 ANNALS OF THE SOUTH AFRICAN MUSEUM
Corset, P. S. 1958. Larvae of certain East African Trichoptera. Rev. Zool. Bot. afr. 58, 203-213.
Harrison, A. D. 1958. The effects of sulphuric acid pollution on the biology of streams in the
Transvaal, South Africa, Verh. int. Ver. Limnol. 13, 603-610.
Harrison, A. D., KELLER, P. & Dimovic, D. 1960. Ecological studies on Olifantsvlei, near
Johannesburg. Hydrobiologia 15, 89-134.
Harrison, A. D., KELLER, P., & LompBarp, W. A. (in press). Hydrobiological studies on the
Vaal River in the Vereeniging area (Part 2). The chemistry, bacteriology and inverte-
brates of the bottom muds. Hydrobiologia.
JacguEMarRT, S. 1957. Trichoptera des lacs Kivu et Edouard. Exploration Hydrobiologique des
Lacs Kivu, Edouard et Albert, 3 (2), 67-129.
Kimmins, D. E. 1957. Notes on the Psychomyidae (Trichoptera) from the African mainland
(south of the Mediterranean region), with particular reference to the genera Ecnomus and
Psychomyiellodes. Trans. R. ent. Soc. Lond. 109, 259-273.
LestacE, J. A. 1921. Trichoptera, Jn Rousseau, E. Les larves et nymphes aquatiques des insectes
d’ Europe. 15 343-964. Brussels: J. Lebégue & Cie.
MarutEr, G. 1943a. Trichoptera. Explor. Parc Nat. Albert Miss. Damas. 11, 3-34.
MarulErR, G. 1943b. Trichoptéres du Congo Belge. Rev. Zool. Bot. afr. 37, 64-88. -
MaruieEr, G. 1958. Trichoptéres du lac Tumba. Bull. (Ann.) Soc. ent. Belg. 94, 302-320.
Mosety, M. E. 1932. New exotic species of the genus Ecnomus (Trichoptera). Trans. ent. Soc.
Lond. 80, 1-17.
Mosety, M. E. 1935. New African Trichoptera. Ann. Mag. nat. Hist. (10), 15, 221-232.
NIELSEN, A. 1957. A comparative study of the genital segments and their appendages in male
Trichoptera, Biol. Skr. 8, (5), 1-159.
Umer, G. 1931. Trichopteren von Afrika. Disch. ent. £. 1931, 1-209.
Umer, G. 1957. Kocherfliegen (Trichopteren) von den Sunda-Inseln, III. Arch. Hydrobiol.
Suppl. 23, 109-477.
VAILLANT, F. 1953. Deux Trichoptéres nouveaux du Sahara central. Bull. Soc. zool. Fr. 78,
149-157:
Ann. 8. Afr. Mus., Vol. XLVI Plate XVI
O*l mm
Ecnomus oppidanus Barnard. Holotype ¢: A, lateral view of genitalia; B, internal process
of X (lateral) ; C, internal processes of X (dorsal) ; D, dorsal view of genitalia; E, ventral view of
genitalia; F, posterior view of genitalia. Paratype 3: G, internal processes of X; H, dorsal view
of aedeagus and attached spines. g from Sandspruit, Transvaal: J, genitalia, lateral view
(VAL 456 G); K, internal process of X, lateral view, left side; L, internal processes of X, dorsal
view; M, internal process of X, right side.
Ann. S. Afr. Mus., Vol. XLVI Plate XVII
t
t
+
t
k
t
’
#
i
z
Or} mm XK
Ecnomus oppidanus Barnard. 3 from Lake Chrissie (VAL go06 X (1)): A, lateral view of
genitalia; B, dorsal view of genitalia (left internal process omitted). 3 from Friedenheim Farm
dam (LOW 157 A): G, dorsal view of genitalia, outer lobes of X tucked in. f from Friedenheim
Farm dam (LOW 168 A): D, internal processes of X, lateral view. Probable 29 of E. oppidanus
Barnard: 2° from Lake Chrissie (LOW 179 B (3)): E, ventral view of contracted genitalia;
F, ventral view of expanded genitalia; G, lateral view of expanded genitalia.
Ann. S. Afr. Mus., Vol. XLVI Plate XVIII
aS
/
ee NS
Ecnomus kimminsi sp. n. Hoiotype g from Vaal River (VAL 860 F (2)): A, lateral view
of genitalia; B, dorsal view of genitalia (with setate papilla from within lobe of X further
enlarged) ; C, ventral view of genitalia; D, dorsal view of aedeagus; E, lateral view of internal
process of X; F, lateral view of aedeagus. Larva from Klein Vaal (VAL 1022 M): G, dorsal
view of thoracic tergites and head; H, anal claw (twice magnification of G); J, dorsal view of
labrum; K, L, ventral view of right and left mandibles; M, ventral view of maxillae and labium
(tips of maxillary palp and maxilla further enlarged), with gular sclerite etc.
fe
oe
oo
= -
Ann. S. Afr. Mus., Vol. XLVI Plate XIX
Stee
Ecnomus kimminsi sp. n.: A, fore- and hind-wings of holotype ¢g. E. oppidanus Barnard,
probable larva: B, head and thoracic nota, dorsal view; C, labrum; D, right and left mandibles,
ventral view. E. thomasseti Mosely, larva: E, head and thoracic nota, dorsal view; F, labrum;
G, right and left mandibles, ventral view; H, process of trochantin; J, pupal case, with sieve
plate further enlarged; K, ventral view of 2 genitalia, with tips of papillae further enlarged.
(Note: Scale of wings and pupal case indicated, otherwise scales used same as in the other plates.)
Ann. S. Afr. Mus., Vol. XLVI Plate XX
|-O mm :
Ecnomus kunenensis Barnard: A, holotype g, lateral view. Ecnomus sp. A, 2: B, ventral
view of genitalia. Ecnomus sp. C, 2: C, ventral view of genitalia. Ecnomus natalensis Ulmer, 9:
D, ventral view of genitalia; d, sclerotized part of inner plate. Ecnomus sp. B, 2: E, ventral view
of genitalia; F, dorsal view of head and thoracic nota of larva; G, H, ventral view of right and
left mandibles of larva.
[ CONTINUED FROM INSIDE FRONT COVER
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Bettrdge zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
HARTMAN, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
_ Harman, O. 1948b. The marine annelids erected by Kinberg. Ark. ool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sct. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 41, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
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arranged in chronological order. . . .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalis capensis Schmarda 1861, p. 86, pl. 20, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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oY ANNALS
OF THE
pOUTH AFRICAN MUSEUM
VOLUME XLVI
PART XIX
SOME NEW CADDIS FLIES (TRICHOPTERA) FROM THE WESTERN
CAPE PROVINCE
IV: SOME HYDROPTILIDAE
By
Ke Mer. Scorr
National Institute for Water Research
South African Council for Scientific and Industrial Research
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[ CONTINUED ON INSIDE BACK COVER
SOME NEW CADDIS FLIES (TRICHOPTERA) FROM THE WESTERN
CAPE PROVINCE
IV: SOME HYDROPTILIDAE
By
KM. Sconr
National Institute for Water Research
South African Council for Scientific and Industrial Research
[Accepted November 1962]
(With 3 plates)
CONTENTS
PAGE
Introduction : ; : . 469
Descriptions of species . : PS VAGD)
Summary . : , : "473
References . ‘ : : ae 47o
INTRODUCTION
This is the fourth paper in the present series; in it a new species of Ortho-
trichia from the Great Berg River is described, together with its larva and larval
and pupal cases. The other Hydroptilidae collected along the same river are
commented upon, and the larva of Oxyethira velocipes (Barnard) described.
Methods of work have been the same as described in earlier papers (Scott,
1955, 1958, 1961). Many of the Hydroptilidae referred to were collected during
the survey of the Great Berg River (Harrison and Elsworth, 1958; Harrison,
1958); further specimens have since been taken in the same area. The new
species, Orthotrichia barnardi sp. n., was first correlated by Dr. A. D. Harrison,
from larvae inhabiting the small cases resembling caraway seeds illustrated
by Dr. Barnard in his work on South African caddis flies (Barnard, 1934, p. 391).
Since then further specimens have been bred out, from similar cases taken from
various parts of the Berg River; the resulting imagos all appear to belong to the
same species. The caraway seed case is typical of genus rather than species, as is
usually the case in the Hydroptilidae; a different species has been bred out
from very similar cases elsewhere in South Africa, this will be described in a
later paper. The immature stages of the two species, though superficially alike,
do show recognizable differences. The new species is named in honour of
Dr. K. H. Barnard, who first discovered the pupal cases.
Sincere thanks are again due to Dr. A. D. Harrison (now at the University
College of Rhodesia and Nyasaland) for much of the material, to Mr. D. E.
469
Ann. S. Afr. Mus. 46 (19), 1963, 469-478, 3 pls. SMITHSONIAN
eee
470 ANNALS OF THE SOUTH AFRICAN MUSEUM
Kimmins of the British Museum (Natural History) for his very kind advice, and
to Professor J. H. Day of the Zoology Department of the University of Cape
Town in whose department this work was carried out.
DESCRIPTIONS OF SPECIES
Genus ORTHOTRICHIA Eaton
Eaton, 1873, 141.
McLachlan, 1880, 518.
Orthotrichia barnardi sp. n.
(Plate XXI, a-c; plate XXII, a—m)
A minute, dark, hairy insect with a median longitudinal pale streak down
head, thorax and wings when at rest. Spurs 0, 3, 4. No ocelli. The larvae and
pupae inhabit small, dark, seed-like cases.
Imago ( 3, described when alive, 2 similar) (pl. XXI, a-c; XXII, a—c) : Head
(pl. XXI, Gc) : Eyes large, black; dense tuft of peach-coloured setae on face between
eyes, extending back between bases of antennae; main part of vertex bare, but
apical part with some setae, and the whole entirely obscured by these and the
facial setae and by two dense, bushy tufts of setae which arise from a pair of
large, raised warts on the back of the head. These tufts are mainly pinkish,
with a few black lateral setae. Antennae with 28-30 segments (24-25 in Q),
about half wing length, pinkish fawn with grey annulations (setae thickened
and shaded); two white bands near apex, each of two segments. Maxillary
palps five-jointed, pinkish with some grey setae anteriorly, first and second
joints small, the second with long stiff bristles; labial palps three-jointed,
pinkish. Thorax (pl. XXI, G); prothorax reduced, with a small pair of median
warts and a larger pair of lateral warts; mesonotum with scutum covered with
recumbent setae, postnotum and scutellum with a dense fur of erect setae;
metanotum with a median band of recumbent setae flanked by lateral tracts of
erect hairs; all thoracic setae pinkish except for tufts of black hairs on tegulae
and on antero-lateral corners of pronotum. Legs: very hairy, pinkish, with
some grey setae anteriorly (pinkish when seen from below, grey from above).
Mid- and hind legs with tibiae expanded and bearded. Spurs 0, 3, 4, hairy,
normal, the inner preapical spurs on the hind legs very long. Wings: forewings
very densely hairy, almost entirely covered with erect, somewhat thickened,
blunt-tipped setae; mainly black, with a broad streak of pale peach-coloured
setae along the hind margin, producing a conspicuous pinkish longitudinal
stripe down the middle of the back when the insect is at rest. This stripe starts
from the tufts on the head and narrows gradually, merging posteriorly with the
thick peach-coloured fringe at the tips of the forewings; it is partially inter-
rupted by two blackish patches near the middle. There are also several peach-
coloured streaks on the wings, particularly towards and along the anterior
margin. Fringe pink and black, brindled in appearance in some parts, very
long and dense (wider than wing in widest part). Hind wings greyish with
NEW CADDIS FLIES (TRICHOPTERA) FROM THE WESTERN CAPE PROVINCE 471
extremely long, silky, pinkish-grey fringe (three to four times width of wing).
Abdomen pale green, genitalia brown. In specimens which have dried out for
some time, the colouring fades to a brownish black marked with buff, with a
pinkish tinge in places; abdomen grey-green; setae on legs and thorax pale gold.
In spirit specimens the general appearance is brown and yellowish-gold.
Wings (pl. XXII, a): g forewing 3:0mm., 9 same. ¢ forewing without lobe,
with forks 1, 2, 3: R, with long cross-vein to Rs which makes it appear basally
double; R, does not arise from stem of R,,, as in O. sanya Mosely, O. spinicauda
Kimmins and O. straeleni Jacquemart, but in the normal way. There is a line
of long, stiff hairs on the ventral side of the wing, along the basal half of Cu,;
no black scales along subcosta. § hind wing with fork 2 only. 2 forewing with
venation very similar to that of J, but somewhat obscured by thickened, pitted
bands to which upright setae were evidently attached; similar, narrower bands
also occur between several of the apical veins. Sc shorter than in ¢; R, incom-
plete basally. 9 hind wing also very similar to that of g, but R, incomplete
basally, and cross-vein joins Rs and M,,, instead of Rs and M;; pitted bands also
present between some of the veins, particularly in the apical forks.
6 genitalia (pl. XXI, a-F): gd genitalia of the usual complex asymmetrical type.
Sixth sternite with a tuft of hair covering a spatula-shaped process in the mid-
ventral line; seventh sternite with a pointed process. Ninth segment largely
withdrawn within the eighth. Ninth tergite strongly sclerotized, wider pos-
teriorly than anteriorly, from the side cut away anteriorly, side-pieces being
present in the posterior half only; these sweep round to join the claspers, the
two sides being united ventrally by a membranous area which is bounded
laterally by longitudinal ventral sutures, as Nielsen describes for O. tetensii
(Nielsen, 1957) (see pl. X XI, F). From these side-pieces a pair of asymmetrical
lateral processes arise, each tipped with a long seta, the process on the right
being the longer. The claspers are short, narrow, leaf-like, with an irregular
margin, the left one being larger than the right; each has a curved, finger-like
projection forming the inner tip, and a stout seta on the outer edge. In dorsal
view the ninth tergite is fused onto the tenth, which forms a narrower, lightly
sclerotized hood, widening basally to form a sheath for the aedeagus; the hood
bears a sharp, laterally directed spine near the right side of the apex. This
spine is connected to a long, curved, blunt-ended process beneath the hood.
Another strong, sinuously curved, backwardly-directed spine arises near the
junction of IX and X in the mid-dorsal line; this is conspicuous in lateral views.
The apical spine appears to correspond to the right-hand, laterally bent spine
in O. tetensii, but there is no corresponding one on the left, unless indeed the
curved dorsal spine should represent it, or even possibly the blunt-ended
process beneath the hood. The aedeagus is long and slender, widening anteriorly,
with a spiral titillator coiled loosely round it (pl. XXI, £); when fully extended
the aedeagus may be as long as the entire abdomen, about 1-5 mm.; according
to Nielsen (1957) it is incapable of retraction when once extended. Within the
dorsal hood, above the aedeagus, there are two spine-like parameres; the one on
472 ANNALS OF THE SOUTH AFRICAN MUSEUM
the left side is long and slender, running the full length of IX and X and
projecting slightly beyond the apex of the hood; the one on the right is less than
half as long, arising near the junction of [IX and X and projecting from the
right side close to the curved process of the basal plate. Both appear to be con-
nected with an internal sclerite or sclerites, shown by dotted lines in plate XXI, p.
Below the aedeagus there is an elongated structure apparently connected to
a narrow basal apodeme; it widens and divides posteriorly into two branches,
each tipped with a stout seta. This forked structure can be clearly seen in a
cleared specimen in ventral view (pl. XXI, F), and the two apices are visible
projecting beyond the ends of the claspers. According to Nielsen (1957) the
forked structure may possibly represent the dorsal branches of the coxopodites
of the claspers.
? genitalia (pl. XXII, B, c) : The 2 genitalia show clear relationship to those of
O. spinicauda Kimmins (1958). The sixth sternite has a small, sharply pointed,
median process; seventh sternite normal. Eighth segment, as in sfinicauda, forms
a complete ring, the dorsal and lateral parts with an irregular margin fringed
with stout setae, the sternite forming an asymmetrical subgenital plate, partly
membranous, which appears to enclose a duct; apex of plate strongly sclerotized
and apparently connected to the membranous part of [X dorsal to it. Tergite
of IX forming a lightly sclerotized hood, partly withdrawn into the eighth
segment; sternite membranous, bearing a pair of subtriangular sclerotized
plates which are usually pressed against the dorsal hood; they may, however,
stand away from it, being attached along the proximal margin; proximal and
lateral parts of these plates are more heavily sclerotized than the rest, apical
margin irregular, pale, with a few small setae. The dorsal hood narrows just
beyond the plates, and the terminal portion, rounded and with a small pair of
rod-like cerci and a few setae, appears to represent the tenth segment.
This species resembles most other African species so far described in
possessing a bifid structure beneath the aedeagus, a spiral titillator accom-
panying the aedeagus, and partly fused claspers; like O. aequatoriana Kimmins
and O. alboguttata Jacquemart it lacks the row of black scales between costa and
subcosta in the g forewing. It shows most resemblance to O. spinicauda Kim-
mins, but can easily be distinguished from that species by its much larger size,
wing differences, and, in the ¢ genitalia, by the strongly curved dorsal spine
and the slender, upcurved, lateral processes. The 9 also resembles spznicauda, but
is easily recognizable in ventral view by the brown scale-like sclerotized plates.
Distribution
Case-bearing larvae of this species have been found in the upper part of the
Great Berg River from late winter to early autumn, mainly on stones in current
and on the trailing Scirpus digitatus on such stones, also in slower-flowing reaches
in pools with stony bottoms, and sometimes on marginal vegetation. Pupae
and imagos appeared in the Mountain Torrent Zone (Station 1) in early
autumn (March), and in the foot-hills (Stations 3 and 5) in spring and summer
NEW CADDIS FLIES (TRICHOPTERA) FROM THE WESTERN CAPE PROVINCE 473
(September to December). Larvae were also found much lower down the
river in the stony run at Piketberg (Station 18) in the late summer. No larvae
were found during the winter floods. Early instars are not at present identifiable,
and many even of the smaller case-bearing larvae passed right through the nets
used. Further details of the habitat will be found in Harrison and Elsworth
(1958) and Harrison (1958).
Specimens (tmagos)
All except GBG 770 H were collected from the Great Berg River as larvae
or pupae, either from stones in current or Scirpus in current, and bred out in the
laboratory. GBG 770 H was caught in the field, on the fish-ladder in Tulbagh
Kloof, Klein Berg River (a tributary of the Berg).
GBG 542 A-F, 4 dd, 2 99 (Stn. 1, March 1952, A.D.H.).
GBG 770 H, 1 ¢ (Tulbagh Kloof, 23/10/53, A.D.H.).
MISC 70, 2 §¢ (Stn. 3, December 1954, K.M.F.S.).
MISC 71, 2 99 (Stn. 3, December 1954, K.M.F.S.; one pinned).
MISC 85, 1 ¢ (Stn. 3, December 1954, K.M.F.S.).
MISC 122, 1 g, 2 99 (Stn. 3, October 1956, K.M.F.S.).
MISC 123, 1 3g, 1 @ (Stn. 3, October 1956, K.M.F.S.).
MISC 134, 1 ¢ (Stn. 3, October 1959, K.M.F:S.).
(Catalogue numbers all refer to the collections of the National Institute for
Water Research.)
The holotype 3 (GBG 542 A) and paratypes (GBG 542 B, 3g, and MISC
71, 2) will be placed in the South African Museum, and paratypes (MISC 70,
2 464, and MISC 123 A, 1 9) in the British Museum (Natural History). The rest
of the material, together with larval and pupal specimens, is in the collections
of the National Institute for Water Research.
Immature stages
Larval cases found vary in length from 1-5 to 2-5 mm.; none was found
smaller than this. Case-bearing larvae are in their 5th instar according to
Nielsen (1948), and increase considerably in size in this instar, adding to their
cases as they grow. Similar-looking free-living Hydroptilid larvae have been
seen, probably earlier instars of this species.
Larval case (pl. XXII, £)
The mature larvae live in small, brown, bivalve cases made of silky secre-
tion. These resemble caraway seeds, and each has one pair of dorso-lateral
ridges. The cases show striae, evidently resulting from the mode of formation
from threads of secretion; the line of junction of the two halves of the case is
mid-ventral, where the two ‘valves’ overlap slightly.
Mature larva (pl. XXII, p-L)
Length of larva slightly less than that of case, up to about 2-3 mm. Head
brown, with paler areas round the fairly large eyes; lens-like areas present in
474 ANNALS OF THE SOUTH AFRICAN MUSEUM
cuticle above eyes (pl. X XII, Fr). Clypeus wide, posteriorly triangular, one pair
of slight lateral indentations. Labrum with an asymmetrical median point;
mandibles large, blunt, with a single apical tooth below which is an indentation
bounded proximally by a tooth or point, the broader left mandible with an
inner brush of slender spines. Gular sclerite Y-shaped, a small triangular anal
gular sclerite also present (pl. XXII, L). Pro-, meso- and metanota (pl. XXII, F)
each with a pair of brown sclerotized plates, posterior margins darkened; narrow
dark brown sternites also present (posterior or anal sclerites only). Pronotum
longest, mesonotum shorter, metanotum shortest, each with a fringe of short,
stiff setae interspersed with longer setae along the anterior border. Legs small,
brown, much like those described by Nielsen for O. tetensii; forelegs shorter and
stouter than mid- and hind legs, which are sub-equal, the hind legs being the
longer (pl. XXII, c, n, J). Abdomen large, thick, white, more dorso-ventrally than
laterally compressed, with clear segmentation; segment 1 small, with a few
dorsal setae, largely covered by segment 2, segment 5 largest. Abdomen fits
so closely into the case that the latter must be split open to extract the larva.
No gills, no lateral line. Ninth segment with strongly sclerotized semicircular
brown anal sclerite, bearing eight long marginal setae and two shorter, stout,
curved bristles. Anal appendages small, each with a large basal sclerite and a
minute hooked anal claw (pl. XXII, p, k).
The larva shows clear resemblances to that of O. tetensit Kolbe as figured by
Nielsen (1948) and Ulmer (1957) ; there are, however, various minor differences,
and the cases of O. tetensii have paired lateral ridges, in the fully grown speci-
mens at any rate. There are also resemblances to the larva of Orthotrichiella
ranauana Ulmer (1957). Ulmer distinguishes the larvae of Orthotrichia and
Orthotrichiella on a basis of claw length as related to length of tarsus, and of
antenna length as related to length of left mandible (1957, p. 182). The larva of
Orthotrichia barnardi, however, has not quite the proportions of either, though
nearer to the latter; the foreclaw is slightly shorter than the tarsus, and the
claws of mid- and hind legs only half, or less than half, the length of the tarsi;
length of antenna about a third of the length of the left mandible. The adults,
however, definitely appear to fall into the genus Orthotrichia.
The larva is easily distinguishable from the species figured by Corbet
(1958, fig. 4, ef), as it lacks the dark, patterned metanotum and the dark
mark on the clypeus, and the case has one, not two, pairs of dorso-lateral
ridges. In comparison with the figures of the larva of O. straelent (Jacquemart,
1957, figs. 135-154), the thoracic nota of O. barnardi lack the broad, darkened
bands, and the sclerite on IX is much more strongly sclerotized; the larva of
O. straelent appears to make a plain case without lateral ridges.
Pupal case (pl. XXII, m)
The pupal cases have terminal holdfasts which anchor them firmly to
Scirpus plants or to stones. The larval cases are utilized in their formation, with
additional material closing them at both ends; they are opened by a V-shaped
NEW CADDIS FLIES (TRICHOPTERA) FROM THE WESTERN CAPE PROVINCE 475
dorsal slit when the mature pupa emerges. The larval sclerites are retained in
the end of the pupal case opposite to this opening.
Ross (1944, p. 117) states that the pupae of the Hydroptilidae are very
uniform in structure and no characters have been found to key them to genus;
no description of the pupa is therefore given; it is of the usual Hydroptilid type.
HAydroptila capensis Barnard
Hydroptila capensis Barnard 1934, 391, fig. 51, a7. Harrison and Elsworth, 1958, Tables 19, 21,
24. Harrison 1958, p. 263.
In the Great Berg River, the larvae and pupae of this species occurred
mainly in the foot-hills, often in quiet tributary streams (Stations 1, 9, 10, 13
and 19 on main river, also Station 4 on the Franschhoek stream at La Motte,
in the Dwars River at Groot Drakenstein, in the Klein Berg at the Tulbagh
Barrage, and in the Kuils River near Piketberg), and principally at times of
the year when the rivers were low and running slowly and patches of Spirogyra
had been able to develop. Thus most were found during the summer and
autumn months, when the winter floods were over and the flow had slowed
down. Numbers encountered were not generally large, excepting in early
autumn one year (March 1953) at Wellington (Station 12), where slight organic
pollution had resulted in a blanketing growth of Spirogyra among the stones in
the stickles. The river was low and flow sluggish at the time. It seems that the
limiting factor is current and its effect on algal growth rather than temperature,
because numbers of larvae and pupae were also found in June (early winter),
in this case, however, in a very small tributary brook that runs down behind the
Forest Station at La Motte, where they were living on alga-covered stones in a
small pool sheltered from flooding by banks of sedges.
This species is one of the few that may appear in temporary water in this
country; the Kuils River does not run during the dry season (larvae and pupae
were found there in spring—October), and the author has also found this
species in temporary mountain streams and a recently filled farm dam in the
same area.
H. capensis is typically found in association with Spirogyra, with filaments
of which the larval and pupal cases are covered, but such larvae cannot be
identified as H. capensis on case alone without correlation, at least with mature J
pupae, since specimens bred out in the Transvaal have shown that H. cruciata
Ulmer also covers its cases with Spirogyra (material supplied by Mr. F. M.
Chutter; this will be described in a later paper).
Hydroptila sp.
Hydroptila sp. Barnard 1934, 393, fig. 51; /-
‘Hydroptila sp. A’ Harrison 1958, p. 263.
Small numbers of larvae were also found with cases similar to those
described by Barnard (1934); the bivalve cases are made of secretion covered
476 ANNALS OF THE SOUTH AFRICAN MUSEUM
with minute sand grains. Similar larvae collected in the Transvaal have been
bred out by Mr. F. M. Chutter, the imagos proving to be a new species of
Hydroptila, but it cannot be assumed that the Berg River specimens belong to
the same species without correlation, a difficult matter as the larvae are very
small and far from common. A few were found in backwaters at Station 3,
Berg River, in spring and summer, and among stones in the stickles in autumn.
Oxyethira velocipes (Barnard)
(Plate XXIII, a-x)
Argyrobothrus velocipes Barnard, 1934, 393, figs. 52, a-/.
Argyrobothrus velocipes Barnard, Harrison and Elsworth 1958. Harrison 1958.
Oxyethira velocipes (Barnard), Kimmins 1962.
Distribution
Larvae belonging to this species construct flat, transparent, flask-like cases,
and were found from Station 1 in the Mountain Torrent Zone of the Berg
River right down to the last station before the estuary. They appeared among the
marginal vegetation in spring, summer and autumn, when the river was
slowing down, and sometimes became locally common, showing a sporadic
distribution which evidently depended on amount of shelter and extent of algal
growth among other factors. This might be expected from the feeding habits,
beautifully described by Nielsen (1948). The larvae were identified by correla-
tion in the laboratory, imagos all proving to be O. velocipes (Barnard). As
Barnard did not describe the larva of his species, but only figured the pupal
case, a description of the larva and case is given below.
Larval cases (pl. XXIII, a, 8)
The cases of the younger fifth instar larvae have plain, smooth edges; as the
larva adds to the case, however, it does so in such a manner that each new
section added is evidently commenced some little distance within the old
posterior opening. As a result of this, the cases of older larvae show a series of
strong lateral serrations along each side of the case. In this they differ from the
cases of O. costalis described by Nielsen (1948), except for one aberrant speci-
men (Nielsen, 1948, fig. 54, K). In O. costalis the smooth, plain shape is retained,
and the same case used for pupation. Larval cases of O. velocipes show the
remnants of material used for the ring-shaped fixing belt, just as in O. costalts.
The larvae are orientated in the cases with head to the narrow end with the
small, circular opening; the wide, flat end usually remains closed owing to the
shape of the case, it can however be pushed open.
Larval cases vary in length from 1-3 to 4:5 mm.; the fifth instar larvae
inhabiting them range from 1-3 to 2:2 mm. in length; as they increase in size
the abdomen becomes proportionately larger in comparison with the rest of
the body, always, however, retaining its laterally compressed shape, with the
posterior end curled round ventrally.
a es
NEW CADDIS FLIES (TRICHOPTERA) FROM THE WESTERN CAPE PROVINCE 477
Fifth instar larvae (pl. XXIII, a—-K)
Plate XXIII, A, illustrates a mature larva in its case; as, however, in such
specimens the thorax and ist abdominal segment are largely obscured by the
2nd abdominal segment, a slightly younger 5th instar larva has been chosen to
illustrate details of structure (pl. XXIII, c-x).
Head pale yellowish; eyes fairly large, under lens-like areas of cuticle (pl.
XXIII, p) ; clypeus with one pair of lateral indentations, pointed antero-lateral
corners, and triangular posterior area, three pairs of the bristles on the clypeus
are thick, dark brown, and expanded towards the base. Labrum with a pair of
pointed antero-lateral spines, an apparently soft central projection, and two
larger and eight smaller setae, all of which are decumbent. There are tufts of
setae on the ventral side. Both clypeus and labrum show strong resemblances to
those of O. costalis. Antennae fairly long, with rounded base, slender sensilla,
and long lateral seta. There is one pair of extremely long, slender setae, situated
just above the eyes; the other setae on the head are much shorter and also
slender (except for the thickened ones on the clypeus). ‘The mouthparts usually
seem to be carried tucked inwards. Mandibles (pl. XXIII, #) : the right mandible
is simple, with 3 small apical teeth; the left mandible has one apical tooth, below
which are two broad, projecting, cutting edges, each with about 4 low, blunt
teeth. There appears to be a deep cavity between these, from which arises a
row of stout spines; the dorsal cutting edge is shorter than the ventral, its base
being scooped out to show a penicillus of slender hair-like spines. These two
lots of spines presumably correspond to the two brushes of setae in the left
mandible of O. costalis, described by Nielsen. Gular sclerite (pl. XXIII, J), a
scooped-out triangle with transverse thickenings; a small triangular anal gular
sclerite also present. Thorax: Pro-, meso- and metanota lightly sclerotized, pale
yellowish, the two latter narrower than the former, with pointed, darkened,
lateral corners and few setae; pronotum wider, with rounded anterior corners
and more setae. Legs: forelegs short, mid- and hind legs long and slender, sub-
equal, not, however, increasing in size as the abdomen grows during the fifth
instar; arrangement of setae as illustrated (pl. XXIII, E, F, Gc); legs pale
yellowish to brownish in colour, the mid- and hind legs with a brownish band
across the femora. Abdomen whitish, strongly laterally compressed, segmentation
obvious; in the mature larva segment 2 hides most of segment 1 and the meso-
and metanota. Ninth segment with a pale yellowish sclerite and a few strong
setae. Anal prolegs fused to form 1oth segment; small anal claws present; each
claw appears to consist of one large hook and four smaller ones, but the smallest
are difficult to distinguish clearly. No gills, no lateral line.
Pupal case (pl. XXIII, t)
The pupae are also found in flat, transparent cases; these have, however,
smooth, plain edges like those of the youngest fifth instar larvae, and appear
to be freshly made, not altered larval cases. The cases are anchored to a leaf
by four discs made of secretion, two at each end, and the rounded opening
478 ANNALS OF THE SOUTH AFRICAN MUSEUM
is closed by a fluffy plug of secretion as in O. costalis, not by a domed cap as in
O. incana (Ulmer, 1957, fig. 277). The other (wide) end is also closed off, by a
narrow strip of the same material, making an oval space in which the pupa
lies. The pupae are orientated the opposite way round to the larvae, with the
head at the wide end of the case. The larval sclerites are found within the case,
at the narrow end.
SUMMARY
A new species of Orthotrichia, O. barnardi sp. n. (Trichoptera: Hydroptili-
dae), is described from South Africa, together with its larval stages. The larva of
Oxyethira velocipes (Barnard) is also described, and an account given of Hydrop-
tilid material collected from the Great Berg River, Western Cape Province.
REFERENCES
BARNARD, K. H. 1934. South African caddis-flies (Trichoptera). Trans. roy. Soc. S. Afr. 2%,
291-394.
Corset, P. S. 1958. Larvae of certain East African Trichoptera. Rev. Zool. Bot. afr. 58, 3-4,
203-213.
Eaton, E. A. 1873. On the Hydroptilidae, a family of the Trichoptera. Trans. ent. Soc. Lond.
1873, 141.
Harrison, A. D. 1958. Hydrobiological studies of the Great Berg River, Western Cape Province.
Part 2. Trans. roy. Soc. S. Afr. 35, 227-276.
Harrison, A. D., & Etswortn, J. F. 1958. Ibid., Part 1. Trans. roy. Soc. S. Afr. 35) 125-226.
JACQUEMART, S. 1956. Trois Orthotrichia nouveaux du Lac Kivu (Trichoptera, Hydroptilidae).
Bull. Inst. Sci. nat. Belg. 32, 9, 1-6.
Jacquemart, S. 1957. Trichoptera des Lacs Kivu et Edouard. Exploration Hydrobiologique des Lacs
Kivu, Edouard et Albert (1952-1954), 3, 65-129.
Kiunins, D. E. 1957. New and little-known species of African Trichoptera. Bull. Brit. Mus. (Nat.
Hist.) Ent. 6, 1-37.
Kiwmins, D. E. 1958. New species and subspecies of Odonata and on some Trichoptera from
S. Rhodesia and Portuguese East Africa. Bull. Brit. Mus. (Nat. Hist.) Ent. 75 349-368.
Kimmins, D. E. 1962. New African caddis-flies (Order Trichoptera). Bull. Brit. Mus. (Nat. Hist.)
Ent. 12, 83-121.
Macponatp, W. W. 1950. The larvae of Mystacides azurea L., Cyrnus flavidus McLachlan and
Oxyethira simplex Ris (Trichoptera). Proc. R. ent. Soc. Lond. (A) 25, 1-3, 19-28.
MaruleER, G. 1943. Trichoptera. Explor. Parc. nat. Albert Miss. Damas. 115 3-34.
McLacuian, R. 1880. A monographic revision and synopsis of the Trichoptera of the European fauna.
518. London & Berlin.
Mose ty, M. E. 1948. Trichoptera collected by Miss R. H. Lowe at Lake Nyasa. Ann. Mag. nat.
Hist. (12) 15 45-47.
NIELSEN, A. 1948. Postembryonic development and biology of the Hydroptilidae. Biol. Skr.
5s (1), 1-200.
NIELSEN, A. 1957. A comparative study of the genital segments and their appendages in male
Trichoptera. Biol. Skr. 8, (5), 1-159.
Ross, H. H. 1944. The caddis-flies, or Trichoptera, of Illinois. Bull. Ill. nat. Hist. Surv. 23, 1-326.
Scott, K. M. F. 1955. Some new caddis flies (Trichoptera) from the Western Cape Province—I.
(Some Sericostomatidae.) Ann. S. Afr. Mus. 41, 367-380.
Scott, K. M. F. 1958. Ibid.—II. (Some Leptocerinae.) Ann. S. Afr. Mus. 44, 39-52.
Scott, K. M. F. 1961. Ibid.—III. (More Leptocerinae.) Ann. S. Afr. Mus. 46, 15-33.
Umer, G. 1951. Kécherfliegen (Trichopteren) von den Sunda-Inseln, Teil I. Arch. Hydrobiol.
Suppl. 19, 1-528.
Umer, G. 1957. Ibid. Teil III. Arch. Hydrobiol. Suppl. 23, 109-470.
Ann. S. Afr. Mus., Vol. XLVI Plate XX1
~.
w= l
wo na Os es OS NS UO ea
<< eS ee ee ee
we en
Orthotrichia barnardi sp. n., 3 genitalia: A, dorsal view; B, ventral view; C, left latera
view; D, right lateral view; E, aedeagus in dorsal view showing spiral titillator; F, claspers in
ventral view to show position of furcate structure; G, dorsal view of head and thoracic nota.
(In all cases except E and F the spiral titillator and furcate structure have been omitted where
seen by transparency. In C and D the position of the eighth and part of the seventh segments
are shown in outline.)
Ann. S. Afr. Mus., Vol. XLVI Plate XXII
Orthotrichia barnardi sp. n.: A, fore- and hind-wings of $; B, dorsal view of 2 genitalia;
C, ventral view of 9 genitalia; D, whole larva, lateral view; E, larva in case (half magnification
of D); F, dorsal view of head and thoracic nota of larva, with small part of margin of pronotum
further enlarged; G, H, J, fore-, mid- and hind-legs of larva; K, ventral view of terminal segments
of abdomen of larva; L, ventral view of head of larva, with antenna and right and left mandibles
further enlarged (labium and palps omitted for clarity); M, empty pupal case, attached to
Scirpus leaf.
Ann. S. Afr. Mus., Vol. XLVI Plate XXIII
2]
Oxyethira velocipes (Barnard): A, mature 5th instar larva in case; B, young 5th instar
larva in case; C, lateral view of large but not quite mature 5th instar larva, extracted from case,
anal claw further enlarged; D-K, parts of same larva, further enlarged (D, head; E, F, G, mid-,
hind and forelegs; H, left and right mandibles in dorsal view; J, gular sclerite and submental
sclerite; K, clypeus and labrum); L, pupal case.
[CONTINUED FROM INSIDE FRONT COVER
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrége zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
Hartman, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
Izuxa, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYM Y. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. 2nd ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.—All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name... .
B. Form of bibliographic references to synonymic names.—The first reference following
any name in the synonymy should be to the earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalis capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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INDEX
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When reference is made to a separate book, give in the order listed:
Author’s name; his initials; date of publication; title, in italics; edition, if any;
volume number, if any, in black type arabic numerals; pagination, if reference is
to part of book only; place of publication; name of publisher.
When reference is made to a paper forming a distinct part of another book or
collection, give:
Name of author of paper; his initials; date of publication; title of paper; ‘In’
italicized; name of author of book; his initials; title of book, in italics; edition, if
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place of publication; name of publisher.
[CONTINUED ON INSIDE BACK COVER
TOOTH REPLACEMENT IN THE CYNODONT
THRINAXODON LIORHINUS SEELEY
By
A. W. Crompton
South African Museum, Cape Town
(With 17 figures in the text)
CONTENTS
PAGE
Introduction , ; Amy ie
Material . ; . 480
Replacement of individual teeth . . 481
Structure of the crowns of the postcanines . 483
Discussion
Extension of growth series of Thrinaxodon . 497
Posterior migration of the postcanine row. 501
Replacement cycle > , . 502
Differentiation of the postcanine row eG
Notes on the postcanine row of gomphodont
cynodonts . ; » 515
Summary . : : , a RTO
Acknowledgements . 520
References . . , . 520
Key to lettering : . 52x
INTRODUCTION
The very mammalian appearance of the dentition of such theriodont
reptiles as the cynodonts invited the assumption that these reptiles replaced
their teeth as mammals do and that they had milk and permanent dentitions
(Broom, 1913) rather than the indefinite series of replacing teeth occurring in
typical reptiles. But Parrington (1936) has shown that in the cynodont Thrinaxo-
don alternately numbered postcanines were replaced at different times. In the
eight skulls at his disposal Parrington could show that each of the postcanines
was replaced at least once. He divided the material into a younger group in
which postcanines nos. 1, 3 and 5 had been replaced shortly before death or
were actually undergoing replacement, and an older group in which post-
canines 2, 4 and 6 had been replaced shortly before death, and he assumed that
the tooth position of the Ist postcanine in the younger group was homologous
with the tooth position of the Ist postcanine in the older group. Parrington
noted that some of the specimens have six upper postcanines whereas others
have seven, but he gave no explanation for this. He claimed that the 7th post-
479
Ann. S. Afr. Mus. 46 (20), 1963, 479-521: 17 figs.
a
BST TUIIOH JAN J 194
480 ANNALS OF THE SOUTH AFRICAN MUSEUM
canines were unlikely to have had predecessors and were erupting for the first
time. Also Parrington described the mandibular postcanines in one specimen
in which he noted that the 7th and 8th postcanines had five cusps and the 5th
four cusps, instead of the customary three.
Subsequently Brink (19556) has described a very young specimen of
Thrinaxodon, found in close association with a large specimen which has seven
teeth. He formed the opinion that the postcanines were replaced several times
during life because the teeth of the young specimen were so much smaller than
those of the large specimen. I have (Crompton, 1955) discussed tooth replace-
ment in Cynognathus zone and Middle Triassic cynodonts, notably Scalenodon.
In Scalenodon postcanine teeth were found to have been added at the back of
the postcanine row during life and a few were lost in front. Fourie (1963) has
obtained similar results in Diademodon. Recently I (Crompton, 1962) showed
that in the bauriamorph E£riciolacerta the replacing teeth developed in pits in
the alveolar bone lingually to the roots of the functional postcanines and that
alternately numbered teeth were replaced at different times.
MATERIAL
Four specimens of Thrinaxodon have recently been beautifully prepared
with the aid of acetic acid by the staff of the British Museum (Natural History).
These are B.M.N.H. R3731, R5480, R511a and R511. The first three were
studied by Parrington (his specimens E, G and I respectively). I studied three
of the British Museum specimens, viz. R3731, R511a and R511. Several
specimens in the collections of the South African Museum have also been
prepared with the aid of acetic acid. Consequently much additional information
on the structure of the tooth crowns and tooth replacement has come to light.
For the purpose of this paper a small skull of Thrinaxodon housed in the
Universitats Institut fiir Palantologie u. historische Geologie in Munich and
the very young specimen of Thrinaxodon housed in the Bernard Price Institute
for Palaeontological Research in Johannesburg were studied. For convenience
the specimens studied have been lettered A to I, as follows:
A = Bernard Price Institute for Palaeontological Research Catalogue No.
274.
B = Universitats Institut fiir Palaontologie u. historische Geologie, Munich.
= B.M.N.H. R3731 = Parrington’s Specimen E.
1D. SA. RS 77
E =S.A.M. K 380.
F = B.M.LN.S. Rais,
G=sA.M YE. 296;
H = B.M.N.H. R511a = Parrington’s Specimen I.
Po = Sea ee SG:
The structure of the postcanine teeth indicates that this sample may contain
more than one species of Thrinaxodon. This is discussed on pp. 511.
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 481
REPLACEMENT OF INDIVIDUAL TEETH
Postcanines: A shallow longitudinal groove (long.g., fig. 2 A and 6 A and D)
is present in the maxilla and mandible lingual to the alveolar borders of the
postcanines and the canine. This groove presumably housed the dental lamina.
Replacement teeth commenced their development in shallow pits (p) in the
floor of this groove in the dentary and in the roof of this groove in the maxilla.
Initially these pits were separated by a layer of bone from the alveoli of the
A Rear ee
dost Reacts
Fic. 1. Yhrinaxodon liorhinus.
A, right maxillary dentition of Specimen A.
B and C upper dentition of Specimen B.
Key to lettering p. 521.
482 ANNALS OF THE SOUTH AFRICAN MUSEUM
functional postcanines which lie labial to them. These pits are well preserved
in the acid-prepared specimens and in many cases replacing teeth are preserved
in the pits (rep.t., fig. 2 B). The functional teeth are usually firmly held in
place by a ring of attachment bone around the neck of the tooth. The attach-
ment bone (a.b.) is well preserved around the neck of several postcanines
studied and is especially well preserved around the 2nd and 4th upper post-
canines of specimen E (fig. 5 A). A ring of small foramina indicates the Jine of
fusion between the attachment bone and the tooth itself. The various stages of
the growth of a replacing tooth between its initial development in a pit and its
eventual eruption are well shown in the acid-prepared specimens. As the
replacing tooth increased in size the pit containing it also increased in size and
opened into the alveolus of the functional tooth. The replacing tooth migrated
labially and the root of the functional tooth was resorbed to make space for it.
Consequently, shortly before the replacing tooth erupted little remained of the
root of the functional tooth. At this stage the remaining part of the functional
tooth was secured solely by the ring of attachment bone around its neck at the
alveolar border. In many specimens the functional tooth was lost at this stage
by absorption of the neck of the tooth in a horizontal plane in line with the
outer surface of the ring of attachment bone so that nothing visible remains
externally to indicate the previous presence of a functional tooth except a ring
of foramina in the attachment bone around the remnant of the tooth root.
(See fig. 1 B.) This phenomenon accounts for the gaps in the tooth row in the
right maxillae of Parrington’s specimens A and F. Before the replacing tooth
erupted the alveolus was considerably enlarged by the resorption of the attach-
ment bone and the new tooth erupted into an alveolus far larger than the neck
of the replacing tooth. Consequently the younger teeth that erupted shortly
before death were loosely held in their respective alveoli, e.g. 1st, 3rd, and 5th
in the maxilla of specimen C and 3rd, 5th and 7th in the mandible of specimen
C (fig. 2). The older teeth, e.g. 2nd and 4th in the maxilla of specimen C
(fig. 2 A and B), were firmly held in place by attachment bone.
A replacement cycle identical to that of Thrinaxodon appears to have been
present in the bauriamorph Ericiolacerta (Crompton, 1962) and was also
present in Cistecephalus zone cynodonts that have been studied by the author.
The material studied for this paper supports Parrington’s (1936) conclu-
sion that the odd- and even-numbered postcanines erupted at different times.
This is clearly seen in most specimens, e.g. in the maxilla of specimen C (fig. 2 A)
the 1st, 3rd and 5th erupted shortly before death and were loosely held in large
alveoli whereas the 2nd and 4th were tightly held and have pits for replacing
teeth in the maxilla lingual to them. However, it appears that all the odd-
numbered (or even-numbered) teeth did not erupt simultaneously; but that
the posterior replacing teeth erupted before the anterior teeth of the same set.
This is well shown in the right mandible of specimen H (fig. 10 D). The 3rd,
5th and 7th teeth are loosely held in the alveoli and erupted shortly before
death. The 7th tooth was fully erupted, the 5th partially erupted and the 3rd
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 483
was still completely contained in its alveolus. In terms of alternate replacement
a replacing tooth would be expected to be present in the first alveolus. How-
ever, the old functional tooth is still present in this position, but a deep pit for
a well-developed replacing tooth is present adjacent to the old 1st functional
postcanine. In the left mandible (fig. 10 C) the condition is similar to that in
the right, except that the old functional Ist postcanine has been lost and a
partially displaced replacing tooth is present in the first alveolus. This evidence
suggests that the odd-numbered replacing postcanines erupted in the following
order: 7th, 5th, 3rd, 1st. In the maxilla of specimen F (fig. 6 A) the replacing
tooth for the 6th postcanine (r.pc.6.) is larger than that for the 2nd (r.pc.2.)
and that of the 4th is intermediate in size between the 2nd and the 6th. The
younger postcanines, i.e. 1st, 3rd and 5th of this maxilla, that are loosely held
in their alveoli appear to have erupted in the following order: 5th, 3rd, 1st.
A similar phenomenon can be observed in most of the specimens studied with
the exception of the lower postcanines of specimen G (fig. 9) where the rule of
alternate replacement seems to break down over a short distance.
In most reptiles that actively replace their teeth a wave of replacement
passes along each alternately numbered tooth series either from back to front
or front to back (Edmund, 1960, 1962). In Thrinaxodon the wave passes from
back to front.
Canines: Replacing canines are present in both the upper and lower jaws
of most specimens of Thrinaxodon studied. The replacing canines commenced
their replacement in distinct pits in the maxilla and dentary. In the specimens
studied the pits in the maxilla (specimen E, fig. 5 A) are always found antero-
lingual of the functional canine whereas the pits in the dentary (specimen E,
fig. 5 B) are always found postero-lingual of the functional tooth. As the replac-
ing canine enlarged, the pit containing it fused with the main alveolus (speci-
men D, fig. 3 A and B) and the replacing canine migrated to lie in front of the
functional canine in the upper jaw and behind the functional canine in the
lower jaw (specimen F, fig. 6 B and D). Fourie, however (1963), has described
a specimen of Thrinaxodon in which the upper replacing canine lies behind the
functional canine.
Incisors: The incisor region is not well preserved in most of the specimens
studied. The replacing incisors appear to have developed in pits behind the
functional ones. These pits, where preserved, are confluent with the alveoli of
the functional teeth. In the premaxilla of specimen F (fig. 6 A and B) replacing
teeth are present behind all four incisors. It is not possible to determine the
order of replacement of the functional incisors.
STRUCTURE OF THE CROWNS OF THE POSTCANINES
In a few instances Parrington (1936) was able to describe additional cusps
on the lingual side of the crowns of upper postcanines. In the acid-prepared
specimens the whole structure of the crowns of the teeth can be studied, and it
484
ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 2. Thrinaxodon liorhinus, Specimen C.
A and B upper dentition, C and D, lower dentition and
E, lingual view of the lower 6th postcanine.
Key to lettering p. 521.
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 485
has been found that in many cases quite complex patterns are present which
differ considerably from the simple crown pattern usually associated with
Thrinaxodon.
On the basis of the pattern of the crowns of the lower posterior postcanines,
it is possible to divide the specimens studied into three ill-defined groups.
Specimens A, B or I, however, cannot be assigned to any of these groups as the
structure of their lower postcanines is not known. The members of the three
groups form a consecutive series of increasing size (see table I, page 500).
Group I consists only of specimen C.
Group II consists of specimens D, E, F and G.
Group III consists of specimen H.
Group I
Lower Postcanines. Specimen C' (fig. 2 C, D and E)
Seven lower functional postcanines are present. A remnant of a tooth
(r.pc.) is present anterior to the 1st functional postcanine and an unerupted
postcanine (a.pc.) which does not appear to have had a predecessor is present
behind the last functional postcanine.
The crown structure of the more recently erupted 3rd, 5th and 7th post-
canines is different from that of the older 2nd, 4th and 6th. In each series, i.e.
the odd-numbered, 3rd, 5th, 7th and the even-numbered 2nd, 4th and 6th,
the crown pattern becomes increasingly complex in a posterior direction. Of
great interest and importance is the fact that each of the more recently erupted
teeth has crowns with a simpler pattern than the older teeth anterior or posterior
to it. The tooth with the most complex crown is the 6th postcanine
(fig. 2 E). It consists of three cusps aligned antero-posteriorly; a main cusp
(m.c.), an anterior accessory cusp (a.a.c.) and a posterior accessory cusp
(p.a.c.). Because the anterior accessory cusp is larger than the posterior acces-
sory cusp the anterior margin of the main cusp is shorter than its posterior
margin. Anterior to the anterior accessory cusp and posterior to the posterior
accessory cusp two smaller cusps are present. These have for convenience been
designated as anterior and posterior cingular cusps (a.c.c., p.c.c.) respectively.
It is, however, extremely difficult to determine whether the anterior and pos-
terior cingular cusps do in fact develop upon a definite cingulum. In some cases
this appears to be the case, but in other cases these cusps appear to be quite
distinct from the cingulum. A well-developed cingulum is present on the
lingual surface of the crown. It supports a series of small cusps. ‘The largest are
twin cusps that lie either side of the midline. A deep valley separates the anterior
and posterior twin cusps. Two small cuspules are present anterior to the twin
cusps and one is present posterior to them.
The crown pattern of the 4th and 2nd postcanines is similar to that of the
6th except that accessory and the anterior and posterior cingular cusps and
the cusps on the lingual cingulum are not so well developed. The crowns of the
486 7 ANNALS OF THE SOUTH AFRICAN MUSEUM
younger 7th, 5th and 3rd postcanines in the odd-numbered series also consist of
three cusps (anterior accessory, main and posterior accessory) aligned antero-
posteriorly. The anterior and posterior cingular cusps are situated low down on
the lingual surface of the anterior and posterior accessory cusps and they do
not lie anterior or posterior to the latter as they do in the 6th postcanine. In the
odd-numbered teeth no cusps appear to be present on the lingual side between
the anterior and posterior cingular cusps.
Upper postcanines (fig. 2 A and B)
As in the lower postcanines there is a great difference between the older
and younger postcanines, but this is not as marked as in the lower jaw. The
crowns are in many cases damaged.
Six functional postcanines are present. The even-numbered erupted later
than the odd-numbered. The 6th postcanine has apparently been shed and an
unerupted postcanine is visible above it. The crown of the 4th postcanine,
which is a member of the older series of teeth, consists of three antero-posteriorly
aligned cusps, anterior and posterior cingular cusps and a series of cusps upon
a lingual cingulum. The cingulum of the crown of the second postcanine is
very poorly developed and supports only an anterior cingular cusp. In the
younger 5th and grd postcanines anterior and posterior cingular cusps are
present but no cusps are present on the lingual cingulum.
Group II
Lower postcanines. Specomen D (fig. 3 B and C and 4B)
Seven functional postcanines are present. A remnant of a tooth root
(r.p.c.) is present anterior to the 1st. A small, apparently non-functional tooth
is present behind the 7th postcanine (pc.8.). This tooth is completely covered
laterally by the anterior border of the coronoid process. As in specimen C,
the crowns of the older teeth are different from those of the younger teeth.
The sixth postcanine (fig. 4B), belonging to the older series, has three
cusps (anterior accessory, main and posterior accessory) aligned antero-
posteriorly. The main cusp and the anterior accessory cusp are about the same
size and the posterior accessory is slightly smaller. The posterior cingular cusp
lies directly behind the posterior accessory cusp whereas the anterior cingular
cusp lies slightly antero-lingually of the anterior accessory cusp. As a result the
crown, when viewed from above, has a wide, blunt anterior region and a
tapering posterior region. A large and prominent cingular cusp (c.c.) is present
on the lingual surface below the junction of the main and anterior accessory
cusps. A small poorly defined cuspule is present posterior to this cusp. A poorly
developed cingulum (cing.) is present between the two lingual cingular cusps
(c.c.) and the posterior cingular cusp. In the fourth postcanine the main cusp is
considerably larger than the accessory cusps. The anterior cingular cusp is
fairly well developed, but the posterior cingular cusp is small and the cingulum
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 487
and cusps upon the cingulum are very poorly developed. When viewed
from above, the crown has a wide anterior region and a tapering posterior
region, as in the 6th postcanine. The 4th postcanine is shorter but wider than
the 6th.
In the second postcanine the crown is dominated by the main cusp and
the anterior and posterior accessory cusps and the anterior and posterior
rep.c. p.
A a
rep.t.
rep.c.
.
(
PAKS:
Fic. 3. Thrinaxodon liorhinus, Specimen D.
A, lingual view of the upper postcanines,
B, lingual and C, labial view of the lower postcanines.
Key to lettering p. 521.
cingular cusps are very small. The lingual cingulum is absent. In crown view
the tooth is more circular than the 4th postcanine. Within the even-numbered
postcanines there is, therefore, in a posterior direction a progressive increase
in the size of the accessory and cingular cusps and a progressive decrease in the
size of the main cusp relative to the size of the accessory cusps. In addition
posteriorly the teeth become progressively more longitudinally ovate.
Similar tendencies can be observed in the odd-numbered teeth of the
same specimen, but the structure of the crown is slightly different. Unfor-
tunately the crown of the 7th postcanine is partially destroyed. The remaining
portion is similar to the 5th except that the cingulum and cingular cusps are
absent and it is more longitudinally ovate.
ANNALS OF THE SOUTH AFRICAN MUSEUM
488
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489
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY
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490 ANNALS OF THE SOUTH AFRICAN MUSEUM
In the 5th postcanine the crown is dominated by the main cusp and the
accessory cusps are small. In contrast to the even-numbered teeth the posterior
cingular cusp does not lie directly behind the posterior accessory cusp, but
postero-lingually to it. A well-defined anterior cingular cusp is absent; in its
place is a series of small cuspules arranged to form a short ridge (a.cing.)
running downwards in a posterior direction. Because the posterior cingular cusp
is well developed and the anterior cingular cusp poorly developed, the crown
has a wide (transverse) posterior region and a slightly tapered anterior region,
i.e. Opposite to the conditions in the even-numbered teeth. A small cuspule
(a.l.c.) called the anterior labial cusp is present on the labial surface of the
tooth below the junction of the main and anterior accessory cusps. The 3rd
postcanine is similar to the 5th except that the main cusp is relatively larger
and the remaining cusps relatively smaller. The cuspule on the labial surface is
absent.
It is clear from the above description that the crowns of the odd-numbered
series (3, 5 and 7) have the same basic structure and that this is quite distinct
from the structure of the crowns of the even-numbered series (2, 4 and 6).
In each series, however, the teeth become more complex in a posterior direc-
tion. Thus not only do the odd- and even-numbered teeth replace at different
times in specimen QC, but in addition the crown patterns of the two series are
distinct from one another.
The 8th postcanine is an enigma. It is smaller than any of the other lower
postcanines with the exception of the first. ‘The main cusp dominates the crown.
The only part of the cingulum developed is the anterior cingular cusp. It is
discussed on p. 509. The last upper postcanine of this specimen is also of a
simpler construction than the penultimate tooth. This is the only specimen
in the series in which this phenomenon was observed.
Specimen E (fig. 5 B, GC, D)
This specimen is larger than D, but the individual teeth are smaller than
those of D.
Seven functional lower postcanines are present. The remnant (r.pc.) of a
tooth is present anterior to the Ist postcanine. Partially erupted 8th post-
canines were present on both sides, but have been lost from the specimen.
These teeth appeared to have had no functional predecessor as there are no
indications of a root remnant in their alveoli. Unfortunately only the 6th and
7th postcanines are well preserved. The crowns (fig. 5 CG and D) of these two
teeth are characterized by a mixture of the features present on the crowns of
the 6th and 7th postcanines of specimen D. In both, the accessory cusps are
nearly as large as the main cusps. In the 6th postcanine the anterior cingular
cusp lies almost directly in front of the anterior accessory cusp, but the posterior
cingular cusp lies slightly postero-lingually to the posterior accessory cusp.
Two well-developed cingular cusps (c.c.) are present on the lingual surface at
the base of the anterior accessory cusp. These are followed posteriorly by three
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TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY
Se OEE
492 ANNALS OF THE SOUTH AFRICAN MUSEUM
minute cuspules. A well-developed cusp is present on the lingual surface of the
base of the posterior cingular cusp. This tooth is almost identical with the 6th
postcanine of specimen D. In the 7th postcanine the anterior cingular cusp is
presumably represented by a well-developed cusp antero-lingual to the anterior
accessory cusp. A single large cingular cusp (c.c.) is present below the junction
of the anterior cingular cusp and anterior accessory cusp. No further cusps are
1CM.
pc.1.
Fic. 6. Thrinaxodon sp., Specimen F.
A and B, upper dentition; C and D, lower dentition.
Key to lettering p. 521.
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 493
present behind this cusp between it and the posterior cingular cusp. A well-
developed cusp is present on the labial surface (a.l.c.) antero-labial to the apex
of the anterior accessory cusp when seen in crown view. The 5th postcanine
apparently had a high dominant main cusp. The anterior cingular cusps are
arranged to form a ridge not unlike a similar structure of the 3rd and 5th
postcanine of specimen D.
Specimen F (Fig. 6C and D and fig. 7)
The lower postcanines of specimens F and G are more similar to one
another than they are to those of specimens D and E, but because of numerous
similarities all four specimens have been placed in Group II.
Gicic ~~ \t AC
aCCa-a Ss : Wn
‘ ul Ay
(RAW
fc
Fic. 7. Thrinaxodon sp., Specimen F.
A and B, lingual and crown views of the 7th lower postcanine.
Key to lettering p. 521.
Seven functional postcanines are present in the right mandible (fig. 6 D).
They are preceded by the remnant of a tooth (r.p.c.) and an unerupted and
partially formed tooth (a.p.c.) is present in a crypt behind the last postcanine.
The crowns of the individual postcanines are not sufficiently well preserved to
permit a detailed comparison of the crown structures of odd and even post-
canines, but the differences do not appear to be as marked as in specimens C
and D. As in other specimens there is a progressive complication of the crowns
in a posterior direction. The crowns are characterized by well-developed
cingular cusps. These are particularly well preserved in the 7th functional
postcanine (fig. 7). In this tooth the accessory cusps are large (a.a.c., p.a.c.),
but not as large as the main cusp. An anterior cingular cusp is present antero-
lingual to the anterior accessory cusp. It is followed by two large cingular
cusps (c.c.). The posterior portion of the cingulum consists of a series of six
cuspules which, with the exception of one, increase progressively in size in a
posterior direction. Well-developed cuspules are present on the labial surface
of the crown at the base of the anterior and posterior accessory cusps (a.l.c.,
p.l.c.). With the exception of the cusps on the posterior portion of the cingulum
this tooth is almost identical to the 7th postcanine of specimen E (fig. 5 C
and D).
ANNALS OF THE SOUTH AFRICAN MUSEUM
494
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TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 495
Specimen G (fig. 9)
The jaw contains eight functional postcanines. The 8th, 5th and 3rd
appear to have erupted shortly before death. The 6th tooth is missing. The
remaining teeth are all badly worn. This is the only specimen in which post-
canines show advanced signs of wear and in which replacement does not
appear to be strictly alternate.
The crown of the 5th postcanine (fig. 9 B and C) is almost identical to the
7th functional postcanine of specimen F (fig. 7) except that the posterior
cingular cusp (p.c.c.) lies lingual to and not posterior to the posterior accessory
‘cusp. The 8th postcanine of specimen G is identical to the 5th except that it
lacks the posterior labia] cusp and is more longitudinally ovate. The anterior
part of the cingulum of the grd postcanine does not support the three
large cusps that are present in this position in the 5th and 8th postcanine, but
consists of a series of six small cusps arranged to form a ridge which terminates
anteriorly at the base of the anterior accessory cusp. The posterior cingulum
supports a single well-developed cusp (p.c.c.) postero-lingual to the posterior
accessory cusp. This tooth is similar to the 5th functional postcanine of specimen
D (fig. 4B). In postcanines nos. 8, 5 and 3 the main cusp becomes progres-
sively larger in an anterior direction and the accessory cusp progressively
smaller. The 2nd and 4th postcanines although badly worn are characterized
by large anterior cingular cusps and poorly developed posterior cingular cusps.
In this respect they are not unlike the 6th postcanines of specimen D (fig. 4 B)
and Specimen E.
Upper postcanines, Specimen D (figs. 3 A and 4 A)
Without exception the upper postcanines have crown patterns less complex
than those of the corresponding lower teeth. Six functional postcanines are
present in specimen D. The 6th is preserved partially erupted. The 1st and grd
appear to have erupted shortly before death. The 5th already has a large pit
for a replacing tooth at its base. The Ist postcanine (fig. 4 A) consists of a large
main cusp, which is slightly recurved; the anterior accessory cusp is missing,
but a small posterior accessory cusp (p.a.c.) is present. A cingulum and cingular
cusps are absent. The 3rd postcanine has a large main cusp which is flanked by
smaller anterior and posterior accessory cusps. A small posterior cingular cusp
is present on the lingual surface at the base of the anterior accessory cusp. No
other cingular cusps are present. The 5th postcanine is similar. The 6th post-
canine is almost identical to the Ist postcanine except that it possesses a very
small anterior accessory cusp. The 2nd postcanine appears to have both anterior
and posterior cingular cusps.
Specimen E (fig. 5 A)
Six functional postcanines are present. The 1st, 3rd and 5th are only
partly erupted indicating that they erupted shortly before death. The structure
of the crowns of the postcanine teeth are identical to those of specimen D.
ANNALS OF THE SOUTH AFRICAN MUSEUM
496
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TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 497
Specimen F (fig. 6 A and B)
Six functional postcanines are present in specimen F.. The Ist, 3rd and 5th
erupted shortly before death and well-developed replacing teeth are present in
pits at the bases of the older 2nd, 4th and 6th postcanines. The crypt for an
additional tooth is present behind the 6th postcanine. The structure of the
crowns of the upper postcanines is identical to those of specimens D and E.
Specimen G (fig. 8)
Six functional teeth are present. In the even-numbered series the 2nd
and 4th have well-developed pits for replacing teeth above them, but the
replacing tooth for the 6th had already erupted before death. The 2nd and
3rd teeth are almost identical in structure and possess well-developed anterior
and posterior cingular cusps. The 4th is badly worn. The 5th and 6th lack
well-developed posterior cingular cusps. The 6th has two well-developed cusps
on the labial side of the tooth (p.l.c., a.l.c.) one above the anterior and one
above the posterior accessory cusp.
Group III
Lower postcanines. Specimen H (fig. 10 G and D)
The lower postcanines of this specimen are characterized by poor develop-
ment of the cingulum and cingular cusps. In this respect they are quite different
from the lower postcanines of Groups I and II. The lower postcanines of
specimen H closely resemble the upper postcanines of this and other specimens.
Seven functional postcanines are present. The crown of the 7th consists of the
characteristic main cusp, anterior and posterior accessory cusps, and anterior
and posterior cingular cusps. In the 5th and grd the crown structure is similar
except that the main cusp becomes progressively more dominant in an anterior
direction.
Upper postcanines. Specimen H (fig. 10 A and B)
Six functional postcanines are present in the right maxilla. The cingular
cusps are very poorly developed and appear to be entirely absent on most of
the postcanines.
Discussion
Extension of Growth series of ‘Thrinaxodon’
The present study has confirmed Parrington’s (1936) conclusion that
alternate replacement took place in the postcanine teeth of Thrinaxodon. Parring-
ton showed that it was possible to divide the specimens of Thrinaxodon that he
studied into two groups; a younger group in which the Ist, 3rd and 5th maxillary
postcanines were younger than the remaining ones and an older group in which
the 2nd, 4th and 6th maxillary postcanines were younger than the others.
498 ANNALS OF THE SOUTH AFRICAN MUSEUM
The skull lengths of Parrington’s younger group vary between 61 mm. and
79 mm. (see table I). One of Parrington’s specimens, viz. his E (C in the
present investigation), was prepared in acid and it has been confirmed that the
Ist, 3rd and 5th maxillary postcanines were the younger. Four additional
skulls falling within this size range were studied. These are specimens D, E,
F and G. In all of them the Ist, 3rd and 5th postcanines were the younger.
The skull lengths of these specimens are given in table I. Parrington concluded
that the 1st, 3rd and 5th teeth were replaced at least once and that the and,
1CM
A ol pc.2 rep.c. 6
CO 2.
nem
1CM
pers:
Fic. 10. Thrinaxodon sp., Specimen H.
A and B, upper and C and D, lower dentitions.
Key to lettering p. 521.
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 499
4th and 6th were also replaced at least once. Parrington’s two stages are
diagrammatically shown in figure 11 A. In this figure they are labelled II
and III. Fully erupted teeth are drawn in black below the alveolar border and
-unerupted replacing teeth in dotted lines above the alveolar border. Only the
portion below the alveolar border of partially erupted teeth is shown in black. .
Parrington’s younger
group
Parrington’s older
group
Fic. 11. Tooth replacement in Thrinaxodon.
A, diagram to illustrate simple alternate replacement concluded by Parrington
to have been present in Thrinaxodon. Additional growth stages I and IV have
been added.
B, diagram to illustrate a possible way in which posterior migration of the post-
canine row could have been achieved.
In figure 11 A, stage II, the 1st, 3rd and 5th are partially erupted and in
figure 11 A, stage III, the 2nd, 4th and 6th of stage II have been replaced and
partially erupted teeth are present in these alveoli. Specimens both larger and
smaller than those available to Parrington were available for the present
study. See table I.
500 ANNALS OF THE SOUTH AFRICAN MUSEUM
TABLE I
Designation of Length of skull
Designation of specimens in (tip of snout Youngest teeth
specimens in Parrington’s to posterior end
present paper (1936) paper of sagittal crest) I 2 A 5 Omen
36 mm. x x x
B 58 mm. x x x
B 61 mm. x x x
C 63 mm. x x x
Cc E 68 mm. x x x
F 70 mm. x « x
D 73 mm. < x x
E 78 mm. x x x
G 78 mm. x x x
F 78 mm. x x x
G 80 mm. x x x
H 82 mm. x x x
H I 85 mm. x x x
I 86 mm. x x x
In specimen A (fig. 1 A) the end, 4th and 6th postcanines were younger
than the Ist, 3rd and 5th. This skull is incomplete, but as reconstructed by
Brink (19550) it could not have had a skull length of much more than 36 mm.
It is the youngest specimen available and in comparison with older specimens
the teeth are minute. It has seven functional postcanines. This fact is discussed
on p. 502.
In specimen B (fig. 1 A and B) (skull length 58 mm.) the 2nd, 4th and 6th
postcanines were also the youngest. The difference in the size of the corre-
sponding teeth in A and B suggests that several growth stages intervened
between specimens A and B. In specimens with skull lengths between 61 mm.
and 80 mm. the ist, 3rd and 5th teeth were the younger. From 80 mm. to
85 mm. the end, 4th and 6th were the younger. In a damaged Thrinaxodon
skull (referred to as specimen I) in the collection of the South African Museum
with a skull length of approximately 86 mm., the Ist, 3rd and 5th teeth appear
to have been the younger.
The additional material shows that the series available to Parrington
covered only part of the growth series and that the individual postcanines
were replaced several times. The additional growth stages are shown in figure
11 A, stages I and IV. Stage I represents specimens A and B in which the end,
Ath and 6th are the youngest. The older teeth in stage I have been numbered
1 and the younger teeth 2. By the time stage IV is reached the teeth numbered
1 have been replaced twice and, therefore, the teeth in the corresponding
positions have been numbered 3. The teeth numbered 2 in stage I are replaced
once and the teeth in the corresponding position in stage IV are, therefore,
numbered 3. It is, however, apparent from an analysis of the postcanine teeth
of the specimens available that in Thrinaxodon the replacement pattern is more
complex than the simple alternation shown in figure 11 A. Further detailed
consideration of the specimens is therefore necessary.
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 501
Posterior migration of the postcanine row
The number of postcanines in the jaws studied varies slightly. In many
cases remnants of postcanines are present in front of the functional postcanines
and in some cases unerupted or partially erupted teeth are present behind the
functional row. In some cases these teeth do not appear to have had predeces-
sors. This information is summarized in table II. The number of functional
teeth appears to be fairly constant. All the maxillae, with the exception of
specimen A, have six functional teeth and all the mandibles with the exception
of specimens D and G have seven functional postcanines.
TABLE II
UPPER POSTCANINES Rare LOWER POSTCANINES
Anterior No. Unerupted Anterior No. Unerupted
tooth functional posterior _ Total tooth functional posterior Total
remnants p.c. p.c. remnants p.c. p.¢.
A 7 7
B 6 6
C 6 ORT renin) 7 (1) 9
D 6 6 (1) 8 9
E 6 LO (1) 7 (1) 9
[8 6 (1) 7 (1) 7 (1) 9
G 6 (1) 7 8 8
H 6 (1) 7 7 7
The constant number of functional postcanines in most specimens, the
frequent occurrence of a tooth remnant anterior to the functional row and the
presence behind the postcanine row of an unerupted postcanine which does
not appear to have had a predecessor, suggest that during growth the anterior
postcanines were lost and that new postcanines were added posteriorly. In
mandibles of specimens C, E, F and G (figs. 2, 5, 6 and g) it would be expected
in terms of alternate tooth replacement that the 1st functional tooth would be
a younger tooth erupted shortly before death or at least that it would have a
well-developed replacing tooth in a pit at its base. This is not the case. In all
four specimens the first functional tooth is firmly held by attachment bone and
there is no indication of a replacing tooth at its base. The fact that the first
functional tooth was not about to have been replaced in these four specimens
suggests that in a more advanced growth stage the first functional tooth became
the tooth remnant anterior to the postcanine row which is preserved in some
specimens. This fact supports the conclusion that postcanines were lost anteriorly
and added posteriorly during growth. In other words, the functional row of
postcanines migrated backwards during growth. This appears to be so arranged
that the number of functional postcanines remained roughly constant during
growth. An explanation of this nature could account for the varying postcanine
count attributed to Thrinaxodon in the literature. For example, if in the lower
jaw the new tooth behind had erupted sufficiently to be considered part of the
502 ANNALS OF THE SOUTH AFRICAN MUSEUM
functional row the number would be given as eight, if not as seven. In figure
11 B, a series of growth stages are shown in which teeth are progressively lost
in front and added behind. This has been superimposed upon the simple
alternate replacement shown in figure 11 A. Although this model appears
to be theoretically sound it only agrees with the state of eruption of the post-
canines in some of the specimens, to wit those in which the 2nd, 4th and 6th
postcanines are replacing. If teeth were added and lost in the manner suggested
in figure 11 B, the state of the postcanine row at each growth stage would
appear to be identical, i.e. it would always appear that the 2nd, 4th and 6th
teeth were replacing because teeth are added and lost at the same rate. This is
clearly not the case as specimens are also available in which the Ist, 3rd and
5th teeth are replacing.
Posterior migration of the postcanine row appears to be a characteristic
of many of the advanced cynodonts, e.g. Scalenodon (Crompton, 1955); South
American gomphodont cynodonts (Patterson and Olson, 1961) and Diademodon
(Fourie, 1963).
There is ample evidence that the canines replaced several times. As the
replacing canines were larger than their predecessors the new teeth invaded
the region occupied by the anterior postcanine. It appears that in order to
compensate for the loss of anterior postcanines further teeth were added at
the back of the tooth row.
Replacement cycle
In figure 12 an attempt has been made to fit the postcanine rows of the
specimens studied into an orderly sequence. This series indicates how posterior
migration of the postcanine row was achieved in such a way that in some speci-
mens the odd-numbered teeth are replacing and in others the even-numbered
teeth are replacing. The upper postcanines are shown in figure 12 A and the
lower postcanines in figure 12 B. In this figure teeth drawn in dotted lines
above the alveolar border in the upper jaw and below it in the lower jaw had
not yet erupted.
The first growth stage (I) in the series in figure 12 A is a diagrammatic
representation of the upper postcanine row in specimen H (fig. 10). The end,
4th and 6th teeth are in the process of erupting, but the 6th is further erupted
than the 2nd or 4th (replacement wave). Replacement teeth are present above
the grd and 5th postcanines, but not above the 1st. An unerupted tooth is present
behind the 6th in position 7. There was no functional predecessor for this tooth.
It is larger than the replacing tooth at position 5.
In specimen A (stage 1 A on the right of fig. 12 A) seven teeth are present,
of which the 2nd, 4th and 6th are the youngest. It is possible that the state of
the postcanine row in specimen A is similar to that of H except that the
unerupted postcanine in position 7 has erupted and is functional.
In a later growth stage (fig. 12 A, II) the 1st functional postcanine which
had no replacement tooth in stage I has been shed. The partially erupted 2nd,
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 503
4th and 6th postcanines of stage I are now fully erupted. Because one post-
canine has been lost the 2nd, 4th and 6th of stage I become the ist, 3rd and
5th of stage II. Consequently the replacing teeth that were developing in stage I
above the grd and 5th replace the previous teeth and erupt in stage II as the
end and 4th. The unerupted additional tooth (position 7) of stage I erupts as
the 6th in stage II. Consequently although in both stages I and II the and,
4th and 6th teeth are the youngest, the teeth referred to are not the same in
both stages.
Replacing teeth are developing above the rst, 3rd and 5th teeth in stage
II. Stage II appears to correspond with specimen B. In this specimen there is
no indication of an unerupted tooth behind the functional postcanine row. As
the functional 1st, 3rd and 5th in specimen B are considerably larger than the
corresponding teeth in specimen A, several growth stages probably separated
these two specimens. This point will be considered again later.
In stage III the replacing teeth developing above the Ist, 3rd and 5th
postcanines in stage II have erupted. In stage II replacing teeth are erupting
in the 2nd, 4th and 6th alveoli. In stage III these are fully erupted. An
additional tooth is developing behind the functional row and the development
of a replacing tooth for the 5th has commenced. The state of eruption and
replacement of the postcanines in this stage corresponds exactly to that present
in specimens E and F (figs. 5 and 6). Specimens C, D and G appear to
be slightly in advance of stage III because the replacing tooth for the 6th
postcanine has erupted and replaced the old functional tooth in this position.
This is illustrated in stage III a on the right of figure 12 A. Stage III is followed
by a stage IV. Apart from the fact that the first functional tooth is now one
position further back, this stage is identical to I, thereby completing the cycle.
The replacement cycle (fig. 12 B) in the lower jaw is basically the same as
that concluded to have been present in the upper jaw. The only real difference
is that the number of functional teeth is seven instead of six.
In growth stage I (fig. 12 B) the 1st, 3rd, 5th and 7th teeth are in the
process of erupting. Replacing teeth are developing below the 2nd, 4th and 6th
postcanines. This stage corresponds with the postcanine row of specimen H.
In the right mandible of this specimen the old functional first tooth had not
been shed, but it was shed on the left. This tooth has been indicated by a dotted
outline in stage I. In specimen H there is no indication of an unerupted post-
canine behind the last functional tooth. Specimen D (fig. 3 B) corresponds with
stage I in regard to the state of eruption of the first seven postcanines, but a small
8th tooth is present behind the postcanine row. The state of eruption of the
postcanines in specimen D is shown diagrammatically in stage I A (on the
right-hand side of fig. 12 B). This specimen is discussed in more detail below.
In stage II the replacing teeth which were developing below the and, 4th
and 6th in stage I are erupting and replacing teeth are developing for the 3rd,
5th and 7th postcanines, but not for the first postcanine. An unerupted tooth
is developing behind the 7th postcanine. The 1st, 3rd, 5th and 7th teeth that
904:
ANNALS OF THE SOUTH AFRICAN MUSEUM
DVS
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY
‘IIMOT “gq pue szoddn ‘vy
‘souruvojsod oy} [Je Jo s[9Ao yuous0¥"]doy ds UOpOXDULLY [.
9) &
506 ANNALS OF THE SOUTH AFRICAN MUSEUM
were partially erupted in stage I have in stage II erupted further. The 7th
is fully erupted, but the Ist is only partially erupted. This stage is not repre-
sented in the material studied. In stage III the partially erupted 1st postcanine
of stage II is fully erupted and retained. The other odd-numbered, functional
teeth (3rd, 5th, 7th) of stage II have, however, been lost in stage III and their
replacing teeth which were developing in stage II are now erupting. The
retention of the Ist postcanine in stage III is presumably correlated with this
tooth having no successor and being younger than the other functional odd-
numbered teeth in stage II. Replacing teeth are developing below the end,
4th and 6th postcanines and a partially erupted tooth is present behind the
functional row. This stage corresponds with the postcanine row in specimens
C, E and F (figs. 2, 5 and 6). In these specimens the remnant of a tooth is visible
anterior to the functional postcanine row. This is discussed below. Specimen
G (fig. 9 A) is similar to stage III in the state of eruption of the first five post-
canines, but the partially erupted 8th postcanine shown in stage III is fully
erupted.
The next stage, IV, is identical to I, except that the first functional tooth
is one position further back. The ist postcanine of stage III which has no
replacing tooth, is retained as a tooth remnant. This remnant would presumably
have been retained in the next three stages of the replacement cycle.
Edmund (1960, 1962) has made a thorough study of tooth replacement in
reptiles. He has shown that the spacing between hypothetical impulses which
initiate the development of teeth and which travel along the free margin of the
dental lamina can account for the fact that in some reptiles the replacement
pattern is strictly alternate, whereas in others waves of replacement pass from
back to front or front to back along alternately numbered tooth series. Back
to front waves are present in all the specimens of Thrinaxodon studied. Edmund
has shown that the basic units in the dentition of all reptiles are <ahnreihen.
He has suggested that Bolk’s term ‘Odontostichi’ be dropped. <ahnrethen
consist of a series of teeth that are either the ist, or the 2nd, or the 3rd, etc.,
teeth developed from successive tooth positions, e.g. all the first teeth to develop
at each successive tooth position form the Ist <ahnrethe, all the second teeth,
the end <ahnreihe, and so on. Development of teeth at each tooth position is
iniated by an impulse travelling backwards along the free margin of the dental
lamina. The spacing between successive impulses determines the spacing
between successive <ahnrethen. Edmund has shown that the functional dentition
in reptiles is the product of several <ahnreihen. In order to obtain a complete
dentition many members of each <ahnrethen must be discarded or resorbed,
for example, as Edmund has shown in a dentition consisting of 21 positions,
*... 15 members of the first <ahnrethe are discarded before the first anlage of
position twenty-one is laid down, and many members of the first seven Zahnrethen
must be discarded before the first mature tooth is produced at position twenty’.
The type of replacement which took place in Thrinaxodon is basically the
same as that which takes place in living reptiles. Therefore, the postcanine row
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 507
of Thrinaxodon also probably consisted of the products of several <ahnrethen.
It should, therefore, be possible to account for cycle of replacement concluded
to have taken place in the postcanine row of Thrinaxodon in terms of Edmund’s
theory of the development of the functional dentitions of reptiles.
In figure 13 an attempt has been made to reconstruct the ontogenetic
stages which led up to the development of the functional lower postcanine row.
These are illustrated in figure 13 A to G. Subsequent changes in the post-
canine row which were concluded in the section above to have taken place are
shown in figure 13 H to L. The embryonic stages were deduced by working
back from figure 13 H in terms of Edmund’s theory.
In a hypothetical early stage, figure 13 A, the first impulse to travel
along the free margin of the dental lamina is shown at position no. 1. It has
initiated the development of the first tooth at this position. The first impulse is
indicated as an arrow followed by the number 1. In a later stage (fig. 13 B)
the impulse has moved to tooth position 2 and initiated the development of a
tooth at this position. The tooth at position 1 is larger. In figure 13 C the first
impulse has passed position 3 where it initiated the development of a tooth.
A second impulse, 2-5 tooth positions behind* the first impulse, has initiated
a second tooth at position 1. In a later stage, figure 13 E, this second impulse
is 2°5 tooth positions in front of tooth position 1 and a third impulse com-
mences in position 1. In figure 13 G the third impulse has initiated the develop-
ment of the 3rd tooth at tooth position 3. Three teeth have, therefore, been
formed at teeth positions 1, 2 and 3. The second impulse is 2-5 tooth positions
in advance of the third impulse and two teeth have been formed in positions
4 and 5. The first impulse has initiated a tooth at tooth position 8 and one
tooth has, therefore, been formed at tooth positions 6, 7 and 8.
In figure 13 H, the 4th impulse has initiated the development of a tooth at
position 2, 2°5 tooth positions behind the 3rd impulse. The 4th impulse did
not initiate the development of a tooth at position 1. The 1st impulse has
faded out and no further teeth are initiated at this stage behind tooth position 8.
In figure 13 H the alveolar border is indicated. The parts of the functional
teeth above the alveolar border have been drawn in black. Unerupted teeth
have been drawn in dotted outline. The teeth drawn in outline only above the
functional teeth are the teeth that were resorbed or shed during development.
Figure 13 H corresponds to stage III of the replacement cycle shown in figure
12 B, i.e. similar to the arrangement of the postcanines in specimens C, E, and
F. In figure 13 H the functional postcanine row is built up of the teeth belonging
to the first three Zahnreihen (Z1, Z2 and Z3). Five teeth have been lost from the
first <ahnreihe and three from the 2nd Zahnreithe. Development of the 4th
Kahnrethe (Z4) has commenced. The rst, 2nd and 3rd functional teeth are
the third teeth to have developed in these positions and are part of the 3rd
Kahnrethe. The 4th, and 5th are the second teeth to have developed in this
*The word ‘behind’ is used with reference to the direction of migration of the hypothetical
impulse. Since the latter is moving from back to front, ‘behind’ means ‘anterior to’.
508 ANNALS OF THE SOUTH AFRICAN MUSEUM
| 3 2) vi 9
AA
Blelala Alo tE
fa
Agel
; a i ra
ete |
7 aRale AN a) o G
ay
(\
(\
Fic. 13. Thrinaxodon sp.
Diagram to illustrate hypothetical development stages
(A-G) leading up to a functional lower postcanine row (H)
and the subsequent replacement cycle (I-L). Based upon
Edmund’s theory.
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 509
position and form part of the second <ahnrethe. The 6th and 7th teeth are the
first to have developed in these positions and form part of the first <ahnrethe.
In a later growth, figure 13 1, the 1st functional tooth of figure 13 H is
lost and no new tooth is present in this position. Therefore, the tooth in position
2 in figure 13 I is the first functional tooth. Figure 13 I is the same as stage IV
or I in figure 12 B. In this stage the condition of the postcanine row is as in
specimen H. In figure 13 I, the 4th impulse has initiated a tooth in position 3
(i.e. beneath the 2nd functional tooth). The 3rd impulse has initiated a tooth in
position 5 and the 2nd impulse has initiated a tooth in position 8. Figure 13 J
is equivalent to stage II in figure 12 B. Figure 13 K corresponds to stage III
in figure 12 B and the 2nd, 3rd and 4th impulses have moved one tooth position
backwards. Figure 13 K is a repetition of figure 13 H. In the former the 2nd
impulse has faded away and the 5th impulse has initiated a tooth at tooth
position 3 (functional 2). Figure 13 L is a repetition of figure 13 I. In figure
13 L the partially erupted tooth in position 3 is the first functional tooth. Thus
Edmund’s theory of successive impulses which initiate teeth and travel along
the free margin of the dental lamina can account for the replacement cycle
for the postcanine series concluded to have taken place in the lower jaw of
Thrinaxodon.
We must now return to consider specimen D, which is peculiar in possessing
eight functional lower postcanines. Furthermore the extra tooth, instead of
being large, is small and simple. On the basis of a scheme of the type shown in
figure 13, it is possible to account for these peculiarities. It can be assumed that
in the development of specimen D, when a stage similar to figure 13 H was
reached, the first impulse did not fade away but initiated a tooth at position 9.
A tooth initiated in this stage would become functional in a later stage similar
to figure 13 I. With the exception of the additional tooth, the state of eruption
of the individual postcanines in specimen D is similar to that shown in figure
13 I. It is possible to account for the simple crown pattern of this tooth if it 1s
assumed that its crown was formed behind the point where the morphogenetic
gradient for ‘molarization’ was at its maximum (see p. 513).
If Edmund’s scheme is followed it is also possible to account for the
replacement cycle concluded to have taken place in the upper postcanines.
These are shown in figure 14. The first stage shown (fig. 14 A) corresponds
exactly with figure 13 G. It is unnecessary to repeat the stages leading up to
figure 14 A as these correspond to those shown in figure 13. For convenience
the teeth in figure 14 are orientated pointing downwards, to indicate that they
are upper teeth. The postcanine row in figure 14 A consists of six functional
teeth and an unerupted tooth behind. This row consists of the products of three
Kahnrethen (Zi to Z’). The 2nd impulse has initiated a tooth at position 5 and
the 3rd impulse a tooth at position 3. The stage represented in figure 14 A
is the same as stage I in figure 12 A. In a Jater stage, figure 14 B, which corre-
sponds to stage II in figure 12 A the 1st functional tooth has been shed and no
new tooth has been initiated in this position. The rst tooth initiated by the 4th
510 ANNALS OF THE SOUTH AFRICAN MUSEUM
impulse is present at tooth position 2. In figure 14 C, which corresponds with
stage III in figure 12 A, the 2nd impulse has initiated a new tooth in position 8,
and the 3rd a tooth in position 5 and the 4th a tooth in position 3. Figure 14 D
is identical to figure 14 A, except that the tooth row has moved one tooth
position backwards. After the stage shown in figure 14C the 2nd impulse
Les eh ne ey ue 10)
Fic. 14. Thrinaxodon sp.
Diagram to illustrate the application of Edmund’s theory
of tooth replacement to the replacement cycle for the
upper postcanines.
appears to fade away. In figure 14 E to I the replacement cycle is continued.
In figure 14 E the 5th impulse has initiated a tooth in position 3. In figure
14 H the 6th impulse has initiated a tooth in position 4.
It is thus possible to account for the state of eruption and replacement of
individual postcanines and the posterior migration of the postcanine row in
several specimens of Thrinaxodon if it is assumed that:
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY HLT
(1) regular impulses, 2:5 tooth spaces apart, travel caudad along the free
margin of the dental lamina;
(2) each new impulse commences one tooth position behind the tooth position
where the previous impulse commenced.
The sequence of a replacement in successive growth stages shown in figure 12
was hased upon the state of eruption of the individual postcanines in a number
of actual specimens. It is important that this sequence agrees with the sequence
based upon Edmund’s theory and shown in figures 13 and 14.
Once the replacement cycle is determined it is possible to estimate the
minimum number of times each postcanine was replaced. The youngest
specimen known (specimen A) appears to correspond with the stage shown in
figure 14 A. The ist and 2nd teeth of this stage are apparently the third teeth
to have developed in these positions, the 3rd and 4th are the second, the 5th
and 6th, the first at their respective positions. The state of eruption of the
postcanines of specimen B corresponds with the stages shown in figure 14 B,
E and H. As there is a considerable difference in the size of the individual
postcanines of specimens A and B (see fig. 1) it is not possible for specimen B
to have been the immediate successor of a specimen similar in size to that of
specimen A. Therefore, several growth stages probably separate specimens
A and B. If specimen A is equated with figure 14 A it is reasonable to equate
specimen B with figure 14 E. Figure 14 F corresponds with specimens QC, D, E,
F and G and Parrington’s specimens E, F, G (see table I). Specimen H and
Parrington’s I corresponds with figure 14 G. A specimen corresponding with
figure 14H is not known. Specimen I corresponds with figure 14 I. In figure
14.1 the 1st functional tooth corresponds with tooth position 4. The postcanine
row in figure 14 I is made up of the members of the grd, 4th, 5th and 6th
Kahnrethen (Z3—-Z*). The 1st functional postcanine (i.e. the tooth at tooth position
4) is the 6th tooth to develop in this position, the 2nd and grd functional are
the 5th to develop in these positions, the 4th functional is the 4th tooth to
develop in this position, the 5th functional tooth is the 3rd tooth to develop in
this position and the 6th functional the 1st to develop in this position.
This means that the 1st functional tooth of an old specimen has been
replaced at least five times, the 2nd and 3rd, four times, the 4th, three times, the
5th, twice, and the 6th not replaced. The fall off in the number of replacements
in a posterior direction is correlated with the posterior migration of the post-
canine row.
Differentiation of the postcanine row
A characteristic of Thrinaxodon is the increase in the complexity of the
crown pattern in a posterior direction. The anterior postcanines are ‘canini-
form’ in appearance and the crown is dominated by the main cusp whereas
the posterior teeth have a complex ‘molariform’ pattern not unlike that of some
early mammals. It has already been shown (Crompton, 1963) that Thrinaxodon
512 ANNALS OF THE SOUTH AFRICAN MUSEUM
was one of the earliest of the mammal-like reptiles which was capable of a
sustained bite across the postcanines. The power of this bite was greatest across
the posterior region of the postcanine row and it is possible to correlate the
complex postcanines in this region with this fact. The simple pattern of anterior
postcanine teeth was possibly correlated with their position close to the canine.
If the anterior teeth had complex crowns they would have interfered with the
stabbing action of the canines to a greater extent than simple ‘caniniform’
postcanines.
In most of the specimens studied the postcanines do not all form a graded
series in which each postcanine is slightly more molariform than the tooth
immediately in front of it. Usually it is possible on the structure of the crowns
divide the postcanine row into two series, one consisting of partially erupted
Fic. 15. Thrinaxodon sp.
Posterior migration and differentiation of the lower postcanines.
postcanines and one consisting of the fully erupted postcanines. The series are
alternately numbered, i.e. the teeth numbered 3, 5 and 7 form one series and
the teeth numbered 2, 4, 6 form the other series. In each of these series there is a
progressive complication of the crown structure in a posterior direction. One
of the major differences between the crown structure of the two series is that
the partially erupted teeth have simpler crown patterns (or were more canini-
form) than the fully erupted teeth immediately anterior or posterior to them.
During growth postcanines are lost from the front of the row and others are
added behind. Consequently if differentiation of the postcanine row into simple
postcanines anteriorly, and complex postcanines posteriorly, is to be maintained
during growth each tooth must be replaced by a tooth with a simpler crown
pattern. This is illustrated in figure 15. The stage of the replacement cycle
shown in figure 15 A corresponds to figure 13 H and that shown in figure 15 B
corresponds to that shown in figure 131. The tooth developing below the
second functional in figure 15 A would become the 1st functional tooth in the
later stage, figure 15 B. It must therefore be more caniniform than the tooth it
replaces. This explanation could account for minor differences between adjacent
teeth, but in fact the differences are far greater than could be accounted for in
this way. It has been noted above that it is possible to divide the row up into
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 513
‘two series. Teeth belonging to one series have different crown patterns from
teeth belonging to the other series. The differences between the two series in
specimen D are described in detail on p. 486.
The complication of the teeth in a posterior direction suggests that they
develop under the influence of a morphogenetic gradient, in the manner which
has been suggested by Butler (1939) for mammalian dentitions. Butler postulates
Fic. 16. Thrinaxodon sp.
Diagram to illustrate posterior migration and
changes in the morphogenetic gradient respon-
sible for the structure of the lower post-
canines.
a short steeply graded field of ‘caninization’ determining the form of the canine
tooth, succeeded by and slightly overlapping a longer, more gently graded
molarization field, affecting the entire postcanine row. The teeth immediately
behind the canine will therefore show little molarization since they develop
under the influence of the low anterior end of the molarization gradient and
are also slightly affected by the caninization field. Further back the canine field
vanishes and the molarization field becomes increasingly strong, so that the
514 ANNALS OF THE SOUTH AFRICAN MUSEUM
teeth become progressively more molarized from front to back of the postcanine
row. In figure 16 the morphogenetic gradient of the molarization field in a
series of growth stages has been shown diagrammatically. For simplicity sake
the caninization field is not shown. It can be imagined as lying directly in
front of the 1st postcanine and fading out rapidly, so that its influence does not
extend beyond the Ist postcanine. During growth the gradient migrates in a
posterior direction. In figure 16 A the gradient commenced at tooth position
1. In figure 16 D the gradient had shifted to tooth position 2. In the intervening
stages a gradual shift of the gradient in a posterior direction presumably took
place.
In specimens C, D, E and F the most molariform teeth in the lower jaw
have well-developed cingular cusps and the main cusps are about the same size
as the accessory cusps. In the larger specimen G the most molariform tooth is
less molariform than those in specimens QC, D, E and F. In this tooth the cingular
cusps are well developed, but the main cusp is relatively larger. In the larger
specimen H, the most molariform tooth is less molariform than that of specimen
G. The main cusp is large and the cingular cusps are poorly developed. This
suggests that the morphogenetic gradient is very steep in early stages, but levels
off in later growth stages. In the four growth stages shown in figure 16, this
feature is shown diagrammatically. The replacing teeth for ist, 3rd and 5th
teeth in stage A develop when the gradient is steep and commences at tooth
position 1, whereas the replacing teeth for the 2nd, 4th and 6th teeth develop
at a later stage, B, when the gradient is further posterior and is less steep.
The replacing teeth developing in stages A and B form the functional post-
canines in stage C. The structure of the crowns of Ist, 3rd and 5th in stage C
increases in complexity in a posterior direction, but despite this their crown
patterns are distinct from those of the and, 4th and 6th. It may be possible to
correlate this with the fact that the two series developed under different gradients
occupying different positions. Take, for example, the 3rd, 4th and 5th functional
postcanines in stage C. The 4th tooth is formed in stage B. The strength of the
gradient at this point in stage B has been indicated and labelled Y. The 3rd
and 5th functional teeth of stage C are formed in stage A. The strength of the
gradient affecting these teeth is indicated and labelled a and b. As a and 3d are
both larger than » the 3rd and 5th teeth will be more molariform than the
fourth.
This conclusion is highly speculative and well-prepared additional material
is required to confirm these results. If it is correct it would explain why a
recently erupted tooth has a simpler and different crown pattern than the
postcanines anterior and posterior to it. It is of some interest that the changes
in the ontogeny of Thrinaxodon are of essentially the same type as Butler (1939)
suggests have been operative during mammalian phylogeny.
In the descriptive section the specimens were divided into three groups.
This division was based on the structure of the postcanine teeth in the mandible.
The different crown patterns are correlated with skull length, and it is therefore
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 5)
possible that these differences are related to changes which the morphogenetic
gradient undergoes during growth. Consequently these differences probably
have no taxonomic importance. Until this is confirmed or denied no attempt
should be made to place the different groups in different species.
NOTES ON THE POSTCANINE ROW OF GOMPHODONT CYNODONTS
Diademodon is characterized by a differentiated postcanine row (Brink,
1955a, 1957; Crompton, 1955). Three distinct regions can be recognized in
both upper and lower jaws—an anterior, central and posterior. The three to
four postcanines of the anterior region have simple conical crowns with a slight
development of a cingulum on the lingual edge. These teeth are small in com-
parison with the remaining postcanines. The central region consists of about
seven postcanines with transversely ovate gomphodont crowns. In this region
wear of the crown surfaces decreases progressively in a posterior direction.
The size of the crowns increases progressively in a posterior direction. The
posterior region consists of about four teeth. In this region the crowns become
progressively more longitudinally ovate in a posterior direction. Fourie (1963)
has referred to the crown patterns of these four teeth as ‘intermediate gompho-
dont’, ‘intermediate sectorial’, ‘large sectorial’ and ‘small sectorial’.
Diademodon specimens covering a wide size range are known. In all speci-
mens the postcanine row is differentiated in the same way and in all the speci-
mens the postcanines of the central region are characterized by a progressive
decrease, in a posterior direction, of the wear of the crown surfaces and a pro-
gressive increase, also in a posterior direction, of size. Brink (1957) has noted
‘.. . that replacement would tend to cause an irregular series of differently
sized teeth’. An irregular series of postcanines in the central region has never,
to the author’s knowledge, been observed in Diademodon.
The teeth of the central region are considerably larger in older specimens
of Diademodon than in younger specimens. Tooth replacement must therefore
have occurred in this region. Replacement must have taken place in such a way
that the postcanines of both the central and posterior regions always formed a
graded series. How was this achieved? In order to answer this question it 1s
necessary to review the evidence of replacement of the postcanines in
Diademodon.
Fourie (1963) has found evidence for the loss of anterior postcanines during
growth. He has also found unerupted postcanines which had no predecessor
behind the functional postcanines. He has concluded that during life the post-
canine row migrated posteriorly. The number of functional teeth remained
constant as teeth were lost and added at roughly the same rate. Broom (1913)
has reported replacement of some of the anterior conical postcanines by new
conical postcanines. Fourie will also describe several specimens showing replace-
ment in this region.
In a specimen of Diademodon housed in the Geologish-Palaontologisches
Institut und Museum in East Berlin the right maxilla contains two conical
516 ANNALS OF THE SOUTH AFRICAN MUSEUM
anterior postcanines. ‘These are followed by a small, greatly worn transversely
ovate postcanine. This tooth is the first of the central region consisting of post-
canines with transversely ovate crowns. In the left maxilla three and not two
anterior postcanines with sharply pointed conical crowns are present. The
posterior of the three appears to have erupted shortly before death and shows
no sign of wear. There is no doubt that numerically this tooth corresponds with
the greatly worn transversely ovate postcanine in the right maxilla. This sug-
gests that replacement in the left maxilla was in advance of that in the right
and also that a simple conical postcanine replaced a transversely ovate post-
canine. The author (1955) has reported the replacement of the 6th postcanine
(second postcanine of the central region) in Diademodon. The replacing tooth
appears to be more conical than transversely ovate.
a (Rocco ae) (LB
Sek cy) lO) 5h ce es one a) Te ee 0S
| } eh i
T | Bus 48 bays Ge : i gu Gs ie w
Fic. 17. Diademodon sp.
Diagram to illustrate replacement and posterior migration of the postcanine
row.
The author (1955) has also claimed that in two specimens of Diademodon
one of the upper longitudinally ovate sectorial postcanines in the posterior
region was replaced by a tooth with a gomphodont crown pattern. This claim
was based upon the fact that a longitudinally ovate foramen which follows
immediately after an alveolus for a gomphodont tooth is present in these speci-
mens below the unerupted gomphodont postcanine. This foramen was inter-
preted as an alveolus for a sectorial postcanine. Brink (1957) has challenged
this view and has offered an alternative interpretation for the longitudinally
ovate foramina behind the functional postcanine row. He has claimed that
‘. , . these openings could equally well be interpreted as ordinary gubernacular
canals’. I find this interpretation difficult to accept for two reasons: (1) in most
specimens of Diademodon the postcanine row is terminated by sectorial post-
canines; (2) replacement of sectorial teeth by gomphodont teeth at the back
of the postcanine row has also been reported in Crvicodon (Crompton, 1955)
and in Trirachodon (Parrington, 1960). Fourie (1963) has confirmed the replace-
ment of sectorial teeth by gomphodont teeth at the back of the row in Diademo-
don. He has shown that the ‘intermediate gomphodont postcanines’ are replaced
by ‘gomphodont’; the ‘intermediate sectorial’ by ‘intermediate gomphodont’
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY Rly
and, ‘large sectorial’ by ‘intermediate sectorial’, a ‘small sectorial’ by ‘large
sectorial’.
In figure 17 the information listed above has been applied to a growth series
of Diademodon. In the first growth stage I the postcanine row consists of fifteen
teeth. The 1st to the 4th are conical, the 5th to the 11th are gomphodont.
In a posterior direction the latter increase progressively in size and wear
decreases progressively. The i2th is intermediate gomphodont, the 13th
intermediate sectorial, the 14th large sectorial and the 15th small sectorial.
There is evidence that the 1st and 2nd postcanines were lost, that the 3rd—6th
and 12th—-15th postcanines were replaced and that teeth were added pos-
teriorly. If teeth are lost anteriorly and added posteriorly then it follows that if
the differentiation of the postcanine row is to be maintained that:
(1) the grd and 4th postcanines must have been replaced by conical teeth;
(2) the 5th and 6th postcanines must have been replaced by conical teeth;
(3) the 12th postcanine must have been replaced by a gomphodont tooth;
the 13th by an intermediate gomphodont; the 14th by intermediate sec-
torial and the 15th by a large sectorial;
(4) the new tooth at the back of the row must be a small sectorial type.
All this is in agreement with the available evidence. If replacement of this
nature is repeated differentiation of the postcanine row will be maintained
throughout the period of growth. The crowns of the central region will always
form a graded series because as greatly worn anterior teeth of the series
are replaced by conical teeth new unworn gomphodont teeth are added
behind.
The description above accounts for the maintenance of a fully differentiated
postcanine row, and it is interesting to speculate on the growth stages that led
up to the formation of a fully differentiated postcanine row. The youngest
Diademodon specimen that has been described (Fourie, 1963) already has a
fully differentiated row. In this specimen, however, there is no evidence that
teeth had been lost anteriorly. The progressive increase of the wear of the
crowns of the central region suggests that during growth these teeth were added
one after the other after the animal had started to use its teeth. This implies that
Diademodon commenced active life with only a few teeth. This also suggests that
the posterior region of the row was only added after a full complement of
gomphodont teeth had erupted. As mentioned above the youngest available
specimen does not appear to have lost anterior teeth. This may suggest that
replacement only commenced after a complete postcanine dentition had
erupted.
The above discussion of the growth of the postcanine row is highly specu-
lative, but the available evidence on the growth of the postcanine row of
Middle Triassic cynodonts tends to confirm at least some of the coriclusions
518 ANNALS OF THE SOUTH AFRICAN MUSEUM
reached above. A good growth series of the jaws of Scalenodon has been described
(Crompton, 1955).
The youngest available specimen of Scalenodon has five postcanines. These
are all transversely ovate and wear decreased in a posterior direction. Conical
anterior postcanines are not present. During growth postcanines were added
progressively behind to increase the length of the postcanine row. The largest
specimen has eleven postcanines. Sectorial teeth were only added at an advanced
growth stage. These were replaced by transversely ovate teeth. Teeth were lost
in front during growth. This process appears to have commenced at an early
stage, but appears to have been slower than addition behind.
Olson and Patterson (1961) have stated that in undescribed South Ameri-
can cynodonts teeth were also lost anteriorly and added behind. Kiihne (1956)
has made a similar observation in the tritylodontid Oligokyphus.
In no gomphodont cynodonts is there any indication of alternate replace-
ment in the postcanine row. It appears rather that postcanines are added
progressively from front to back and that some of these teeth are progressively
replaced. In Dzademodon some of the anterior and posterior postcanines are
replaced, in Scalenodon only posterior postcanines are apparently replaced and
in Oligokyphus none of the cheek teeth were replaced.
In most placental mammals teeth are laid down progressively from front
to back. Edmund (1960) has pointed out that as the deciduous dentition and
the molars are the first teeth to be formed in the positions they occupy, they
constitute a single <ahnrethe. ‘The permanent incisors, canines and premolars
constitute a second <ahnrethe. In Thrinaxodon and other early cynodonts the
products of several <ahnrethen function simultaneously and alternate replace-
ment is present. The absence of alternate replacement in gomphodont cynodonts
and the progressive addition of teeth suggest that gomphodont cynodonts
paralleled to some extent later development in mammals. Large sections of the
functional postcanine row of Diademodon appear to form part of a single <ahn-
reihe since teeth were added progressively without anterior replacement. ‘Teeth
numbered 1 to 15 in figure 17 (I) may constitute a single <ahnrethe. The teeth
that replaced them may constitute a second <ahreihe of which the central
members had been suppressed.
SUMMARY
1. Tooth replacement is described in eight specimens of the cynodont
Thrinaxodon liorhinus Seeley. It is shown that replacing teeth developed in pits
lingually to the functional teeth. During growth the replacing teeth migrated
labially to lie below the crowns of the functional teeth. It is concluded that the
replacement cycle of individual teeth is identical to that reported for Ericzo-
lacerta (Crompton, 1962).
The replacement cycles of the entire postcanine row (both upper and
lower) are deduced from the state of eruption of the individual postcanines of
the specimens studied.
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY 519
2. Parrington’s (1936) observation that the teeth replaced alternately is
confirmed. It is shown that a wave of replacement passed from back to front
along both the odd and even numbered postcanines.
3. The loss of anterior postcanines and the addition of further teeth at the
back of the row resulted in a posterior migration of the functional postcanine
row. This was probably correlated with the increase in size of each successive
replacing canine.
The stages at which teeth were lost anteriorly and added posteriorly is
illustrated.
It is shown that Parrington’s (1936) series of Thrinaxodon only covered
part of the growth period. Older and younger specimens are described.
4. It is shown that it is possible to interpret the replacement cycle in
Thrinaxodon in terms of Edmund’s (1960) interpretation of reptilian tooth
replacement. It is shown that in Thrinaxodon erupting postcanines usually had
simpler (less molariform) teeth than the fully erupted postcanines immediately
anterior or posterior to them. It is suggested that the posterior migration of the
postcanine row can only partially explain this phenomenon.
It is suggested that the postcanines of Thrinaxodon developed under the
influence of a morphogenetic gradient (Butler, 1939). Teeth developed under
the influence of the anterior portion of the gradient were ‘caniniform’ and
those developed under the posterior portion of the gradient ‘molariform’. It is
suggested that in the early stages the gradient was steep, but that in older
specimens the gradient levelled off and moved progressively caudad. It 1s
concluded that the younger teeth in each specimen (i.e. partially erupted)
developed under the influence of a different gradient than the older teeth in the
same specimen. It is concluded that this could account for the difference in
crown structure between old and new postcanines. It could also account for the
fact that teeth developed at the same time would have had the same basic
crown structure.
5. Tooth replacement in gomphodont cynodonts is discussed. It is shown
that in Diademodon the postcanine row is divided into three regions. The post-
canines of the central region always form a structurally graded series. It is
shown that during growth teeth were lost anteriorly and added posteriorly;
it is shown that the postcanine row continued to be differentiated in the same
way despite the posterior migration of the postcanine row during growth. It is
concluded that replacement was limited to the anterior and posterior regions
and to the anterior and posterior members of the central region. No replacement
took place along the remainder of the central region.
It is concluded that during growth teeth were added progressively in
Diademodon and that large sections of the postcanine row of Diademodon con-
stituted a single <ahnreihe. It appears that in this aspect Diademodon paralleled
later mammals.
520 ANNALS OF THE SOUTH AFRICAN MUSEUM
ACKNOWLEDGEMENTS
I am deeply indebted to Dr. E. White, F.R.S., of the British Museum for
permission to study and describe the Thrinaxodon skulls and also for the generous
facilities which he and his staff placed at my disposal during my visit to the
British Museum in 1960. I wish also to record my thanks to Professor R. Dehm
and Dr. A. 8. Brink for permission to study and describe the Munich and
Johannesburg material. I wish to record my thanks to the C.S.I.R. of South
Africa, the West German Government and the Royal Society and Nuffield
Foundation for travel and study grants that enabled me to undertake the work
described in this paper.
The South African Museum specimens were beautifully prepared by
Mr. C. Gow.
I am extremely grateful to Dr. F. R. Parrington, F.R.S., for permitting
me to study his collections and for reading and criticizing the manuscript of
this paper.
I am deeply indebted to Dr. R. F. Ewer for reading the first draft of this
paper and for making numerous suggestions and corrections.
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.
REFERENCES
Brink, A. S. 1955a. A study of the skeleton of Diademodon. Palaeont. Afr. 35 3-39.
Brink, A. S. 1955b. Note on a very tiny specimen of Thrinaxodon liorhinus. Palaeont. Afr. 35 73-78.
Brink, A. S. 1957. Speculations on some advanced mammalian characteristics in the higher
mammal-like reptiles. Palaeont. Afr. 4, 77-06.
Broom, R. 1913. On evidence of mammal-like dental succession in cynodont reptiles. Bull.
Amer. Mus. nat. Hist. 32, 465-468.
But Ler, P. M. 1939. Studies of the mammalian dentition.— Differentiation of the postcanine
dentition. Proc. zool. Soc. Lond. 109, 1-36.
Crompton, A. W. 1955. On some Triassic cynodonts from Tanganyika. Proc. zool. Soc. Lond.
125, 617-669.
Crompton, A. W. 1962. On the dentition and tooth replacement in two bauriamorph reptiles.
Ann. S. Afr. Mus. 46, 231-255.
Crompton, A. W. 1963. On the lower jaw of Diarthrognathus and the origin of the mammalian
jaw. Proc. zool. Soc. Lond. 140, 697-753.
Epmunp, A. G. 1960. Tooth replacement phenomena in lower vertebrates. Contr. R. Ont. Mus.
Life Sci. Div. No. 52, 1-190.
Epmunp, A. G. 1962. Sequence and rate of tooth replacement in the Crocodilia. Contr. R. Ont.
Mus. Life Sct. Div. No. 56, 1-42.
FourtE, S. 1963. Tooth replacement in the gomphodont cynodont Diademodon. S. Afr. F. Sct.
59, 211-213.
Ktune, W. G. 1956. The Liassic therapsid oligokyphus. London: British Museum.
PARRINGTON, F. R. 1936. On the tooth-replacement in theriodont reptiles. Phil. Trans. (B) 226,
I2I-142.
PARRINGTON, F. R. 1960. Private communication.
PATTERSON, B., AND Oxson, E. C. 1961. A triconodontid mammal from the Triassic of Yunnan.
International colloquium on the evolution of lower and non-specialised mammals, p. 129-191. Brussels:
Paleis der Academién.
TOOTH REPLACEMENT IN THRINAXODON LIORHINUS SEELEY
Key to LETTERING
anterior accessory cusp
attachment bone
anterior cingular cusp
anterior portion of lingual cingulum
anterior labial cusp
additional postcanine
alveolus for first postcanine
alveolus for 8th postcanine
cingular cusp
cingulum
longitudinal groove for the dental lamina
main cusp
pit for replacing tooth
posterior accessory cusp
posterior cingular cusp
Ist postcanine
8th postcanine
posterior labial cusp
pit containing developing crown of a replacing canine
replacing canine
replacing incisor
redundant or suppressed postcanine
crown of the 6th replacing postcanine
521
[ CONTINUED FROM INSIDE FRONT COVER
References thus appear as follows:
AUGENER, H. 1913. Polychaeta. In Michaelsen, W., ed. Beitrdge zur Kenntnis der
Meeresfauna Westafrikas. 2, 67-625. Hamburg: Friederichsen.
EKMAN, S. 1953. <oogeography of the sea. London: Sidgwick & Jackson.
HartTMAN, O. 1948a. The polychaetous annelids of Alaska. Pacif. Sci. 8, 1-58.
Hartman, O. 1948b. The marine annelids erected by Kinberg. Ark. Zool. 42, 1-137.
IzukA, A. 1912. The errantiate Polychaeta of Japan. 7. Coll. Sci. Tokyo. 30, art. 2,
1-262.
Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa.
Ann. Mag. nat. Hist. (10), 11, 487-509.
SYNONYMY. Arrangement according to Schenk, E. T. & McMaster, J. H.:
Procedure in taxonomy. and ed. Stanford, Stanford university press, 1948. Bibliogra-
phic references modified.
‘I. Synonymy arranged according to chronology of names.—All published scientific names
by which a species has been previously designated (subsequent to 1758) are listed
in chronological order, with bibliographical references to all descriptions or descrip-
tive citations following in chronological order after each name. .
B. Form of bibliographic references to synonymic names.—'The first reference following
any name in the synonymy should be to the earliest citation of that name. This
should be followed by references to all subsequent citations of the same name,
arranged in chronological order... .’
Bibliographical references modified to consist of author’s name, date of citation,
pagination and illustrations (plates and figures).
Example : —
Eulalia (Steggoa) capensis Schmarda
Eulalis capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1903, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
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Vol. II. 1900-1902 Zoology and Geology (excl. Parts 1-3, 5, 7-8) R 2.75
III. 1903-1905 Zoology (excl. Parts 1-2, 5) mt 3.35
IV. 1903-1908 Palaeontology . 8.08
V. 1906-1910 Geology, Palacontolosy:, Zoology, Anthropology (exe.
Parts 1-2, 5, 7-9) 1.54
VI. 1908-1910 Zoology (excl. Part 1, Index) 5.48
VII. 1908-1913 Palaeontology (excl. Parts 1-4) 1.26
IX. 1911-1918 Botany (excl. Parts 1-2) 6.03
X. 3911-1914 Zoology (excl. Parts 1-3) , 10.54
XI. rg911-1918 Zoology (excl. Parts 2, 7, Index) 6.73
XII. 1913-1924 Palaeontology and Geology Tt.71
XIII. 1913-1923 Archaeology and ee 8.55
XIV. 1915-1924 Zoology 8.04
XV. 1914-1916 Zoology 10.26
XVI. 1917-1933 Botany 8.57
XVII. 1917-1920 Zoology 8.45
XVIII 1921 Zoology 10.33
XIX. 1924-1925 Zoology 8.10
XX. 1924-1926 Zoology x 6.27
XXII. 1925-1928 Palaeontology . 4.85
XXIII. 1925-1926 Zoology 4.25
XXIV. 1929-1938 Anthropology anal Ethnology (cae Part ae 6.00
XXV. 1927-1928 Zoology we st 4.70
XXVI. 1928 Zoology 3.60
XXVII. 1929 Anthropology 3.60
XXVIII. 1929-1932 Palaeontology .. 6.30
XXIX. 1929-1931 Zoology 5.80
XXX. 1931-1935 Zoology 8.88
INDEX _ of papers, authors and subjects published t in rele L XXX 0.23
XXXI. 1934-1950 Palaeontology (excl. Parts 1-2) 4.20
XXXII. 1935-1940 Zoology 7.62
XX XIII. 1939 Zoology 5-05
XXXIV. 1938 Zoology 5-75
XXXV. 1956 Zoology 14.4.0
XXXVI. 1942-1948 Zoology 6.13
XXXVII. 1947-1952 Archaeology 4.33
XXXVIII. 1950 Zoology 9.00
XXXIX. 1952 Zoology 6.55
XL. 1952-1956 Botany 3.18
XLI. 1952-1955 Zoology 9.60
XLIT. 1953-1956 Palaeontology 8.10
XLII. 1955-1957 Zoology and Palacontolosy 4.80
XLIV. 1957-1959 Zoology and Palaeontology 6.64
XLV. 1959-1960 Zoology and Palaeontology 11.33
XLVI. 1961- Zoology and Palaeontology (Parts 1— io 8.19
Copies may be obtained from—
The LIBRARIAN, SoutH ArFricaAn Museum, CAPE Town.
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