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ANNALS

OF THE

SOUTH AFRICAN MUSEUM

VOLUME VII.

ANNALS

OF THE

SOUTH AFRICAN MUSEUM

VO L UME VII.

DESCRIPTIONS OF THE PALyEONTOLOGICAL MATERIAL

COLLECTED BY THE OFFICERS OF THE GEOLOGICAL SURVEY

OF CAPE COLONY AND OTHERS.

WOODS
MOL.E,
MA

PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

AND

THE GEOLOGICAL COMMISSION OP THE COLONY OP THE

CAPE OF GOOD HOPE

BY WEST, NEWMAN & Co., LONDON
1913.

TKUSTEES OF THE SOUTH AFEICAN MUSEUM.

The Right Hon. JOHN XAVIER MEURIMAN, M.L.A.

THOMAS Mum, C.M.G., LL.D., M.A., F.B.S., F.K.S.E., Superintendent-General

of Education.
JOHN WILLIAM JAGGEH, M.L.A., F.E.Stat.S.

SCIENTIFIC STAFF OF THE SOUTH AFEICAN

MUSEUM.

Loots ALBERT PERINGUEY, D.Sc., F.Z.S., F.E.S., Director.

WILLIAM FREDERICK PURCELL, B.A., Ph.D., Keeper of Land Invertebrates, except

Insects.
ARTHUR WILLIAM ROGERS, D.Sc., F.G.S., Keeper of the Geological and Minera-

logical Collections.

ROBERT BROOM, D.Sc., M.D., Keeper of the Palfeontological Collections.
E. P. PHILLIPS, M.A., F.L.S., Assistant in Charge of the Herbarium.
K. H. BARNARD, B.A., Assistant in Charge of Fish and Marine Invertebrate

Collections.
S. H. HAUGHTON, B.A , Assistant in Charge of Geological Collections.

LIST OF CONTBIBUTOKS.

PAGE

C. W. ANDREWS.

Description of a New Plesiosaur (Plexiosanrus capensis, sp. nov.) from the

Uitenhage Beds of Cape Colony 309

E. BROOM.

The Fossil Fishes of the Upper Karroo Beds of South Africa 251

Notice of Some New South African Fossil Amphibians and Reptiles 270

On a large Extinct Species of Biibalis '279

On Evidence of a Large Horse recently Extinct in South Africa 281

On the Shoulder Girdle of Cynognathns 283

An Attempt to Determine the Horizons of the Fossil Vertebrates of the

Karroo 285

On the Dinosaurs of the Stormberg, South Africa 291

On a New Species of Propappus, and on the Pose of the Pareiasaurian

Limbs 323

On a Species of Tylosanrus from the Upper Cretaceous Beds of Pondoland 332

On a New Type of Cynodont from the Stormberg 334

On Some Points in the Structure of the Dicynodont Skull 337

On the Manus and Pes of Parciasatinis 353

On a New Mesosaurian Reptile (Noteosaurus africanus) 358

A Revision of the Reptiles of the Karroo 361

F. L. KITCHIN.

The Invertebrate Fauna and Palaeontological Relations of the Uitenhage

Series 21

W. D. LANG.

Polyzoa and Anthozoa from the Upper Cretaceous Limestone of Need's

Camp, Buffalo River 1

H. WOODS.

Echinoidea, Brachiopoda, and Lamellibranchia from the Upper Creta-
ceous Limestone of Need's Camp, Buffalo River 13

LIST OF PLATES.

III.

IV.

V.

VI.

f Filisparsa ramosa, iVOrbigny.

Filisparsa fragilis, Marsson.

Diastopora compressa (Goldfuss).

Idmonea virgula, d'Orbiijny.

Crisina cenomana, d'Orbigiiy.

Crisina excavata, d'Orbigiiy.

Crisina marginata, d'Orbigny.
L-! Tervia dorsata (von Hagcnoic).

Tervia gibbera, Gregory.

Entalophora virgula (von Hagcnoii-).

Entalophora con ju gat a (yon Reims').

Entalophora echinata (Rumer).

Spiropora verticillata (Goldfuss).

Entalophora madreporacea (Goldfuss).
[ Coptosoma capense, sp. nov.

f Serpula cf. concava (J. Sow.).
Pecten (Syncyclonema) orbicularis, J. Sou-.
Pecten (Camptonectes) cottaldinus, d'Orbiijny
Pecten (Camptonectes) projectus, Tatc.
Pecteii (Chlamys) cf. subacutus, Linn.
Lima (Acesta) obliquissima, Tatc.
II.- Lima (Mantellum) neglecta, Tate.
Perna atherstoni, Sharpe.
Mytilus uitenhagensis, sp. nov.
Modiola baini, Sharpe.
Nucula uitenhagensis, sp. nov.
Grammatodon jonesi (Tatc}.
Unio uitenhagensis, sp. nov.

Trigonia ventricosa (Krauss).
Trigonia kraussi, sp. nov.
Trigonia rogersi, sp. nov.

Trigonia rogersi, sp. nov.
Trigonia holubi, sp. nov.

Trigonia herzogi, sp. nov.
Trigonia rogersi, sp. nov.

Trigonia vau, Sharpe.
Trigonia stowi, sp. nov.

Vlll

List of Plates.

PLATE

/

VIL-;

VIIL.

Trigonia stowi, sp. nov.

Trigonia conocardiiformis (Krauss).

Cardita nuculoides, Tate.

Astarte (Eriphyla) pinchiniana, Tate.

Anthonya lineata, sp. nov.

Tancredia schwarzi, sp. nov.

Thetironia papyracea (Sharpe).

Thetironia oblonga, sp. nov.

Trapezium? tatei, sp. nov.

Meretrix uitenhagensis, sp. nov.

Meretrix uitenhagensis, sp. nov.
Mactra? dubia, sp. nov.
Pleurornya baini, Sharpe.
Thracia sp.

Turbo atherstoni, SJiarpe.
Turbo rogersi, sp. nov.
Turbo rainutulus, sp. nov.
Natica uitenhagensis, sp. nov.
Natica rogersi, sp. nov.
Natica? mirifica, sp. nov.
Actffionina atherstoni (Sharpe).
Limnsea remota, sp. nov.
Phylloceras rogersi, sp. nov.
Bochianites glaber, sp. nov.
Meyeria schwarzi, sp. nov.

Holcostephanus wilmante, sp. nov.
Holcostephauus cf. baini (Sharpe).
Holcostephanus rogersi, sp. nov.
Meyeria schwarzi, sp. nov.

Holcostephauus cf. baini (Sliarpe).
Holcostephanus rogersi, sp. nov.
Holcostephanus modderensis, sp. nov.
Meyeria schwarzi, sp. nov.

Holcostephanus uitenhagensis, sp. nov.

Hybodus africanus, n. sp.
Ccelacanthus africanus, Broom.
Ceratodus ornatus, n. sp.
Hydropessum kannemeyeri, g. et sp. nov.

Helichthys draper!, S.-Woodicard.
Helichthys brown!, g. et sp. nov.
XIII. -i Oxygnathus brown!, n. sp.
Cleithrolepis minor, n. sp.
Pholidophorus brown!, n. sp.

IX.

X.

XL

XII.

XIV.

XV.

Gryponyx africanus, g. et sp. nov.

Gryponyx africanus, g. et sp. nov.
Massospondylus carinatus, Owen.
Massospondylus harries!, sp. nov.
.Etonyx palustris, g. et sp. nov.

List of New Generic Names.

PLATE

XVI. Massospondylus harriesi, sp. nov.

IMassospondylus harriesi, sp. nov.
^Etonyx palustris, g. et sp. nov.
Geranosaurus atavus, g. et sp. nov.

XVIII. Plesiosaurus capensis, sp. nov.

XIX.- f
VVT - Propappus rogersi, sp. nov.

A.A1. (

Tylosaurus capensis, sp. nov.

IX

XXII.

Tritheledon riconoi, g. et sp. nov.

DATE OF ISSUE OF THE PARTS.

Part 1, July 2, 1908.
Part 2, September 24, 1908.
Part 3, April 28, 1909.
Part 4, July 5, 1911.
Part 5, July 12, 1912.
Part 6, March 20, 1913.

LIST OF NEW GENEKIC NAMES
INTRODUCED IN THIS VOLUME.

^Etonyx, n. g. (Dinosauria), Broom

Bauria, n. g. (Cynodontia), Broom

Eccasaurus, n. g., Broom

Geranosaurus, n. g. (Dinosauria), Broom

Gryponyx, n. g. (Dinosauria), Broom

Gyposaurus, n. g. (Dinosauria), Broom

Heleophilus, n. g. (Mesosauria), Broom . . .
Helichthys, n. g. (Actinopterygii), Broom. . .
Hydropessum, n. g. (Actinopterygii), Broom
Noteosaurus, n. g. (Proganosauria), Broom .
Tritheledon, n. g. (Cynodontia), Broom

PAGE
304
272
276
306
29-4
293
277
254
266
358
334

INDEX OF GENERA AND SPECIES.

A.

PAGE

ACANTHODISCUS, Uliliij 207

ACESTA, H. ( A. Adams 71

ACIVEONINA, d' Orb it) >nj 176

acutus (Heleophilus), Broom .... 277

^ELUKOSAUKUS, Owen 363

^ELUROSUCHUS, Broom 364

.ETONYX, Broom 304, 364

africanus (Belemnites), Tate .... 27

africauus (Capitosaurus), Broom.. 271

africanus (Ccelacanthus), Broom. . 253

africanus (Gryponyx), Broom .... 294

africanus (Hybodus), Broom .... 252

africanus (Noteosaurus), Broom .. 358

ALOPECODON, Broom 363

ALOPOSAURUS, Broom 363

AMMMONITES 25

angulosa (Idmonea), d'Orbiguy .. 6

ANTHODON, Owen 362

ANTHONVA, Gabb 137

antipodum (Berenicea), Tate .... 33

ARCH.EOSUCHUS, Broom ;i62

ARCTOGNATHUS, Broom 363

ARCTOSUCHUS, Broom 363

ARNOGNATHDS, Broom 363

ASTARTE, J. Soicerby 26, 128

atavus (Geranosaurus), Broom .. 306
atherstoni (Actaeonina), Sharpe 37, 17C
atherstoni (Holcostephanus), Sharpe

27, 28, 31, 37, 187

atherstoni (Perna), Sharpe 75

atherstoni (Pinna), Sharpe 76

atherstoni (Psammobia), Sharpe.. 154

atherstoni (Turbo), Sharpe 164

B.

baini (Holcostephanus), Sharpe

27, 28, 31, 197

baini (Modiola), Sharpe 85

baini (Mytilus), Sharpe 33

baini (Pareiasaurus), Seeleij 353

baini (Pleuromya), Sharpe 157

BAURIA, Broom 272, 364

BELEMNITES, Lamarck 210

BEKEXICEA, Lamarck 33

PAGE

BOCHIANITES, Lory 181

borcherdsi (Cyprina), Tate 150

bronni (Astarte), Krauss 26

bronni (Seebachia), 43, 47

browni (Euskelesaurus), Huxley . . 292

browni (Helichthys), Broom 254

browni (Massospondylus), Seeleij 293
browni (Oxygnathus), Broom .... 259
browni (Pholidophorus), Broom .. 267
BUBALIS, Hamilton-Smith 279

C.

calypso (Idmonea), d'Orbigny .. 5

CAMPTONECTES, Meek 65

capense (Coptosoma), Woods .... 13

capensis (Ceratodus), S.-Woodicard 253

capensis (Equus), Broom 281

capensis (Gyposaurus), Broom .. 293

capensis (Ormosaurus), Lydekker 292

capensis (Plesiosaurus), Andrews 309
capensis (Semionotus),S. -Woodward 2C2

capensis (Tylosaurus), Broom .... 332

caperata (Patella), Tate 163

CAPITOSAUROS, Ulilnster 271

CARDITA, Bnigtiiere 127

carinatus (Massospondylus), Owen 291

CARYOPHYLLIA, Stokes 11

cassiope (Trigonia), Tate .... 32, 125

cenomana (Crisina), d'Orbigny .. 5

CERATODTJS, Agassiz 253

CHELYPOSAURUS, Broom 363

CHLAMYS, Bolten 69

CISTECEPHALUS, Oicen 363

CLEITHROLEPIS, Egerton 264

CfELACANTHUS, Agassiz 253

complicata (Crassatella), Tate .. 26, 27

compressa (Diastopora), Goldfuss 3

concava, cf. (Serpula), J. Sowerby 63

conjugata (Entalophora), v. Eeuss 9
conocardiiforniis (Trigonia), Kranss

23, 27, 35, 42, 119
cottaldinus (Pecten), d'Orbir/ny

37, 48, 65

CRASSATELLA, Lamarck 26

CRIOCERAS, d'Orbigny 27

Index of Genera and Species.

XI

PAGE

CEISINA, d'Orbiqny 4

CRYPTOCYNOPON, Sceley 363

CYNOCHAMPSA, Owen 363

CYNODRACO, Owen 363

CYNOGNATHUS, Sceley 283, 363

cynops (Bauria), Broom 272

CYNOSCCHUS, Owen 363

CYPRINA, Lamarck 149

D.

decurrens (Tervia), Pocta 7

DELPHINOGNATHCS, Seeh'y 362

dentata (Gervillia), Kraunn .... 43, 47

DIADEMODON, Sceletj 364

DI^LURODON, Broom 363

DIASTOPORA, Lamoiiroitx 2

DICTYOPYGE, Egerton 262

DICYNODON, Owen 339, 363

disticha (Idmonea), Michclin .... 5

dominicalis (Gastrochffina), Sliar/ie 162

dominicalis (Pholadomya), Sharpe 33

dorsata (Tervia), v. Hayenow .... 7

draperi (Helichthys), S .-Woodward 257

dubia (Mactra?) Kitchin 156

E.

ECCASAURUS, Broom 276, 362

echinata (Entalophora), Rtimcr .. 9

EMYDOPS, Broom 303

ENDOTHIODON, Owen 363

ENTALOPHORA, Lamouroux 8

EOSUCHUS, Watson 364

EQUUS, Linnams 281

EEIPHOSTOMA, Broom 363

ERIPHYLA, Stoliczha 128

ERYTHROSUCHUS, Broom 364

EUNOTOSAUEUS, Seelei/ 364

EUPARKERIA, Broom (ined.) 366

EUSKELESAURUS, Huxley 292, 304

excavata (Crisina), d'Orbigny 6

EXOGYRA, Say 16, 77

extoni (Cleithrolepis), S.- Wood ward 264

F.

FILISPARSA, d'Orbigny

formosa (Dictyopyge), Broom . .
fragilis (Filisparsa), Marsson . .
fungatus (Hamites), Brongniart

G.

GALECHIEUS, Broom
GALEOPS, Broom
GALEPUS, Broom

1

262

2

25

362
362
362

PAGE

GALESAURUS, Owen 363

GASTROCH.ENA, Spengler 162

GERANOSAURUS, Broom 306, 364

gibbeni (Tervia), Gregory 7

glaber (Bochianites), Kitchin .. 38,181

GLANOSUCHUS, Broom 363

goldfussi (Trigonia), Tatc :32

GOJIPHOGNATHUS, Seelei/ 364

GONIOJIYA, J. I.. Agassiz 159

GORGONOPS, Owen 363

GRAMMATODON, Meek it Haydcn . . 88

VJIIOTRTANIA, Kpeyer 26

GRYPONYX, Broom 294, 364

GYPOSAURCS, Broom 293, 364

H.

HAMITES, Park 25

harries! (Massospondylus), Lroom 299

HELEOPHILUS, Broom 277

HELEOSAURUS, Broom 364

HELEOSUCHUS, Broom 364

HELICHTHYS, Broom 254

herzogi (Astarte), Goldfitxs.. 26, 47,128
herzogi (Lyrodon), Cfoldfttss .... 25
herzogi (Trigonia), Goldfuss 34, 40, 101

HOLCOSTEPHANHS, Newnayr 184

holubi (Trigonia), Kitchin. . 35, 40, 103

HORTALOTARSUS, Seelcij 293

HOWESIA, Broom :i64

HY^NASUCHUS, Broom 363

HYBODUS, Agassiz 252

HYDROPESSUM, Broom 266

I.

ICTIDOGNATHUS, Broom 363

ICTIDOPSIS, Broom 363

ICTIDOSUCHUS, Broom 363

IDMONEA, Lamouroux 4

imbricata (Exogyra), Krauss 27, 37, 43,77

imbricata (Placunopsis), Tatc .... 32

intermedius (Hamites), J. Howerby 25

J.

jonesi (Grammatodon), Tatc
jonesiana (Ostrea), Tatc .. .

48, 88
31

K.

kannemeyeri (Ceratodus), Sceley. . 253
kannemeyeri (Hydropessum), Broom 266
kannemeyeri (Trematosaurus),

Broom 270

KAROOMYS, Broom 364

kolbei (Dicynodou), Broom 338

Xll

Index of Genera and Species.

PAGE

kraussi (Cucullea), Tatc 43

kraussi (Trigonia), Kitchin.. 34, 49, 95

L.

LACAZELLA, M.-Chalmas 15

LIMA, Brutjuiere 71

LIMN^EA, Lamarck 178

lineata (Anthonya), Kitchin .... 137

longlandsiana (Astarte), Tatc .... 128

LYCOBHINUS, Broom 363

LYCOSAUBUS, Oiccn 363

LVCOSUCHUS, Broom :163

LYBODON 25

LYSTROSATJRUS, Cope 363

M.

MACTRA, Linnaeus 156

madreporacea (Entalophora), Gold-

/H.vS 10

MANTELLUM, Bolten 74

marginata (Crisina), d'Urbigiiy .. 6

MASSOSPONL-YLUS, Owen . . '291, 293, 364

MELIKODON, Broom 364

MERETRIX, Ijumarck 151

MESOSAURUS, Gervais 364

MESOSUCHUS, Watson 364

MEYERIA, HP Coy 212

MICROGOMPHODON, Secley 304

minor (Cleithrolepis), S. -Woodward 266

minutulus (Turbo), Kitchin 168

mirifica (Natica ?), Kitchin 174

modderensis (Holcostephanus)

Kitchin 202

MODIOLA, Lamarck 85

MOSCHOPS, Broom o62

MYTH/US, Linnaeus 82

N.

NATICA, Lamarck 171

neglecta (Lima), Tate 37, 74

NEITHEA, Droult 17

NOTEOSAUROS, Broom 358, 364

NOTOCHAMPSA, Broom 307, 364

NUCULA, Lamarck 87

nuculoides (Cardita), Tate 127

NYTHOSAURUS, Owen 363

PAGE

0.

obliquissima (Lima), Tate

oblonga (Thetironia), Kitchin ..
orbicularis (Pecten), J. Sowerby

ORINOSADRUS, Lydekkcr

ornatus (Ceratodus), Broom

71
146

65
292
253

OSTREA, Linnaeus 16, 77

OSTREA (EXOGYBA), T. Kay 16, 77

OUDENODON, Oic en 338

OXYGNATHUS, Egerton 259

P.

PALACRODON, Broom 364

PALIGUANA, Broom 364

palustris (^Etonyx), Broom 304

papyracea (Thetironia), Sharpe .. 142

PAKDOSUCHUS, Broom 363

PAREIASAURUS, Owen 353, 362

PATELLA, Linnaeus 163

PECTEN, Midler 65

PECTEN (NEITHEA), Droult 17

PELOSUCHUS, Broom 362

PERNA, Jintguiere 17, 75

PHOLIDOPHORUS, Ayassiz 267

PHYLLOCERAS, Suess 179

pinchiniana (Astarte), Tate .. 33,135

pinchiniana (Serpula), Tate 64

PINNA. Linnaeus 76

PLACUNOPSIS, Morr. & Lycctt 32

PLESIOSAURUS, Conybeare 309

PLEUROJIYA, Agassiz 157

PLEUROTOMARIA, Def ranee. 163

prisons (Bubalis), Broom 279

prisons (Eccasaurus), Broom .... 276

PRISTERODON, Huxley 363

PRISTEROGNATHUS, Xeeley 363

PROCOLOPHON, Owen .... 362

PRODICYNODON, Broom 363

projectus (Pecten), Tate 48, 66

PROPAPPUS, Seeley 323, 362

PROTEROSUCHUS, Broom 364

PSAMJIOBIA, Lamarck 154

PTYCHOMYA, Agassiz 26,27

E.

ramosa (Filisparsa), d'Orbigny .. 2

REIIOXDIA. <labb 26

remota (Limnaaa), Kitchin 178

riconoi (Tritheledon), Broom .... 334
rogersi (Holcostephanus), Kitchin 201

rogersi (Natica), Kitchin 173

rogersi (Phylloceras), Kitchin .... 179

rogersi (Propappus), Broom 323

rogersi (Trigonia), Kitchin .. 34, 42, 99

rogersi (Turbo), Kitchin 167

rubidgeanus (Pecten) 32

rugulosa (Cyprina), Sharpe .... 37, 149

S.

SAUROSTERNON, Huxley 362

SCALOPOSAURUS, Owen 363

Index of Genera and Species.

xin

PAGE

SCAPANODON, Broom 363

schenki (Astieria), Ptivloir 202

schwarzi (Meyeria), Kitchin .... 212
schwarzi (Tancredia), Kitchin.. 38, 139
SCYLACOGNATHUS, Broom (ined.) . . 366

SCYLACOPS, Broom (ined.) 366

SCYLACOSAURUS, Broom 363

SCYMNOGNATHUS, Broom 363

SCYMNORHINUS, Broom (ined.) .... 366

SCYMNOSAURUS, Broaiii 363

SEEBACHIA, Neumai/r 26

SEMIONOTUS, Agassiz 2C2

SERPULA, Linnaeus 63

SESAMODON, Broom 364

simplex (Ileptotubigera), de Loriol 4
skirtopodus (Hortalo tarsus). Seelcy 293

SOLECURTUS, de Blainville 155

spinosissimum (Crioceras), Neumayr

27, 31

SPIROPORA, Lamouroux 10

stowi (Trigonia), Kitchin . . 41, 42, 115
subacutus cf. (Pecten), Lamarck.. 69
subanceps (Ammonites), Tate .... 27
subjurensis (Placunopsis), Tate .. 32
SYNCYCLONEMA, Meek 65

T.

TAMBOEMA, Seeley 362

TANCREDIA, Lycett 139

TAOGNATHUS, Broom 363

TAPINOCEPHALUS, Owen 362

tatei (Trapezium?), Kite/tin .... 148

tatei (Trigonia), Neumayr 27,34,

38, 40, 125

TAUROPS, Broom 362

tenuis (Helichthys), Broom 258

TERVIA, Jullien 6

THAMNASTR.EA, Lesanvage 62

THECIDEA (LACAZEI.LA), M.-Chalmas 15

THECODONTOSAURUS, Broom 364

THELEGNATHUS, Broom 362

PAGE

THETIRONIA, Stoliczka 142

THRACIA, Leach 160

TIGRISUCHUS, Owen 362, 363

TITANOSUCHUS, Owen 303

TRAPEZIUM, v. Miililfeldt 148

TREMATOSAORUS, Bra-tin 270

TRIBOLODON, Seeley 364

TRIGONIA, Bruguiere . 91

TRIRACHODON, Seeley 364

TRITHELEDON, Broom 334, 364

TROCHOSAURUS, Broom 363

TURBO, Linnaus 164

TYLOSADRUS, Marsh . 332

U.

uitenhagensis (Holcostephanus) ,

Kitchin 206

uitenhagensis (Meretrix), Kitchin 151

uitenhagensis (Mytilus), Kitchin. . 82

uitenhagensis (Natica), Kitchin .. 171

uitenhagensis (Nucula), Kitchin . . 87

uitenhagensis (Unio), Kitchin .... 89

UNIO, Retzius 89

V.

vau (Trigonia), Sharpe .... 29, 41, 42,

48, 49, 110
ventricosa (Trigonia), Krauss 27, 29,

34, 40, 46, 49, 91

verticillata (Spiropora), (.ioldfuss 10
virgula (Entalophora), r. Hagenow 8
virgula (Idmonea), d'Orbigny .... 4

W.

wilmanae (Holcostephanus),

Kitchin 37, 195

(1)

1. Polyzoa and Anthosoa from the Upper Cretaceous Limestone of
Need's Camp, Buffalo River. By W. D. LANG, M.A., F.G.S.,
British Museum (Natural History).

Plate I.

THE rock containing the Polyzoa here described, is so full of them
in places that they compose the rock nearly to the exclusion of
its other components. Such pieces closely resemble in general ap-
pearance specimens of a rock composed mainly of Polyzoa from the
Dariian of Paxoe, in the island of Seeland, Denmark. Moreover, in
both rocks the surfaces of the specimens are very ill-preserved,
being rough, with the details ill-defined, as though the specimens
were covered with a thin incrustation. And in both the large
proportion of erect, cylindrical Polyzoa is remarkable.

The species described below range from the Neocomian to the
Danian, and the majority are restricted to the Chalk. In so far
as it is possible to assign a definite horizon to the deposit on the
evidence of the Cyclostome Polyzoa only, it appears that its age is
Senonian or Danian.

DIASTOPOBnm

GENUS FILISPAESA, d'Orbigny.

Gregory :;: places this genus among the Diastoporidae because of
its resemblance to Proboscina. He says it may be regarded as
" a group of ProboscincB with an erect habit " ; and goes on to show
how some species of Filisparsa correspond with some of Proboscina.
But he also points out that Pergens t placed the genus in the
Idmoneidae. The nature of the zoarium and distribution of the
zooecia are such that the genus occupies an intermediate place

* J. W. Gregory, B.M. Cat. Cret. Bryozoa, vol. i., 1899, p. 67.
t E. Pergens, Rev. de Bry. du Cret. fig. par d'Orb., Bull de la Soc. Beige de
GeoL, iii., 1890, Mem., p. 339.

1

2 Annals of the South African Museum.

between the two families ; and thus it shows the probability of
the present classification of the Cyclostome Polyzoa being highly
artificial. But until the distribution and life history of each
described form has been determined, it is impossible to form a
natural classification of the whole. Meanwhile, the present
artificial classification must be used for lack of better.

FILISPABSA KAMOSA, d'Orbigny.

Plate I., fig. 1.

1853. Filisparsa ramosa, d'Orbigny, Bry. Cret., p. 819, pi. 756
figs. 18-22.

Zoariurn of erect, flattened, solid branches, in one plane. Zooecia
only opening on one face of the zoarium. Apertures tend to arrange-
ment in V-shaped bands across the zoarium, but their distribution
is somewhat irregular ; four or five in a series ; distance between
apertures of the same series one-half times to once the diameter
of an aperture ; between apertures of neighbouring series twice to
four times. Zooecia immersed.

There are three fragments of this species, one of which is
figured.

Distribution. Turonian and Senonian.

FILISPABSA FBAGILIS, Marsson.

Plate I., fig. 2.

1887. Filisparsa frayilis, Marsson, Bry. Eiig. ; Pal. Abh., vol. iv.,
p. 35, pi. iii., fig. 8.

Zoarium of erect, cylindrical, or slightly flattened solid branches,
in one plane. Zooecia only opening on one face of the zoarium.
Apertures arranged in single series across the zoarium ; four or
five in a series ; distance between apertures of the same series
one-half times or less the diameter of an aperture ; between those
of neighbouring series twice-and-a-half to thrice. Zooecia immersed,
but the boundaries between them are more or less apparent as
ridges. One fragment out of seven is figured.

Distribution. Senonian.

GENUS DIASTOPOEA, Lamouroux.

Diastopora is here used, as in the British Museum Catalogue, for
Diastoporidse with an erect uni-, bi-, or even multi-laminar zoarium,
consisting of fronds or of very flattened branches.

Polyzoa and Antliozoa from Upper Cretaceous Limestone. 3

DlASTOPORA COMPRESSA (Goldfuss).

Plate I., fig. 3.

1827. Ccriopora compressa, Goldfuss, Petref. Germ., p. 37, pi. xi.,

fig. 4.
1851. Ditaxia compressa, von Hagenow, Bry. Maastr. kr., p. 50,

pi. iv., fig. 10.

Zoarium of erect bilaminar fronds. Zocecia immersed, and their
boundaries not apparent, or very slightly so. Apertures irregularly
arranged ; the distance between the apertures in a proximal-distal
direction is about once the diameter of an aperture; that in a
direction at right angles to the last is once or less than once the
diameter of an aperture. This definition does not distinguish D.
compressa from Mesenteripora Hyselyi, de Loriol (Urg. Land. : Mem.
Soc. helv. Sci. Nat., vol. xxiii., 1869, p. 40, pi. hi., fig. 1), from the
Urgonian of Neuchatel ; and from the figures it is difficult to see
any difference between these two forms. But fig. 10 m. on pi. iv. of
von Hagenow's work shows a piece of a zoarium with the apertures
elongate in a proximal-distal direction. It is only described in the
legend " Theil der Oberfliiche am oberen Rande," but it looks as if
this were a worn surface and the elongation of the apertures were
due to the angle between the surface and the direction of the zocecia
at that level. It is possible that in M . Hyselyi the direction of the
zooecia would be perpendicular to the surface for some distance, in
which case a worn specimen would show circular apertures.

D'Orbigny's figure of Bidiastopora Campichcana (Bry. Cret.,
1853-4, p. 800, pi. 784, figs. 6 and 8) also resembles Diaspora
compressa and Mesenteripora Hyselyi. But Pergens, who has seen
d'Orbigny's specimens, identifies B. Campicheana with B. acuta,
d'Orbigny, and re-figures the latter (Revision des Bry. Cret. fig. par
d'Orbigny, 1890, pi. xi., fig. 7), showing it to be a form quite distinct
from D. compressa and M. Hyselyi.

Gregory places as synonyms of Diastopora compressa the two
forms figured by d'Orbigny as Mesenteripora compressa and M.
ncocomiensis (Bry. Cret., 1853-4, pi. 756, figs. 10-13 and 7-9) ; but
both these forms have emergent zooecia.

Among the material described the only specimen referable to this
species is a small fragment of the base of a frond. The earliest
zooecia, as would be expected, differ from the later ones ; they are
further apart and slightly emergent. Near the edge of the fragment
they are seen to be close together and immersed, with their boun-
daries slightly shown. The apertures, especially near the edge,

4 Annals of the South African Museum.

where they are slightly worn, are somewhat elongated in a
proximal-distal direction.
Distribution. Senonian.

IDMONEID^E.

GENUS IDMONEA, Lamouroux.

IDMONEA VIKGULA, d'Orbigny.
Plate I., fig. 4.

1851. Lhuonea virgula, d'Orbigny, Bry. Cret., pi. 631, figs. 15-17.
1863. Reptotubigera simplex, de Loriol, Invert. Mt. Sal6ve, p. 127
pi. xvi., fig. 3.

Zoarium of a single zocecium-bearing ridge, without a selvage
short (about thrice as long as broad), rapidly becoming broader
distally. The only specimen of this species has a flat reverse face,
but does not encrust any object. But its general shape and corre-
spondence with d'Orbiguy's figure of Idmonea virgula renders it
probable that it was originally encrusting some perishable object
which has since decayed. This raises the question whether other
Idmoneidte with flat or convex reverse faces, such as Crisina
marginata, d'Orbigny, in some cases may not have been forms
encrusting some object, such as seaweed, which has since
perished.

Distribution. Neocomian and Cenomanian.

GENUS CRISINA, d'Orbigny.

The separation of Crisina from allied Idmoneid genera, especially
from Retecava, is attended with much difficulty.* The genus is
here used as restricted by Gregory,! and distinguished from Tervia
and lieticrisina, which Gregory regards as subgenera. Tervia is
here considered as equal to Crisina in systematic significance.
Crisina thus regarded may be defined as a genus of Idmoneidse,
in which the zoarium is erect, and consists of solid branches, which
are subcylindhcal or moderately compressed ; the reverse face is
well developed, but does not overlie an " axial rod " as in Bctccava;
there is no unpaired median nor multiple median series of apertures
on the obverse face ; the apertures are in single lateral series, and
circular.

* See W. D. Lang, Geol. Mag., 1907, p. 125.

t J. W. Gregory, Brit. Mus. Cat. Cret. Bry., 1899, vol. i., pp. 159, 168, and 176.

Polyzoa and Antliozoa from Upper Cretaceous Limestone. 5

CEISINA CENOMANA, d'Orbigny.

Plate I., fig. 5.
1851. Crisina cenomana, d'Orbigny, Bry. Cret., p. 732, pi. 614,

figs. 1-5.
1845. Idmonea disticha, Michelin, Icon. Zooph., p. 204, pi. Hi.,

fig. 18.
1853. Idmonea calypso, d'Orbigny, Bry. Cret., p. 733, pi. 747,

figs. 10-14.

1890. Idmonea cenomana, Pergens, Eevision des Bry. Cret. fig. par
d'Orbigny, p. 344, pi. xii., fig. 3.

Zoarium subcylindrical, and the reverse face is convex or flat ;
there is no keel on the obverse face ; the zooecia are immersed, and
when the reverse face is flat the zocecial boundaries are clearly
marked ; the lateral series of apertures are quite regular, and just
reach the middle line of the obverse face ; there are about four
apertures in each lateral series ; the apertures are circular, all of
one size ; those of the same series are distant less than one diameter
of an aperture ; those of different series two or three diameters of an
aperture.

This definition distinguishes C. cenomana from all cretaceous
forms of Eetecava as well as from others of Crisina. Those species
of Retecava which most resemble it have a very slight median keel
on the obverse face, except E. Abbotti (Gabb and Horn), (Journ.
Acad. Nat. Sci. Philadelphia, ser. ii., vol. iv., 1860, p. 404, pi. Ixix.,
figs. 45-47), but in this form there is a ridge down the obverse face
of each zooecium.

Two distinct forms are included under the above definition, and
both occur in the material here described. In the first, resembling
d'Orbigny's figure of C. cenomana, the zooecial boundaries are hardly
visible and the reverse face is convex. In the second (pi. xlv., fig. 4),
resembling d'Orbigny's figure of Idmonea calypso, the zooecial boun-
daries are strongly marked and the reverse face is flat. The only
justification for placing the two forms under one name is their
identification by Pergens.* Speaking of C. cenomana, under the
synonyms of which he had placed I. calypso, Pergens says,
" quelquefois les branches sont aplaties a la partie inferieure et
forment ainsi une variete decrite comme 1. calypso." That is,
I. calypso is an earlier growth stage of C. cenomana confined to the
proximal portions of the zoarium. In view of the very fragmentary
character of the specimens usually obtained of these erect Idmoneids,
* E. Pevgens, Eevision des Bry. Cret. fig. par d'Orbigny, 1890, p. 344.

6 Annals of the South African Museum.

it is exceedingly rare to get knowledge of the growth stages of any
one form. But the specimens of C. cenomana here described do not
look like proximal ends of zoaria. It is likely, then, that a form
exists whose adult zoarium over its greater extent has the characters
of I. calypso, d'Orbigny, while these are confined to the proximal
parts of an adult zoarium of C. cenomana.

Distribution of both C. cenomana and I. calypso. Cenomanian.

CRISINA EXCAVATA (d'Orbigny).
Plate I., fig. 6.

1853. Idmonca excavata, d'Orbigny, Bry. Cret., p. 742, pi. 749,
figs. 11-15.

Zoarium subtriangular with a fiat or concave reverse face ; the
lateral series are regular and contain six to eight apertures ; the
apertures are circular; the zocecia are emergent. Idmonea filiformis,
d'Orbigny (Bry. Cret., 1853, p. 744, pi. 750, figs. 1-5), and Idmonea
lata, d'Orbigny (Bry. Cret., 1853, p. 734, pi. 748, figs. 6-10) resemble
this form, but have only three apertures in a lateral series. Idmonea
marginata, d'Orbigny (Bry. Cret., 1853, p. 744, pi. 749, figs. 20-23),
with which Gregory unites this form, differs from it by having
immersed zocecia.

Distribution. Senonian.

CRISINA MARGINATA (d'Orbigny).
Plate I., fig. 7.

1853. Idmonea marginata, d'Orbigny, Bry. Cr6t., p. 744, pi. 749,

figs. 20-23.
1853. Idmonea angulosa, d'Orbigny, Bry. Crt., p. 735, pi. 748,

figs. 11-15.

Zoarium subtriangular with a flat or convex reverse face ; the lateral
series are regular and each contains four or five apertures ; the
apertures are circular. This form is very like G. excavata (d'Orbigny),
but has fewer apertures in a lateral series, and the zocecia are
immersed.

Distribution. Turonian and Senonian.

GENUS TEEVIA, Jullien.

Idmoneidse whose zoarium is erect and consists of solid branches.
There is an unpaired median series of apertures between the two
rows of simple lateral series. Tervia is considered by Gregory as a

Polyzoa and Anthozoa from Upper Cretaceous Limestone. 7

subgenus of Crisina, from which it differs in possessing a simple
median series of apertures on the obverse face. The fact that it is
not always easy to distinguish this series as composed of apertures
distinct from the most median apertures of the lateral series shows
the artificiality of the genus. This point is well exemplified in von
Hagenow's figure of T. dorsata (von Hagenow).

TERVIA DORSATA (von Hagenow).
Plate I., fig. 8.

1851. Idmonea dorsata,von. Hagenow, Bry. Maastr. kr., p. 31, pi. ii.,
fig. 10.

Zoarium subcylindrical or triangular or flattened in a plane parallel
to the obverse and reverse faces ; without dorsal processes ; reverse
face flat or concave ; zocecia three or four in a lateral series, im-
mersed but the boundaries are clearly marked; the distance between
the different series of apertures is one and a half to two and a half
times the diameter of an aperture ; those of the same series are
distant about a quarter the diameter of an aperture. This is the
commonest form among the specimens of Cyclostomes from Need's
Camp.

Distribution. Senonian.

TERVIA GIBBERA, Gregory.
Plate I., fig. 9.

1899. Crisina (Tervia) gibbera, Gregory, Brit. Mus. Cat. Cret. Bry.,
vol. i., pp. 173-4, Text fig. 12.

Zoarium subcylindrical with the reverse face flat and possessing
spine-like projections, bearing no zocecia, termed by Gregory " dorsal
processes"; zooecia about three in a lateral series, immersed, but
the boundaries are just shown ; distance between the different series
of apertures three to four times the diameter of an aperture ; those
of the same series distant about one-half the diameter of an
aperture. Only two specimens have been found among the material
here described.

Distribution. Senonian.

TERVIA DECURRENS (Pocta).

1892. Idmonea decurrens, Pocta, Mech. koryc. Hory. ; Ceska. Ak. Fr.
Jos. Pratze, sect. 2, p. 18, pi. ii., figs. 3-6.

Zoarium subtriangular in cross-section and flat on the reverse side ;
the zocecia are immersed with their boundaries apparent ; apertures

8 Annals of the South African Museum.

two or three in a lateral series ; those of different series distant four
or five diameters of an aperture, those of the same series two to three
diameters of an aperture.

A single specimen is somewhat doubtfully referred to this species.
It is the proximal end of a zoarium, though the first few zooecia
appear to be missing. Beginning as a thin, cylindrical branch, the
zoarium rapidly widens and becomes compressed in a plane parallel
to the obverse and reverse faces, at the same time becoming very
curved, so that in a section along the proximal-distal axis the obverse
face appears convex and the reverse face concave. When the
breadth of the zoarium has reached about one-third of its length,
it divides into two subcylindrical branches ; one of these is broken
off short, but the other continues for about T5 mm., without altering
in shape. It would be interesting to know if on further growth the
branches flatten and widen, thus recapitulating the growth stages of
the first part of the zoarium ; * or whether the first part is really an
encrusting portion, whose support has decayed, which has thrown
up cylindrical branches in the same way as a Phalangella-like base
has been shown by Gregory f to throw up branches of the genus
Crisina. The general zooecial characters are those of Tervia decurrens
(Pocta), in which species it is provisionally placed.

Distribution. Cenomanian.

ENTALOPHOKnm

GENUS ENTALOPHOEA, Lamouroux.

ENTALOPHOBA VIRGULA (von Hagenow).
Plate I., fig. 10.

1840. Ceriopora virgula, von Hagenow, Mon. Kiig. : Neu. Jahrb.,

1840, p. 646.
1851. Pustulipora virgula, von Hagenow, Bry. Maastr. kreide.,

p. 17, pi. i., fig. 3.

Zoarium consisting of smooth cylindrical branches ; the apertures
are irregularly distributed on the zoarium and are very far apart,
their distance apart in a proximal-distal direction being generally
more than four times the diameter of an aperture. There are never
more than two apertures in the breadth of a branch. The zorecia

* See W. D. Lang, Geol. Mag., 1905, pp. 259-260.

f J. W. Gregory, Brit. Mus. Cat. Cret. Bry., vol. i., 1899, p. 159, text-fig. 11.

Polysoa and Antlwzoa from Upper Cretaceous Limestone. 9

are immersed. Entalopliora Pergensi, Gregory,* which the above
definition would also include, has thinner branches and the apertures
still further apart than in E. virgula.

This is the commonest species of Cretaceous Entalopliora and has
been found in most of the localities whence Cretaceous Polyzoahave
been recorded ; but only one specimen has been found among the
material here described.

Distribution. Neocomian-Danian.

ENTALOPHOEA CONJUGATA, von Eeuss.
Plate I., fig. 11.

1872-3. Entalopliora conjugata, von Eeuss, Bry. unt. Quad. Palseon-
togr., vol. xx., part. L, p. 117, pi. xxix., fig. 8.

Zoarium consisting of cylindrical branches ; the apertures are
distant from one another in a proximal-distal direction more than
four times the diameter of an aperture ; the apertures occur in rows
of two or three across the branch ; the zooecia are immersed and
their boundaries may be faintly marked. This form is very like
E. geminata f (von Hagenow), but the zooecia are shorter, the boun-
daries of the zooecia are less clearly marked, and the apertures occur
in groups of three instead of two as in E. geminata. A single
specimen of this species occurs among the material here described.

Distribution. Cenomanian.

ENTALOPHORA ECHINATA (Bomer).
Plate I., fig. 12.

1840. Pustulopora cchinata, Eomer, Verst. Nordd. kr., p. 22, pi. v.,

fig. 23.
1846. Pustulopora ecliinata, von. Eeuss., Verst. Bohm kr., p. 64,

pi. xiv., fig. 4.

Zoarium consisting of cylindrical branches ; the apertures are
distant from one another in a proximal-distal direction about four
diameters of an aperture. There are about four apertures in the
breadth of a branch ; the diameter of the apertures is about 2 mm. ;
the zooecia are immersed, with their boundaries more or less apparent.
This species is very nearly allied to E. madreporacea (Goldfuss),
from which it differs in having the apertures slightly further apart in
a proximal-distal direction, the zocecial boundaries more prominent,

* J. W. Gregory, Brit. Mus. Cat. Cret. Bry., vol. i., 1899, p. 226, pi. x.,
figs. 5, 6, 7.

f Pustulipora geminata, von Hagenow, Bry. Maas. kr., 1851, pi. i., fig. 11, p. 20,

10 Annals of the South African Museum.

and the apertures less regularly arranged. The peristomes, too, are
higher; but in the three specimens of this species among the material
here described the peristomes are worn.
Distribution. Cenomanian to Danian.

ENTALOPHORA MADREPOBACEA (Goldfuss).
Plate I., fig. 14.

1827. Ceriopora madreporacea, Goldfuss, Petref. Germ., p. 35, pi. x.,

figs. 12a and b.
1851. Pustulipora madreporacea, von. Hagenow, Bry. Maastr. kr.,

p. 18, pi. i., fig. 8.

Zoarium consisting of cylindrical branches ; the apertures are
distant from one another in a proximal-distal direction less than four
diameters of an aperture ; there are four to six apertures in the
breadth of a branch ; the apertures are somewhat irregularly arranged
in spirals ; the zocecia are immersed. Three ill-preserved specimens
from among the East London material are doubtfully referred to this
species. All are very short fragments of stout branches, and, in so
far as the zooecial characters can be determined, resemble E. madre-
poracca more closely than any other form.

Distribution. Cenomanian to Danian.

GENUS SPIROPORA, Lamouroux.

Spiropora should probably be united with Entalophora, for in
many species of the latter genus the apertures are arranged
spirally. Spiropora, however, may be defined as resembling
Entaloplwra, but the apertures are arranged in whorls, or, if in
spirals, these are very much looser than in Entalopliora.

SPIROPORA VERTICILLATA (Goldfuss).
Plate I., fig. 13.

1827. Ceriopora verticillata, Goldfuss, Petref. Germ., vol. i., p. 36,

pi. xi., fig. 1.
1851. Cricopora verticillata, von Hagenow, Bry. Maastr. kr., p. 20,

pi. i., fig. 12.

Zoarium consisting of cylindrical branches, having the apertures
arranged in whorls ; there are five to ten apertures in the breadth of
a branch ; the whorls are distant four or five diameters of an aperture ;
the distance between the apertures of a whorl is a half or less than a
half the diameter of an aperture ; the zooecia are immersed, but the

Polyzoa and Antliozoa from Upper Cretaceous Limestone. 11

boundaries between them are clearly marked by thin ridges. Next
to Tervia dorsata (von Hagenow), this is the commonest species of
Cyclostome among the material here described.
Distribution. Neocomian to Danian.

ANTHOZOA.

In the Need's Camp material are several specimens of simple
corals, thickly encrusted with Cheilostome Polyzoa. Transverse
sections show that they belong to the genus Garyophyllia, but the
species cannot be determined owing to their encrusted condition.

(13)

2. Echinoidca, Bracliiopoda, and Lamellibranchia from the Upper
Cretaceous Limestone of Need's Gamp, Buffalo River. By
HENRY WOODS, M.A., University Lecturer in Palaeozoology,
Cambridge.

Plate I. and three figures in text.

AN account of the deposit from which the fossils here described
were obtained is given at the end of this paper (page 425). The
large specimens of Pcrna were found in the hard crystalline lime-
stone ; most of the other fossils came from the polyzoa-limestone.
No species giving decisive evidence of the horizon of the limestone
has been recognised, but the affinities of the forms described seem to
show that the deposit belongs to a late stage of the Chalk.

ECHINOIDEA.

COPTOSOMA CAPENSE, sp. nov.
Plate I., figs. 15-17. Text-figure 1.

Test of medium size, with sub-circular outline, convex above,
flattened below, somewhat concave around the peristome ; height
equal to less than half the diameter ; greatest diameter in the
lower third.

Apical disc rather small, not preserved.

Ambulacra not quite as wide as the interarnbulacra. Poriferous
zones undulating. Pores unigeminal, except on the base of the test,
where they are bigeminal. Plates high, consisting of six fused
plates, with the six pairs of pores forming an arc. Each compound
plate bears a large primary, imperforate tubercle (either very slightly
or not at all crenulate), with a large areola, and a few small tubercles
and granules around the margin. The primary tubercles are largest

14

Annals of the South African Museum.

at the ambitus, and smaller on the dorsal surface and on the base
of the test.

Interambulacral plates high, each bearing a large primary, imper-
forate, slightly crenulate tubercle with an areola of moderate size ;
these tubercles are largest at the ambitus, where they are surrounded
by a narrow band of small tubercles and granules. Above the
ambitus, and on the base of the test, the primary tubercles become
smaller ; above the ambitus the surrounding band of granules
becomes much wider. At the ambitus and continued on to the base
of the test there is a row of secondary tubercles at the external
margin of the area.

Peristome of moderate size.

FIG. 1. Coptosoma capcnse, sp. nov. Ambulacral and interambulacral areas, x 2.

The bigeminal pores extending from the peristome almost to the
margin of the base seem to distinguish this from other species of
Coptosoma. The form of the test is similar to that of the type
species, C. cribrum, Desor, but the granules, especially on the inter-
ambulacral areas, are much more extensively developed.

Portions of three tests have been found, the largest and most
perfect of which is here figured. In all cases the apical disc is
missing.

The only other remains of echinoids found in this deposit are some
strongly ridged spines of Cidaris, and a portion of the test of an
irregular form which may perhaps belong to Cassidulus.

Echinoidca, Brackiopoda, and Lamcllibranchia

15

BRACHIOPODA.

THECIDEA (LACAZELLA), sp.
Text-figure 2.

The ventral valve is deep, more or less conical but sometimes
rather irregular, and either slightly curved or nearly straight. The
area is large and triangular, with a raised pseudo-deltidium. On the
exterior of the shell are growth-ridges parallel to the margin of the
valve. In the interior of the valve are narrow longitudinal ridges.

A

B

C

D

E

G

FIG. 2. Thecidca (Lacazella), sp. A, B, C, three views of the same specimen ;
D, G, a ventral valve seen from the cardinal side (D) and from the dorsal
surface (G) ; E, P, two dorsal valves. All x 3.

The thin median septum is joined by a short branch on each side
near the cardinal margin. This valve is deeper and less curved than
in most examples of Thccidea vermicularis (Schlotheim), :;c but is not
so deep as in T. longirostre, Bosquet. |

* For figures and references see Geinitz, Das Elbthalgeb. in Sachsen (Palffionto-
graphica, vol. xx., pt. 1, 1872), p. 162, pi. xxxv., figs. 35-38. Bosquet, Brach.
Terr. Cret. Super, de Limbourg (1859), p. 26, pi. Hi., figs. 4-11. Posselt, Brach.
danske Kridtformat. (Danmarks geol. Undersog., N r 6, 1894), p. 53, pi. iii.,
figs. 23-25.

t Bosquet, op. cit., p. 33, pi. iv., figs. 4-6

16

Annals of the South African Museum.

The dorsal valve is oval, slightly convex, with the umbo near the
margin. The internal margin is ornamented with ridges. From
the median septum one or sometimes more branches come off on
each side.

Several examples of this species were found by Mr. Lang when
searching for Polyzoa in the broken-up rock.

LAMELLIBEANCHIA.

OSTREA, Sp.

A small specimen of part of a left valve is probably an example
of 0. vesicularis, Lamarck.* That species occurs in the Ariyakir
Group of Southern India, and is widely distributed in deposits of
Senonian age.

Some other examples of Ostrea, but not sufficiently perfect for
specific determination, have been found.

C

FIG. 3. Oxtrea (Exoyyra'), sp.
valve. Natural size.

FIG. 4. Perna, sp. Left valve. In-
ternal cast of part near the umbo
Eight and of part of the ligamental area.

Natural size.

OSTREA (BXOGYRA), sp.
Text-figure 3.

There are two right valves which resemble 0. (Exogyra] Langloisi,
Coquand, from the Senonian (Santonian) of Algeria and Tunis. f

* Coquand, Monogr. du Genre Ostrea (1869), p. 35, pi. xiii. , figs. 2-10. Stoliczka,
Cret. Fauna S. India, vol. iii. (1871), p. 465, pi. xlii., figs. 2-4, pi. xliii., fig. 1, pi.
xlv., figs. 7-12. Peron, Brachiopodes, etc., de la Tunisie (1893), p. 175. Wanner,
Palasontographica, vol. xxx., pt. 2 (1902), p. 119, pi. xvii., figs. 10-12.

f Peron, op. cit., p. 146, pi. xxiv., figs. 14, 19.

Echinoidea, Brachiopoda, and Lamellibranchia.

17

PECTEN (NEITHEA), sp.

An imperfect specimen which appears to be part of a left valve of
Neithea is present in the collection.

PEENA, sp.
Text-figures 4-6.

There are twelve internal casts of a large sub-quadrate Perna.
In some cases portions of the shell are preserved and show a
prismatic structure similar to that seen in P. Ricordeana, d'Orb.

FIG. 5. Perna, sp. Internal cast. Left valve and anterior view of both valves.
The posterior and the ventral margins are imperfect, x |.

The largest specimen has a height of about 150 mm., and its hinge-
line is about 85 mm. long. One example shows the cast of the
ligament area and grooves. This species resembles P. valida
(Stoliczka) * from the Ariyalur Group (Senonian) of Southern India,

* Cret. Fauna S. India, vol. iii. (1871), p. 409, pi. xxii., fig. 1.

2

18

Annals of the South African Museum.

V

and also P. cereviciana, Petho, :;: from the Senonian of Cereviic
(Hungary). But more perfect specimens are needed before an
exact determination can be made. The ligament grooves are less
numerous and more widely separated than in P. cereviciana, but

j? IG 6. Perna, sp. Internal cast. A considerable portion of the posterior part
of the shell appears to be missing, x .

the flattened part of the shell extending ventrally from the umbo is
similar in both species.

* Die Kreide- (Hypersenon-) Fauna d. Peterwardein. Gebirges (Palseonto-
^raphica, lii., 1906), p. 234, pi. xvii.

Echinoidea, Bracliiopoda, and Lamcllibranchia. 19

NOTE ON THE LOCALITY.

These fossils were obtained by Mr. J. W. Woods, of East London,
from Need's Camp, on the Buffalo Biver, twenty miles from the sea
and at an altitude of about 1,200* feet. Mr. Woods says the lime-
stone is " exposed in two open workings about two miles apart, and
the one about fifty feet higher than the other. In the lower the
greatest thickness is six feet and seems to be composed entirely of the
remains of small shells, echinids, cup-corals, and various spines. . . .
In the upper quarry the limestone is mainly crystalline and hard,
and seems to be a mass of shell of mussel type : a few ' shark's '
teeth have been found in it. We had a section there of eight feet
to examine, and how much more exists cannot be stated as the
workings have not gone deeper.

" The areas where the deposits occur are completely enclosed by
dolerite: they may be described as two level ten-acre lots surrounded
by boulders ; and were, I presume, old shore-basins or lagoons,
walled in by the igneous dyke, where the molluscs lived and died,
or into which they were gradually swept by the tide."

The interest of this fauna is very great because it occurs at a place
more than half-way between the Pondoland Upper Cretaceous out-
crops and the main area of the marine Lower Cretaceous in
Uitenhage. No fossils from similarly situated beds have been
described, though limestones like that from the top quarry at
Need's Camp are known from the neighbourhood of Sand Flats
at about 1,200 feet above sea-level.

A. W. KOGERS.

* This is the figure given by Mr. Woods ; it is probably too great by 100 feet,
as in Gamble's list of heights the Need's Camp beacon is said, on the authority of
Capt. W. Bailey, B.E., to be 1,144 feet above sea-level.

(20)

EXPLANATION OF PLATE I.

POLYZOA AND ECHINOIDEA FROM THE UPPER CRETACEOUS OF NEED'S CAMP,
BUFFALO RIVER, CAPE COLONY.

1. Filisparxa rainosa, d'Orbigny. x 12. Obverse face of part of a zoarium,

showing tendency of apertures to be arranged in V-shaped rows (p. 408).

2. Filisparsa fraqilis, Marsson. x 12. Obverse face of part of a zoarium

(p. 408).

3. Diastopora compressa (Goldfuss). x 12. A young zoarium, showing the first-

formed zocecia to be slightly emergent, while the more distal are im-
mersed (p. 409).

4. Idmonea virgula, d'Orbigny. x 12. Obverse face of zoarium (p. 410).

5. Crisina cenoniunn, d'Orbigny. x 12. Obverse face of a specimen resembling

d'Orbigny's figure of Idmonea calypso, showing strongly-marked zooecial
boundaries (p. 411).

6. Crisina excavata (d'Orbigny). x 12. Obverse face of part of a zoarium.

In the proximal part the lateral series of apertures of each side are
practically opposite, and so appear to form continuous bands across
the zoarium. Distally is a front view of the right-hand series only, and
those of the left-hand are seen in profile (p. 412).

7. Crisina mnnjinata (d'Orbigny). x 12. Obverse face of part of a zoarium viewed

rather from the right side, so that only the edges of the left-hand series
of apertures appear ; the alternate arrangement of the series of apertures
is therefore not obvious in the figure (p. 412).

8. Tervia dorsata (von Hagenow). x 12. Obverse face of part of a zoarium

(p. 413).

9. Tervia yibbera, Gregory, x 12. Obverse face of part of a zoarium viewed

from the left-hand side. The end of a " dorsal process " appears at the
top left-hand side of the figure, springing distally from the reverse face
(p. 413).

10. Entalophora virgula (von Hagenow). x 12. Part of a zoarium (p. 414).

11. Entalophora conjugata (von Reuss). x 12. Part of a zoarium. Some of the

apertures appear to be arranged in whorls or spirals ; but these are only
rows of three, and the series do not continue on the half of the branch
not shown (p. 415).

12. Entalophora echinata (Romer). x 12. Part of a worn zoarium (p. 415).

13. Spiropora verticillata (Goldfuss). x 12. Part of a zoarium (p. 417).

14. Entalophora madraporacea (Goldfuss). x 12. Part of a worn zoarium

(p. 416).
15-17. Coptosoma capense, sp. nov. Natural size. (p. 419).

Ann.S.Afp.Mus.Vol.VII.

PL I.

, '

17

T.A. Brock del.

Need's Camp Fossils.

West, Newman collo.

(21)

3. The, Invertebrate Fauna and Palceontological Relations of the
Uitenhage Series. By F. L. KITCHIN, M.A., Ph.D., F.G.S.

CONTENTS.

PAGE

I. INTRODUCTION 21

II. THE AGE OF THE FAUNA.

(a) Summary of Previous Work ... ... ... ... 25

(b) Additional Evidence for Age ... ... ... 30

III. COMPARISON WITH EXTRA-EUROPEAN FAUNAS.

(a) Possible Traces of a Belated Fauna in the South-

west of Madagascar ... ... ... ... 39

(b) Comparison ivith the Fauna of the Oomia Group in

Cutch ... ... ... ... ... ... 39

(c) Traces of a Related Fauna in the Godavari District

and in Hazara (N. W. Himalayas) ... ... 46

(d) Neocomian in German East Africa ... ... ... 46

(e) Neocomian in South America ... ... ... ... 49

IV. THE DISTRIBUTION OF THE UITENHAGE FAUNA IN EELA-

TION TO SOME THEORETICAL QUESTIONS ... ... 51

V. DESCRIPTIONS, CRITICAL NOTES, AND LISTS OF FOSSILS 60

VI. SUMMARY OF EESULTS 227

VII. LIST OF WORKS CITED 233

I. INTEODUCTION.

THE marine invertebrate fossils collected in 1900 by Mr. A. W.
Eogers and Mr. E. H. L. Schwarz from the Uitenhage Series, and
forwarded to me by Mr. Eogers on behalf of the Geological Commis-
sion, were found at localities near Uitenhage and below this on the
Zwartkop's Eiver, and in the valley of the Sunday's Eiver near and
below Dunbrodie. Further materials have been sent from the South

3

22 Annals of the South African Museum.

African Museum, principally comprising specimens presented by
E. Eubidge and C. A. Fairbridge, and I have also received a small
but interesting collection of Mollusca found by Miss M. Wilman in
the Uitenhage strata exposed in the Coega neighbourhood. In addi-
tion to the marine fossils obtained by Messrs. Eogers and Schwarz,
some shells of the genus Unio were collected in the Bezuidenhout's
Valley below Blue Cliff.

In 1905 Mr. Eogers devoted some time to a further examination
of the Uitenhage beds in the valleys of the Sunday's Eiver and
Coega Eiver, and made an additional collection of fossils ; he also
obtained a few specimens from an exposure of Uitenhage beds at
Brentford, in Knysna.* The results of an examination of these
supplementary materials have been incorporated in the following
account, which has been withheld from publication for this purpose.

In view of the extensive manner in which the fauna of the Uiten-
hage Series has already been made known, particularly by the
writings of Krauss, Sharpe, Tate, and Neurnayr, it is not surprising
to find that among the specimens collected during the preliminary
survey in the Uitenhage and Port Elizabeth Divisions comparatively
few new forms are represented. It could be gathered, however,
from the published account of this survey,! that more prolonged
search would in all probability add materially to an already lengthy
list, and, in fact, the specimens obtained by Mr. Eogers in 1905 well
bear this out, for they include a larger proportion of new species
than that shown by the collections made in 1900. J

In the following pages I have discussed the present state of our
knowledge regarding the palaeontological relations of the fauna, so
far as concerns the invertebrate remains, and then proceeded to the
description of the new materials submitted to me, while adding
critical remarks and supplementary descriptions relative to some of
the forms already known. Opportunity is also afforded by the pre-
sent study to describe and figure two Uitenhage ammonites which
are represented by specimens preserved in the collection of the
Geological Society of London, though not included in the material
sent to me from Cape Town ; it is useful to have some account of
these, both for the sake of comparison with other members of the

* A new locality for Uitenhage marine fossils ; see Schwarz (2), pp. 50, 74. [The
numbers in parentheses after the names of authors, in this and subsequent refer-
ences to literature, correspond with the titles of works bearing the same number
under the names of those authors in the bibliographical list with which this memoir
concludes.]

t Rogers and Schwars (1).

t For an account of the survey of 1905, see Kogers (2).

The Invertebrate Fauna of the Uitenhage Series. 23

same genus which occur in the Uitenhage beds and in order to
broaden the foundation for future work.

From statements made in the Eeports mentioned above, it seems
that in our present state of knowledge no satisfactory subdivision of
the Uitenhage beds is possible, and that the members of the series
are so variably developed that no consistent nomenclature is to be
hoped for until detailed mapping can be carried out. Of the sub-
divisions hitherto employed, the uppermost member of the series,
the Sunday's Eiver or Marine Beds, has yielded the majority of the
Mollusca which are discussed in this paper. The underlying
"Wood Bed" series has also furnished a few species of marine
molluscs in addition to Unio and remains of fossil plants. The so-
called Enon Beds represent the lowest division of the formation in
this district. Messrs. Eogers and Schwarz, while indicating the
local significance of the adopted subdivisions, draw attention to the
limited value of this nomenclature, and mention facts which clearly
show the contemporaneous variation of facies in the series. Thus,
to the north of Uitenhage, the Marine Beds appear to be synchronous
with part of the local conglomerate of " Enon " character, and it is
pointed out that at Plettenberg's Bay also, the Sunday's Eiver Beds
are partly replaced by conglomerate resembling that of the Enon
Beds, but here yielding Trigonia conocardiiformis, one of the most
characteristic fossils of the Marine Beds.*

The question of the age to be assigned to the Uitenhage Series, as
is well known, has called forth strikingly different opinions from
various authors. It was suggested by Stow in 1871 t that the want
of unanimity among the earlier writers might have been due to care-
less collecting and the mingling of specimens obtained from different
horizons. This author therefore made his own collections with due
regard to the localities and the individual bands in w r hich he found
the fossils to occur, and he attempted a correlation of the fossiliferous
beds exposed in sections at various places on the Sunday's and
Zwartkop's Eivers. It appears highly probable, however, from the
palaeontological evidence alone, that no very considerable extent of
time is represented by the w r hole of the beds which yield marine
fossils, and there is nothing to show that the different opinions
respecting the age of the series have been arrived at in consequence
of any radical change in the character of the fauna itself in its dis-
tribution through the strata. That any such marked change can be

* Schwarz (1), pp. 53, 61 ; Kogers and Schwarz (1), p. 5 ; Eogers (1), pp. 282-
296. See also Eogers (2), pp. 13, 15.
t Stow (1).

24 Annals of tlie South African Museum.

traced is not evident from the table of sections given by Stow.*
There are, no doubt, minor differences, and local distribution accord-
ing to facies in the successive fossiliferous bands, which may be
more clearly revealed when the district comes to be mapped in
detail ; but there is certainly nothing so far-reaching in this respect
as to have formed grounds for the differences of view expressed, for
instance, by Tate, who ascribed an Oolitic age to the fauna, and
Neumayr, who referred these beds to the Neocomian.f

The divergent conclusions arrived at by the earlier authors appear
rather to have resulted from the different interpretations put upon
the same peculiar association of forms when viewed in comparison
with the limited standards of European type. The principal difficulty
seems to have lain in the fact that none of the fossils could be cor-
rectly identified with those of formations studied in other regions,
and the facies of the fauna, taken as a whole, did not seem to show
such agreement with that of any known assemblage as to give it the
decided stamp which might serve to put the question of age beyond
dispute. This matter has been so frequently dealt with, and its
bearings are so well known, that it may seem superfluous to dwell
upon it here at any length ; but it may be useful to recapitulate
briefly the successive opinions expressed, and in some cases the
grounds upon which they were based, before proceeding to the more
thorough comparison of the fauna with its extra-European equiva-
lents which recent knowledge has rendered possible.

I take this opportunity of expressing my indebtedness to Mr.
A. W. Rogers for the helpful manner in which he has furnished me
with information relating to the collections ; to Dr. A. Smith Wood-
ward, Mr. G. C. Crick, and Mr. E. B. Newton for facilitating
reference to literature and specimens in the British Museum
(Natural History) ; and to Mr. W. Rupert Jones for his ready
assistance during rny repeated examinations of the extensive collec-
tion of Uitenhage fossils in the museum of the Geological Society of
London. My best thanks are also due to Prof. J. W. Gregory
for examining a specimen of Thamnastrcsa submitted to him, and to
Mr. C. D. Sherborn for assistance in a few matters relating to
bibliography ; to Mr. G. W. Lamplugh, Mr. H. A. Allen, and Mr. H.
Woods, I am indebted for several useful suggestions.

* Stow (1), fig. 3. See further remarks on this subject at the end of Section II.
of the present paper.

f In this connection see Neumayr's remarks on Stow's work ; Holub and
Neumayr (1), p. 270, footnote 6

The Invertebrate Fauna of the Uitenhage Series. 25

II. THE AGE OF THE FAUNA.

(a) Summary of Previous Work. In 1837, Hausmann recorded
the occurrence of some shells obtained by Hertzog from strata in
the Sunday's Eiver district, to which he ascribed a Lower
Cretaceous age. These included a Hamites, which was compared
with H. intermedius J. Sow. and H. funatus Brongn., and a Trigonia
which was thought to resemble T. dcedalca Park.* Goldfuss after-
wards described and figured two of Hausmann's shells under the
names Lyrodon herzogi and Cytherea herzogi, and also considered
them to be of Greensand age.f

The examination of a small collection of lamellibranchs obtained
by F. Krauss in 1839 from strata exposed on the Zwartkop's Eiver,
led that author to the conclusion that they indicated a Lower Green-
sand horizon,| and in another paper Krauss furnished excellent
descriptions and figures of these shells, maintaining a similar view
concerning their age.

In 1851 a collection of fossil plants and molluscs, obtained by
E. Eubidge on the Sunday's Eiver, was exhibited before the British
Association at Ipswich by Colonel Portlock, who remarked that the
shells were apparently of Jurassic age, while Dr. Harvey's examina-
tion of the plant remains was thought to corroborate this view. ||

In his well-known paper published in 1856, A. G. Bain 5! referred
the Uitenhage fossils with doubt to the Lias, basing this opinion
upon the prevalence of a supposed Liassic form, " Gryphcea incurva."
The shell mistaken for this, however, was Exogyra imbricata, pre-
viously described by Krauss, which, as we shall see, bears only a
superficial resemblance to the Gryphcea mentioned, and is in reality
closely comparable with certain Lower Cretaceous forms of Exogyra.
Appended to Bain's paper were D. Sharpe's descriptions of the
Secondary fossils collected by Atherstone and Bain from localities
on the Sunday's and Zwartkop's Eivers. The forms described,
principally Mullusca, led Sharpe to the conclusion that they most
nearly resembled European species of the Middle and Lower
Oolites ; he compared his Ammonites atherstoni with A. macro-
cephalus and A. herveyi, while believing Ammonites baini to be
related to A. humphresianus and other Lower Oolitic forms.

* Hausmann (1), p. 1457.

t Goldfuss (1), Band II., p. 202, pi. 137, fig. 5(1837); p. 239, pi. 149, fig. 10
(1840).

I Krauss (1), pp. 129, 130. Krauss (2).

|| Portlock (1). H Bain (1).

26 Annals of the South African Museum.

In 1867, a paper by Ealph Tate," in which many new forms were
described and figured, added largely to our knowledge of the
Uitenhage fossils, and this author was led to some remarkable con-
clusions from his study of the fauna. He believed the assemblage
to indicate a Jurassic age, and stated that he thought it to represent
the fauna of the Oolitic rocks of Europe, and to approximate to that
of the Great Oolite. It seems clear that in instituting this com-
parison, he made use of some molluscan types little adapted to serve
the purposes of a critical correlation, and he was at the same time
misled by several quite erroneous identifications. He also mis-
understood the affinities of the cephalopods and the significance of
certain Trigonice which alone might have been expected to form
obstacles to his conclusions. It is here scarcely necessary to do
more than refer to the curious generalisation arrived at by Tate
concerning the relation of these supposed Jurassic deposits to the
Jurassic strata of Europe, namely, that the " Oolites " of South
Africa 'are the representatives of the whole of the Jurassic rocks of
Europe with the exception of the Upper Oolites, and illustrate an
intermingling of palasontological types which are analogous to, or
identical with, those distributed in successive zones in Europe.

In his monograph on the Cretaceous lamellibranchs of Southern
India, Stoliczka i made some reference to Uitenhage forms. He
evidently believed Tate's Crassatella complicata to belong to the
genus Ptychomya, and he ascribed Astarte herzogi Krauss to
Speyer's genus Grotriania. He further expressed the opinion that
Krauss's Astarte bronni might belong to the Cretaceous genus
Eemondia Gabb, and thought that in addition to these and Trigonia
ventricosa, several other Uitenhage shells show a Cretaceous rather
than a Jurassic aspect ; attention was drawn to the great similarity
between Trigonia ventricosa (Krauss) and the Cretaceous T. tuber -
culifera Stol., from Southern India. While we shall see that
Stoliczka rightly recognised some of the Uitenhage forms to exhibit
Cretaceous affinities, he was in error in ascribing Astarte herzogi to
the genus Grotriania, and, as afterwards pointed out by Neumayr,
wrongly supposed Astarte bronni to belong to the genus Eemondia.
A. bronni is so distinctly characterised that Neumayr proposed for
it the new generic name Seebachia a fact which appears to have
been overlooked by Stanton, who in 1897 still tentatively included
it in the genus Itcmondia.\

One of Tate's Uitenhage species, the so-called Crassatella com-

* Tate (1). f Stoliczka (2), pp. 286, 294, 315 (1871).

J Stanton (1).

The Invertebrate Fauna of the Uitenhage Series. 27

plicata, was later also recognised by Dames to be a representative
of the genus Ptychomya,* and this seemed to point to a later age for
the strata from which it was obtained than that assigned by Tate.
Dames was further led to the belief in the Neocomian age of the
Uitenhage beds by a fragment of an ammonite (sent by Krauss to
L. von Buch) which he thought to be identical with Ammonites
astierianus d'Orb.

In the concluding chapter of his monograph on the British Fossil
Trigonice, Lycettf referred briefly to the TrigonidR of the Uitenhage
Formation, and expressed his opinion that some of the most
characteristic of these point decisively to a Cretaceous age ; he
also showed that the alleged occurrence of T. goldfussi, which Tate
had used as evidence for a Jurassic age, rested on an erroneous
determination.

The whole question of the age of this fauna was afterwards well
handled by Neumayr, J who subjected Tate's work to some criticism.
Neumayr set on one side many molluscan types as of little signifi-
cance in a comparative study, and concluded that a costate Trigonia
(T. tatei Neum.) alone exhibited a marked Jurassic character. On
the other hand, he considered a number of forms to represent
essentially Cretaceous types. Such were Holcostephanus atherstoni
(Sharpe) ; Holcostephanus baini (Sharpe) ; Crioceras sriinosissimum
(Hausm.) Neumayr ; Trigonia ventricosa (Krauss) ; Trigonia cono-
cardiiformis (Krauss) ; Ptyclwmya complicata (Tate); and Exogyra
imbricata Krauss. He suggested that Tate's Ammonites subanceps,
which was thought by Tate to resemble the Jurassic A. anceps Kein.,
might really represent the young of Crioceras spinosissimum. It
was admitted by Neumayr that Belemnites africanus Tate, which
Tate placed in the group Canaliculati and considered to afford
strong evidence for an Oolitic age, bears a strong resemblance to
the Jurassic forms B. canaiiculatus Schloth. an'd B. magnificus
d'Orb. ; but at the same time he drew attention to the existence of
a belemnite in the Lower Cretaceous of North Germany which
seemed to share some of the characteristics of B. africanus. In a
later paper, written after an examination of Tate's original specimen
in the collection of the Geological Society, Neumayr || definitely
separated B. africanus from the Canaliculati, and included it in his
group of the Absoluti, which, as he remarked, extend in their
occurrence up to the Aptian ; hence the conclusive nature of the

* Dames (1). f Lycett (3), p. 230 (1879).

{ Holub and Neumayr (1). Misquoted " implicata " by Neumayr.

|| Neumayr (4).

28 Annals of the South African Museum

evidence for age furnished by the occurrence of B. africanus in the
Uitenhage beds was no longer to be recognised." Neumayr's
argument for the Lower Cretaceous age of this fauna appears indeed
to be overwhelmingly strong, particularly in the evidence of the
Cephalopoda, all of which he found to possess near allies in European
Neocomian forms. Regarding the two species of Holcosteplianus ,
Neumayr pointed out that their resemblance to the Jurassic types
with which they had previously been brought into comparison was
only of a superficial character, depending alone on a certain outward
similarity of habit.

Pavlow f has since confirmed this view, after studying South
African specimens, and has placed Holcosteplianus atherstoni and
H. baini in his generic or sub-generic division Astieria, which
comprises numerous Neocomian types ; he even goes so far as to
identify Holcosteplianus atherstoni with H. psilostomus Neum. and
Uhlig, { from the Hilsthon of North Germany, and to the same form
he ascribes a shell from the Neocomian of Speeton.

In view of the fact that the plant remains found in the Uitenhage
beds had previously been thought to point rather to an Oolitic than
a Cretaceous age, and having regard to the division of opinion con-
cerning the affinities of the invertebrate fauna, Messrs. Rogers and
Schwarz, in 1901, were led to adopt the provisional conclusion that
the Uitenhage Series may be assigned to the Upper Jurassic. In
stating this, they mention that the fauna and. flora have been con-
sidered to resemble in some degree those of the Jurassic series in
Cutch. It is well known that the resemblance between certain
lamellibranchs of the Uitenhage beds and those of the Oomia Group
in Cutch has been frequently remarked upon, but it can no longer
be maintained that the marine Oomia strata are in reality of Jurassic
age, at least so far as concerns the Triyonia-beds ; but this is a
point to which we may presently return, and one which we may
consider in greater detail.

In more recent works Mr. Rogers || gives us a comprehensive
account of the Uitenhage Series, and now considers these beds to
represent a portion of the Cretaceous system. The same view
is adopted by Drs. Hatch and Corstorphine in their " Geology of
South Africa" (1905). With reference to the flora, Prof. A. C.

* See also Neumayr (5).

t Pavlow and Lamplugh (1), pp. 492-497 (134-139 in authors' copy).

I Neumayr and Uhlig (1), p. 149, pi. xxxii., fig. 2.

Eogers and Schwarz (1), p. 17.

|| Eogers (1), pp. 281-318; Kogers (2), pp. 15-33, 45.

The Invertebrate Fauna of the Uitenhage Series. 29

Seward, in his memoir on the subject," has shown that the Uiten-
hage plants "include types in part characteristic of Wealden and
in part indicative of Jurassic floras " ; but he believes that the
balance of evidence derived from the plants is in favour of a
Wealden age.

We thus see that recent authoritative opinions have lent strong
confirmation to the views of the earliest writers who made a study
of this fauna. Other works in which less critical reference has been
made to the age of the Uitenhage Series may now be briefly noticed.

In 1857, W. G. Atherstone suggested the partly Jurassic and
partly Cretaceous age of the Uitenhage beds,t but Andrew Wyley,
in 1859, placed the Enon Beds so low as the New Red Sandstone,
and correlated the Sunday's Eiver Beds with the Jurassic (Oolites).:,:

Feistmantel evidently shared Tate's view of an Oolitic age when
he tried to show affinities between certain Mollusca from the Uiten-
hage beds and others from the Oomia Group in Cutch in order to
prove the Lower Oolitic age of the Oomia fauna and bring about
harmony between the evidence for age furnished by the plants and
animals from those beds. In a correlation of the Indian and African
]Mesozoic formations the same author also tabulated the Uitenhage
Series as Jurassic. || G. W. Stow divided the formation into a
"Lower Jurassic" and an "Upper Jurassic" series. *i In 1878
W. T. Blanford, basing his view upon a study of Tate's and Stow's
papers, concluded that the beds containing Hamitcs yielded too large
a number of Middle Jurassic forms for reference to the Neocomian ;
but he believed them to represent a very high Jurassic horizon,
while pointing out that Trigonia ventricosa and T. van were still
higher/ 1 "

In 1880 Griesbach classed the Uitenhage beds as Jurassic, ft while
the same view was adopted by T. Rupert Jones in 1884 I j and by
Moulle in the following year.^ Even so recently as 1897, Futterer
has accepted Sharpe's and Tate's conclusions. || || On the other hand,
Giirich.V, Schenck, :;:: :;: and Molengraaff f f f have referred these beds

* Seward (1), p. 46. t Atherstone (1), pp. 584, 588.

I Wyley (1). [I have not seen Wyley 's Report, but his correlation is set forth
in Tate (1), p. 172, and Corstorphine (1), Appendix.]

Feistmantel (1). || Feistmantel (4), pp. 54, 59, 84.

1 Stow (1). ** W. T. Blanford (1), p. 118.

ft Griesbach (1), pp. 90, 93. + + Jones (1) ; Jones (2), p. 737.

Moulle (1), p. 216. Illl Futterer (1), p. 625.

ITU Giirich (1). ** Schenck (1), p. 231.

ftt Molengraaff (1). [I have not been able to see this work, but quote the
statement and reference on the authority of Newton (2), p. 146.]

30 Annals of the South African Museum.

to the Lower Cretaceous ; but in a more recently published tabular
correlation of the geological formations of the Transvaal and Cape
Colony, Molengraaff has ascribed them with a note of interrogation
to the Middle Jurassic." In 1896 E. B. Newton published a useful
summary of previous work on the Cretaceous conchology of South
Africa,! and furnished a complete list of the known Mollusca from
the Uitenhage beds, which he classed as Neocomian. In their text-
books de Lapparent J and Kayser have followed Holub and Neu-
mayr in assigning a Neocomian age. Passarge, in his work on the
Kalahari, |j has tabulated the Uitenhage Series with the Upper
Jurassic and Lower Cretaceous. Lemoine believes it to be of
Lower Cretaceous age.^ 7

(b) Additional Evidence for Age. We see from the foregoing that
a comparison of the Uitenhage Mollusca with European forms
seems to point definitely to a Lower Cretaceous age, for it will
be conceded that the evidence of the Cephalopoda must be allowed
to carry the greatest weight in influencing a decision. The repre-
sentatives of Holcostephanus (sensu stricto), Hamites and Crioceras
already recorded, supply in themselves sufficiently striking evidence,
and notable additions to these are other forms of Holcostcphanns, as
well as representatives of Acanthodiscus and Bochianites included
in the collections submitted for examination and described in these
pages. Two other species of Holcostephanus (sensu stricto), hitherto
undescribed, are represented in the collection of the Geological
Society of London. One of these is apparently allied to H. ather-
stoni (Sharpe), but has greater lateral compression as well as other
distinctive characters. It closely resembles H. psilostomus Neum.
and Uhlig, of the European Neocomian, and is the "compressed
variety" of H. atherstoni mentioned by Pavlow, :;::: who aptly sug-
gested close relationship to a shell from the Neocomian of Spain,
figured by Nickles as Holcostephanus hispanicus. The significance
of such an assemblage of Cephalopoda in the Uitenhage beds cannot
be mistaken, quite apart from the bivalve forms to be mentioned
below and a representative of the Crustacean genus Meyeria. In
Europe the known species of Holcostephanus (sensu stricto) are
almost wholly, if not entirely, confined to strata of Upper
Valanginian and Lower Hauterivian age.

In his paper on the Uitenhage fauna E. Tate ft gave a tabular list

* Molengraaff (2), p. 119. t Newton (2).

I de Lapparent (1), p. 1267. Kayser (1), p. 444.

|| Passarge (1), p. 82; see also p. 597. IF Lemoine (1), pp. 383, 389.

** Pavlow and Lamplugh (I), pp. 492, 496, (134 and 138 of authors' copy).

ft Tate (1), p. 166.

The Invertebrate Fauna of the Uitenhage Scries. 31

of some of the more important fossils, together with their supposed
extra-African allies and analogues, which, with one exception, were
thought to be Jurassic. An examination of this list shows, however,
that a different construction must now be put upon the comparisons,
as may be demonstrated by the mention of a few examples. In
addition to Holcostephanus atlierstoni and H. baini, which were
erroneously thought to be allied to well-known Oolitic forms, there
is Tate's Ammonites subanceps, a single specimen, regarding which
that author wrote : " It is doubtful whether this be not a mere variety
of A. anceps Eeinecke, of the Middle Oolites." As already men-
tioned, Neumayr thought that it might represent a stage in the
individual growth of Crioceras spinosissimum, which he supposed
to have had the whorls in contact at such an immature period.
This may be a correct view, but it seems equally probable that
we are dealing with an immature example of some species of
Hoplites (sensu lato). Tate's specimen, of which his published
figures convey a very imperfect and even misleading impression,
shows great similarity to certain forms which have been ascribed
by Sayn to Hoplites arnoldi (Pict. and Camp.), and by Toucas
to H. botellce Kilian. These were thought by Pavlow to have
been wrongly identified, and are united by him under the name
Hoplites lieteroptyclius, which has more recently been included by
Uhlig in his narrower generic group Solgeria. A somewhat similar
form has been described and figured by Bogoslowsky under the
name Hoplites aff. arnoldi Pict., and this is said to occur with
Belemnites lateralis Phill. above the " Ejasan-Horizont " in Eussia
(Mostja Eiver). : ' : Hoplites subanccps is more closely and less
coarsely ornamented than this, but there are points of agreement
in the type of sculpture. Further observations on "Ammonites"
subanceps are given in the remarks which follow the description
of a specimen ascribed to Acanthodiscus sp., in the following pages.

Ostrea jonesiana Tate was compared by Tate with 0. costata Sow.
from the Oolites ; but it is in reality a true Exogyra quite compar-
able with shells of the type of E. suliplicata Eoem.,t from the
Hilsconglomerat of North Germany.

Pectcn projectus was compared with P. lens Sow., but such a com-
parison must have little value when we remember that species of
Pecten with similar broad characteristics recur at various horizons
and are not confined to the Jurassic rocks. Pecten projectus, in

* Bogoslowsky (1), pp. 112, 138; Taf. vi., fig. 7.

f F. A. Roemer (1), Nachtrag, p. 25, pi. xviii., fig. 17; F. A. Eoemer (2), p. 47
(1840).

32 Annals of the South African Museum.

fact, compares more satisfactorily with some of the Cretaceous
forms.

The comparison of Pecten rubidgeanus with P. subspinosus Schloth.
cannot be pronounced at all fortunate, while we shall find that Lima
obliquissima shows a closer agreement of characters with Cretaceous
and later forms than with the European Jurassic shells cited by
Tate.

Placunopsis subjurensis Tate, which was compared with P. jurensis
Boern., may perhaps be an Anomia, and in some respects closely
resembles A. pseudoradiata d'Orb., * from the Aptian. Placunopsis
imbricata Tate is compared in the table with P. substriata Lye.,
from the Oolites. P. scmistriata (Bean) was probably the English
shell with which it was intended to institute comparison, since this
name alone is mentioned in Tate's description on page 154 of his
paper. P. imbricata, however, is without radial ornamentation, but
is characterised by well-spaced concentric ridges which recall the
similar ornamentation of Anomia neocomiensis d'Orb., t though
occurring in less emphasised form. The illustration of P. imbricata
gives a very poor idea of the shell, but an examination of the
specimen dispels the notion of a close comparison with known
Jurassic forms.

Trigonia cassiope Tate is not to be united with any known Jurassic
costate Trigonia, and was accordingly renamed T. tatei by Neumayr.
It is well distinguished by the elongated escutcheon, the very
delicate marginal carina, and the fine character of the ornamentation
on the area and escutcheon. Trigonia goldfussi Tate is an immature
shell, and in all probability illustrates the youthful stage of one of
the large and robustly ornamented Pseudo-quadratae which form
such a striking feature in the Uitenhage fauna. It certainly does
not represent either Trigonia goldfussi Agassiz or T. painei Lye.
( = T. goldfussi Morr. and Lye.), with which it was identified. Tate
himself, on page 159 of his paper, said : " Hereafter, however, it
may be found to be the young of a distinct species." It is therefore
the more regrettable that the error of quoting this European Jurassic
shell from the Uitenhage beds should have continued to be perpe-
trated. Stow I repeatedly used the name " T. goldfussi " in his
paper, but I have been unable to ascertain to which of the Trigonia
he referred.

No importance can be attached to the comparison of such types as

* d'Orbigny (4), p. 84 ; Woods (3), vol. L, p. 27, pi. v., figs. 1-3 (1899).
t d'Orbigny (3), p. 754, pi. 489, figs. 1-3 (1848).
t Stow (1), pp. 497-514.

The Invertebrate Fauna of the Uitenhagc Series. 33

Pinna atherstoni Sharpe and P. sharpei Tate with Jurassic forms,
for it is evident that these representatives of the genus lack any
strong distinctive characters which alone would render them of
value in an estimate of geological age.

Nytilus baini Sharpe was compared with M. sowerbiamis d'Orb.,
from the Oolites of Europe. It is a Modiola which certainly has
characters of form and ornamentation closely similar to those which
distinguished M. sowerbiana (d'Orb.) and M. perplicata (Etallon) ;
but this type of Modiola is by no means confined to Jurassic rocks,
having also a widely distributed representative in the Cretaceous
M. flagellifera (Forbes), to mention a well-known example.

Plioladomya dominicalis Sharpe was thought to resemble Liassic
and Oxfordian forms, but it may equally well be brought into com-
parison with Lower Cretaceous representatives ; for example, the
European Aptian shell ascribed, rightly or wrongly, to P. pedernalis
Roem.*

Astarte pincliiniana Tate was brought into comparison with

A. pumila Goldf. and A. excentrica Morr. and Lye., from the Great
Oolite of Europe. The African shell differs in important respects
from these, and, as will be pointed out below, it probably belongs to
the sub-generic group Eripliyla and is certainly most closely com-
parable with Cretaceous forms.

Berenicca antipodum Tate was believed by Tate to be related to

B. striata Haime, from the Lower Lias of France, but has been
regarded by Prof. J. W. Gregory t as a close ally of the Cretaceous
B. gracilis (Milne Edwards).

Two Uitenhage Serpuhe were identified by Tate with S. filaria
Goldf. and S. plicatilis Miinst., from the Inferior Oolite of Europe.
The specimens ascribed to S. plicatilis represent an adherent form
sharply carinated on the back, with a weaker carinal angle on either
side and a circular aperture, but the true S. plicatilis Goldf. has only
a single central, longitudinal keel. S. qmnquangularis Goldf. J
(Upper Jurassic) stands much closer in its characters, and a Serpula
scarcely distinguishable from this occurs in the Lower Cretaceous of
Europe, and may be found named S. quinquangularis in some collec-
tions. S. filaria Goldf. is a solitary, wholly adherent form, com-
mencing with a coiled stage. So far as can be seen from the
imperfect African specimen named S. filaria by Tate, this is a
clustering form consisting of crowded, narrow cylindrical tubes, and

* Pictet and Kenevier (l),p. 60, pi. vi., fig. 7, 1855 ; Moesch (1), p. 93, pi. xxxiii.,
fig. 2. f Gregory (1), p. 112.

\ Goldfuss (1), Band i., p. 230, pi. 68, fig. 8 (1831).

34 Annals of the South African Museum.

it is not distinguishable from the English Lower Cretaceous Serpula
that have been variously referred to Serpula filiformis J. de C. So\v,
and S. plexus J. de C. Sow. We thus see that in the case of the
two Trigonia and the two Serpula which Tate identified with Euro-
pean Jurassic types, the determinations were so far erroneous that
they are now bereft of the significance which was attached to them
as evidence for the age of the fauna.

We may further examine the evidence for age afforded by some
of the more conspicuous bivalves, and firstly I may briefly refer to
the representatives of the genus Trigonia, which in themselves lend
such a characteristic aspect to the fauna. Although the testimony
of these is perhaps not quite so conclusive as Lycett supposed, still,
the balance of evidence furnished by members of this genus may be
safely relied upon to give a fair indication of age, even when con-
sidered apart from associated forms, and the material we are dealing
with in the present instance is fortunately of an exceptionally favour-
able character. In the first place there is Trigonia ventricosa,
T. kraussi, and T. rogcrsi, all possessing the characters of the section
Scabrae a division which, so far as we know, is elsewhere only
represented in rocks of Cretaceous age. These might be considered
to be balanced by the occurrence of T. tatei, a typical member of the
section Costatae, but although this division of the Trigonia has its
maximum development in the Oolitic rocks, it is also represented in
strata of Lower Cretaceous and even later age.

Trigonia peninsularis Coquand,* from the Aptian of Spain, cer-
tainly shows characters of degeneration not shared by T. tatei, but
the Oomia strata of Cutch have yielded two normal representatives
of the Costatae, which, together with some degenerate derivatives of
this section, are accompanied by Trigonia: which exhibit a decidedly
Cretaceous aspect. Another unmodified costate form, T. anguste-
costata Behr. f occurs in the Argentine Eepublic in strata which
are regarded by Behrendsen as Upper Cretaceous, while a typical
member of this section has been described by E. Ascher from the
Grodischter beds (Hauterivian) of Silesia. }

Trigonia herzogi (Goldf.), another very characteristic Uitenhage
form, was believed by Lycett to belong to the Quadratae, which, so
far as known, are exclusively Cretaceous. Steinmann, however,
pointed out that Trigonia herzogi differs in several important par-

* Coquand (1), p. 129, pi. xxiii., fig. 3.

f Behrendsen (1), p. 6, pi. 3, fig. 7.

I Ascher (1), p. 159 [25] , p. xiii. [ii.], fig. 10.

Steinniann (2).

The Invertebrate Fauna of the Uitenliage Scries. 35

ticulars from all known Quadratae, and found that the special features
which distinguish this Uitenliage shell from the Jurassic Clavellatae
on the one hand and the Cretaceous Quadratae on the other, are
shared by Trigonia transitoria Steinm. from Lower Cretaceous rocks
in Bolivia, Chili, and the Argentine Republic. He therefore sug-
gested a new group-name, the Pseudo-quadrate, to comprise these
two TrigonicB, which were the only examples known to exhibit these
peculiar distinguishing characters. Trigonia neuquensis Burckhardt,*
which occurs with T. transitoria in Neocomian rocks at Las Lajas
on the Eio Agrio (Argentine), falls under the same heading, while
T. mamillata I from the Oomia strata in Cutch must also be referred
to this group. Still another form, described in these pages (T. Jwlubi),
is clearly referable to the same category. Although the five members
of the Pseudo-quadratae known to us are all similarly differentiated
from the typical Clavellatae, it is not certain that they are all so
closely related to one another as might at first sight appear to be
the case ; some, at least, may possibly have been independently
evolved from clavellate ancestors. :[ Nor is it by any means certain
that they bear any close and direct relationship to the true Quadratae,
and though definite indications on this point fail us, it is quite
probable that the Quadratae and Pseudo-quadratas represent parallel
and independent groups of derivatives arising from Jurassic Clavel-
latse. It is therefore evident that the Trigonia of the group Pseudo-
quadratae cannot yet be considered in themselves to furnish quite
such reliable guidance to geological age as the true Quadratae, but at
the same time their general aspect, by comparison with the Quadratas,
strongly suggests a Cretaceous age, while the known South American
representatives must, in fact, be regarded as Neocomian. Some of
the most important features wherein the Quadratae differ from the
Clavellatae are also exemplified in the Pseudo-quadratae. Lycett
laid great emphasis on the presence or absence of sculpture on the
escutcheon in distinguishing between the Jurassic Clavellatae on the
one hand, and the Cretaceous Clavellatae and Quadratae on the other,
and in regard to this feature all the Pseudo-quadratae agree with the
Quadratae in having coarsely nodose ornamentation on the escutcheon.
Further remarks on the points of agreement between these groups
are appended to the description of Trigonia holubi.

Trigonia conocardiifonnis (Krauss) was thought by Lycett to fall

* Burckhardt (2), p. 74, Taf. xiv., figs. 4-6.
t Kitchin (1), p. 100, pi. ix., figs. 8, 9 ; pi. x., figs. 1-3.

I Kemarks on this subject will be found to follow the description and the dis-
cussion of the relationships of Trigonia liolubi, in these pages.

36 Annals of the South African Museum.

within the section Scabrae, but a near relationship to members of
this division is scarcely to be inferred from the adult characters
displayed by this peculiar form, while a study of the young shell
does not plainly reveal its affinities. A general comparison, how-
ever, shows that this Trigonia differs from the Jurassic Clavellataa
in a manner somewhat analogous to that which distinguishes the
Scabrae, and a very closely similar and probably intimately related
form occurs in the Neocomian of the Argentine Republic .

On the whole, so far as this Trw/ojw'a-assemblage is comparable
with European forms, it must certainly be considered to display a
Cretaceous rather than a Jurassic character, though when regarded
collectively, it is without counterpart in the European area. The
significance of other Trigonics included in this fauna will be discussed
presently.

Still further indications of geological age are to be derived from
a comparison of the Uitenhage lamellibranchs with European types.
One of the shells submitted to me, which proves to be identical with
a form previously described by Sharpe, who did not recognise its
true generic position, is an example of the well-characterised genus
Thetironia, which in Europe is w r idely distributed in Lower
Cretaceous and higher strata, though it has not been recorded from
rocks of an earlier age. A second species of Thetironia is also
included in the collection. Solecurtus is another genus not known
to occur in rocks older than Cretaceous, and a representative of this
must now be added to the list of Uitenhage molluscs. The speci-
mens sent to me for examination also include examples of Pccten
which can only be identified with the Cretaceous P. orbicular is
J. Sow. and P. cottaldinus d'Orb., while a third form may be most
aptly brought into comparison with P. subacutus Lam. A repre-
sentative of the Cretaceous genus Anthonya must also be recorded
in this connection.

Gervillia dentata Krauss, though belonging to the group of
G. aviculoides,* typically represented in Jurassic rocks (G. deecki
Freeh ; G. aviculoides Sow.), has a close counterpart in G. anccps
Desh. and G. sublanceolata d'Orb., in the Lower Cretaceous of Europe.
A form closely similar, and perhaps identical, occurs also in the
Neocomian of German East Africa. The long lateral tooth which
Krauss thought to be so highly distinctive of G. dentata as almost
to justify the establishment of a new genus, is a normal feature of
the group, and is well developed in G. anceps.\ Another common

* Freeh (1). t Freeh (1), pp. 612, 613.

The Invertebrate Fauna of the Uitenhage Series. 37

and characteristic Uitenhage shell is Exoyyra imbricata Krauss, and
this belongs to a general type well exemplified in the Lower
Cretaceous strata of Europe and South America. Lima neglecta
Tate, belongs to the sub-generic division Mantellum, and most
closely resembles forms which occur in the Lower Cretaceous of
Europe. Eeference has already been made to Ptychomya complicata
(Tate), which is a typical representative of a genus widely dis-
tributed in the Lower Cretaceous rocks, where it first makes its
appearance.

It is abundantly evident, then, that many of the most important
and distinctive molluscan forms, amongst those with which we are
dealing, give similar indications of geological horizon. There can
be no doubt, indeed, that the marine Uitenhage fauna is of Neocomian
age, though it is perhaps not possible to arrive at a very precise and
positive estimate of equivalence with the divisions of the Neocomian
in Europe, owing to the want of close coincidence in the faunas as
developed in such widely separated regions. But Holcostephanus
atherstoni (Sharpe) and H. wilmance, sp. nov., if not actually repre-
sented in Europe, have very close allies there in the Valanginian
and Hauterivian, and the Uitenhage Marine Beds may, in my
opinion, be certainly correlated with these divisions. Further, it
seems to me highly probable that the Marine Beds represent not
more than the strata at the top of the Valanginian and the base of
the Hauterivian.

The possibility that the lowest marine strata may be of greater
age than Neocomian is not only most remote, but is contradicted
by the palaeontological evidence, so far as this goes. Fossiliferous
strata, yielding a marine molluscan fauna characterised by the
prevalence of Gasteropoda and the remains of oysters, occur at
Dunbrodie (Sunday's Eiver) associated with plant-bearing beds, and
are in the lower part of the Uitenhage Series ; according to Messrs.
Eogers and Schwarz they may be classed with the so-called "Wood
Bed " series. Amongst the fossils collected here are Actceonina
atherstoni (Sharpe), Cyprina rugulosa Sharpe, and a Pecten which
I have identified as Pecten cottaldinus d'Orb. While Actceonina
atlierstoni is found also in the highest part of the Marine Beds
on the Zwartkop's Eiver and with the characteristic TrigonicB on the
Sunday's Eiver, Stow recorded the occurrence of Cyprina rugulosa
with similar associates. Although the Wood Bed series was not
seen in the Zwartkop's Eiver Valley below Uitenhage, the lowest
part of the Marine Beds in the Zwartkop's Eiver section, exposed
in a clay-pit near Eawson Bridge, yielded Actceonina atherstoni and

4

38 Annals of the South African Museum.

shells most probably referable to Bochianites glaber sp. nov. ; the
latter has close affinities with European Neocomian forms and
occurs at a higher level in the Marine Beds one mile from Eawson
Bridge on the main line, up side. It is also worthy of remark that
the lowest marine strata found, either at Dunbrodie or in the
Zwartkop's Eiver Valley, yielded no single form which suggests
stronger affinities to Upper Jurassic than to Lower Cretaceous
types ; but those very forms, such as Trigonia tatei Neum. or
Tancredia schwarzi sp. nov., which if considered alone might with
some reason have been thought to point to a Jurassic age, occur in
the higher part of the Marine Beds, associated with characteristic
Mollusca of undoubted Neocomian type.

During his visit to the Sunday's Eiver district in 1905, Mr. Eogers
found additional evidence to show that the whole of the Uitenhage
beds there exposed were deposited with comparatively great rapidity.
He found that forms which are represented in the strata by abundant
individuals occur much more generally distributed throughout the
whole thickness of beds than was at first suspected. Some of the
most typical species of the marine facies (for instance, the familiar
TrigonicB) were observed to occur, together with layers of lignite,
at the very base of the Sunday's Eiver Beds, where Mr. Eogers had
expected to find forms characteristic of the Wood Bed facies. These
marine forms occur also 300 feet higher in the series, and since the
lower beds of the series were seen to be assuming, in some degree,
the aspect of the Wood Bed development, the discovery of the
familiar marine forms in them occasioned some surprise. Mr.
Eogers noted the occurrence of Hamites near the mouth of the
river, where Stow obtained it at first, but he also found it high
up the river in a rock which showed resemblance to the strata of
the Wood Bed series. He concludes from his observations during
this second survey, that Stow must have been led through insufficient
collecting to assign a too restricted vertical distribution to various
species. Mr. Eogers believes that from the same cause his own
records are likely to prove faulty, and that exhaustive collecting
\vould still more fully demonstrate the extensive vertical distribution
of many of the forms throughout the beds exposed. :;:

The above considerations lend great support to the results of a
comparative study of the fauna, namely, that no forms are known
to occur in these beds which give definite indications of the presence
of more than the equivalent of a single palteontological stage.
Allowing, then, that the cephalopods, supported in no small degree

* For the published account of the 1905 survey, consult Eogers (2), pp. 15-33.

The Invertebrate Fauna of the Uitenhage Series. 39

by the lamellibranchs, afford satisfactory data for a comparison with
European standards, we may safely conclude that no portion of the
Uitenhage Series represents a period of time earlier or later than
the Neocomian. It must be said, indeed, that the almost entire
restriction of Holcostephanus, sensu stricto (= Astiena Auctorum),
to the upper part of the Valanginian and lower beds of the
Hauterivian in Europe * suggests much narrower limits, when we
consider how important a place is taken by members of this genus
in characterising the cephalopod-fauna of the Uitenhage beds.

Leaving, now, the consideration of evidence which leads to these
conclusions, we may proceed to compare the Uitenhage fauna with
those occurrences in extra-European regions which, in greater or
less degree, bear the imprint of a similar facies.

III. COMPABISON WITH EXTRA-EUROPEAN FAUNAS.

(a) Possible Traces of a Related Fauna in the Soiith-west of
Madagascar. Douvill has drawn attention to the occurrence
of some lamellibranchs obtained by Lieutenant Boutonnet from
deposits situated in the Fiherenga Valley, in the basin of the
Isakondry River, east of Tullear.f Mention is made of a large
Trigonia, said to be analogous to some of the Trigonice of the Oomia
beds in Cutch and of the Uitenhage Series, and with this is asso-
ciated a shell referred by Douville to the genus Pycnodonta, and said
by him to be closely comparable with Exogyra imbricata Krauss
(also referred to Pycnodonta by Prof. Douville). On the evidence of
these fossils, the strata containing them are considered by Douville'
to be of Cretaceous age, and Lemoine classes them provisionally as
Lower Cretaceous. The account of these fossils so far published
is unfortunately meagre, and it must here suffice to have drawn
attention to the occurrence of forms in Madagascar which may
possibly indicate the presence of a fauna of Uitenhage character.

(6) Comparison with the Fauna of the Oomia Group in Cutch.
Striking data for correlation are to be found among the lamelli-
branchs which constitute so large a part of the Uitenhage fauna,
and it becomes apparent that some of the conspicuous elements
which help to lend a definite character to the assemblage, while
finding no exact parallel in the European area, have an unmis-
takable counterpart in the fauna of the Oomia Trigonia-beds. This

* In North Germany, for instance, the restriction of these forms to such narrow
limits is well marked. See von Koenen (3), pp. 4, 8, 9.

t Douville (2), p. 388; Douville (4), p. 215 ; Lemoine (1), p. 176.

40 Annals of the South African Museum.

agreement is the more remarkable from the fact that in seeking
additional data for comparative study among the cephalopods of the
Oomia beds, the indications of community furnished by the lamelli-
branchs are found to obtain no positive support. The cephalopods
described by Waagen* from the Oomia Group, with the exception of
the belemnites, do not comprise representatives of any of the genera
obtained from the Uitenhage beds, and moreover, when brought into
comparison with European forms, seemed to Waagen to indicate a
Tithonian or Portlandian age.

Stoliczkaf first directed attention to the great similarity between
a Trigonia collected by Wynne during the geological survey of
Cutch,^ and the South African T. ventricosa (Krauss), and the identity
of these was afterwards confirmed when the Oomia shell was definitely
referred to T. ventricosa by Waagen, Peistmantel,|] and W. T.
Blanford.1i Feistmantel, in 1876, ::::;: noted the close similarity be-
tween Trigonia herzogi from the Uitenhage beds and a Trigonia from
the Oomia Group which I have recently described under the name
T. mamillata. As already pointed out, T. mamillata is comparable
with T. herzogi and the South American T. transitoria in the manner
in which it exhibits characters somewhat intermediate between those
of the Clavellatae and the Quadratae, but it bears a still closer
resemblance to T. Jiolubi sp. nov. Owing to their large and massive
shells, and their well- characterised sculptural plan, the members of
this group of Trigonia form a prominent feature in the faunas in
which they occur, and supply significant data in the present com-
parative study.

The basis for comparison is further strengthened by the association
of true Costatae with members of the Scabrse, both in the Oomia
and the Uitenhage strata a fact to which I have already referred.
T. parva accompanies T. ventricosa in Cutch, and although T. tennis,
another costate form, has not been recorded from any of the
localities where T. ventricosa is found, it occurs in beds at least not
older than those from which the remaining Oomia Trigonia have
been obtained. In the Uitenhage Series we have T. tatei, which,
although not closely comparable with T. tennis, and distinguished by
much coarser ribbing of the flanks, also shares some of the charac-
teristics by which the Oomia form is contrasted with most of the

* Waagen (1). f Stoliczka (2), p, 315 (1871).

+ Wynne (1), pp. 225, 231. Waagen (1), p. 237.

|| Feistmantel (2), p. 164; (3), p. xxxvii.

1f Medlicott and Blanford (1), p. 261 ; (2), p. 224. See also Kitchin (1), p. 104.

"* Feistmantel (1), p. 116.

The Invertebrate Fauna of the Uitenhage Series. 41

Costatae from the Jurassic rocks of Europe. These distinguishing
features are the elongated oval figure, the delicate sculpture of the
marginal carina and area, and the elongated form and fine ornamen-
tation of the escutcheon ; they lend an aspect of similarity which
cannot be overlooked, especially when these lingering examples of a
typically Jurassic section are brought into comparison with the
majority of European representatives.

Other Trigonice which serve in most conspicuous manner to
characterise the faunas under consideration, and certainly at first
sight appear to furnish the strongest corroborative data in a correla-
tion, are the members of the group of Trigonia v-scripta in the
Oornia beds and the group of T. vau in the Uitenhage Formation.
The similarity of one of the Oomia Trigonice to T. vaic Sharpe seems
to have been first pointed out by Feistmantel, : ' : and W. T. Blanford f
shortly afterwards suggested that a shell from the Oomia Group,
which was ascribed by Feistrnantel to the genus Goniomya, might in
reality represent the African T. van. Which of three Oomia forms
was here and else where \ referred to in this connection is uncertain,
but it is possibly the one which I have described under the name T.
v-scripta, although the less well-known T. dubia in reality bears a
much closer resemblance to T. van, particularly in the outline.
When dealing with the Oomia Trigonice, I pointed out that T.
v-scripta, when adult, more closely approaches an undescribed
Uitenhage form which is associated with T. van. This is T. stowi,
described in these pages, and it is certainly closely allied to T. van ;
the youthful characters of shape and ornament are practically
identical in these two types, but a marked divergence is observed to
accompany progressive growth, and the adult forms are perfectly
distinct. The characters of the youthful period in T. v-scripta are
also practically the same as in T. recurva, with which it is associated
in the Oomia beds, and in this case likewise, wide divergence is
exhibited as the adult characters are required. But the marked
differences which exist between the essential features in the youthful
stage of the two Uitenhage and the two Oomia shells respectively
induced me, when describing the Cutch Trigonice, to distinguish
between the " Group of T. van " and the " Group of T. v-scripta."
Subsequent examination of further material has strengthened the
belief that we are here dealing with a case of convergent development,
and that close alliance is by no means to be inferred from the

* Feistmantel (1), p. 116. f W. T. Blanford (1), p. 118.

I Medlicott and Blanford (1), p. 261.

Kitchin (1), p. 70, pis. vii., figs. 6-8; viii., figs. 1-3.

42 Annals of the South African Museum.

similarity in the adult stage in members of these two groups,
however peculiar and striking may be the characters which appear
to unite them, and at the same time to differentiate them from all
other sections of the genus with which we are acquainted. Since
the members of these two groups appear to illustrate mere homceo-
morphy, their value as evidence in the correlation of the faunas
becomes very much reduced, and if dissociated from the forms which
accompany them, they could not well be considered to afford proof
of contemporaneity. But whatever be the causes that determine the
evolution along converging lines in shells which, by their youthful
characters, betray a heterogenetic origin, we may in this case safely
infer from their occurrence with an association of forms in so many
respects similar, that they acquired their common characters at
approximately the same time.

Before concluding this comparison of the Uitenhage and Oomia
Trigonice, attention may be directed to certain broad features of
general habit which in some measure lend a distinctive aspect to
several members of the genus in the faunas under discussion ; and
it will be noticed that while these features serve in great degree to
imprint a facies on the assemblage which brings it into contrast
with European occurrences, the same broad distinguishing characters
are not confined to one section of the genus, but are shared by
members of stocks not intimately related. There is the tendency to
great posterior elongation of the shells, and in some cases a
siphonal gape ; the obliteration of the carinae with disappearance of
a definite demarcation between flank, area, and escutcheon ; the
dwindling and disappearance of sculpture on the area ; and in
several instances the situation of the urnbones relatively far from
the anterior extremity. In the Oomia beds these points are illus-
trated in varying degree in certain degenerate derivatives of
Costatae, and in the group of T. v-scripta ; in the Uitenhage beds
they are exemplified in T. vau, T. stowi, T. rogersi, and T. conocardii-
formis. In both Trigonia vau and T. dubia the parallelism with the
genus Gomomya cannot be overlooked, and I have previously
suggested that if complete shells of these could be procured, they
would be found to gape at the siphonal end ; this idea is now
supported by a specimen of T. stoivi sent to me from the South
African Museum, which is almost uninjured at the siphonal border,
and which plainly exhibits a gaping habit.

We do not find further aids to comparison amongst the few Oomia
Mollusca which have already been described, and a detailed account
of the remaining lamellibranchs collected by Wynne and Stoliczka

Tlie Invertebrate Fauna of the Uitenhagc Series. 43

from these beds is still in course of preparation ; but I am indebted
to the Director of the Geological Survey of India for permission to
utilise the Indian specimens at present in my keeping for the purpose
of this correlation. Several Oomia types at once suggest most
strongly their affinity to Uitenhage forms, and these are the
following. An Exogyra occurring in Oomia beds at the Trummo
Eiver is certainly not distinguishable from individuals of E. imbricata
Krauss, and may at any rate be thus provisionally named ; an
Astarte found in association with Trigonia ventricosa and other
characteristic Oomia forms very closely resembles Astarte Jierzogi
Krauss, though it is not identical with it ; there are fragments of a
large, coarsely-ribbed Cuciillaa which, so far as can be seen, shares
all the distinguishing features of the strongly characterised Gucullcea
Jcraussi Tate ; a large Gervillia very closely resembles G. dentata
Krauss, and is probably identical with it ; * and lastly, the Oomia
beds have yielded specimens referable to the genus Seebachia, other-
wise only known by Seebackia bronni (Krauss), from the Uitenhage
Series. Two of these individuals from Cutch very closely resemble
S. bronni, and it is not improbable that they are identical with it.

The closely similar character of these lamellibranch-faunas is
clearly shown when we place side by side the identical, proximate,
and analogous types, as follows :

Qitenhage Beds. Oomia Beds.

Exogyra imbricata Exogyra imbricata

Cucullcsa kraussi Cucidlcsa kraussi

Gervillia dentata Gervillia dentata ?

Astarte herzogi Astarte sp., near herzogi

Trigonia ventricosa Trigonia ventricosa

Trigonia holubi }

Trigonia herzogi } (Pseudo-quadrat*) Trigonia mamillata

Trigonia of the vau Trigonia of the v-scripta

group group

Seebachia bronni Seebachia bronni ?

A more critical and detailed study of the Oomia lamellibranchs
may possibly reveal further connecting links, but two inferences may
already safely be drawn from the general agreement observed to
exist between these geographically widely separated faunas. Firstly,
despite the absence of clues to correlation derivable from the Cepha-
lopoda, we may conclude that the faunas were approximately con-

* See also W. T. Blanford (1), p. 118.

44 Annals of the South African Museum.

temporaneous ; and secondly, that the lines of intercourse between
the two areas were probably much more direct than was formerly
believed to be the case.

This correlation appears therefore to be in conflict with Waagen's
conclusions concerning the age of the marine Oomia beds, which
were thought to be Portlandian, but a brief consideration will show
that this difficulty is more apparent than real. Doubts might per-
haps be expressed as to the validity of a comparison based solely
upon lamellibranch types, and it must be admitted that in many
similar cases it would be right to place greater confidence in the
evidence of cephalopods ; but in the present instance the lamelli-
branchs compared are peculiarly well characterised, and indeed com-
prise no single form known in the European area. On the soundness
of the evidence which these well-marked types afford, and upon which
the above conclusions are founded, I think no doubt can reasonably
be cast.

The Mollusca of the marine Oomia beds, shown by Stoliczka to occur
in the lower part of the group, cannot be said to give such definite
indications of geological age as are to be derived from a study of the
Uitenhage fauna. The ammonites from these lower Oomia beds
were considered by Waagen to show close affinity with Upper Jurassic
forms in Europe, and he believed some of the Trigonics to corroborate
fully his view concerning the Portlandian age." But a detailed
study of the TriyonicB has shown that this belief was not well founded,
and the relationship of any of the Oomia forms to the Portlandian
Gibbosae is at the best a matter for conjecture. I have provisionally
referred one of these Trigonics to the group Gibbosae on the strength
of a certain broad similarity of characters, while at the same time
recognising the possibility that this form may represent an aberrant
derivative of some costate stock. Judged by analogy with Trigonia
retrorsa, which can only be regarded as a degenerate costate type, the
relationship of T. spissicostata to the Gibbosae is extremely doubtful,
but nothing more definite on this point can be said until material can
be collected in a sufficiently favourable state of preservation to throw
light on the nature of the youthful stage. Other Oomia Trigonia
which exhibit characters simulating those of the Portlandian Gibbosae
have been shown to be allied to the section Costataa, and nothing
quite comparable with these diversely modified derivatives is known
in Jurassic rocks, though the late adult stage of T. peninsnlaris Coq.,
from the Aptian of Spain, shows an analogous obliteration of sectional
features which was regarded by Lycett to indicate degeneracy. The

* Waagen (1), p. 233 (1875).

The Invertebrate Fauna of the Uitenhage Series. 45

value of Trigonia mamillata as an indication for age has already been
considered, and the supposition of the Cretaceous rather than the
Jurassic affinities of this form receives emphatic support from the
presence of Trigonia ventricosa and T. pulclira, both members of the
section Scabrae. Though claiming corroboration of his views from
Trigonice which he supposed to be related to Portlandian forms,
Waagen did not state that these were found in actual association
with his Oomia ammonites ; and the fact that none of the critical
ammonite-species upon which he relied in his correlation is recorded
from any of the localities such as Goonaree, Oomia, or Huroora,
where the Trigonia-beds are well developed, gives room for the
suggestion that the ammonites and lamellibranchs may not represent
horizons of quite the same geological age. Further, if w y e examine
Waagen's descriptions and figures of the four ammonites which were
thought to represent European forms, it may perhaps be allowed
that too much reliance has been placed on the evidence they were
supposed to afford. Two were referred only with doubt to the
respective European species ; of the remainder, one represents a
type of Perisphinctes which, so far as can be judged from the
description and figure, does not justify the definite conclusions drawn
by Waagen concerning its relation to a European Portlandian form.
The other, a single specimen referred to the Tithonian Perisphinctes
eudichotomus Zittel, is so preserved that the lobe-line is not visible,
and a comparison of Waagen's figure with Zittel's original specimen
in the Palaeontological Collection of the State at Munich shows that
the Indian form is rather thicker and more involute, though the
agreement is otherwise good.

It is scarcely necessary to dwell at any length on the question of
the alleged discrepancy between the evidence of the plant and animal
remains in settling the age of the Oomia beds, since this matter has
already been so fully dealt with. The plants of the Oomia group
were obtained from strata for the most part above the marine beds
which yielded the cephalopods and lamellibranchs, but they led
Feistmantel to refer these beds to the Middle Jurassic. Even should
a revision of the Oomia plants prove the correctness of Feistmantel's
view that the flora exhibits Oolitic affinities, there are many reasons
why the evidence of the marine fauna must be allowed to outweigh
that of the plants in a correlation with European stratigraphical
standards.*

Putting on one side the ammonites, the exact bearings of which
on this question are somewhat doubtful, it may be said that there is

* W. T. Blanford (2).

46 Annals of tlie South African Museum.

nothing in the fauna of the Oornia Trigonia-beds to contradict the
results reached by a study of the Uitenhage Mollusca, and a con-
sideration of all the available evidence seems to justify the conclusion
that here also we are dealing with the part equivalent of the
Neocornian in Europe.

(c) Traces of a Belated Fauna in the Godavari District and in
Hazara. Traces of the Uitenhage-Oomia lamellibranch-fauna, as
principally indicated by the presence of Trigonia ventricosa, have
also been found to occur in an outlier of the Tripetty beds about
24 miles north-east of Coconada near the south-east coast of the
peninsula of India. :;: Trigonia ventricosa is said to be here
accompanied by Trigonia smeei, a characteristic fossil of the
Oomia Group in Cutch, though it seems possible that this may be
T. crassa, an Oomia form of similar type. I The small collection
of fossils from near Coconada was examined and named by Stoliczka,
and it comprises Inoceramus and a few other lamellibranchs as well
as Helicoceras and other remains of cephalopods.

Trigonia ventricosa has also been recorded from strata exposed in
the Margalla Pass in Hazara (N.-W. Himalayas), where it is said to
be found in profusion.;]: The bearings of these occurrences in a dis-
cussion of the broad question of distribution will be considered below.

(d) Neocomian in German East Africa. The Lower Cretaceous
fossils collected during W. Bornhardt's journeys in German East
Africa (1895-1897), and described by G. Miiller, are of special
interest in the present connection, because they exhibit in some
measure a Uitenhage facies and also furnish strong links with the
molluscan fauna of the Oomia Group. The remains of Cephalopoda
are unfortunately very scanty and again fail us as a basis of com-
parison, but important links are found in some of the lamellibranchs,
which include well-characterised Trujonia. Foremost amongst these
is Trigonia ventricosa, occurring at a locality -8 km. north of the
Nkundi stream, 29 km. north-west of Kiswere, in strata ascribed to
the Lower Neocomian and brought into correlation with the Uitenhage
beds by Dr. Miiller. Trigonia bcyschlagi G. Miiller, which here
accompanies T. ventricosa in great abundance, appears to have no
counterpart in the Uitenhage beds, but it clearly belongs to the same
category of modified Costatae as T. smcei\\ and T. crassa*>\ from the

* King (1), p. 229 ; Feistmantel (2), p. 164; Feistrnantel (3), p. xxxvii.

t Kitchin (1), pp. 42, 43.

I Wynne (2), p. 125 ; Medlicott and Blanford (1), p. 503.

G. Muller (1). || J. de C. Sowerby (3), pi. Ixi., fig. 5.

[ Kitchin (1), p. 44 ; pi. iv., figs. 46, pi. v., figs. 1-3.

The Invertebrate Fauna of the Uitenhage Series. 47

Oomia beds in dutch ; the resemblance to T. crassa is indeed very
striking, though I have elsewhere stated reasons why these two
forms cannot be considered identical.* But T. bcyschlagi is certainly
either closely related to T. crassa or illustrates a stage of removal
from the normal ancestral costate plan, on some parallel line, quite
comparable with that exemplified by the Oomia form. It was
probably in consequence of insufficient acquaintance with Trigonia
smeci that Miiller failed to recognise the true relationships of this
peculiar shell ; the points of similarity to the Uitenhage shell See-
bachia bronni (Krauss), to which he called attention, are merely
superficial, involving the general outward habit only. Uhligt has
expressed the opinion, however, that the grounds for the generic
separation of these two forms are hardly convincing ; but he can
scarcely have compared the figure showing the dentition of the right
valve of Trigonia beyschlagil with Neurnayr's excellent correspond-
ing illustration of Seebachia. A glance at these figures proves
beyond question that Miiller was right in assigning his shell to the
genus Trigonia, and now it is interesting to find the clue to its
narrower relationships by a comparison with the Oomia forms, a
detailed account of which had not appeared at the time when Miiller
wrote.

Amongst the fossils collected at a locality in the district of Ntandi,
35 km. west of Mtshinga, which Miiller considered to indicate a Middle
Neocomian horizon, a large Gervillia, apparently identical with
G. dentata Krauss, serves as a connecting link to both the Uitenhage
and Oomia faunas. There is also a Ptychomya (Ptychomya hauchc-
cornei Miill.) which, though distinct from P. complicata (Tate), may
possibly be nearly related to it. Trigonia bornJiardti Muller, from
the same locality, is only so far comparable with the Uitenhage-
Oomia Trigonice, that in common with some of these it exhibits a
marked posterior elongation, with absence of sculpture from the area
and the posterior part of the flank in the adult shell, in which also
the area has ceased to be demarcated from the flank. An Astarte
from the Oomia Trigonia-beds compares very closely with the shells
from Ntandi described by Miiller under the name Eriphyla stuhl-
manni, particularly in the character of the hinge and the very deep
lunule, though it differs somewhat in the outline and ornamentation ;
Astarte herzogi from the Uitenhage beds also belongs to the same
division of the genus.

* Kitchin (1), p. 121. f Uhlig (3).

I G. Muller (1), pi. zix., fig. 3.

Holub and Neumayr (1), pi. ii., fig. 4ft.

48 Annals of the South African Museum.

Still greater interest attaches to a Triyonia which strongly recalls
the peculiar types belonging to the group of T. v-scripta in the
Oomia Triyonia beds and the Uitenhage group of T. vau. This
is T. killini Mull., found in strata ascribed to the Upper Neocomian
at a locality 23 km. west-south-west of Mtshinga, and it possesses
characters which appear to indicate relationship to one of the groups
mentioned rather than to the members of other groups with which
Dr. Miiller has also brought it into comparison. Triyonia kiihni, it
is true, seems to be distinguished from T. vau as well as from T.
v-scripta and its allies both by its less equilateral form and the
persistent ornaments of its escutcheon ; but to judge from the
description, and especially from the figure of an imperfect specimen"
which shows the convexity of the anterior profile and the crowded
ribs of the frontal series obliquely crossing the growth lines just as
in the Oomia T. recurva there can be little doubt about the position
of this shell relative to the main divisions of the genus. Whether it
is more closely allied to the group of T. v-scripta or to T. vau and
T. stowi sp. nov., cannot be ascertained until more perfect specimens
can be obtained and the sculpture of the youthful growth-stage
examined.

Of less significance, perhaps, is the occurrence of Pecten striato-
punctatus Eoem., which was found w r ith Gervillia dentata, and Area
uitenhaycnsis Miiller, which accompanies Triyonia beyschlayi. They
may be brought into near comparison with Pecten pro jectus Tate and
Area jonesi Tate, though little importance could be attached to such
types as these, if taken alone. Another Pecten, from the same
locality as that from which P. striato-punctatus was obtained, was
considered by Miiller to represent very probably P. cottaldinus
d'Orb., which also occurs in the Uitenhage beds. The relationship
of the Oomia fauna to that desci'ibed from German East Africa is of
course what we might expect to find, having regard to the manner
in which a close connection between the Upper Jurassic faunas of
East Africa and Cutch has been demonstrated by those who have
studied the Cephalopoda.

Enough has been said to indicate the position taken by these East
African occurrences in a widely distributed Neocomian fauna of
southern type, and to show that such connecting links as are
available, no less than the general aspect which the fauna, wherever
it is developed, derives from the prevalence of peculiarly charac-
terised Triyonice, plainly serve to unite the molluscan assemblages
of these three remotely separated districts in the eastern hemisphere.

* G. Miiller (1), Taf. xxv., fig. 8.

The Invertebrate Fauna of the Uitenhage Scries. 49

Some indications of a western extension of the same type of fauna
are to be recognised on the South American continent.

(e) Neocomian in South America. Amongst the fossils collected
in 1899 by the late J. B. Hatcher from the Pueyrrydon Series in
Patagonia" are several lamellibranchs of distinctive type which
permit of close comparison with some of those above discussed.
They have been described and figured by Stanton,! who was led
to ascribe a Lower Cretaceous age to the Pueyrrydon Series, though
he was unable to establish a narrower correlation with the forma-
tions of other regions. The most striking resemblances to Uitenhage
forms are seen in two Trigonice, from the Belgrano beds, T. sub-
rentricosa Stanton, a member of the section Scabrae, and T. hetero-
sculpta Stanton, believed by Stanton to be allied to T. van Sharpe.
Trigonia subventricosa certainly approaches very closely to T. vcn-
tricosa, particularly in the wide spacing and robustness of the
anterior varices. The differences in the sculpture pointed out by
Dr. Stanton appear to be less emphasised when this Patagonian
form is brought into comparison with the large and more coarsely
ornamented T. Jcraussi sp. nov., which occurs with T. ventricosa. But
if the relations of height to length exhibited by the figured specimen
of T. subventricosa be those which characterise the average shells
of that form, then there is good ground for the separation of these
three members of the Scabrae, which at the same time appear to be
united by similar broad features of peculiar form and ornament.
They are the only known examples of this particular extreme type,
and probably represent a group in this southern developmen
parallel with that of which T. scabricola Lycett and T. aliformis
Park, are typical members in Europe.

The ornamentation of T. lieterosculpta is of such a kind as to
recall instantly the conspicuous shells of the groups of T. v-scripta
and T. vau. The principal points of resemblance are the manner in
which steeply inclined ribs of two series on the flank meet to form
an angular pattern, and the absence of sculpture from the area and
the posterior portion of the flank in the adult shell. T. lietcroscutyta
is certainly not to be brought into comparison with the European
Undulatae, but its whole aspect suggests a close connection with the
group of T. vau. Its near relationship to T. vau and T. stowi is
clearly to be inferred from a comparison of the youthful characters,
which are very similar, and affinity with these rather than with the
Indian group of T. v-scripta is exactly what we might expect to
find, having regard to the geographical relationships. The adult
* Hatcher (1). f Stanton (3).

50 Annals of the South African Museum.

T. heterosculpta differs, however, from its African allies by the less
posterior elongation of the shell.

Here again, in the Belgrano beds, we find an associated Gervillia
of large dimensions, which does not appear to be widely removed
from G. dentata Krauss, while a large Astarte (A. perlata Stanton)
with short and deeply excavated lunule, may be said to have its
parallel in the Ooniia beds of Cutch, if we confine the comparison
to broad outward features. At a somewhat lower horizon in the
Pueyrrydon Series (the Gio beds) Ostrea tardensis Stanton, which
occurs abundantly, represents a type of Ostrea or Exogyra which
has much in common with Exogyra imbricata Krauss ; and to judge
from the description of 0. tardensis, it seems likely that these two
forms may really be very closely comparable. A special value can-
not, of course, be attached to the comparison of such types as these
if studied alone, but the resemblance gains significance when viewed
in connection with the other Mollusca above discussed. It will be
seen therefore, that although the basis of comparison is somewhat
slender, yet the occurrences above noted in no slight measure
suggest the closest alliance with the niolluscan fauna of the South
African Neocomian ; and it may be added that there is nothing in
the Pueyrrydon fauna itself to contradict the assumption that we
are dealing with a western part-equivalent of the Uitenhage
development.

A search for further traces of the characterising elements of the
Uitenhage fauna in the Lower Cretaceous deposits of South America
reveals evidences of a very suggestive description. The Lower
Cretaceous beds in Bolivia and Chili which have yielded Trigonia
transitoria Steinmann * are probably of similar age and seem to
represent the more northerly equivalents or part-equivalents of the
Pueyrrydon Series. Trigonia transitoria has also been found to
occur abundantly at the Arroyo Triuguico and at Quili Malal in
the Argentine Republic, in strata ascribed by Behrendsen 1 to the
Neocomian. It occurs, moreover, with significant associates in the
Trigonia-beds of Lower Neocomian age exposed on the left bank
of the Rio Agrio, opposite to Las Lajas (Argentine Republic) ; J
these beds yield another member of the Pseudo-quadratse (T.
neuquensis Burckhardt) and a Trigonia very closely resembling the
South African T. conocardiiformis, as well as an Eriphyla in some
respects comparable w r ith the South African Astarte (Eryphyla)

* Steinmann (1), p. 260; Steinmann (3); R. A. Philippi (1), p. 63 ; Paulcke (1),
p. 297. t Behrendsen (1), p. 25.

| Burckhardt (1), p. 21. Burckhardt (2), pp. 72-75, 111, 112.

The Invertebrate Fauna of the Uitenhage Series. 5]

herzogi. Dr. Stanton has suggested that a Trigonia quoted by
Behrendsen* as " Trigonia cf. aliformis Park." from supposed Upper
Neocomian beds at the Arroyo Pequenco may possibly be identical
with the Patagonian T. subventricosa. It is accompanied by an
oyster which Behrendsen identified with Exogyra couloni, and found
to agree well with the figures of Ostrea couloni cited by Bayle and
Coquand t from the Neocomian at Arqueros in Chili. The Chilian
shells ascribed to 0. couloni bear a considerable resemblance to the
Uitenhage Exogyra imbricata Krauss, with which Coquand even later
identified them,J and accompanying this oyster in Chili is Trigonia
delafossei Bayle and Coquand, the resemblance of which to T. ven-
tricosa has been elsewhere remarked upon. Associated with these
forms is a Crioceras identified by Bayle and Coquand with the
European C. duvali Lev., thus recalling the analogous association
of Trigonia ventricosa, Crioceras spinosissimum and Exogyra imbri-
cata in South Africa. Of the above-mentioned Mollusca, Trigonia
delafossei, T. transitoria, T. ncuquensis, and T. cf. conocardiiformis
suggest very strongly their relationship to Uitenhage and Oomia
forms. It should be noted also that Philippi has described Trigonia
from localities in Chili, which share the characters of peculiar
sculpture and siphonal elongation shown by the group of T. van
Shavpe, while T. eximia E. A. Philippi, from the Tinguirica valley
in Chili, appears to be closely related to T. conocardiiformis (Krauss)
and Burckhardt's T. cf. conocardiiformis.

IV. THE DISTKIBUTION OF THE UITENHAGE FAUNA
IN EELATION TO SOME THEOEETICAL QUESTIONS.

We may now briefly consider in what measure the correlation
of the Uitenhage Neocomian fauna with the similar assemblages in
German East Africa and in Cutch bears upon the theory of an Indo-
African land barrier during early Cretaceous times ; and further, we
may inquire how far the facts concerning the dispersal of these
Mollusca have significance in relation to Neumayr's theory of distri-
bution according to climatic zones, as deduced from a study of the
Cephalopoda. The one question is in reality largely bound up with
the other, and since in both cases the inquiry is of a so purely
palasontological character, it may be of advantage briefly to review
the evidence now available. Moreover, it is advisable to glance at

* Behrendsen (1), p. 418. t Bayle and Coquand (1), p. 37.

I Coquand (2), p. 158.

Lycett (3), p. 120 (1875) ; Kitchin (1), p. 108 ; Paulcke (1), p. 296.

52 Annals of the South African Museum.

these aspects of the distribution as now known to us, because
Neumayr himself attached very great importance to the testimony
of the Uitenhage fauna in supporting both the above theories. But
at that time it was not known that the links connecting the bivalve-
faunas of the Uitenhage and Oomia strata were so numerous or
complete as they are now proved to be, while the existence of
a related molluscan assemblage in German East Africa was un-
suspected.

Although Neumayr was led to the theory of a great equatorial
enclosed sea (" Ethiopian Mediterranean ") chiefly by a comparative
study of faunas of essentially Jurassic character, yet he felt justified
in utilising the Uitenhage fauna also as an aid in this palaeogeographic
reconstruction.* He believed that the radical differences between
the Mollusca of these beds and the more northerly East African
occurrences of Upper Jurassic age were an indication that the
faunas had for some time lived in distinct areas separated by a land
barrier, and from this he thought to derive support for a theory the
truth of which had seemed to be attested by evidence of a different
character.! In spite of weighty opposition,]: this theory of a land
connection extending from the Indian peninsula through Madagascar
to South Africa at the beginning of Cretaceous times has continued
to find favour, and it is only comparatively recently that the signifi-
cance of the palasontological evidence has again been called in
question. In view of the presence of some traces of the Uitenhage
fauna in the Godavari district and also in Cutch and Hazara,
Neumayr supposed that a connection between the equatorial and
southern waters must have existed in the form of a strait, the
situation of which he believed to be most probably about the present
Gangetic plain. The late Dr. W. T. Blanford held a similar view ;
he summarised the main arguments in favour of the theory,
suggesting that a shallow water connection near India, situated
very possibly to the eastwards, though not precisely as Neumayr
supposed, would account for the northerly dispersal, and that if this
were later converted into land, the "progressive diminution of
European species in the three stages of the S. Indian Cretaceous
beds would be explained by the increasing effect of isolation."
Professor Suess also speaks of these traces of the Uitenhage fauna
in Cutch and in the Salt Eange as possible indications of an

* Neumayr (3) ; Neumayr (6), pp. 259, 261, 295, 296, 529.

f Stow (1), p. 546; H. F. Blanford (1), pp. 534-540; Medlicott and Blanford
(1), pp. xxxix., Ixviii., Ixxii., 297.

I Wallace (1), pp. 422-427. W. T. Blanford (3), pp. 98, 99.

The Invertebrate Fauna of the Uitenhage Series. 53

encroachment of the southern development into the northern
region." With reference to the same subject, Mr. E. D. Oldham
wrote : " This barrier does not seem to have been absolutely
continuous throughout the Jurassic period, or there may have been a
mode of communication round the north of the Peninsula of India
by which some migration took place, and so the presence of a few
Cutch species, which are also found on the east coast of India and
in South Africa, is accounted for." f

The belief in this supposed land mass extending through
Madagascar and cutting off an equatorial basin from the colder
southern ocean, has been thought to receive the strongest support
from a comparison of some belemnites of Neocomian age found at
localities in the north-west of Madagascar.} Neumayr found that
while these belemnites belong to groups which in their distribu-
tion are essentially associated with the equatorial and South
European development, the single belemnite (B. africanus Tate)
then known from the Uitenhage beds is not closely comparable
with any of these, but belongs to a group distributed in the northern
hemisphere only in the boreal region and in the northern part of the
temperate zone. Prom this he drew the conclusion that these
representatives of contrasted groups of belemnites belong to faunas
of radically different type, which flourished in separate geographical
regions. Much has been made of this piece of evidence, which, so
far as it goes, is admittedly very suggestive. The late Dr. W. T.
Blanford || more recently drew attention to some independent
evidence of another kind which indirectly lends support to the
theory of an Indo-African land barrier in pre-Tertiary times, but this
only bears on the general question of the existence of a barrier,
without any possible reference to its state of completeness during
any part of the Cretaceous period. Dr. F. Kossmat 1i has shown
that so far as the distribution of the faunas of Ariyalur (Senonian) age
was known at the time when he wrote, the evidence was in favour of
a barrier separating the waters of the Mediterranean province (with
its easterly extension) from the South Indian ocean. He concluded
from a careful comparative study of the Cephalopoda that the fauna
represented in the Trichinopoly-Pondicherry districts had inter-
course with the European area only by way of Natal and to the
west.

The distribution of the much older Uitenhage fauna must certainly

* Suess (1), p. 536. t In Medlicott and Blanford (2), p. 211.

I Newton (1), p. 333. Neumayr (5).

|| W. T. Blanford (4). 1[ Kossmat (1), Kossmat (2).

5

54 Annals of the South African Museum.

lead us to a somewhat different conclusion concerning the relation of
the equatorial and the southern ocean at that period. In order
to form an approximately correct picture, we must at least compare
faunas of similar age, for it is probable that migration from one area
into another may take place exceedingly rapidly when measured
by geological standards, and that a fauna may thus become quickly
modified as regards some of its most salient characteristics.
Dr. Kossmat has brought forward evidence to show that in the
migration of Upper Cretaceous cephalopods " the distribution of
the species did not require any geologically measurable time." *
In contrasting the Uitenhage fauna with any occurrences on the
western shore of his equatorial sea, Neumayr was not able to
compare with contemporaneous faunas, except perhaps that of the
Belemnite-beds in the north-west of Madagascar. But in this case
the possible grounds of comparison were of the most slender
description, since in the Neocomian of Madagascar the known fauna
consisted solely of belemnites ; and hence the very hazardous nature
of the generalisations which Neumayr was led to make as a result of
the comparison. It is very probable that the great contrast between
these faunas may owe its strength in no slight measure to differences
of local facies, a factor which does not seem to have been sufficiently
taken into account ; f the Belemnite-beds of Madagascar probably
represent deeper water conditions, in contrast to the littoral character
exhibited by the Uitenhage molluscs. Moreover, it is easy to
conceive that the lines of communication between the two areas
were in reality somewhat indirect, without necessarily accepting
Neumayr's extreme interpretation of the facts. It may be admitted
that the arguments drawn from evidence furnished by distribution at
an earlier and a considerably later age decidedly favour the view
that in Neocomian times a ridge extended from India to South
Africa ; this, however, may have been only partially elevated into
land, and one or more intervening tracts of shallow water would suffice
to allow passage from north to south. Pavlow J and Uhlig have
pointed out the very close affinity between species of Holcostephanus
from the Uitenhage beds and H. schenki (Opp.) from the Spiti
Shales, while Kossmat has also remarked on the close relationship
between Uitenhage fossils and Lower Neocomian forms in the Salt

* Kossmat (3), pp. 78-81.

f Since these sentences were written (in 1903), Lemoine has expressed the same
opinion; see Lemoine (1), p. 391.

I Pavlow and Lamplugh (1), p. 493 (p. 135 of authors' separate copy).
Uhlig (4), p. 132.

The Invertebrate Fauna of the Uitenhage Series 55

Range.* He thinks this cannot readily be explained except by
an oceanic connection separating India and Africa. In addition, the
question of the existence of very strong palasontological links con-
necting the Uitenhage fauna with that of the marine Oomia beds
in Cutch, concerning which some doubts might at one time have
been held, and were, in fact, expressed, may now be considered
to have been definitely settled, and this cannot be lightly set on one
side. Further, the recent discovery in the Uitenhage beds of
belemnites which are related to the Neocomian Hastati of Europe,
a group represented in the north-west of Madagascar, has undoubted
significance, and helps materially to invalidate Neumayr's ingenious
argument.

There is, indeed, no evidence that the means of communication
lay only by way of the east coast of the Indian peninsula and
thence to Cutch, and while the occurrence of Trigonia ventricosa
near Coconada might seem to suggest that the submergence of the
ridge was not so great as to preclude the migration of some littoral
types, the discovery of a Neocomian fauna in German Bast Africa
showing relationship to that of the Uitenhage Series, throws some
fresh light on the question and appears to require a more direct line
of communication. This may most reasonably be supposed to have
existed in some nearer passage, situated between Africa and India,
as above indicated. The seeming absence of the Uitenhage cepha-
lopods or of the genus Seebachia from the Neocomian of German
East Africa may very probably be due to our imperfect knowledge
of the fauna in this less frequented district. The Cephalopoda of
the Uitenhage Series, although perhaps rarer in their occurrence
than the lamellibranchs, are well known because these beds have
been more thoroughly and more frequently searched than those in
German East Africa.

We may here note Dr. G. Mliller's suggestion f to the effect that
Stow's record of Uitenhage lamellibranchs from the Zambesi, J
which was discredited by Holub, may after all prove to have
been authentic. Until this record can be substantiated, however,
the suggestion may perhaps be without special significance as
pointing to the direction in which a fauna of intermediate
character may be sought for. The fact that it is not in the
nearer East African Neocomian, but in the more remote Oomia
strata that the higher percentage of Uitenhage forms is found to
occur, may possibly indicate that the passage between north and

* Kossmat (3), p. 78, footnote. f G. Miiller (1), p. 571.

I Stow (1), p. 505, in editorial footnote. Suess (1), p. 541, note 28.

56 Annals of the South African Museum.

south was not situated in close proximity to the present African
continent. On this point nothing definite can at present be said,
and it must be admitted that recent advances in our knowledge of
the Mesozoic rocks in Madagascar and on the east coast of Africa can
only serve as a warning against premature speculation on such a
subject. As regards Jurassic deposits, we now know that, contrary
to formerly held belief, these are not confined to the north-west
coast region of Madagascar, for strata of Oxfordian age have been
shown to be present in the south-west part of the island, in the
basin of the Eiver Isakondry, east of Tullear.* Professor Douville
considers that in these deposits the contrast in faunistic and litho-
logical facies to the strata of corresponding age in the north-west of
the island may be accounted for by different conditions of sedimen-
tation, and is not to be ascribed to deposition in separate basins. t
As remarked on a previous page, deposits which may perhaps
be equivalent or partly equivalent to the Uitenhage Series,
also occur in the Isakondry basin. Prof. Douville has given
brief notices of the fossils found in these beds by Lieutenant
Boutonnet, and has drawn attention to the occurrence here of a
large Trigonia which, he says, recalls the Trigonice of the Oomia
beds in Cutch and the Uitenhage beds in South Africa. Associated
with this is a shell which closely resembles Exogyra imbricata
Krauss (referred to by Prof. Douville under the generic name
Pycnodonta}.\ The same region in Madagascar has furnished a
Cenomanian fauna which is said to show relations to the corre-
sponding faunas both of Europe and Southern India. Concerning
the conditions at a later period, the discovery of fossiliferous
Senonian deposits at Fanivelona and Marohita on the east coast
of Madagascar is very significant. The fossils found on the Eiver
Sakaleou, 10 km. from the coast and 30 km. north of Mahela,
are stated by Prof. Boule to include forms which show clear rela-
tionship to the fauna of Ariyalur type in Southern India and also to
the Senonian of Baluchistan. || Some of Prof. Boule's remarks on
this subject are as follows : " Hitherto it has been admitted that the
eastern coast of Madagascar is lacking in all sedimentary deposits of
the Secondary era, and this belief has played an important role in
the theories expressed by various scientists ; Oldham, Neumayr,
Suess, Kossmat, etc., on the former distribution of land and sea and
concerning the existence, during the Secondary era, of a continent

* Boule (2), p. 131. t Douville (3), p. 435.

I Douville (2), p. 388; Douvilte (4), p. 215; Lemoine (1), p. 176.

Boule (1), p. 184. || Boule (2), pp. 132, 133.

The Invertebrate Fauna of the Uitenhage Series. 57

uniting Africa with India (Lemuria of the zoologists). This hypo-
thesis appears to be justified in the case of the Triassic epoch, since
there are close relationships, both from the palaeontological and
stratigraphical point of view, between the deposits of India and
of the south of Africa (fauna with dicynodont reptiles ; Glossopteris
flora) ; but the hypothesis can no longer be applied to the Jurassic
epoch, for various reasons which would take too long to enumerate
here. As to the Cretaceous epoch, the discovery, on the east coast,
of the fossils mentioned above, compels us to admit that our great
colony was already an island. The affinities of these fossils with
those of the west as well as with those of the east of India support
the same conclusion." Haug has confirmed the analogies between
the Upper Cretaceous faunas of India and Madagascar, but he con-
siders that the discovery of Senonian deposits on the east side of
Madagascar does not weaken the hypothesis of an Indo-Malagasy
continental mass.f In discussing M. Haug's paper, Prof. Boule
maintained that the theory of a land barrier is now very difficult to
uphold, and thought that recent discoveries in Madagascar greatly
weaken the theory.]: Again, with M. Thevenin, he has written as
follows : "If it has truly existed, the Indo-Malagasy continent
must have been reduced then to a long Indian peninsula or to
a suite of islands situated on positions where one to-day observes
depths of 6,000 metres."

Turning now to the African mainland, it may be remarked that
from the coast, as far south as Delagoa Bay, an Aptian fauna has
been described which, while later than the Uitenhage fauna, and
consequently not to be closely compared with it, contains cephalo-
pods exhibiting the closest relationship to those of the Aptian with
Acanthoceras martini in Southern Europe. || This in itself appears
to be sufficient to throw doubt upon the existence of a barrier near
the African coast at that period. Further, the recent discoveries of
an Upper Cretaceous fauna in Mozambique, showing clear affinities
with the Southern Indian development (Utatur and Ariyalur stages),^
seem quite to disprove the existence of a permanent barrier even at
the approach of that later time which we have hitherto thought to
furnish the most sure evidences of separation. It is apparent that

* See also Boule (3) ; Boule (4), pp. 684, 685; Douville (4), p. 21f> ; Boule (5) ;
Boule and Thevenin (1), p. 59 ; Lemoine (1), p. 232.
f Haug (1), p. 397. { Haug (1), p. 398.

Boule and Thevenin (1), p. 59. See also Woods (4), p. 348.
I! Kilian (2) ; Kilian (3).
J Choffat (1) ; Choffat (2) ; Choffat (3) ; Lemoine (1), p. 396.

58 Annals of the South African Museum.

although the known evidence seems to be on the whole destructive
of Neumayr's view regarding the isolation of the Uitenhage fauna,
yet any attempt to reconstruct precisely the geographical relations
of the region under consideration in early Cretaceous times must
still remain purely within the realm of speculation.

One point at any rate is clear, that the argument based upon the
sharp contrast between the Neocomian fauna of Cape Colony and
comparable occurrences situated on the African continent to the
north of the supposed separating ridge, can no longer be utilised.
We are now able to compare contemporaneous faunas in which, so
far as the lamellibranch element is concerned, the validity of the
comparison is not impaired by differences of facies due to local con-
ditions ; and the supposed contrast is no longer perceivable. The
apparently almost complete restriction of the Uitenhage cephalopods
to the southern district in Africa may still seem to require some
further explanation, but should scarcely occasion more surprise than
the fact that only one species from the Neocomian Belemnite-beds
of north-west Madagascar is included in the East African fauna
described by Miiller, although both inhabited the equatorial waters.
As a further example of the risk of making premature deductions
from the apparent dispersal of cephalopods, it may be noted that
not one of the cephalopod-genera of the Uitenhage area has been
recorded from the Pueyrrydon series in Patagonia, although the
occurrence of these forms there would be exactly in accord with
a distribution regulated by the principles upheld by Neumayr. The
evidence of the other Mollusca seems, indeed, to lend every support
to the view that a continuous shore-line extended between South
Africa and South America, and this idea has gained further
justification from the recent work of C. Burckhardt.

This brings us to inquire whether the Uitenhage ammonitoids are
in reality so peculiarly restricted in their geographical distribution as
at first appears. It is certainly remarkable that no definite traces of
them have been found in the African equatorial regions," while close
allies are known from Western Europe and the Himalayas. Indeed,
it may be said that these cephalopods exhibit in the main a
distinctly " Middle European " facies, and thus at first sight
seem to lend support to Neumayr's distribution theory.! We
have seen that amongst the lamellibranchs, leaving out of account

* A very badly preserved ammonite found at one of the localities visited by
Bornhardt has been referred to the genus Holcostephanus. Whether this indicates
the presence of Uitenhage forms, which appears not improbable, further collecting
alone may be expected to decide. t Neumayr (2).

The Invertebrate Fauna of the Uitenhage Series. 59

many apparently cosmopolitan forms, some of the most characteristic
and highly specialised types follow a very different distribution. It
might perhaps be supposed that members of this class, owing to
their relatively passive habit, were more inured to varied environ-
ment than the cephalopods, while these, on the other hand, possibly
equipped with better facilities for exercising choice of station, were
more delicately adjusted to conditions of temperature, ood-supply,
and other special characters of environment. Nothing, however, is
known regarding the exact mode of life of these cephalopod-types ;
but while the evidence so ably handled by Neumayr seemed for
some time inevitably to urge the acceptance of his fascinating theory,
a body of facts has since been brought to knowledge which must
surely indicate that climatic conditions played at the most an insig-
nificant part in regulating the dispersion of ammonites, and that
distribution of land and sea is in reality to be recognised as the most
potent determining factor.* It is therefore scarcely conceivable that
in the case of the Uitenhage fauna a distribution of land and sea
which permitted the migration of well-characterised lamellibranch-
forms over such a wide area, could have offered any obstacles to a
similar geographical range for some, at least, of the cephalopod types.
It must be remembered, too, that members of the true Holco-
stephanus (Pavlow's Asticria) have been described from the Mazapil
district in Mexico.!

It becomes plainly apparent that with the limited evidence as yet
available, the only reasonable course is to suspend judgment con-
cerning the exact significance of the Uitenhage Cephalopoda in this
question of distribution, and a definite pronouncement will only be
justifiable when our knowledge of the Indian and African Neocomian
faunas is more complete. In view of the facts to which reference
has already been made, it is necessary to abandon the supposition
that these cephalopods were excluded from the equatorial waters ;
but in the meantime there is nothing to show whether the absence
of these forms from the Neocomian of German East Africa and
Cutch is merely apparent, in which case the fact of their occur-
ence there may become established by extended search, or whether
it is real, and to be accounted for by causes of a local character. In
this connection it is well to bear in mind that in Europe certain
Cephalopoda of the Chalk, though living under conditions which
might be expected to have assured exceptional uniformity of environ -

* Kossmat (2), p. 53; Nikitin (2); Burckhardt (2), pp. 115-135; Solger (1),
pp. 220-221 ; G. Boehni (2) ; Tornquist (1), p. 285 ; Ortmann (1) ; Stanton (2)
Burckhardt (3), p. 179. t Burckhardt (3), p. 183-185.

60 Annals of the South African Museum.

ment, were nevertheless peculiarly local in their occurrence, even
within a relatively restricted area. The greater, then, is the neces-
sity for caution in generalising from scanty data regarding the
insufficiently explored littoral Neocomian deposits with which we
are dealing.

V. DESCRIPTIONS, CRITICAL NOTES, AND LISTS OP

FOSSILS.

In the records of occurrence accompanying the following
descriptions, it will be understood that when no collector's name
is mentioned, and no other indications are given, the specimens in
question were obtained by Mr. A. W. Rogers and Mr. E. H. L.
Schwarz in 1900 or by Mr. Rogers in 1905. The numbers quoted
in parentheses refer to numbers printed on labels affixed to the
specimens. All the specimens obtained in 1900 by Messrs. Rogers
and Schwarz, and some also sent to me from the collection in the
South African Museum, are labelled with plain numbers printed on
yellow paper. Those collected in 1905 by Mr. Rogers have blue
labels bearing a number followed by an alphabetical letter. A few
of the specimens from the South African Museum and all those
comprising the collection obtained by Miss M. Wilman at Coega
are without numbers.

With the exception of Holcosteplianus baini, H. modderensis, and
H. wilmancB, all the fossils fully dealt with in these pages are repre-
sented in the collections submitted to me by Mr. Rogers ; but it need
scarcely be remarked that these only include examples of about two-
thirds of the species of invertebrates known to occur in the Uitenhage
Series, and it may be added that the published accounts of several
types figured or described by Sharpe and Tate call for critical
revision at some future time. To deal exhaustively with all the
invertebrate forms hitherto recorded from the Uitenhage beds is
beyond the scope of this memoir ; and since a single compiled list
of the whole fauna, including names taken without criticism from
other works, would be open to obvious objection, it will be found
that lists of species with localities given at the close of this descrip-
tive section, contain names which, except those of the ammonites just
mentioned, refer only to specimens entrusted to me by Mr. Rogers.
Additional notes of occurrences, however, frequently accompany the
following descriptions and comparisons, and are based on specimens
preserved in the collection of the Geological Society of London or in
the British Museum (Natural History). A few forms not dealt with

The Invertebrate Fauna of the Uitenhage Scries. 61

in detail below have already been briefly noticed in the second section
of this paper, in so far as it seemed necessary to draw attention to
their significance in reference to the question of geological age : the
names of these, and the remainder of the invertebrate fossils of the
Uitenhage Series which have hitherto been recorded, are brought
together in the supplementary list and brief accompanying notes
with which the present section of this memoir concludes.

In those few instances in which Uitenhage forms are referred to
European species, I have refrained from burdening the subjoined
accounts with full synonymic lists, but have given a reference to the
original description and, where possible, to a recent work in which
fuller guidance to the literature of the species may be obtained.
Throughout the following pages, however, nomenclatural references
are restricted to those works which contain such information as to
insure, in my belief, the truly synonymic value of the citations.

In the description of the Gasteropoda a conventional orientation
is employed for the sake of clearness, and when use is made of the
terms "above" and "below" in this connection it is assumed that
the specimen be held with the apex directed vertically upwards.
The terms " nepionic " and " neanic," occasionally employed with
reference to the young or immature stages in some of the lamelli-
branchs described below, are so well known and so frequently used
as to call for no explanation here/" 1 '

For an account of the available geological information in connec-
tion with the occurrence of the specimens obtained in 1900 reference
should of course be made to the Report by Messrs. Rogers and
Schwarz,! but this may be supplemented by a brief note contained
in a letter written to me by Mr. Rogers, to the following effect :
" The specimens from Dunbrodie and Blue Cliff are from the lower
beds of the Uitenhage Series, those from the Clay Pit near Rawson
Bridge from the lowest marine beds in the Zwartkop's River section.
Those from the Grass Ridge, Uitenhage, and from the kloofs near
Red House and Picnic Bush belong to the highest beds we found,
and the specimens from the Graaff-Reinet railway section and those
from the main line up-side of Rawson Bridge are from the middle
portion of the marine beds." I arn informed that the collection
obtained at Coega by Miss Wilman was made at a locality on the
farm of that name, though outside the river valley, while the label
"Coega River" refers to a section in the river valley on the same
farm, but probably on a rather lower horizon in the Marine Beds.

* See Hyatt (1) ; Jackson (1), p. 293 ; Buckman and Bather (1) ; Hyatt (2), p. 94.
t Rogers and Schwarz (1) ; also Rogers (1), pp. 281-292.

62 Annals of the South African Museum.

In the case of the specimens from the collection of the South African
Museum labelled " Sunday's Eiver," no more detailed account of
locality is available, but these are all typical Mollusca of the Marine
Beds or Sunday's Eiver Beds. Of all the fossils examined, the only
forms which indicate fresh-water conditions are the Unio from the
Wood Beds of the Bezuidenhouts Eiver, below Blue Cliff station, and
a single specimen of Limncea from the section at Buck Kraal, Sunday's
Eiver. Whatever may be the exact relation of the fossiliferous beds
exposed below Dunbrodie (occurring below a band containing plant
impressions) to the main part of the " Wood Bed " series, the fauna
from Dunbrodie is essentially a marine one : as already stated in the
second section of this paper, there is no reason on palaeontological
grounds to suppose that this small marine assemblage, characterised
by the prevalence of Gasteropoda, is materially older than the fauna
yielded by the Trw/oma-bearing Marine Beds.

The collection obtained by Mr. Eogers from the Marine Beds of
the Coega Eiver and Sunday's Eiver valleys, in 1905, includes some
interesting additions to the fauna. For full information concerning
the occurrence of these and commoner associated forms, the recent
report by Mr. Eogers should be consulted.* This work adds
materially to our knowledge of the Sunday's Eiver Beds, particularly
with regard to the vertical distribution of many of the Mollusca in
the extensive sections examined ; but the results in no way strengthen
the hope that a scheme of zonal subdivision of the Marine Beds may
soon be within reach of attainment. On the contrary, the evidence
obtained by Mr. Eogers, as already mentioned, seems to emphasise
the striking similarity in the aspect of the fauna throughout the
whole thickness of beds exposed.

The few specimens found by Mr. Eogers in a patch of marine
Uitenhage beds at Knysna Estuary f only represent the product of
a hasty search, and it is believed that careful collecting at this new
locality may yield good results.

CLASS ANTHOZOA.

GENUS THAMNASTR^EA J Lesauvage.

THAMNASTE^A sp.

There are several specimens of a Thamnastrc&a from the locality
between milestones 24^ and 24f on the railway between Uitenhage

* Rogers (2), pp. 22-33. f Schwarz (2), pp. 50, 74.

J For remarks on the correct application of this generic name see Gregory (2),
p. 131.

The Invertebrate Fauna of the Uitenhage Series. 63

and Graaff-Eeinet (345-349). These occur in a hard limestone
matrix, and the coral itself is replaced by crystalline carbonate of
lime and is in a condition unfavourable for detailed study. The
corallum is relatively slender, subcylindrical and branching in form.
The calicinal centres are situated at about 2 mm. from one
another.

Prof. J. W. Gregory has seen a specimen of this form, and con-
siders that it may be correctly assigned to the genus Thamnastr&a.
In view of the scanty material so far obtained, and its imperfect con-
dition, a fuller description and comparison with described species
may be deferred for the present.

CLASS ANNELIDA.

GENUS SEEPULA Linnams.

SERPULA cf. CONCAVA (J. Sowerby).

Plate II., figs. 1, la.

Cf. 1814. Vermicularia concava J. Sowerby, The Mineral Conchology

of Great Britain, vol. i., p. 125, Tab. 57, figs. 1-5.
,, 1836. Vermetus concavus J. de C. Sowerby, Trans. Geol. Soc.

Lond., ser. 2, vol. iv., p. 343, pi. xviii., fig. 10.
,, 1884. Vermicularia concava E. Damon, Geology of Weymouth,

Portland and Coast of Dorsetshire, 2nd edition, p. 128,

fig. 48.

Description. The tube is wound in the form of a very flat spire,
widely umbilicated, with the apical part not projecting above the
outside whorl when viewed in lateral profile. The whorls are in
close contact, or slightly embracing. The tube is of slightly flat-
tened cylindrical or oval section, the flattening at right angles to
the axis of the spire. After the third whorl the tube ceases to be
coiled and becomes free. The surface of the tube is marked by
numerous irregular transverse wrinkles and furrows.

Dimensions.

Greatest diameter across the whorls 9 mm.

Greatest external diameter of the tube at the close

of the coiled stage 3 ,,

Occurrence. A single specimen was found between milestones

64 Annals of the South African Museum.

24| 24f on the railway between Uitenhage and Graaff-Beinet (332),
in the railway cutting.

Remarks. The specimen described in all probability has not
attained its full growth, and the manner in which the tube for
about the last 2 mm. of its length has grown free from the coil
suggests that we have only the commencement of an uncoiled stage
illustrated in this individual. The subsequent growth of the tube
was probably in a more or less nearly straight line as in Serpula
concava, and it is impossible to say what dimensions the detached
portion of the tube might attain.

In general aspect this Serpula very closely resembles S. concava
(Sow.) (Upper Greensand), and agrees in the dimensions of the
coiled stage, but without a sufficient number of specimens for critical
comparison nothing further can be said regarding its actual affinities.
A similar form from the Neocomian of France was figured by Ley-
merie as Serpula lituola.* In the general habit and mode of growth
there is also resemblance to S. datnesi Noetling, f from the Ceno-
manian boulders of North Germany, but the African form differs in
the smaller number of whorls, the smaller dimensions, and the
thinner walls of the tube.

SERPULA PINCHINIANA Tate.

1867. Serpula (Vermilia) pinchiniana R. Tate, Quart. Journ. Geol.
Soc., vol. xxiii., p. 163, pi. ix., figs. IQa-c.

The large cylindrical tubes of this form are sometimes seen
attached to massive lamellibranch-shells such as Exogyra imbricata
Krauss, or one of the pseudo-quadrate Trigonia. The adherent
stage is irregular in the direction of its growth : it sometimes follows
a sinuous course, and in some cases the tube is bent sharply
back upon its path and grows over itself. The generally smooth
surface of the tube is occasionally marked by irregular annular
thickenings.

Mr. Rogers collected specimens from the Cliff on Buck Kraal,
Sunday's River, attached to Trigonia herzogi (122h), and at the left
side of the Coega Valley, half a mile down from the railway (458g,
on Trigonia holubi}. Another example is attached to a valve of
Exogyra imbricata from the Sunday's River (303), belonging to the
collection of the South African Museum.

* Leymerie (3), pi. 6, fig. 5.

t Noetling (1), p. 10 [206], Taf. i., figs. 8-10.

The Invertebrate Fauna of the Uitcnhage Scries. 65

CLASS LAMELLIBRANCHIATA.

GENUS PECTEN 0. F. Miiller.*
SUB-GENUS SYNCYCLONEMA F. B. Meek.

PECTEN (SYNCYCLONEMA) ORBICULARIS J. Sowerby.
Plate II., figs. 2, 3.

1817. Pecten orbicularis J. Sowerby, Mineral Conchology of Great

Britain, vol. ii., p. 193, tab. 186.
1902. Pecten (Syncyclonema) orbicularis H. Woods, Mon. Cret.

Larnell. Engl., vol. i., part iv., p. 145, pi. xxvii., and

text-fig. 1. (Palaeontographical Society.)

Two specimens sent to me from the South African Museum appear
to differ in no respect from typical examples of this widely distributed
form. One of the specimens, a single valve, has a delicate concentric
banding extending over the whole surface a character by which it may
be recognised as a right valve. With the exception of its imperfect
preservation in the umbonal region and its somewhat smaller dimen-
sions, this specimen agrees very perfectly with the right valve
figured by Mr. Woods in pi. xxvii., fig. 11 of his monograph (see
above). The other specimen appears smooth to the naked eye, but
shows minute and delicate, faint concentric markings under the lens.
This represents a left valve, and it has the anterior ear preserved.
Length 26'5 mm.; height 28 mm.

A small specimen (a left valve) referable to the same species was
found by Mr. Eogers in the Coega Eiver Valley. Length 13 mm. ;
height 15 mm.

Locality. Sunday's Eiver (279, 280). Coega Valley, on the left
side of the valley, half a mile down from the railway (453g).

SUB-GENUS CAMPTONECTES F. B. Meek.

PECTEN (CAMPTONECTES) COTTALDINUS d'Orbigny.

Plate II., fig. 4.

1847. Pecten cottaldinus A. d'Orbigny, Paleont. Fran9., Terr. Cret.
vol. iii., p. 590, pi. 431, figs. 7-11.

* For a full account of the generic and sectional divisions of Pecten see Verrill
(1) ; for further references to recent literature on this subject consult Woods (3),
vol. i., p. 145 (1902).

66 Annals of the South African Museum.

1902. Pecten (Camptonectes) cottaldinus H. Woods, Mon. Cret.
Lamell. Engl., vol. i., part iv., p. 156, pi. xxix., figs. 1-3.
(Palaeontographical Society.)

A specimen collected by Messrs. Eogers and Schwarz agrees so
closely with Pecten cottaldinus that it can only be considered identical.
It is a right valve in which, unfortunately, the anterior ear is the
only portion which retains the shell wholly preserved ; the remainder
of the valve is for the most part in the form of a cast. Here and
there, where some of the inner layers of the shell are adhering to
the cast, a faint and broad concentric waving is seen ; this, however,
does not seem to be markedly impressed upon the cast and appears
to die out towards the posterior margin. It probably stands in
relation to shell structure and not to surface sculpture, for the
faint folds are too broad to correspond with the concentric surface
markings which are exhibited by European specimens. The specimen
agrees with P. cottaldinus in all characters of proportion and outline,
as well as in the shape and relative size of the ears and the deep
byssal notch.

Dimensions. Length 38 mm. ; height 42 mm.

Occurrence. Dunbrodie, in the cliff below the old school-house on
the right bank of Sunday's Eiver (305, 306).

Remarks. This form is well distributed in the Neocomian (princi-
pally the Hauterivian) and Aptian of Europe, and it has also been
recorded by G. Muller from the Neocomian of German East Africa.

PECTEN (CAMPTONECTES) PROJECTUS Tate.
Plate II., figs. 5, 5a.

1867. Pecten projectus E. Tate, Quart. Journ. Geol. Soc., vol. xxiii.,
p. 155, pi. ix., fig. 6.

Siqiplementary Descriptive Note. The ears are very unequal and
the byssal notch deep. The delicate ornamentation of the valve
surface presents a very variable aspect in different states of preser-
vation, and under different conditions of illumination and enlarge-
ment. In frequent instances it appears to consist of fine, radiating,
impressed linear striae, diverging from the middle line and increasing
by bifurcation when traced towards the valve margins. When the
surface is not very perfectly preserved, these lines may appear to be
separated by plain flattened interspaces, while the lines themselves
may exhibit an imperfect punctate structure. Where the surface is
perfectly preserved, however, the ornaments are observed to be of

The Invertebrate Fauna of the Uitenliage Series. 67

a much more complex nature. If, in a suitably preserved adult
specimen, the surface half-way down the valve and below this be
examined under slight magnification and with favourable illumina-
tion, it is seen that the raised interspaces or minute flattened ribs
themselves exhibit a peculiar punctate structure on that side of the
rib nearest to the lateral margin of the valve. This is developed in
such a way that the outer margin of each little rib is cut into by a
series of minute wedge-shaped indentations, which leave more
prominent projecting portions of the rib standing out somewhat
after the manner of the hydrothecaa on a graptolitic stipe. In some
cases the successive wedge-shaped grooves may almost cross the rib,
but for the most part the margin of the rib which is nearest to the
middle line of the valve is straight and entire.

Another peculiarity of the ornamentation is seen in some cases in
the manner in which, when the radial striae are traced upwards from
the valve-margin towards the place of convergence at the middle
line, by favourable illumination they may be observed to be continued
upwards across the middle line, giving rise to a slight intercrossing
here.

In the left valve the anterior ear is large and rectangular, the
posterior ear small. This valve has a rather more equilateral
aspect than the right valve, which is sometimes very markedly
inequilateral.

Dimensions. (1) (2) (3)

Height 13 . 19 21 mm.

Length 12 . 17 20

No. (3) is Tate's figured type specimen.

Occurrence. Eailway cutting between milestones 24^-24f on the
Uitenhage to Graaff-Reinet railway (297, 350) ; also obtained by Miss
M. Wilman at Coega Eiver. Tate says : " Collected by Dr. Eubidge
in a soft, yellowish-grey, sandy limestone at the Zwartkop River
Heights, and at Prince Alfred's Rest ; in both localities it is
frequent."

Eemarks. The notched sculpture of the ribs, described above,
may be well observed when the preservation is exceptionally good.
It appears only in faint traces in a portion of the surface of one
of the specimens examined by Tate, and in other individuals less
well preserved the markings only consist of radiating linear striae,
obscurely punctate in places, with smooth interspaces. A right
valve from the Coega River, which agrees closely with Tate's figured
type, has the surface so well preserved that the intimate details of
sculpture exhibited call for the above additional descriptive note.

68 Annals of the South African Museum.

The delicate notched structure is also well shown in specimens from
the above-mentioned locality near Uitenhage.

Tate brought this shell into relationship with Pecten lens Sowerby,
a Jurassic form ; and although it shares with this and with the
Portlandian P. morini de Loriol :;: some of the principal features which
characterise these familiar Jurassic types, yet it may equally well be
brought into comparison with Cretaceous forms. The Jurassic
shells mentioned, besides differing in the outline, have simply striae
with a punctate structure, but no encroachment of this sculpture into
the interspaces between the linear striae. Pecten suprajurensis
BuvignierJ an Upper Jurassic form, approaches much more closely
to P. projcctus in general character and outline, possessing a similar
bulging frontal profile, but is likewise merely ornamented by delicately
punctate linear striae with smooth interspaces, attaining, moreover,
much larger dimensions. Pecten buclii Roem.J also Upper Jurassic,
is well distinguished by its almost equilateral outline and by the minute
transverse striations which cross the interspaces between the radial
linear strias.

Pecten striato-punctatus Roemer, from the Neocomian and Aptian
of Europe, is more equilateral in outline and is more delicately
ornamented ; the punctate structure is much more minute and
confined wholly to the striae, leaving the very narrow intervening
spaces smooth. ||

Pecten curvatus Geinitz,*' though having greater relative height
than P. projectus, sometimes seems to approach the somewhat in-
equilateral form of this, though in figures of other shells which have
been united with P. curvatus** the valves are almost equilateral.
Although Geinitz figured the magnified ornamentation of the valve-
surface so as to produce an aspect of sculpture widely different from
that given for instance by Eeuss, yet it appears from the descriptions
that punctate linear striae with smooth interspaces characterise these
forms.

Nilsson's P. virgatus, of Upper Chalk age from Sweden, ft is

* de Loriol and Pellat (1), p. 107, pi. x., fig. 6.

I Buvignier (1), p. 24, pi. xix., figs. 21-23.

} F. A. Roemer (1), pi. xiii., fig. 8 (1836), and Nachtrag, p. 27 ; de Loriol, Royer
and Tombeck (1), p. 389, pi. xxii., figs. 12, 13.

F. A. Roemer (1), Nachtrag, p. 27; d'Orbigny (3;, p. 592, pi. 432, figs. 4-7
(1847); Woods (3), vol. L, p. 157, pi. xxix., figs. 4-6 (1902).

|| See also figures given by de Loriol under the name P. arzierensis; Loriol (3),
pi. iv., figs. 3-5. [ Geinitz (1), p. 16, Taf. iii., fig. 13.

" Geinitz (2), Theil i., p. 193, Taf. 43, fig. 15 (1872); Theil ii., Taf. 10, fig. 1,
(1872) ; Reuss (1), part ii., p. 28, Taf. 39, fig. G (as P. diraricatiis).

ft Nilsson (1), p. 22, Tab. ix., fig. 15.

The Invertebrate Fauna of the Uitenliage Scries. 69

more equilateral and more coarsely ornamented than P. projectus,
and it has been shown, moreover, by Hennig * that Nilsson's
unsatisfactory figure is misleading; this shell really has relatively
strong ribs with linear interspaces, increasing by division and
insertion and not diverging from the middle line in the manner
represented by Nilsson. The delicate transverse striae which cross
the radial ornaments are scarcely comparable with the notching
of the "ribs" shown by P. projectus. The shell from the Gosau
beds described by Zittel under the name P. virgatus^ is almost
equilateral, and is much higher in outline than P. projectus.

The shell from the Cretaceous of the east end of Maud Island
(Queen Charlotte Islands), ascribed to Camptonectes curvatus Geinitz
by "Whiteaves,! is somewhat similar to P. projectus, to judge from the
figure of a right valve, but it is more equilateral in form.

A Pecten from the Neocornian in German East Africa was
described by G. Miiller under the name P. striato-punctatus, but
the specimen figured is so preserved that a comparison with P.
projectus is difficult, although it appears to differ from this in being
more equilateral.

Pecten (Camptonectes} pueyrrydoiiensis Stanton,[| from the Bel-
grano beds (Lower Cretaceous) of Patagonia is another shell of
similar type, but differs at least in the more equilateral form of the
right valve.

Pecten kameruncnsis von Koenen,* from Lower Cretaceous strata
in the Cameroons, differs from P. projectus by the more equilateral
form and the rather coarser ornamentation, with stronger lateral rib-
development. Von Koenen describes how the furrows appear here
and there divided by swellings into little grooves, but says that this
stands in connection with the crossing of lines of growth a relation
not to be observed in the minute ornamentation of P. projectus.

SUB-GENUS CHLAMYS J. F. Bolten.

PECTEN (CHLAMYS) cf. SUBACUTUS Lamarck.

Plate II., figs. 6, 6a.

Cf. 1819. Pecten subacutus J. B. de Lamarck, Hist. Nat. des Anim.
sans Vert., vol. vi., p. 181.

* Hennig (1), p. 41, pi. 2, figs. 28, 33. f Zittel (2), p. 109, pi. xvii., fig. 8.

I Whiteaves (3), p. 242, pi. 32, fig. 4. G. Miiller (1), p. 550, Taf. 24, fig. 7.

1 1 Stanton (3), p. 12, pi. iv., fig. 1.

II von Koenen (1), p. 20, Taf. iii., figs. 14, 15.

70 Annals of the South African Museum.

Cf. 1847. Pecten subacutus A. d'Orbigny, Pal^ont. Frang.,Terr. Cret.,

vol. iii., p. 605, pi. 435, figs. 5-10.

,, 1872. Pecten subacutus H. B. Geinitz, Das Elbthalgeb. in
Sachsen, part i., p. 195, pi. xliv., fig. 5 (Palaeonto-
graphica, vol. xx.).

,, 1902. Pecten (Chlamys) subacutus H. Woods, Mon. Cret. Lamell.
Engl., vol. i., part iv., p. 169, pi. xxxi., figs. 7-9.
(Palseontographical Society.)

Descriptive Note. A single specimen sent from the collection of
the South African Museum seems to approach very closely to P. sub-
acutiis. The shell is ovate in outline, prolonged dorsally, with an
acute apical angle (about 80). The inflation is slight, with the left
valve a little more convex than the right.

The valves are ornamented by about 22 strong, angular ribs with
sharp summits. Near the inferior margin the summits of the ribs
are separated from one another by a distance of about 2 mm. On
parts of the shell the ribs are studded with numerous blunt, spinose
projections. Where the surface is well preserved the ribs and inter-
spaces are seen to be ornamented by very delicate, transverse linear
markings, most clearly visible in the interspaces. Near the inferior
margin these become obscurely developed and are masked by
coarser ridges and grooves of accretion. The hinge-line and ears
are unfortunately not preserved, but having regard to the other
characters there is no reason to suppose that they differed essentially
from those of P. subacutus.

Dimensions. Height (about) 42 mm. ; length 35 mm.

Occurrence. Marine Beds of Sunday's Eiver (304).

Bemarks. Pecten subacutus is characteristic of the Cenomanian
in Europe, but this African shell approaches more closely to it
than to any form from the Lower Cretaceous with which I am
acquainted. In the imperfect material at disposal, no features
are apparent which would justify a definite separation from the
European form, yet in the absence of the ears in this specimen
and the lack of further comparative material, the question of identity
must remain open.

Pecten urgoncnsis de Loriol,* common in the Urgonian of Grand
Saleve, is a shell of very similar type, so far as can be judged from
de Loriol's description and figures ; but apparently its scaly orna-
ments are larger and more prominent, and the surface is perhaps
more coarsely ribbed.

* de Loriol (2), p. 389, pi. C, figs. 25, 26.

The Invertebrate Fauna of the Uitenliagc Series. 71

GENUS LIMA J. G. Bruguiere.

SUB-GENUS ACESTA H. and A. Adams.

LIMA (ACESTA) OBLIQUISSIMA Tate.

Plate II., fig. 7.

1867. Lima obliquissima E. Tate, Quart. Journ. Geol. Soc., vol. xxiii.,
p. 156, pi. ix., fig. 5.

Supplementary Descriptive Note. The shell has great height in
comparison with length. The posterior outline is gently rounded,
the anterior profile straight. The straight anterior border passes
at its lower end by a somewhat sudden curve into the rounded
inferior border. The valves are most convex in the umbonal half and
near the anterior side ; they become flatter posteriorly and inferiorly.
Anteriorly, the flank passes abruptly, though without marked cari-
nation, into an extensive, flat frontal face, truncating the shell. This
frontal area is in no degree sunk or concave in form. The umbones
are acute and sharp, terminal at the anterior extremity of the
hinge-line; the superior border of the shell is straight and very
short, and truncates the shell to form a right angle with the
anterior border. The anterior ear is rudimentary or not developed ;
the posterior ear is much reduced, and not definitely demarcated
from the flank.

The ornamentation consists of delicate linear radial grooves,
very crowded near the umbo, gradually diverging until, at a distance
of 40 rnm. from the umbo, they may be separated by interspaces
2 mm. broad. At a distance exceeding 30 mm. from the umbo, these
lines are almost or quite absent from the middle part of the flank,
though well impressed on the lateral parts. The lines follow a
more or less irregular and wavy course and are sometimes suddenly
deflected to right or left when traced down from a prominent
concentric growth-line. When well preserved, the linear orna-
ments show a minutely punctate structure, most perfectly developed
in the umbonal half of an adult shell. The interspaces are very
gently convex or are flat, but are without sculpture. On the frontal
area the ornaments are of a more pronounced character, and may
take the form of imbricating ridges with the edges directed away
from the valve-margin. They radiate from the umbonal region
and pass along the frontal face, gradually approaching the frontal
valve-margin and forming an acute angle with this. On the frontal
area, therefore, the ornaments are very obliquely crossed by the
lines of accretion.

72 Annals of the South African Museum.

Dimensions.

Greatest measurement radially from the umbo ... 64 mm.
Greatest length measurement, at right angles to

the last 40 ,,

Greatest depth of a single valve 10

Occurrence. Eailway cutting between milestones 24-J-24f on the
line from Uitenhage to Graaff-Reinet, about three miles from
Uitenhage (346) ; also stated to occur at Grass Ridge. Specimens
in the collection of the Geological Society are labelled " Sunday
River " (Rubidge) and "Zwartkop River" (H. Longlands). Tate's
record of locality is " in a yellow shell-rock from the Zwartkop
River sandstone, with Placunopsis undulata and fossil wood
(Rubidge)." Fine specimens sent to me from the South African
Museum are from the Sunday's River. Mr. Rogers obtained this
form on the left side of the Coega Valley, half a mile down from the
railway (455g).

Remarks.- Tate unhappily brought this shell into comparison with
two British Jurassic forms, L. rigidula (Phill.) from the Cornbrash,
and L. ovalis (J. Sow.) from the Great Oolite. Lima rigidula* is of
quite another type ; it has a widely different outline, is comparatively
coarsely ornamented, and has well-developed anterior and posterior
ears. L. ovalis, \ more oblique and anteriorly produced, with
minutely delicate and crowded linear ornaments, can scarcely be
brought into close comparison.

Lima obliquissima in reality may be most closely compared with
those shells to which has been applied the sub-generic name Accsta,l
represented in Cretaceous and later strata, and existing at the
present day. The characters whereby this group of forms is distin-
guished from Plagiostoma and other divisions of the genus have
been clearly set forth by E. Philippi in his analysis of Lima.
Although I have as yet been unable to ascertain the position and
form of the ligament pit in L. obliquissima, the close agreement in
other features can leave little or no doubt that this shell must be
united with Acesta, if this sub-generic group be adopted at all.
These features are principally seen in the great relative height of the
valve ; the anteriorly little-produced outline ; the anterior and ter-
minal position of the umbones ; the great reduction of the anterior
ear and the imperfectly demarcated posterior ear. The fine linear

* Phillips (1), p. 116, pi. vii., fig. 13.'

t J. Sowerby (1), tab. 114, fig. 3 (1815) ; Morris and Lycett (1), Part 2, p. 29,
pi. iii., fig. 5.

I H. and A. Adams (1), p. 558. E. Philippi (1), p. 630.

The Invertebrate Fauna of the Uitenhage Series. 73

sculpture of the flank, it is true, recalls that of some Plagiostoma,
but the shell is in other respects strongly contrasted with that
group. Concerning the value of the sub-generic name Acesta, par-
ticularly in the application to fossil forms, opinions have widely
differed. It was accepted by Stoliczka* and by Zittel.f and latterly
by Philippi, while regarded by Fischer as only representing a
section of Lima.\ This last author merely quotes the living type
species, L. cxcavata. G. Boehm, however, has expressed the
opinion that the sub-genus Acesta cannot be upheld, pointing
out that the original diagnosis is inaccurate ; that the name is
unpractical in its application to fossil forms, in many of which
the direction and position of the ligament pit, upon which the
exact determination depends, cannot be investigated ; and further,
that the distinctive features of the proposed sub-genus are of slight
importance. The first of these objections can have little weight,
merely depending, as pointed out by Philippi, upon an original
error of orientation. There is something to be said for the last
point, perhaps, and it is clear that amongst fossil representatives
of the genus there are some forms which illustrate in varying
degrees the inequilateral character and the development of the
anterior auricle. The name, however, though possibly not standing
for a natural group, may for the present be conveniently em-
ployed to indicate the apparent affinities of the forms to which it
is applied.

L. obliquissima shows a striking outward similarity to the typical
living Lima (Acesta) excavata (Fabr.) Chemn.,|j though differing
from this by the relatively more elongated outline, the shorter hinge-
line, and the more restricted posterior cardinal expansion of the flank.
A Neocomian shell having great resemblance in form, to L. obliquis-
sima is Lima undata (Desh.) Leym.,*' but it is distinguished by its
longer hinge-line and stronger ornaments. In the shortness of the
superior margin and the character of the surface ornamentation,
Lima orbignyana Math.* :: shows closer agreement, but it is dis-
tinguished from L. obliquissima by the more perfectly demarcated
posterior ear, the concave outline behind this, the concave anterior
area and the extension of linear ornaments over the whole flank.
Matheron's figure shows a distinctly developed projecting anterior

* Stoliczka (2), pp. 413-415. f Zittel (5), p. 26 (1881).

{ Fischer (1), p. 941 (1886). G. Boehm (1), p. 625.

|| E. Philippi (1), Taf. xxiv., fig. 5.

IT Leymerie (2), p. 10, pi. 8, fig. 8; d'Orbigny (3), p. 528, pi. 414, figs. 9-12
(1847). ** Matheron (1), p. 182, pi. 29, figs. 3, 4

74 Annals of the South African Museum.

auricle, though this is represented as much more reduced in the
figure given by d'Orbigny.

A Lima from the Lower Cretaceous of England, ascribed by W.
Keeping * to Eoemer's L. longa, shows great similarity in general
form and outline. It differs, however, from the African shell by the
relatively longer hinge-line and more extensive posterior ear, and
although appearing to vary considerably in regard to the surface
ornamentation, is characterised by a more pronounced type of radial
sculpture ; L. longa is further distinguished by the possession of
great height measurement in proportion to its length.!

There is closer agreement, again, between L. obliquissima and
L. tenuitesta Whitfield, J from Upper Cretaceous rocks in Syria.
The two are practically identical in outline and the relative length
of the superior border, but L. tenuitesta has the linear striae of the
surface more numerous and closely crowded at a given distance
from the umbonal apex.

SUB-GENUS MANTELLUM J. P. Bolten.

LIMA (MANTELLUM) NEGLECTA Tate.

Plate II., figs. 8, 8a.

1867. Lima neglecta E. Tate, Quart. Journ. Geol. Soc., vol. xxiii.,
p. 156, pi. vii., figs. 4a, 4&.

Three specimens, all left valves, are referable to this species. The
ribs are acutely ridged, with sharp summits. The middle of the
angular interspace is sometimes occupied by a fine linear ridge, and
occasionally a second one may be present. These are stronger than
the numerous, delicate, linear ridges which are developed on the
sloping sides of the main ribs.

A specimen 20 mm. in length has a greatest diameter, measured
at right angles to the length, of about 12 mm.

Occurrence. On the left side of the Coega Valley, half a mile down
from the railway (448g, 449g, 453g). The specimen described by
Tate, numbered 11,013 in the collection of the Geological Society of
London, came from the Sunday's River.

Remarks. Tate's figure of this species is very inadequate, and
does not give a good idea of the shell. The appearance of two sets

* Keeping (1), p. 112, pi. v., fig. 6.

t F. A. Eoemer (1), p. 79, Taf. xiii., fig. 11 (1836) ; (2), p. 57 (1841). Since the
above lines were written, Mr. H. Woods has published an account of this shell and
considers that the Upware specimens were rightly identified with the German form :
Woods (3), vol. ii., p. 25 (1904).

I Whitfield (1), p. 390, pl.iv.A, figs. 1, 2.

The Invertebrate Fauna of the Uitenhage Series. 75

of ribs directed away from a dividing line, so clear a feature in the
figure, is quite erroneous. In the specimen itself, an appearance of
dichotomy occurs only in one rib, and this seems to be due to an
accident of preservation. There is a crack in the shell, and a
portion of the valve on one side has become pushed slightly over
the surface on the other side of the crack. From an examination of
the specimen it is difficult to say with certainty whether this injury
took place during the life of the animal, or subsequently, but the
resulting irregularity in the sculpture was in any case not a normal
character.

There is a close general resemblance between L. (Mantellum)
neglecta and L. parallela J. de C. Sow. ; :;c but in Lima ncglecta
the oblique elongation is greater and the posterior ribs are more
crowded and delicate. Lima (Mantellum) gaultina Woods t is more
closely similar to L. neglecta in outline, but its antero-ventral border
is more narrowed, and the fine linear rib in the interspace between
the main ribs is lacking.

GENUS PEENA J. G. Bruguiere.
PERNA ATHERSTONI Sharpe.
Plate II., fig. 9.

1856. Perna atherstoni D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,
vol. vii., p. 193, pi. xxii., figs. 4, 4a.

Occurrence. This shell was found at Dunbrodie (Sunday's Eiver),
in the cliff below the old school-house (282, 305, 320), and at
Walton's Farm, below Dunbrodie (311). It was recorded by Sharpe
from the same place (Geelhoutboom) " in greenish grit with
fragmentary shells."

Remarks. These specimens from Dunbrodie were thought to
represent very probably a new form, but a comparison with Sharpe's
original specimens of P. atherstoni shows that they differ from these
in no essential particular. One of the Dunbrodie individuals, it is
true, shows a slight difference in outline ; its antero-inferior border,
at a distance of 15 mm. from the umbonal apex, has a broad in-
dentation or inward curve in profile, somewhat more strongly
developed than in the original of Sharpe's figure 4. Other specimens
which I have examined exhibit this feature in less degree, and it
appears to be a character which is subject to some individual

* For full description and references to literature see Woods (3), vol. ii., part 1,
p. 28, pi. v., figs. 14, 15 (1904).

f AVoods (3), vol. ii., part 1, p. 31, pi. v., figs. 16-20 (1904).

76 Annals of the South African Museum.

variation. The convex fold of the valve, passing back from the
unibo, is marked off from a small, flattened, marginal portion of
the valve between the fold and the antero-inferior margin near
the umbonal end, in clearer manner as a rule than is depicted in
Sharpe's figure. In this respect the figure is misleading, for the
fold is well developed in the original specimen. In another point,
also, the illustration leaves something to be desired; the postero-
inferior border, restored by a dotted line, should in reality have been
represented as a rounded curve similar to that followed by the outline
of the same part in Perna mytiloides Lam., and should not have
been drawn in the angular or sub-angular form which lends such
a false aspect to the outline of the shell as depicted in Sharpe's
work.

P. atherstoni is characterised by its oblique figure and the rela-
tively great length of the hinge-line. In these features it recalls
P. mytiloides Lam., from the Upper Oolites of England," which in
general aspect it much resembles ; but in P. atherstoni the shell is
less sharply pointed and produced at the urnbo, is less inflated
anteriorly, and is more obliquely elongated. There are further
points of difference which it is unnecessary to recount.

GENUS PINNA Linnaeus.
PINNA ATHEESTONI Sharpe.

1856. Pinna atherstoni D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,
vol. vii., p. 193, pi. xxii., fig. 1.

Occurrence. Railway cutting between milestones 24i-24f on the
railway from Uitenhage to Graaff-Reinet, about three miles from
Uitenhage (329, 330), where it is very abundant in a bed of nodular
limestone.

Remarks. It has been noted by Messrs. Rogers and Schwarzthat
this shell, associated with Holcosteplianus atherstoni, marks the
most constantly recognisable zone in the Marine Beds of the Zwart-
kop's River valley.

Pinna atherstoni was compared by Sharpe and Tate with P.
hartmanni Zieten, from the Lias of Europe, and it certainly agrees
more closely with P. hartmanni, as figured by Goldfuss.t than with
the generality of Cretaceous forms, most of which are more elon-
gated and slender in outline. The original figure given by Zieten \

* Damon (1), pi. ii., fig. 5.

t Goldfuss (1), Band ii., p. 164, Tab. cxxvii., fig. 36 (1837).

t Zieten (1), p. 73, Tab. 55, fig. 5 (1833).

The Invertebrate Fauna of the Uitenhage Series. 77

has, however, a very different aspect from that of Goldfuss ; it
apparently has a narrower apical angle, and the surface ornaments
are so delineated as to be scarcely comparable with those of P.
atherstoni. The resemblance of P. atherstoni to the Liassic form
usually known by Zieten's name is clearly a fact of no importance
in the question of the age of the Uitenhage fauna. It may be noted
that a specimen of Pinna from the Perna-bed of Atherfield, Isle of
Wight, preserved in the Sedgwick Museum at Cambridge, has form
and proportions closely similar to those of P. atherstoni. The
specimen is unfortunately ill-preserved and unfitted for a de-
tailed comparison. A similarly broad form has also been recorded
from the Buda Limestone, the upper part of the Lower Cretaceous
series of Texas.*

GENUS OSTEEA Linnasus.

OSTREA SP.

Specimens of Ostrca (305, 306, 336) which occur in the cliff below
the old school-house at Dunbrodie (right bank of Sunday's River),
in association with Perna atherstoni, Pecten cottaldinus and Gastro-
cliCBiia dominicalis, are in a very poor state of preservation and
consist chiefly of fragments. Valves of flattened form measure
upwards of 40 mm. from the umbo to the opposite border and about
30 mm. from margin to margin in a direction perpendicular to this.
The outline of the valve becomes broadly pointed at the umbonal
end. Other imperfect individuals are represented by valves of
smaller dimensions, and these appear to be less elongated and more
convex. None of the valves suffices for specific determination, but
it is probable that two forms are represented.

GENUS EXOGYEA T. Say.

EXOGYRA IMBRICATA KraUSS.

1843. Exogyra imbricata P. Krauss, Amtlicher Bericht iiber die

zwanzigste Versammlung der Gesellsch. deutsch.

Naturforscher und Aerzte zu Mainz im Sept., 1842,

p. 129.
1850. Exogyra imbricata P. Krauss, Nov. Act. Acad. Caes. Leop.-

Carol. Nat. Cur., vol. xxii., pt. 2, p. 460, Tab. 50, figs.

2a-2d.
1856. Gryplicea imbricata D. Sharpe, Trans. Geol. Soc. Lond.,

ser. 2, vol. vii., p. 197, pi. xxiii., figs. 3a, 3b.

* Shattuck (1), p. 19, pi. vii.

78 Annals of the South African Museum.

1867. Ostrea imbricata E. Tate, Quart. Journ. Geol. Soc., vol.xxiii.,

p. 154.

1904. Pycnodonta imbricata H. Douville, Bull. Soc. Geol. France,
ser. 4, tome iv., p. 215.

Occurrence. This form is widely and abundantly distributed in
the Marine Beds of the Zwartkop's, Sunday's, and Coega Eiver
valleys. It was met with at almost all the exposures in the
Zwartkop's valley visited by Messrs. Kogers and Schwarz. Ex-
amples sent to me are from the railway cutting between milestones
24| 24f on the railway from Uitenhage to Graaff-Eeinet (312),
and others were collected by Miss M. Wilman at Coega. Speci-
mens sent from the collection of the South African Museum are
from the Sunday's Eiver (300, 301, 302, 303).

Remarks. A striking character of the late adult and senile
condition of E. imbricata consists in the prodigal manner in which
shell substance is added at the margins of the valves, particularly
noticeable in the smaller (right) valve in the neighbourhood of the
hinge. Here, with its margin consisting of many coarse, imbri-
cating laminae, this valve may attain a thickness exceeding 30 mm.
The ligament pit is seen to be strongly curved near the apex, but
straight in the portion representing the later growth-stages.

This abundant and characteristic shell has given rise to some
differences of opinion concerning its generic position within the
Ostreidae, which, however, may be sufficiently accounted for by the
great variability of form exhibited by different individuals, and by
the fact that the shell presents features which are recognised as
leading characteristics of both Exogyra and Gryphcea. For this
reason Tate used the name Ostrea in the broad sense, stating his
opinion that the species in question " affords a good illustration of
the mere sectional value (which cannot at all times be employed) of
the groups Exogyra and G-rypli&a." In many instances, it must
be admitted, apparent passage forms seem to render difficult the
definite separation of the three genera, so easily distinguishable
when represented in their more characteristic aspects. * Jackson
has made some luminous observations on the relation of these three
genera and on the influence of the character of attachment by
cementation in producing modifications in members of the Ostreidae.t
He believes that Gryphcea and Exogyra " are probably aberrant
members of the Ostreidas, not typical forms in the line of evolution
of the group."

* Leymerie (Ifl). See also Peron's remarks on these genera, Peron (1),
pp. 107-9. t Jackson (1), sections v. and vii.

The Invertebrate Fauna of the Uitenhage Series. 79

It is highly probable that the shells classed as Gryphaa do not
represent a homogenetic group, but are polyphyletic in origin, in-
cluding repeated offshoots from an ostrean stock. While, in the
progress from Ostrca to Gryphcea, there is diminished duration of
the attached habit, so that the typical Gryphaa is fixed by cementa-
tion of the left valve only during the young stage, examples are not
wanting in which the highly specialised characters of Gryphcea are
seen to become modified by the acquirement once again of more
prolonged attachment. As an instance of this, reference may be
made to the shells which, at the close of the deposition of the
Oxford clay in England, seem to have largely replaced the familiar
Grypluza dilatata J. Sow. In these modified forms, the attached
valve did not become free until the neanic stage had been com-
pleted or the adult stage had been well entered upon, and the shell
therefore perforce retained a relatively flatter and more ostreiforni
aspect.' 1 ' Exogyra imbricata might be thought, on cursory exami-
nation, to bear no slight analogy to these : the area of attachment
has very frequently a similar relation, in point of dimensions, to
the whole fully-grown valve, and it is only on the cessation of
attachment that the individual acquires the manner of growth of a
Gryplicsa and develops the arcuate form which led Sharpe to institute
comparison with Liassic shells. It must be realised, however, that
in this African form we have an illustration of the passage from a
more complete to a less persistent duration of attachment, in the
life of the individual, for a study of the youthful stage shows beyond
doubt the exogyrate ancestry, and this may be clearly seen in many
specimens in which the nature of the youthful characters did not
become masked by the modifications incidental to fixation. The
duration of attachment varied very much in different individuals,
and no doubt often depended upon the nature and form of the object
to which the young shell adhered. In some cases a relatively large
area of attachment may retain the impress of some foreign surface,
such as that of the large and coarsely ribbed Cucidlcea kraussi
Tate, or a pseudo-quadrate Trigonia, in a manner which largely
obscures the true nature of the umbonal region. Other individuals,
again, seem to have secured themselves to some less suitable or
stable surface, and at an earlier stage to have entered upon the
period of freedom. In these, as in some which have been attached
to an even surface, the characters of the fixed stages are clearly seen
to be those of a true Exogyra, and present the strongest contrast to

* Compare also figure of Gryphcea alligata from the Corallian of Nattheim
Quenstedt (1), p. 752, Tab. 91, fig. 25 (1857).

80 Annals of the South African Museum.

the features of the nepionic and neanic stages of Grypli&a as illus-
trated, for instance, in the admirable descriptions and figures given
by Hill and Vaughan of members of the genus occurring in the Lower
Cretaceous strata of Texas." When we see how closely the manner
of growth in the adult stage of E. imbricata often simulates that of
a true Gryphaa with simple ostrean ancestry, it must be admitted
that the parallelism is very striking.

It would be difficult to indicate within narrow limits the true
relationship of Exogyra imbricata, but it is evident that its nearest
morphic counterparts are essentially characteristic of Lower Creta-
ceous rocks. Thus, a very close resemblance exists between this
form and some of the shells classed by Leymerie under the name
Exogyra sinuata and E. subsinuata. Leymerie first regarded all
these as varieties of Exogyra sinuata,}- and included forms to which
numerous specific names had previously been applied. He subse-
quently separated E. subsinuata as a species distinct from E. sinuata,
and used several varietal designations of both. Eegarding the
value of the nomenclature employed by this author or by Coquandj:
in dealing with the forms comprised within this perplexing group,
I am at present unable to attempt the critical and detailed study
which alone would warrant the expression of any judgment ; in like
manner, the varying use of the name Exogyra couloni by different
authors has given rise to such a degree of confusion that, in insti-
tuting comparisons, it must suffice to make reference to actual
figures or descriptions, leaving aside for the time being all question
of the value or correctness of the specific names attached to certain
of these.

Krauss rightly recognised the broad relationships of E. imbricata
and believed it to be nearly related to " Grypli&a couloni Defr.,"
from which, however, he found it to differ in the following points :
" It is larger, more thick-shelled, longer and narrower ; has a keel
narrower at the end (though always rounded), more concave laminae
lying much higher one upon the other, and a stronger umbo bent
further over the opercular valve, than in Gryphcea couloni Defr."
Some of the shells figured by d'Orbigny as Ostrea couloni show
no slight resemblance to selected individuals of E. imbricata, but are
narrower than the generality of the Uitenhage specimens, and the
imbricating lamellae of the larger valve are more prominent and
irregular. The broad, plicated specimens figured by d'Orbigny
under the same name on plate 466 of his work cannot be brought

* Hill and Vaughan (1). f Leymerie (1).

J Coquand (2). d'Orbigny (3), pi. 467, figs. 1-3 (1848).

The Invertebrate Fauna of the Uitenhage Scries. 81

into comparison. The shells figured by Coquand * as Ostrea couloni
include narrow forms, more elongated and more decidedly carinated
than E. imbricata, as well as a broad type, developing plications
of the larger valve in the later adult stage, which is in no respect
closely comparable with the form we are considering.

Sowerby's Exogyra sinuata,\ though itself very variable, is
typically widely separated from E. imbricata by the strong carina-
tion of the larger valve and the widely expanding form of this, with
the much less arcuate habit of growth and the considerably greater
adult dimensions. Some French forms figured by Leymerie and
brought into relationship with Sowerby's species, though perhaps
erroneously, afford, on the other hand, material for a close com-
parison with E. imbricata. Thus, Leymerie's E. sinuata, var.
elongata J can be fairly closely matched, while extreme individuals
of E. imbricata, in their narrow outline and very arcuate growth,
nearly resemble that author's figure of E. subsinuata, var. aquilina.
The shell from the Neocomian of the Haute-Marne figured by Bayle ||
as Aetostreon aquilinum (Leym.), again, closely resembles very narrow
and curved specimens of the African form.

Ostrea tardensis Stanton, H from the Lower Cretaceous Gio beds
of Patagonia, may, in selected individuals, somewhat closely
approach E. imbricata, but its near relationship to this must be
considered doubtful ; it seems, however, to represent in some
respects an analogous type. The exogyrate character of the
umbonal region is only feebly indicated, according to Dr. Stanton,
who says that the " lower valve is very thick and very convex,
obscurely carinate, with the beak more or less twisted laterally, but
not distinctly coiled." It differs also by frequently acquiring a
marked crescentic form in the adult. If the Patagonian shell had
a true exogyrate ancestry, which appears probable, the early
characters seem to have so far disappeared as to indicate that
relationship to E. imbricata is probably somewhat remote.

The shell from the Neocomian of Arqueros in Chili described by
Bayle and Coquand as Ostrea couloni, '"''' and afterwards brought into
association with a heterogeneous company under the collective name

* Coquand (2), p. 180, pi. 65, fig. 10 ; pi. 71, figs. 8-10; pi. 74, figs. 1-5; pi. 75,
figs. 1-6, 22.

f J. Sowerby (1), vol. iv., Tab. 336 (1822). [As Gryphaa.]

I Leymerie (2), p. 17, pi. 12, fig. 2.

Ibid., pi. 12, figs. 6, 7.

|| Bayle (1), pi. cxl., figs. 3-5.

<f Stanton (3), p. 11, pi. i., figs. 1, 2 ; pi. ii., figs. 1, 2.

* Bayle and Coquand (1), p. 37, pi. vii., figs. 1, 2.

82 Annals of the South African Museum.

" Ostrea aquilina" by Coquand,* closely resembles many specimens
of E. imbricata in the form and inflation of the valves and in the
degree of curvature of the umbonal region ; but it differs by the
more marked carination of the larger valve. Eemes has considered
both the Chilian shell and Exogyra imbricata Krauss to be identical
with Sowerby's Exogyra sinuata,\ but this is certainly an error.
The same author, however, has figured an Exogyra from Stramberg, |
ascribed by him to Exogyra subsinuata Leym., to which selected
individuals of E. imbricata, in which the surface of attachment is
relatively large, may show considerable resemblance.

An exogyrate shell from the Bajocian of Abyssinia was ascribed by
Douville to E. imbricata Krauss, but this was an erroneous identi-
fication, as already pointed out by G. Miiller.|| The resemblance
of the small specimens figured by Douville to selected young
individuals of E. imbricata is indeed not a distant one, but many
of the Uitenhage specimens are attached by a very much larger
surface, and, moreover, attain a colossal size in comparison with
Douville"s types, if these represent the adult stage. The variation
in E. imbricata is so great that a more detailed and critical com-
parison could not be entered upon without a substantial suite of
specimens of the Abyssinian shell.

A shell from the Isakondry basin in Madagascar (presumably
from strata of Lower Cretaceous age) is stated by Douvill6 to bear
a close resemblance to Exogyra imbricata Krauss. 11

GENUS MYTILUS Linnaeus.

MYTILUS UITENHAGEKSIS sp. nov.

Plate II., figs. 10, 11, lla.

Desertion. The shell is slender and elongated in outline, sharply
pointed in front, and antero-ventrally truncated. The hinge-margin
is straight, and passes backwardly by a curve into the obliquely
sloping posterior margin, which is only very slightly convex in
outline until the posterior end of the valve is reached. In profile,
the antero-ventral margin is straight, passing from the acutely
pointed umbo to the posterior angle of the valve, where it abruptly
cuts the curved outline of the posterior margin.

The antero-ventral area is flat, and its surface is perpendicular

* Coquand (2), p. 158. | Eenies (1), p. 216.

J Eemes (1), pi. xxi. (iv.), tigs. 2, 21>.

Douville (1), p. 230, pi. xii., figs. 8, 9. || G. Miiller (1), pp. 569, 570.

IT Douville (2), p. 388; Douville (4), p. 215; Lemoine (1), p. 176.

The Invertebrate Fauna of the Uitenhage Series. 83

to the plane of symmetry of the valves. The junction of this portion
of each valve with the very gently convex upper portion is acutely
angular, giving rise to a sharp carinal ridge which, however, becomes
very slightly blunted towards the posterior end of the valve.

The shell is devoid of sculpture, but its smooth upper surface is
marked by numerous delicate growth-lines which form a sweeping
curve between the hinge-margin and the carinal ridge. The antero-
ventral area is sometimes marked by coarser, rounded, obliquely
running ridges of accretion. Traces of the narrow, elongated,
fossilised ligament have been observed.

Dimensions.

Length of the antero- ventral margin 37 mm.

Length of the cardinal margin 16 ,,

Greatest breadth of flank between the carinal ridge and the

posterior valve-margin 13 ,,

Greatest width of the antero-ventral area in one valve 7 ,,

Occurrence. Found at Dunbrodie, Sunday's Eiver (319). Speci-
mens collected by Atherstone and preserved in the British Museum
(Natural History) are labelled "Cuylers." Also collected by Mr.
Eogers from the lowest beds on Zoet Geneugd, right bank of
Sunday's Eiver (64h), and in the Coega Valley, one mile up the
line from Coega station (479g).

Remarks. This is doubtless the form noticed by Krauss :;: in the
lowest fossil-bearing beds on the left bank of the Zwartkop's Eiver
below Uitenhage, in association with Trigonia herzogi, T. conocardii-
formis, and T. ventricosa. Krauss speaks of a Mytilus " with a sharp
ridge running from the umbo to the hinder extremity and a flat
truncated surface extending from this ridge to the ventral margin."
The shell was capable of attaining dimensions exceeding those
of the larger specimen figured here. A specimen in the Geological
Society's collection, incomplete at the posterior end, must have
originally measured 50 mm. in length ; the maximum breadth of
its flat area is 11 mm. in each valve. Two imperfect large valves
now preserved in the British Museum (Nat. Hist.), which form part
of a collection at one time placed in the Museum of Practical
Geology, probably had a length of fully 45 mm. when complete.
The antero-ventral area measures about 9 mm. in breadth in one
of these, while the greatest breadth of the flank, measured by a line
perpendicular to the carinal margin, is 18 mm. The area is perfectly
flat and the carinal ridge acutely sharp.

In the absence of knowledge regarding the interior of the shell,

* Krauss (2), p. 443.

84: Annals of the South African Museum.

this form can at present only be looked upon as a representative
of the genus Hytilus. No doubt, indeed, concerning its generic
position could have been reasonably suggested, had not a detailed
study of the externally very similar Mytilus lanceolatus J. de C.
Sowerby, led Mr. H. Woods to ascribe that shell to the genus Dreis-
sensia," a determination which was based upon the presence of an
umbonal septum. A strong case was made out by Mr. Woods for
the existence of marine forerunners of the modern Dreissensia, an
inhabitant of brackish and fresh waters, and he was also supported
by the opinion of M. Cossmann, who has found the genus to be
associated with marine forms in the Upper Eocene of the Paris
basin. In view of close external similarity to Driessensia lancco-
lata it would not be surprising if the shell here described were
eventually also shown to be furnished with an umbonal plate,
but until this can be proved it will be necessary to retain the name
Mytilus.

Compared with Dreissensia lanceolata (= Mytilus lanceolatus J.
de C. Sow.)t from the Lower Greensand and Upper Greensand and
Blackdown Beds of England, M. uitenliagensis is seen to agree closely
in general form, but to be distinguished by several points of detail.
In the African shell the antero-ventral area is flat throughout its
extent, which is not the case in the European form ; in M. uiten-
hagensis the carinal ridge is more sharply acute in the umbonal
region than in many examples of D. lanceolata. Further distin-
guishing features possessed by M. uitenliagensis are the relatively
slightly shorter hinge-line, the less steeply sloping outline of the
posterior margin, the relative narrowness of the flank between the
carinal ridge and the posterior margin, and the absence of con-
centric ornaments at any stage of growth. The shell from the
Cretaceous of Shingle Bay, Skidegate Inlet (Queen Charlotte
Islands), described by Whiteaves under the name Mytilus lanceo-
latus, I may or may not be identical with the European form.
To judge from the description and figures, it shows considerable
similarity to M. uitenliagensis, though appearing to differ by the
imperceptible passage of the hinge-border into the posterior border,
the curved outline of the carinal margin, and the concavity of the
antero-ventral area.

Several forms described from the Senonian Greensand of Aachen

* Woods (3), vol. i., p. 110 (1900).

f J. de C. Sowerby (1), vol. v., p. 55, Tab. 439, fig. 2 (1823); H. Woods (3),
vol. i., p. 110, pi. xviii., figs. 13-15, pi. xix., figs, 1-11 (1900).
} Whiteaves (3), p. 236, pi. 31, figs. 7, la.

The Invertebrate Fauna of the Uitcuhage Series. 85

by J. Muller,* and united by Holzapfel i under the name Septifer
tegulatus, may possibly be identical with Sowerby's M. lanceolatus,
according to the view of Prof. Holzapfel and Mr. H. Woods. While
the points of difference from M. uitcnhagensis are in the main those
noted above in the case of the English shell, the specimens from
Aachen dealt with by Prof. Holzapfel appear to be still more strongly
distinguished by their frequently curved antero- ventral margin and
their well-marked concentric ornament.

In Mytilus tornacensis d'Archiac,} from the Tourtia of Tournay,
the binge-line is relatively longer than in M. uitenhagensis, while the
antero-ventral area is not flat and the carina is much less sharp ; there
are also markings on the shell at right angles to the lines of growth.
A form bearing greater similarity to M. uitenhagensis is Mytilus
triangularis J. B6hm, from the Tr/t/oma-sandstone of Lebanon. ||
This has a similar elongated outline, sharp carinal ridge and
flattened antero-ventral area ; but it is apparently well distin-
guished by the manner in which the hinge-margin and posterior
margin pass imperceptibly by a curve into one another when viewed
in profile. In M. triangularis, too, the flank between the carina
and the posterior margin is slightly narrower, while Dr. Bohm's
figure 116 represents the outline of the carina as somewhat curved.
In order to illustrate how great a sameness of characters may
persist or recur in mytiloid shells, it is only necessary to refer to
such a form as M. aviothensis Buv.H from the Lias of Avioth, Breux.
This closely resembles M. triangularis, from the Syrian Cretaceous,
in most outward characters, and differs from M. uitenhagensis
chiefly in those points which distinguish M. triangularis also,
though exhibiting a slight convexity of the antero-ventral area not
developed in these Cretaceous forms.

GENUS MODIOLA Lamarck.
MODIOLA BAINI Sharpe.
Plate II., figs. 12, 12a.

1856. Modiola bainii D. Sharpe, Trans. Geol. Soc. Loud., ser. 2,
vol. vii., p. 193, pi. xxii., figs. 2, 3.

Supplementary Descriptive Note. As an addition to Sharpe 's
description, it may be noted that the oblique lateral " keel " is only

* J. Miiller (1), p. 35. f Holzapfel (1), p. 218, Taf. xxv., figs. 1-9.

J d'Archiac (1), p. 307, pi. xv., fig. 3. J- Bohm (2), p. 218, Taf. vii., fig. 11,
1 1 For remarks on the age of this sandstone see Eehbinder (1), p. 87.
IT Buvignier (1), p. 24, pi. xvi., figs. 35-37.

7

86 Annals of the South African Museum.

developed as a rounded folding of the valve. Above this the surface
is evenly convex, while below it the valve is flattened. An un-
crushed specimen therefore shows in transverse section an outline
different from that depicted by Sharpe in his figure 3b. It is true
that the upper portion of the valve also becomes much flattened in
the posterior part of an adult specimen, but only near the siphonal
margin. The ribs, starting above at the valve-margin, are directed
obliquely backwards, and before the lateral fold is reached they
bifurcate, at the same time curving sharply with the convexity back-
wardly directed. An occasional extra, curved rib is here intercalated.
Just below the lateral fold the ribs unite again in twos or threes and
pass forward in the form of a number of fine, unequally spaced
ridges which occupy the lower flattened portion of the valve and run
roughly parallel to one another and to the lower margin. In addi-
tion to these ridges, the valve below the oblique lateral fold is
ornamented by a series of very numerous and crowded, delicate,
vertically running raised stripes or wrinkles. This vertical wrink-
ling is very regular, and occupies the whole of the surface below the
lateral fold, crossing the horizontal ridges at right angles. The
wrinkles, which number upwards of forty within a space of 10 mm.,
are developed both on the ridges and interspaces.

Occurrence. Collected in the Zwartkop's Eiver valley, one mile
north-east of Eawson Bridge (281). Eecorded by Sharpe from the
Sunday's Eiver at " C. Eoe's drift," and by Stow from above Modder
Drift and from M'Loughlin's Bluff, on Sunday's Eiver.

Remarks. The specimens figured by Sharpe, in the collection of
the Geological Society, show the vertical wrinklings on the lower
portion of the valve, though no mention was made of these. Well-
preserved specimens exhibit these structures very clearly, and show
that they form a definite part of the sculpture.

The group to which this shell belongs is more characteristic of
Jurassic than of Cretaceous rocks. M. sowerbiana (d'Orbigny)
( = M. plicata Sow.)," a typical Oolitic form, has a much more
numerous division of the ribs above the lateral fold, and is rather
more curved in outline. M. perplicata (Etallon),! from the Upper
Jurassic of Europe, is similar in the bifurcation of its ribs ; but it
attains a greater height in relation to length, and has also a more
curved outline. The descriptions and figures of M. perplicata J

* d'Orbigny (4), vol. i., pp. 282, 312; J. Sowerby (1), vol. iii., Tab. 248, fig. 1 (1819).
t Thurmann and Etallon (1), p. 223, pi. xxix., fig. 8 (1862) ; Loriol and Pellat
(2i, p. 156 (312), pi. xviii., figs. 19, 20.

J See also Loriol, Koyer, and Tombeck (1), p. 348, pi. xix., figs. 10, 11.

The Invertebrate Fauna of the Uitcnhage Series. 87

make it clear that the vertical striping on the lower part of the flank
is developed in that form also ; the same occurs in M. mcdus
(d'Orbigny), an allied shell from the Kimeridge.

A shell of very similar type to M. baini is M.flagellifem (Forbes),"
from the Upper Cretaceous of Europe and Southern India ; but an
examination of the type specimens in the Geological Society's collec-
tion shows that these have the ribs dividing chiefly into groups of
three, and the lower part of the flank entirely lacks any traces of the
fine vertical stripes.

A fragment of a similarly ornamented, elongated Modiola from the
Cenomanian of the Sarthe, was considered by d'Orbigny to be speci-
fically distinct from others previously described, and was figured by
Gueranger t as Mytilus sarthcnsis. It may be clearly seen from
that author's photographic illustration that this differs from M. baini
by the relatively greater disparity in the number of large and small
ribs ; in the French shell there ai'e three small ribs to each of the
major ribs.

Modiola rubidgei (Tate),} from the Uitenhage Marine Beds, also
bears a general resemblance to M. baini. In M. rubidgei, however,
the ornaments on the upper part of the valve consist of relatively
few, gently rounded folds in place of the well-defined narrow rounded
ribs in M. baini; in addition, these folds are themselves ornamented
by finer linear markings which pass backwards along them and then
curve forward to run parallel to the lower margin. In M. rubidgei
there is also an absence of vertical striping on the lower part of the
valve.

GENUS NUCULA Lamarck.

NUCULA UITENHAGENSIS Sp. nOV.

Plate II., fig. 13.

Description of a Single Specimen. The shell is oval in outline,
short and high posteriorly, moderately inflated. The cardinal
margin slopes down forwards from the umbo with a gently convex
outline, and passes by a curve into the rather short anterior margin.
The posterior margin falls steeply from the umbo and passes by a
rather sharp curve into the long, gently convex inferior margin.
The greatest height is at the umbo. The umbonal region is rounded
and broad, and not strongly prominent. From the umbo a very
faintly developed, blunt ridge of the valve-surface passes forward,

* Forbes (2), p. 152, pi. xvi., fig. 9 ; Woods (3), vol. L, p. 99, pi. xvii., figs. 1, 2
(1900).

t Gueranger (1), p. 17, pi. xxiii., fig. 1. J Tate (1), p. 157, pi. ix., fig. 11.

88 Annals of the South African Museum.

cutting off a narrow, flattened, or slightly concave area contiguous
with the cardinal margin. The greatest inflation occurs at about the
middle of the valve.

The surface bears numerous closely spaced, delicate, raised, con-
centric linear ornaments, very weakly developed and indistinct in
the umbonal part of the shell, more strongly developed in the
inferior half of the valve. These linear ornaments are not produced
with perfect regularity, but show occasional inequalities in their
spacing, and an occasional coalescence of adjacent lines may be
observed, particularly in the posterior part of the valve.

Dimensions.

Length 17 mm.

Greatest height 11 ,,

Greatest depth of a single valve 5 ,,

Occurrence. Pound by Mr. Eogers in a wash-out 100 feet above
Coega station, at a point one mile north of Coega Hotel (441g).

Remarks. The specimen described has both valves in position,
and is perfect except posteriorly. When the above description was
written the right valve was quite perfect, though the lunular region
(in this genus posterior to the umbo) was somewhat obscured by
very hard matrix, but an attempt to remove this resulted in slight
injury to the valve at the posterior border, so that the margin there
is now not quite perfect. Unfortunately a single specimen only was
obtained, but it seems to be sufficiently well preserved and well
characterised to support the establishment of a new species.

The shell is distinguished by its oval form and rounded, soft out-
lines. In these respects it bears a considerable resemblance to
Nucula obtusa J. de C. Sow., from the Upper Greensand and Black-
down Beds of England."'

GENUS GEAMMATODON F. B. Meek and F. V. Hayden.

GRAMMATODON JONESI (Tate).
Plate II., figs. 14, 14a.

1867. Area (Cucullcea?) jonesi E. Tate, Quart. Journ. Geol. Soc.,
vol. xxiii., p. 161, pi. ix., fig. 9.

Occurrence. Collected by Miss M. Wilman at Coega. This shell
is also recorded by Messrs. Eogers and Schwarz as occurring in the
highest of three beds of sandstone on the Grass Eidge road near
Uitenhage.t The specimens described by Tate occurred "in a

* Woods (3), vol. i., p. 22, pi. iv., figs. 2, 3, 4 (1899).
f Rogers and Schwarz fl), p. 9.

The Invertebrate Fauna of the Uitenhage Series. 89

greenish-grey sandy marl, rather hard and very calcareous, near the
Bridge on the Zwartkop Eiver."

Dimensions. (1) (2) (3) (4)

Length .................. 13 . 18 . 21 22 mm.

Height .................. 9 . 12 . 14 . 13

Depth of a single valve 4 6 ,,

No. (2) is Tate's figured specimen.

Remarks. This form is closely similar in general type to G. sccui'is
(Leym.) :|: (Lower Cretaceous) and G. carinatus (Sow.) t (Gault and
Upper Greensand), and although I have been unable to ascertain
the characters of the hinge, the close agreement in form and orna-
mentation is perhaps sufficient justification for classing this shell in
the same generic group of the Arcidae. In the use of the name
Grammatodon I follow Mr. Woods. J

Compared with G. securis, our shell is found to differ by the con-
siderably finer and more closely spaced ribs on the flank of the left
valve. With regard to the ribbing, there is greater similarity to
G. carinatus, but this, on the whole, is a decidedly more equilateral
shell, although specimens might be selected which in most respects
closely approach G. jonesi. G. carinatus frequently attains much
larger dimensions, with a proportionately coarser development of
the ornaments, than G. jonesi. A shell of similar type, and perhaps
related to G. jonesi, occurs in the Lower Neocomian beds of German
East Africa, which were correlated by G. Mtiller with the Uitenhage
Series. Miiller described and figured this under the name Area
uitenhagensis, and drew attention to the apparent relationship to
Area jonesi Tate. A. uitenhagensis, which is associated with
Trigonia ventricosa (Krauss) and T. beyschlagi Miiller, differs from
Grammatodon jonesi by the fewer number and much greater breadth
of its ribs.

GENUS UNIO A. J. Eetzius.

UITENHAGENSIS Sp. noV.

Plate II., figs. 15, 15,

Description. The shell is of elongated oval outline, posteriorly
produced, and laterally compressed in the postero-inferior region of

* Leymerie (2), p. 6, pi. vii., figs. 6, 7 ; Woods (3), vol. i., part 1, p. 44, pi. vii.,
figs. 14, 15, pi. viii., figs. 1, 2 (1899).

t J. Sowerby (1), vol. L, p. 96, Tab. 44, lower figure (1813) ; Woods (3), vol. i.,
p. 45, pi. viii., figs. 3-8 (1899). I Woods (2).

G. Miiller (1), p. 542, pi. xxv., fig. 5. [This name seems very unhappily
chosen.]

90 Annals of the South African Museum.

the adult. The umbones are situated at almost one-third of the
shell's total length from the anterior extremity. The umbonal
region is relatively very inconspicuous, and of blunt, rounded form.
The shell-substance is for the most part very thick, but becomes
attenuated at the pallial margin. The upper margin in front of the
umbo passes by a curve into the rather short, convex anterior border.
The posterior margin merges into the upper margin and slopes
obliquely back, to pass by an abrupt curve into the inferior
margin.

The greatest height is at the umbo, and the greatest inflation,
which is relatively weak, occurs below the umbonal region, in the
superior half of an adult individual. About the middle of the shell
near the pallial margin the valves appear slightly constricted, as
though laterally pinched in. The surface is marked by numerous
noticeable furrows and ridges of accretion.

Dimensions. (1) (2)

Greatest length 37 . 43mm.

Greatest height 21 . 23 ,,

Greatest depth of a single valve 7 . 8 ,,

Occurrence. In a hard calcareous band in the rocks of Ather-
stone's " Wood Bed " series, on the north bank of the Bezuiden-
hout's Eiver below Blue Cliff station (322, 323).

Remarks. The shells still retain traces of the strong periostracum,
and the elongated external ligament is also in part preserved. This
is apparently the form cited by Messrs. Eogers and Schwarz as
Psammobia,'"'- and such a generic determination might appear to
receive some support from the elongated oval figure of the shell, the
inconspicuous umbones, the relatively compressed aspect of the
valves, and the lengthened external ligament. The valves, how-
ever, are very thick, and this character, as well as the wrinkled
surface and the blunt and obviously corroded umbones, proclaims
another generic position ; moreover, the pallial line is entire. In
the shell-structure, also, although this has become obliterated in its
intimate characters through replacement by calcite, it can be clearly
observed that a line of demarcation separates a thin outer layer,
representing the original prismatic layer, from the relatively thick
inner portion, formerly consisting of nacreous substance.

This form differs from Unio porrcctus J. de C. Sow.,f from the
Wealden, by the less strongly elongated outline and the much less
tapering posterior extremity. Unio antiquus J. de C. Sow.

* Rogers and Schwarz (1), p. 13.

t J. de C. Sowerby (1), vol. vi., Tab. 594, fig. 1 (1828).

The Invertebrate Fauna of the Uitenliage Series. 91

(Wealden) ::: is rather less equilateral, the umbonal region is more
prominent, and the antero-superior border more steeply sloping.

Unio martini J. de C. Sow. (Wealden) f is more equilateral, much
higher and less slender in figure, and probably a more inflated
form.

Unio subsinuata Koch and Dunker,} from the Wealden of North
Germany, is in some respects similar, notably in the form of the
posterior half of the shell and the position of the umbones, but
differs markedly in the horizontally produced superior margin in
front of the umbones. Unio voltzi Koch and Dunker, though
regarded by those authors as distinct from the last, differs in similar
manner from Unio uitenhagcnsis. Unio planus Eoem.|| is more
compressed in form and has a less sloping antero-superior outline.

A comparison with the figures and descriptions of the forms of
Unio from the Jurassic and Cretaceous rocks of North America
does not show that resemblances exist which call for special remark.

GENUS TEIGONIA J. G. Bruguiere.

TRIGONIA VENTRICOSA (Krauss).
Plate III., figs. 1, la.

1843. Lyriodon ventricosa P. Krauss, Amtlicher Bericht iiber die
zwanzigste Versammlung der Gesellsch. deutsch. Natur-
forscher und Aerzte zu Mainz im Sept., 1842, p. 130.

1850. Lyrodon ventricosus F. Krauss (partim}, Nov. Act. Acad.
Caes. Leop. -Carol. Nat. Cur., vol. xxii., pt. 2, p. 456,
Tab. 49, figs. 2c-2/, (exclude figs. 2a, 26).

1871. Trigonia ventricosa F. Stoliczka, Cretaceous Fauna of
Southern India, vol. iii., The Pelecypoda; p. 315,
pi. xv., figs. 9. 9rt. (Palaeontologia Indica.)

1875. Trigonia ventricosa J. Lycett, Mon. Brit. Foss. Trigoniae,
p. 119, and text-figures. (Palaeontographical Society.)

1877. Trigonia ventricosa 0. Feistmantel, Foss. Flora of the
Gondwana System, vol. i., pt. 3, p. 164. (Palseontologia
Indica.)

* J. de C. Sowerby (1), vol. vi., Tab. 594, figs. 3-5 (1828).

t J. de C. Sowerby (2), p. 346, pi. xxi., fig. 17.

J Koch and Dunker (1), p. 58, Tab. vii., fig. 2 ; Dunker and Meyer (1), p. 26,
Taf. xi., figs. 4, 5.

Koch and Dunker (1), p. 59, Tab. vii., fig. 3.

|| F. A. Roemer (1), p. 95, Taf. v.,fig. 14 ; Dunker and Meyer 1), p. 27, Taf. xi.,
fig. 8.

92 Annals of the South African Museum.

1879. Trigonia ventricosa W. T. Blanford, in Medlicott and Blan-

ford, Manual of the Geology of India, pt. 1, p. 261, with
text-figure.

1880. Trigonia ventricosa O. Feistmantel, Foss. Flora of the Gond-

wana System, vol. ii., p. xxxvii. (Palasontologia Indica.)

1893. Trigonia ventricosa W. T. Blanford, in Medlicott and Blan-
ford, Manual of the Geology of India (2nd edition
revised and partly rewritten by E. D. Oldhain), p. 225,
with figure.

1900. Trigonia- ventricosa G. Miiller, Verstein. des Jura und der
Kreide. Deutsch-Ost-Afrika, Band vii., p. 543, Taf. xix.,
figs. 4, 5.

1903. Trigonia ventricosa F. L. Kitchin, Jurassic Fauna of Cutch,
vol. iii., pt. 2, The Lamellibranchiata ; p. 104, pi. x.,
figs. 4-8. (Palaeontologia Indica, Ser. IX.).

1905. Trigonia ventricosa A. W. Rogers, An Introduction to the
Geology of Cape Colony, p. 291, fig. 25 (2).

Occurrence. This widely distributed and very characteristic form
occurs in the Marine Beds at various localities on the Sunday's
and Zwartkop's Rivers. It was collected by Messrs. Rogers and
Schwarz at a kloof on the left side of the Zwartkop's River, east-
north-east of Red House (308), and was found to occur abundantly
in a cliff at " Picnic Bush " ; it is also cited from localities in the
neighbourhood of Uitenhage, on the road to the top of Grass Ridge
and in the railway cutting between milestones 24| 24f on the
Graaff-Reinet railway, about three miles from Uitenhage.* Mr.
Rogers collected it on the left side of Coega Valley, half a mile down
from the railway (463g), and in a small kloof three miles up the left
bank of Sunday's River (14h, 15h). Stow remarked upon the great
abundance of this shell in the cliff at " M'Loughlin's Bluff" or
"Prince Alfred's Rest " on the Sunday's River, f Krauss found it
in the left bank of Zwartkop's River below Uitenhage, associated
with Trigonia herzogi and T. conocardiiformis.

In German East Africa, shells referred to this species have been
found in strata ascribed by G. Miiller to the Lower Neocomian and
brought by him into relationship with the Uitenhage beds, at a
locality a short distance to the north of the Nkundi stream, 29 km.
north-west of Kiswere. There they occur with Trigonia bcyschlagi,
which shows strong resemblance to T. crassa Kitchin, from the
Oomia marine beds in Cutch.

* Rogers and Schwarz (1), pp. 8-10. f Stow (1), p. 502.

The Invertebrate Fauna of the UitcnJiaye Scries. 93

In India, T. ventricosa occurs numerously at various localities in
the marine beds of the Oomia Group in Cutch, notably in the
neighbourhood of Oomia and Goonaree ; it is also stated to have
been found by King in an outlier of the Tripetty beds at Inna-
parazpolliam, to the north-west of Coconada on the south-eastern
side of the Indian peninsula " ; It is recorded, too, from the
Margalla Pass in the district of Hazara, where it is said to occur in
abundance.!

Remarks. Trigonia ventricosa bears a general resemblance to
members of the aliformis group of the section Scabrae, particularly
to T. aliformis Park.,{ with which it was confounded by von Buch,
and to T. scabricola Lycett ; but the relative shortness and great
height of the outline, the very strong anterior inflation, and the
coarse and salient character of the ornaments on the anterior part
of the adult shell, serve to separate it from all known members of
the section in Europe. It has been repeatedly pointed out that
much closer agreement is shown to T. tuberculifera Stoliczka, from
a higher horizon in the Cretaceous rocks of Southern India. The
resemblance appears to be remarkably great, but unfortunately
Stoliczka's description and figures of T. tuberculifera leave some-
thing to be desired. I have elsewhere drawn attention to the fact
that in the figure of the front view of Stoliczka's type |] the ribs of
the right and left valve are represented as though placed opposite to
one another at their frontal terminations ; in T. ventricosa, on the
other hand, as in other members of the section, these ribs alternate
in position, and it seems probable that an error was committed in the
execution of this figure of T. tuberculifera.

A comparison of T. ventricosa with T. kraussi sp. nov. will be
found in the remarks appended to the description of the latter.

T. sub ventricosa Stanton, from the Belgrano beds of the
Pueyrrydon Series in Patagonia,*^ so far as can be gathered from
the description and figures of that shell, is of very similar type, but
seems to be less strongly inflated, and to be of relatively more
elongated form.

T. delafossei Bayle and Goquand,** from the Neocomian of Chili,
also shows a general agreement with T. ventricosa, and may

* King (1), p. 229 ; Feistmantel (2), p. 164 ; Feistmantel (3), p. xxxvii.

f Wynne (2), p. 125 ; Medlicott and Blanford (1), p. 503.

I Parkinson (1), p. 176, pi. xii., fig. 9; Lycett (3), p. 116, pi. xxv., figs. 3-0
(1875). Lycett (3), p. 130, pi. xxvii., figs. 4, 5.

|| Stoliczka (2), pi. xv., fig. 10 (1871). I Stanton (3), p. 18, pi. iv., figs. 19, 20

** Bayle and Coquand (1), p. 37, pi. viii., fig. 27; E. A. Philippi (1), p. 85,
pi. xxxvi., fig. 4 ; Paulcke (1), p. 296, Taf. xvii., fig. 9.

94 Annals of the South African Museum.

perhaps be closely related. It appears, however, to have more
closely spaced and less coarsely nodose ribs. To judge from the
description and figures of T. delafossei given by Paulcke, the
escutcheon is more sharply demarcated from the area than in
T. ventricosa; T. delafossei also does not appear to have attained
such large dimensions. Another South American form, T. nepos
Paulcke, from the Neocomian of Chili, shows to some extent a
similarity to T. ventricosa, especially if the smaller individuals
figured by Paulcke :|: be compared with specimens of similar size ;
T. nepos, however, has more delicate sculpture and is also well
distinguished by the peculiar arrangement of the ribs towards the
frontal face. In the work cited above, Paulcke brings T. nepos and
T. delafossei into close relationship ; yet he perceives in T. nepos an
exceedingly close resemblance to T. baylei Dollfus, t from the Kime-
ridge of Europe, and recognises in the similarity an indication of the
scaphoid ancestry of these Cretaceous forms. It may be pointed
out that the principal character upon which he relies, the existence
of a more or less independent series of anterior ribs, is a feature to
which, alone, such significance can certainly not be attached. It
represents a plan of sculpture which has appeared repeatedly and
independently in various Trigonia- stocks, and in T. baylei and other
typical Jurassic Scaphoideae is associated with well-marked cha-
racteristics which indicate the very high improbability of such
direct relationship as that suggested. In T. ventricosa, T. delafossei,
and T. nepos, just as in the T. aliformis group of Europe, the very
prominent and highly inflated umbonal region, the extreme incurva-
tion of the umbones, the relatively very narrow area and the
crenulation of the shell margin, are characters which are conspicu-
ously developed in these well-marked and specialised groups. In
T. baylei and similar ScaphoideaB, the laterally compressed form,
the weak umbonal incurvation, the nodose carinae and delicately
sculptured area, no less than the relations of frontal to lateral
costae, are also characters of specialisation, and judging by the
evidence of early growth-stages, these shells had ancestry in
simple clavellate forms. A similar though independent ancestry
seems also very probable in the case of the various groups classed
under the broad heading Scabrae.

In the published description of the Trigonia from the Oomia
Group in Cutch, it is stated that the Indian specimens ascribed to
T. ventricosa " although exhibiting great variability, offer no features

Paulcke (1), p. 293, Taf. xvii., fig. 8.

t Dollfus (1), pi. xv. ; Bigot (1), p. 309 [51], pi. xii., fig. 10.

The Invertebrate Fauna of the Uitenhage Series. 95

except their state of preservation by which they may be dis-
tinguished from those occurring in South Africa."' Having due
regard to the range of individual variation shown by the specimens
which occur in the Oomia beds, I believed this statement to be
quite accurate, but a reconsideration of this matter and a renewed
comparison between a larger number of specimens now necessitates
a slight modification. While it is seen that the limits of individual
variation among the Indian specimens cover differences greater
than those to be observed between the average characters shown
by examples of the species from the one region and those from the
other, it should be noted that in India many of these shells exhibit
a somewhat less degree of inflation than that seen in most of the
South African examples. The specimens from Cutch sometimes
have the tubercular ornaments rather more prominently developed,
though in certain cases the effect of prominence seems to have
been enhanced by the mode of preservation of the shells. There
is also a tendency to have the tubercles less closely crowded
together and less regularly moniliform than in the specimens from
South Africa. It is difficult to say in how far these characters
may have been the product of strictly local conditions. There
certainly appears to have been a greater range of individual varia-
tion and greater instability than is shown by the specimens from
the Uitenhage beds. At the same time, the agreement between a
selected series of specimens from the one region and a suite of
individuals from the other is remarkably close, and, all things
considered, I do not think that the points of difference, above
noted, suffice for the establishment of two separate species, or,
indeed, even for the satisfactory recognition of two well-defined
local races.

TRIGONIA KEAUSSI sp. nov.
Plate III., figs. 2, 2a.

1850. Lyrodon ventricosus F. Krauss (partini), Nov. Act. Acad. Cass.
Leop. -Carol. Nat. Cur., vol. xxii., pt. 2, Tab. 49, figs. 2a,
26 (exclude figs. 2c-2/).

Description. The shell is relatively short, anteriorly very high,
with the upper and lower margins rapidly converging posteriorly
towards the very short siphonal margin. The valves are anteriorly
very strongly inflated, but posteriorly compressed. The umbones
are prominent, strongly incurved and markedly recurved, and
situated close to the anterior end. The greatest height is attained

* Kitchin (1), p. 107.

96 Annals of the South African Museum.

at the umbo, and may exceed the total length. The valve is almost
vertically truncated anteriorly and in its anterior part attains such
relatively great depth that a flattened frontal face of considerable
extent becomes developed.

In the youthful stage the ribs of the flank are concentric in
arrangement, extending from the carinal angle to the frontal border.
As growth proceeds, subsequently formed ribs terminate anteriorly
at the frontal border but posteriorly at an increasing distance from
the carinal angle. The ribs at the same time attain a more steeply
inclined direction, and the last-formed rib of the frontal series is
very short and almost vertical. In the early adult stage narrow,
closely spaced ribs extend between the carinal angle and the upper
terminations of the swollen frontal ribs, or may be looked upon as
the attenuated upward prolongations of these since the two sets are
so far in continuity. Successively formed narrow ribs, constituting
a posterior series, have their upward terminations slightly removed
from the carinal angle, and in the posterior half of a fully grown
specimen these crowded ribs of the posterior series entirely occupy
the flank. The ribs of the anterior series are very prominent and
coarsely tubercular, and may be elevated to a height of 7 or 8 mm.
above the surface of the interspaces. These ribs are very widely
spaced, the interspaces reaching 8 mm. in breadth in the adult stage ;
on the frontal face the ribs sweep round in a broad curve towards
the anterior valve-margin. The development of the earlier-formed
ribs is not continued in the later growth-stages, so that in a large
specimen the frontal face is partly without sculpture in its upper
part, towards the valve-border. The ribs of the posterior series are
at first vertically directed, but the majority of them have a marked
backward inclination when traced downwards. They are relatively
weak, and are separated by interspaces less than 2 mm. in breadth.
The whole surface of the flank is crossed by crowded lines of growth
which are most plainly visible in the wide interspaces between ribs
of the anterior series.

The carinal angle is represented by a blunt, rounded fold in the
adult shell. The area is very narrow, and is without ornamentation
in the adult stage, except a strongly impressed median longitudinal
groove. The escutcheon is relatively broad and is deeply excavated
in form.

Dimensions.

Length (approximate) 75 mm.

Height measured from the umbonal region 82 ,,

Greatest depth of a single valve 45 ,,

The Invertebrate Fauna of the Uitenhage Scries. 97

Occurrence. Messrs. Eogers and Schwarz record the occurrence
of this shell ("the large variety of T. ventricosa") * in the railway
cutting between milestones 24^ 24f on the line from Uitenhage to
Graaff-Keinet, about three miles from Uitenhage. The specimen
from the South African Museum, here figured, is probably from the
Sunday's River. Krauss met with this form in the left bank of the
Zwartkop's Eiver below Uitenhage.

Remarks. This characteristic Trigonia, which belongs to the
section Scabras, may be said to exhibit in very exaggerated manner
some of the most striking features which render T. ventricosa
(Krauss) so conspicuous amongst the members of this section.
T. ventricosa, though so well distinguished by its form and ornamen-
tation, is in some respects comparable with the European T.
scabricola Lycett or T. aliformis Park., and was even considered
identical with the latter by L. von Buch.i T. kraussi is so far
removed from European members of the section that a detailed
comparison with these is unnecessary, and it only remains to draw
attention to the characters which differentiate it from T. ventricosa
and the apparently closely allied T. subventricosa which occurs in
the Belgrano beds of Patagonia.

T. kraussi is principally distinguished by its large size and the
very prominent character and wide spacing of the anterior ribs.
Another striking feature is the great anterior inflation and extensive
flattened frontal face. In other respects the resemblance to T.
ventricosa is so great that I was led to ask Mr. Rogers whether he
was aware of the existence of specimens intermediate in character
between the extremes ; his reply, however, was in the negative. So
many specimens of T. ventricosa have been obtained by various
collectors, and the Uitenhage TrigonicB are so well represented in
European museums that we might certainly expect to find inter-
mediate forms, if such existed, in some of the collections. No
specimens connecting the extreme forms appear to have been found,
and amongst the Trigoncs of the Oomia beds in Cutch the representa-
tives of T. ventricosa, met with in abundance, agree with the smaller
typical form in South Africa.

T. kraussi has approximately double the dimensions of T. ventricosa,
but is, perhaps, slightly shorter relatively to height. The number
of anterior ribs formed in the youthful and early adult stages may
perhaps be not widely different, but the spacing of the ribs rapidly
widens in the larger form, and the interspaces may reach 8 mm. in

* Eogers and Schwarz (1), p. 10. f von Buch (1), p. 23.

98 Annals of the South African Museum.

width. In the adult T. kraussi, within a space of 33 mm. measured
backwards along the flank from the lower end of the frontal margin,
three ribs only are included. At a height, measured from the umbo,
which represents the full adult height of T. ventricosa, the anterior
ribs on the flank of T. kraussi are already much more robust and
more widely spaced than in T. ventricosa. The anterior inflation is
also much stronger. These very well-marked differences in characters
developed at a comparable stage in the two forms lead me to
separate definitely T. kraussi from T. ventricosa. It may well be the
case that we are here dealing with two branches of a not far re-
moved ancestral stock, both of which have evolved along similar
lines, but one more rapidly than the other. The more advanced
type may be recognised in T. kraussi, from the fact that it exhibits
all the late adult characters of T. ventricosa while yet in the early
adult stage itself ; the late adult characters of T. kraussi are never
reached in T. ventricosa. It is highly improbable, indeed, that this
giant form merely represents extreme individual variation in T.
ventricosa, and its separation from this seems to me to be
justified.

There is a close general agreement with T. subventricosa
Stanton,* especially in the large dimensions, but the Patagonian
shell is relatively more elongated, has the valves less strongly
inflated, and the flattened frontal face less extensively developed.
In T. kraussi the umbonal region is more narrowed in form and
considerably more prominent, taking Dr. Stanton's figure to repre-
sent a typical specimen of the Patagonian shell.

Both T. ventricosa and T. kraussi are figured by Krauss as repre-
sentatives of the same form, and he makes no mention of the
differences to which I have above alluded, though these are
plainly shown in the figures which accompany his description of
T. ventricosa. In deciding which of Krauss's specimens are to
bear the name T. ventricosa it is well to note that his descrip-
tion more accurately applies to the small specimens, and it
may further be observed that when he spoke of the species as
"particularly numerously" represented he must have referred
to the smaller form. This alone has been abundantly collected
and it has also been frequently spoken of in the literature
under the name T. ventricosa. Moreover, no less than six
authors have figured the smaller shells under the name given
by Krauss.

* Stanton (3), p. 18, pi. iv., figs. 19, 20

The Invertebrate Fauna of the Uitenhage Series. 99

TRIGONIA ROGERSI sp. nov.
Plate III., figs. 3, 3a ; IV., fig. 1 ; V., fig. 2.

Description. The shell is of somewhat elongated form ; it is
anteriorly high and is posteriorly more slender in outline and con-
siderably produced. The valves are moderately inflated in the
anterior half of the shell, and are posteriorly more compressed.
Anteriorly, the surface of the valve curves round towards the frontal
margin so as to give the shell some appearance of anterior flattening
or even truncation, most marked in specimens of large size. The
umbones are situated at about one-quarter of the total length from
the anterior extremity. They are prominent, well incurved, and
slightly recurved. The cardinal margin slopes down gradually,
forming a long, almost straight outline, and passes by a rather
abrupt curve into the short, convex posterior border. In front of
the umbo the valve margin falls steeply at once to form the lengthy
frontal border, gently and regularly convex in outline, which passes
by a regular and broad curve, without break, into the lower border.
This in turn has a gently convex outline, and slopes up gradually
towards the posterior margin. The greatest height falls at the umbo.
The costae of the flank are concentric in arrangement until the
close of the neanic stage, becoming then more and more downwardly
inclined, when traced from their commencement at the carinal
angle. In an adult specimen the ribs in the anterior third of the
shell are not at all curved in form as they pass obliquely down the
flank, but on the frontal face of the valve they curve rapidly so as to
approach the frontal margin horizontally. The successive costse are
more and more steeply directed until at about the middle of the
valve their direction is vertical. Posteriorly to this they have a very
slight backward inclination. The ribs in the anterior, inflated part
of the shell are prominent, strongly nodose or tubercular, and are
separated by interspaces as broad as the ribs themselves ; on the
frontal face the interspaces are much wider. The ribs of the
posterior half of the shell are narrower and less prominent in
character, are only weakly nodose, and are more closely crowded
together. In a large individual, about a dozen large, prominent ribs
may be counted in the anterior part of the valve. In the early adult
period, occasional short rows of delicate nodes of unequal extent may
be intercalated on the surface between the main ribs where these
approach the valve margin on the frontal face. On the surface
of the valves clearly marked lines of growth are well seen as they
cross the interspaces, but are either little conspicuous or absent on

100 Annals of tlie South African Museum.

the tubercles of the ribs. In the largest individuals, characters of
senility are seen in the imperfect rib-formation near the lower
margin, and the replacement of sculpture by simple ridges and lines
of accretion at the frontal margin, where the ribs may then be seen
to terminate at some little distance from the actual margin.

For a distance of about 15 mm. from the urnbonal apex a very
delicately nodose, narrow marginal carina is present. It then
dwindles abruptly and disappears, while the carinal angle becomes
less marked when traced posteriorly and is replaced ultimately by a
gentle and rounded fold of the valve surface.

The area is very narrow anteriorly, and gradually broadens
posteriorly. For a distance of about 15 mm. from the umbonal
apex it is ornamented by delicate, slightly granular, transverse ridges
separated by narrow grooves. The terminations of these ridges at
the carinal angle do not correspond strictly with the upper termina-
tions of the ribs on the flank, but the ridges on the area are rather
more numerous than these. Each of the ridges terminates inwardly
in a delicate, transversely elongated node, a row of which takes the
place of an inner carina. At a distance greater than 15 mm. from
the umbonal apex the area is without sculpture, though two or three
further inner-carinal nodes may be present. The area is divided by
a well-marked longitudinal groove into a narrower, superior, and
broader inferior portion. The groove is continued right to the
posterior margin. At the posterior end of a specimen measuring
84 mm. in length, the area is about 20 mm. in breadth. The
escutcheon is of great size and length, and at the middle of the shell
it is much wider than the area. In its anterior half it is of well-
excavated, concave form. Its ornaments consist of numerous, fine,
beaded ridges or lines of delicate nodes passing from the inner-
carinal nodes rather obliquely backwards across to the cardinal
margin. At 30 mm. from the umbonal apex these ornamenting lines
cease to be formed, and several of them situated posteriorly in the
series do not extend to the cardinal margin. Posteriorly, the inner
carinal ridge of the valve becomes obsolete and the smooth
escutcheon ill-defined. The ligament space is relatively long,
measuring 18 mm. in length in a specimen having a length of
84 mm. In each valve it shows at the cardinal margin a straight
lath-like ridge, becoming gradually thinner when traced back from
the umbo, separated from the escutcheon by a straight, deep, narrow

groove.

The central tooth of the left valve is of relatively great size and
prominence, and its inferior indentation is deep. The inner surface

The Invertebrate Fauna of the Uitenkage Series. 101

of the valve margin is smooth, in the anterior part of the shell at
least. If there is any crenulation at all, it is confined to the posterior
half of the valve, and must be of a weak description.

Dimensions. (1) (2) (3)

Length (estimated) 7s . 84 . 135mm.

Greatest height 53 . 58 92 ,,

Greatest depth of a single valve 19 . 23 . 32 ,,

Occurrence. Obtained by Mr. Eogers from a bare slope W. 30 S.
from the middle of Barkly Bridge, on the farm Olifant's Kop,
Sunday's Eiver (20h) ; also from Coega A'alley, east of the railway,
one mile up the line from Coega station (472g).

Remarks. This is an elongated representative of the section
Scabrse, and, generally speaking, it exhibits the normal characters
of the section. It is remarkable, however, for its very large size and
for the manner in which it shares the character of posterior elonga-
tion with other Tri/jonice from the Uiteuhage beds and from the
Oornia beds of Cutch. This prolongation at the siphonal end, and
the absence of sculpture from the area in the later adult stages are
characters similarly shown in members of the group of Trigonia vau
and in T. conocardi/fon/iis (Uitenhage Series) and in members of the
group of T. v-scripta and certain degenerate Costatas (in the Oomia
beds). There are evidences of degeneracy in all these forms. In the
specimen numbered (2) in the above table of measurements, one valve
is almost complete at the siphonai border, and the form of the valve
at that part suggests that in the complete shell a slight posterior
gape may have been developed, just as in T. stoiui.

In the neanic and early adult stages, T. rogersi bears a consider-
able resemblance to the same stages in T. ventricosa (Krauss). It
differs, however, in details of the sculpture on the area and escutcheon,
and with advancing growth, the characters of outline, degree of
inflation, and ribbing of the shell are so widely distinct in the two
forms as to need no comparison here.

TRIGONIA HERZOGI (Goldfuss).
Plate V., fig. 1.

1837. Lyrodon hcrzogii (Hausmann) A. Goldfuss, Petrefacta

Germanite, Band ii., Lief. 6, p. 202, Tab. cxxxvii.,

fig. 5.
1850. Lyrodon Jierzogii F. Krauss, Nov. Act. Acad. Caes. Leop.-

Carol. Nat. Cur., vol. xxii., pt. 2, p. 453, Tab. 48,

fig. 3.

102 Annals of the South African Museum.

1882. Trigonia herzogi G. Steinmann, Neues Jahrbuch fur Mine-
ralogie, Band i., p. 220, Taf. vii., figs. 1, 2 ; Taf. ix.,
figs. 1, 2.

Occurrence. This is an abundant and characteristic form in the
Sunday's Eiver Beds. It was recorded by Hausrnami from the
neighbourhood of Enon, and by Krauss from the left bank of the
Zwartkop's River below Uitenhage ; also by Stow :;: from various
localities on the Sunday's and Zwartkop's Elvers. A fine specimen
sent to me from the South African Museum is from the Sunday's
Eiver (289). Messrs. Rogers and Schwarz found T. licrzogi on the
north side of the Zwartkop's River at a locality to the west of
Rawson Bridge, and at Picnic Bush; also further up the river on the
road from Perseverance Farm to the Salt Pan, and at Cuyler Manor.
Near Uitenhage, they record this form from two beds of sandstone
exposed in dry watercourses on the ascent of the Grass Ridge road ;
in the railway cutting between milestones 24-| 24f on the Graaff-
Eeinet railway; and on the road to Hillwacht, where the path leaves
the plain. On the Sunday's River it was found near the top of the
white krantz on Wolve Kraal, on the north bank of the river.

The collection made in 1905 by Mr. Rogers includes several
specimens of T. herzogi, which came from the following localities: the
left side of Coega Valley, two miles down from the railway (468g) ;
from the valley east of the railway, one mile up the line from
Coega station (474g) ; the cliff on Buck Kraal, Sunday's River
(116h, 120h, 122h) ; and the cliffs on Zoet Geneugd, Sunday's
River (99h).

Bemarks. Trigonia heriogi is well known from the striking figme
given by Goldfuss, illustrating a fine specimen which was included
in a small collection of Uitenhage molluscs obtained by Hertzog.
These had been previously noticed briefly by Hausmann in his
paper on the geology of South Africa.! The shell is conspicuous
for its large size and elongated outline, its straight and lengthened
cardinal margin and its straight, nodose ribs, backwardly inclined
in the posterior half of a fully grown valve. It was referred by
Pictet and by Lycett to the section Quadrate. An excellent figure
of the shell was given also by Steinmann, who included T. herzogi
with T. transitoria Steinm. in the group Pseudo-quadrat, the
characters of which are briefly discussed below in the remarks
appended to the description of Trigonia holiihi. Steinmann has set
forth in great detail the differences that distinguish T. herzogi from

* Stow (1), pp. 498-505. f Hausmann (1), p. 1458.

TJic Invertebrate Fauna of the Uitcnliage Series. 103

T. transitoria, which occurs in Neocomian strata in Bolivia, Chili,
and the Argentine Republic. T. transitoria shows a closer approxi-
mation of characters to the Clavellatas ; it is not so elongated in
outline, and its strongly nodose ribs have a curved form and a forward
sweep when traced downwards, throughout almost the whole extent
of the flank. In T. transitoria the rows of nodes representing
the marginal and median carinae become joined by transverse rounded
ridges at an earlier growth-stage than in T. herzogi, in which these
rows of nodes, and that representing the inner carina, remain
isolated and distinctly developed until traced farther back towards
the siphonal margin of a fully grown individual. The escutcheon
is smooth in its anterior part in T. transitoria, while ornamented at
an early stage in the African shell; in T. transitoria the escutcheon,
generally speaking, is more sparsely sculptured.

I have elsewhere pointed out the features of distinction between
T. herzogi and T. mamillata, from the Oomia beds of dutch.*
T. mamillata shares the principal characters of thePseudo-quadratae,
but is a shorter shell than T. Jierzogi, and is vertically truncated
in front. In T. mamillata those ribs situated at the centre of the
flank, and posteriorly to this, are slightly curved, and are directed
anteriorly at their lower ends, while the corresponding ribs of
T. lierzocji are backwardly directed and are not curved. On the
area of T. mamillata, the rows of more or less isolated nodes become
united to form smooth transverse ribs nearer to the umbo than in
the African form, in which, also, the lines of nodes and transverse
ridges of the area are rather more closely spaced. In T. mamillata
the tubercles are more closely crowded together in the ribs of the
flank and are, generally speaking, more robust and conspicuous than in
T. herzogi, particularly in relation to the size of the valve. The blunt
transverse ridges of the area towards the posterior end of an adult
specimen are broader and more strongly developed than in T. herzogi.

A comparison with Trigonia holubi is given in the remarks which
follow the description of that form.

Two specimens depicted in the left-hand side of a text-figure
published by Drs. Hatch and Corstorphine f represent T. herzogi,
but very greatly reduced in size.

TRIGONIA HOLUBI sp. nov.
Plate IV., figs. 2, 2<y.

Description. The shell is large and massive, almost oblong in
outline, and usually vertically truncated in front, or with very steep
* Kitchin (1), p. 102. f Hatch and Corstorphine (1), p. 245, fig. 66.

104 Annals of the South African Museum.

forward obliquity ; the length is greater than the height. The nearly
terminal umbones are little conspicuous and weakly incurved. The
cardinal margin forms an almost straight line and slopes back very
gently from the umbo. It forms posteriorly a sub-angular junction
with the relatively long, slightly convex siphonal margin. The
frontal margin is very slightly convex, more seldom quite straight,
in outline, and passes below by a sharp curve or sub-angular junction
into the lower border; this is more convex in profile. The inflation
of the valves is relatively weak ; the greatest height occurs at about
the middle of the valve. A flattened frontal face is well developed
in fully grown specimens. There is no definite carinal angle, but the
area is well demarcated by the limits of its particular sculpture, which
is contrasted with that of the flank.

The sculpture of the flank in the young shell consists of relatively
strong concentric nodular ribs, four of which are already developed
when the shell has attained a height of 10 mm. Subsequently formed
ribs increase rapidly in prominence and robustness and become
steeply inclined, so that after about ten ribs are developed all reach
the inferior margin. The ribs of the adult shell consist of rows of
large and prominent tubercles, some of which reach a diameter
of 5 mm. Most of the ribs have a forward sweep when traced
downwards ; an approximately vertical direction is attained only by
those ribs situated in the posterior half of a fully grown valve, and
these also are slightly curved in form, with the convex side directed
posteriorly. The interspaces are rather narrower than the tubercles
of the ribs. In the early and middle adult stages, the tubercles com-
posing the ribs are well separated from one another on a given rib.
In the late adult and senile stages the tubercles become closely
crowded together and are contiguous. Towards the lower border of
a fully grown individual the tubercles tend to become elongated in a
direction parallel to the shell margin, and they may appear to coalesce
with prominent ridges of growth. With senility the ribs become im-
perfectly developed, while crowded ridges and furrows of growth are
more marked near the pallial margin. In the lower half of the
anterior portion of the flank in an adult specimen some tubercles
are somewhat unevenly dispersed, giving rise to an appearance of
irregularity, with departure from the strictly linear arrangement.
In a fully grown specimen about sixteen ribs are developed on the
flank.

The area in the neanic stage is crossed by transverse ribs con-
tinuous with those of the flank. In the early and middle adult
stages no ribs are developed upon it ; but a line of tubercles limits

The Invertebrate Fauna of the Uitcnhaye Series. 105

the area above, a second row of less well-defined tubercles is situated
just below the longitudinal groove, while a third row of well-spaced
stronger tubercles limits the area below. Prominent transverse ridges
or irregular ribs cross the area in the posterior half of an adult
individual, terminating above and below in the marginal tubercles
and also between them. These ridges make their first appearance
somewhat abruptly, and become more strongly marked and also
more irregular in their form and spacing towards the posterior end
of the area, in a well-grown individual. A few may even pass over
to the flank and be there partly identified with ridges of growth.
The longitudinal groove of the area, which divides it into a
narrower, superior, and broader inferior portion, forms a constriction
in each transverse ridge.

The escutcheon is narrow and relatively very elongated. It is
ornamented by irregular rows of coarse tubercles or prominent ridges
which pass very obliquely across its surface and terminate anteriorly
at the cardinal margin. They are seen to be frequently continuous
with the ornaments of the area. The ligament pit is of narrow and
elongated lanceolate form, and in a large individual extends to half
the length of the escutcheon.

The central tooth of the left valve is very massive and prominent,
with relatively narrow apical angle. The anterior tooth of the right
valve is supported by a well-raised platform. At the posterior end
of the valves the raised ridge on the internal surface which separates
the siphonal channels is situated high up in the siphonal border
and is of relatively great strength and extent ; in a fully grown
individual it can be traced for at least 25 mm. from the margin.
Dimensions.- (1) (2) (3) (4)

Length (about) 98 . 110 . 120 . 132mm.

Greatest height 75 . 78 . 87 . 88

Depth of a single valve 18 . 20 . 22 . 25

Occurrence. Specimens in the collection of the South African
Museum came from the Sunday's Eiver (285 to 288, 290 to 296, 299).
Mr. Rogers collected this form on the left side of Coega Valley, half
a mile down from the railway (458g). An immature specimen from
Brentford, Knysna Estuary (150h), is also probably referable to
T. holubi.

Remarks. The characters of this shell appear to be on the whole
very well defined, while the valves are especially conspicuous both
by their large dimensions and their salient ornamentation. Most of
the specimens submitted to me, in all of which the shell is replaced
by coarsely crystalline calcite, have had the surface considerably

106 Annals of the South African Museum.

corroded by weathering. In these specimens the etching has attacked
the shell along lines of weakness, and has brought the tubercles and
ornamenting ridges into undue relief, which results in an appearance
so deceptive that for some time I was in doubt whether these etched
specimens should not be regarded as a distinct form. A careful
comparison between these individuals and others which have not
suffered from weathering shows that they agree in all essential
characters and that a separation cannot on present evidence be
established. In addition to differences due to preservation, there is
some variation in the sculpture of the area and escutcheon which
may be noted here. The transverse ornaments of the area are in
some individuals more robustly developed than in others ; on the
escutcheon the linear coalescence of tubercles to form parallel,
obliquely running ridges is more complete and regular in some
specimens than in others, and the ridges replace the isolated
tubercles at an earlier stage of growth. There is also some variation
in the arrangement of the tubercles in the lower half of the flank of
adult specimens near the frontal margin. The lower terminations
of the ribs here tend to turn forwards along the more prominent
ridges of growth ; some of the tubercles may appear to blend with
these ridges, which turn up sharply at the frontal face. Another
point in which some variation is shown is in the form of the anterior
profile of the shell. In some specimens there is vertical truncation
in front, in others the outline of the anterior margin slopes forward
somewhat, when traced down from the umbones, so that the foremost
point of the shell is situated near to the junction of the inferior and
anterior margins.

This is the form referred to by Sharpe as a Triyonia allied to
T. lierzogi ; he gave the following description of specimens which
were presented by J. S. Bowerbank to the Geological Society of
London: " Trigonia sp. ('?). This variety (or possibly distinct
species) is closely allied to Trigonia licr.-o<ji, but is more quadrate
in outline than the common variety, the anterior edge being
truncate ; and the costal rows of knobs turn forward as they
approach the ventral border, instead of passing downwards and
backwards. These specimens came from Algoa Bay, and apparently
have been derived from the Sunday River district, from the aspect
of their matrix."

T. holubi clearly cannot be united with either the Clavellatae or
the Quadratae, but its characters are those of the somewhat inter-
mediate kind which led Steinmann to establish the group Pseudo-

* Sharpe (1), p. 202.

TJie Invertebrate Fauna of the Uitenhage Series. 107

quadra tee, to receive Trigonia lierzogi (Goldf.), the well-known
Uitenhage form, and T. transitoria Steinmann, from Lower
Cretaceous strata in Chili, Bolivia, and the Argentine Republic.*
T. neuquensis Burckhardt.i which is found associated with T. transi-
toria in the Neocornian Trigonia-be&s of Las Lajas (Eio Agrio,
Argentine Eepublic), also falls within this division, and another
example of the same group is T. mamillata, which occurs in the
Oomia Trigonia-beAs near Goonaree in Cutch.j These five members
of the Pseudo-quadratae show similar deviation from the characters of
the two sections with which they may best be compared. Like the
Quadratae, they have the escutcheon well ornamented, and the
sculpture of the area is in some respects very similar, especially in
the manner in which transverse ridges, particularly in the late adult
stage, may pass over to the escutcheon on the one hand and the
flank on the other. These characters contribute largely to distin-
guish the group from the Clavellatse, with which, both as regards
the youthful ornamentation and the adult form, it has much in
common. In the Quadratae the area is usually divided by its longi-
tudinal groove into a broader, superior, and narrower inferior
portion ; in the Pseudo-quadratae the upper division is the narrower.
A row of impressions or pits on the inner valve surface, near the
pallial margin towards the posterior end, is present in the Quadratae
but absent in the Pseudo-quadratas. In accordance with the position
of the longitudinal groove of the area, the lower siphonal channel on
the inner valve surface is relatively broad in the Pseudo-quadratae,
and may be almost twice as broad as the upper one.

T. holubi is most closely comparable with the Indian T. mamillata,
to which it shows a striking similarity. In T. holubi, however, the
valve has a rather less convex form, the longitudinal groove of
the area is more strongly marked, and the tubercles of the ribs
in the early and middle adult stages are much less closely spaced.
The most important point of distinction lies in the development of
the sculpture of the area. In T. mamillata the phase in which
irregular transverse costas crossing the area form a dominant feature
is reached at a much earlier stage of growth than in T. holubi.
Thus, in T. mamillata, strongly developed costae appear on the area
soon after the neanic stage is passed and long before half the adult
dimensions are attained. In T. holubi, save for the longitudinal
groove and its accompanying line of tubercles, the surface of the
area is smooth, and only marked by lines of growth, until at least

* Steinmann (2). f Burckhanlt (2), p. 74, Taf. xiv., figs. 4-6.

I Kitchin (1), p. 100, pi. ix., tigs. 8, 9; pi. x., figs. 1-3.

108 Annals of tJic SoutJi African Museum.

half the adult dimensions are attained. The transverse ribs which
are subsequently developed on the area are not so well defined or so
robust as in T. mamillata,

From Trigonia lierzoyi (Goldf.),* T. holnbi is readily distinguished
by the considerably shorter form, the vertical anterior truncation,
the more crowded and much more prominent character of the flank
ornaments, and the later appearance of transverse sculpture on the
area. In the middle of the flank of an adult specimen, the ribs of
T. licrzoiji are straight, while those of T. lioliibi are curved. The
backward slope of the posterior flank ribs in T. licrzorji is another
point of distinction.

There is a marked general similarity to Triamu'a transitoria
Steinmann,! from South America, but the ornamentation differs
considerably. In T. transitoria the tubercles of the costse on the
flank are placed closely against one another through all adult stages
instead of being openly spaced as in the ribs of T. liolubi. In T. tran-
sitoria also, the development of transverse ridges on the area comes
in at a much earlier stage of growth, and, in fact, this ornamentation
is a marked character during a large part of the adult life. There
appear to be other points of difference which it is unnecessary to
enumerate.

T. nciiqucnsis Burckhardt is at once readily distinguished from
T. liolubi by its much shorter and higher outline. Here, also, the
transverse ridges of the area become a marked feature at an earlier
growth-stage than in T. liolubi. In T. neuquensis, the manner in
which these ridges in the posterior part of the shell extend down-
ward on to the flank is much more marked than in the most advanced
stage in T. liolubi.

Note on the Development of tlie Pseudo-quadrate Trigoniae.

A strong contrast between the features of the neanic and adult
stages may be observed very frequently in species belonging to
various sections of this widely conceived genus, but it is seldom that
a transition in characters so marked and abrupt as that shown in
T. liolubi is seen to occur during the adult period. The sudden
acquirement of strong transverse ridges on the area, a character
which is accompanied by a change in the manner of development

* Goldfuss (1), Band ii., p. 202, Tab. 137, fig. 5 (1837).

t Steinmann (1), p. 260, Taf. iii., fig. 3; Steinmann (2), p. 221, Taf. vii.,
figs. 3, 4, Taf. viii., tigs. 1-3; Burckhardt (1), p. 21, pi. xxv., figs. 1-3;
Burckhardt (2). p. 73, Taf. xiv., figs. 1, 2.

The Invertebrate Fauna of the Uitenhage Scries. 109

of the tubercles of the flank costte they become crowded closely
together on a given rib is a very striking feature. In T. mainillata
from Cutch, these characters are also coincident, but they are
acquired at a much earlier period of growth. The members of the
Pseudo-quadratie, in the three continents where they are known
to occur, seem to have been destined to pass through a similar
sequence of developmental phases, but these phases were not
reached by all the species at the same time. Thus, T. mamillata
is in advance of T. Jioliibi. In T. lioltibi, the stage where the area
is demarcated by an upper and lower marginal row of tubercles and
marked by a median longitudinal row, persists until half the adult
dimensions have been reached. In T. mainillata, this trituberculate
stage is passed over rapidly and is superseded in very early adult
life. In T. hcrzoyi it is superseded by the transversely costate stage
(the area alone is still referred to) at a somewhat later period of
development, but still not so late as in T. liolubi. A point of great
interest is that in T. mamillata and T. herzofji, and also in
T. transitoria, the trituberculate stage may be seen to pass into
the transversely costate stage by the progressive transverse elonga-
tion of the tubercles in the three sets, and the gradual coalescence
of the transverse ridges thus formed. In T. liolnbi, this inter-
mediate phase of development is not so noticeable : it is very much
suppressed or is wholly omitted.

The facts here briefly set forth seem to suggest that the group
Pseudo-quadratie may be of an artificial character to some extent
that the forms here included for convenience of classification are not
strictly homogenetic ; and many known parallel instances amongst
molluscs and brachiopods give great probability to the truth of this
idea. We must suppose all these forms to have been descended from
true Clavellatae, just as in the case of the Quadrates of Europe, and
it is a very striking circumstance that the characters which mark
the Pseudo-qiiadratae should appear approximately at the same
geological moment of time in three widely remote continents.
Yet these forms are unknown in the European area. The Post-
jurassic development shown by the Quadratas and Pseudo-quadratge,
each group representing offshoots from one or more clavellate stocks,
took place along parallel though independent lines in the European
area and in the southern development with which we are dealing.
Within each area, again, the development in the supposed several
genetic series (which were departing along the same general lines
fiom the ancestral clavellate characters) may be supposed to have
been in some degree independent.

110 Annals of the South African Museum.

Confining our attention to the Pseudo-quadratae, it is difficult to
believe that the salient characters by which all the members are
distinguished could have been called forth by the ordinary process
of natural selection alone. The acquirement of costation on the
area coincident in direction with lines of accretion ; the encroach-
ment of this simple ornament from the area to the transcrescently
sculptured flank in the late adult and senile stages ; the prodigal
expenditure of shell substance in producing massive sculpture of an
increasingly irregular type in these same growth-stages; and the
evident variation in many characters which indicates a certain
instability of type ; all these point to racial degeneracy. A com-
parison of the Pseudo-quadratse from India, South Africa, and South
America, shows that in all of them this degeneracy was expressed
as the culmination of a definite sequence of developmental phases.
The facts do not warrant the supposition that the members of
the group were wholly independent of one another. Their simi-
larity in leading characters was perhaps due, in varying degree, to
a number of causes : a certain community of ancestry ; continued
intercourse until checked by divergence ; and at the same time the
constant influence of natural selection ; all these may have played
a part. But it seems difficult to escape from the belief that innate
racial tendencies of a very definite kind found their expression in
representatives of these Trigonia-stocks at the same geological
period. Had the members of these genetic series been more
passive, so to speak ; had they been more plastic and more com-
pletely amenable to the call of natural selection, it is scarcely con-
ceivable that environmental conditions should not have occurred
at some earlier time which would be capable of producing closely
similar results. But the advent of the Quadratic in Europe, and of
the comparable though independent Pseudo-quadratse of the south,
took place, so far' as we know, invariably in Neocomian times.

TKIGONIA VAU Sharpe.
Plate "VI., figs. 1, la, 2, 2a, 26, 3.

1856. Tru/oii/a ran D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

vol. vii., p. 194, pi. xxii., fig. 5.

? 1867. Triijonia van, E. Tate, Quart. Journ. Geol. Soc., vol. xxiii.,
pi. vii., fig. 8.

Occurrence. The record of occurrence given by Sharpe was
" Sunday Eiver, in greenish-grey grit, with fragments of wood
and shells ; and in a shelly grit at the Zwartkop Eiver." Speci-

The Invertebrate Fauna of tlic Uitenhage Series. Ill

mens preserved in the collection of the Geological Society of
London are labelled " Sunday Eiver " (Atherstone and Bain),
"Prince Alfred's Eest " (Eubidge), and " Zwartkop Eiver"
(Eubidge). Specimens sent to me from the South African
Museum are from the Sunday's Eiver. Messrs. Eogers and
Schwarz record T. vau from the following localities : in the neigh-
bourhood of Uitenhage, ascending the Grass Eidge road, in the
uppermost of three beds of sandstone exposed in the dry water-
courses, and in the railway cutting between milestones 24^-24f on
the Graaff-Eeinet line; on Sunday's Eiver, in the upper beds exposed
in a white krantz on Wolve Kraal, on the north bank of the river.
In 1905 Mr. Eogers collected specimens 300 yards below Addo
Drift (Tunbridge's), left bank of Sunday's Eiver (40h, 41h, 42h).
An immature specimen (46h) from the same locality, may also
belong to Trigonia vau.

Remarks. A large specimen of T. ran sent to me from the South
African Museum attains a height of 50 mm., measured from the
umbo. The ribbing of the flank ceases to be developed at a distance
of nearly 10 mm. from the inferior margin, in the neighbourhood
of which the shell surface is marked only by ridges and furrows of
growth.

Sharpe gave a very accurate and characteristic figure of this
remarkable form, but the differences which distinguish T. vau from
T. stowi appear up to the present to have escaped observation, and
it is probable that the shells cited under the name T. vau by Stow" ::
are in reality to be ascribed to T. stowi sp. nov. I have had the
opportunity of examining and comparing a considerable number
of excellently preserved specimens of these two Triyonice, and find
that the characters which distinguish them from one another are
very constant. Several fully- grown shells of T. vau sent to me from
South Africa agree very perfectly with the original type and other
individuals in the collection of the Geological Society. The points
of distinction which may be readily observed in the adult shells of
T. vau and T. stowi are set forth in the remarks which follow the
description of the latter. During the youthful growth-stage, how-
ever, these two forms cannot be separated, so that it is uncertain
to which must be ascribed a young specimen obtained by Messrs.
Eogers and Schwarz at Walton's Farm, just below Dunbrodie on
the Sunday's Eiver (307), and another individual measuring 17 mm.
in length, collected between milestones 24i-24f on the Graaff-Eeinet
railway (309). There is the same doubt concerning an immature

* Stow (1), pp. 199-505.

112 Annals of the South African Museum.

specimen from Addo Drift (46b). For the same reason it is uncer-
tain whether the young individual figured by Tate really represents
the true T. ran. Tate's figure, which represents the specimen in
twice the natural size, unfortunately does not give a very satisfactory
picture of the youthful characters of either of these shells.

The close agreement in the characters of the neanic stage reveals
the near relationship of these two forms, and, indeed, such a rela-
tionship might reasonably be inferred from a comparison of adult
characters, which coincide in a remarkable manner as regards the
ribbing of the flank and the posterior elongation. Other shells,
however, which when adult exhibit similar peculiarities of form
and ornamentation, differ widely from T. rail, and its associate
when in the youthful stage. These are the members of the
Triijonia v-scripta group in the marine Ooniia beds of Cutch,*
and a comparison of their youthful characters and those of T. vau
and T. stowi is given in the remarks which are appended to the
description of the latter. T. stowi when adult approaches closely
in many particulars to T. v-scripta itself, but the adult T. vau agrees
much more closely with T. dubia Kitchin, so far as the unsatis-
factorily preserved Indian specimens allow of comparison. In
consequence of imperfect preservation, the nature of the sculpture
in the youthful stage of T. dubia is not known, but it appears
reasonable to assume that it bears closer relationship to the other
members of the group of T. r-xcripta than to T. vau. In the adult
stages, too, points of distinction between T. dubia and T. van are
not wanting. These have been set forth in my account of the
characters of T. dubia f as follows : " In T. vau, the ribs of
the anterior series, though narrow and crowded as in T. dubia,
are seen to be upwardly inclined as they depart from their point
of meeting with the posterior series, and are crossed obliquely by
the lines of growth. In T. dubia, on the other hand, these anterior
costae are placed in a direction quite, or almost, parallel to the
inferior border, and the angle of the lateral V is therefore not
so acute. Moreover, before individuals of T. vau have reached
half their full dimensions, the costaa of the frontal series cease to
be formed close to the frontal border. There arises, therefore, a
space devoid of ornamenting ribs, marked only by ridges and
furrows of growth, and bounded posteriorly by the last-formed
obliquely directed costse of the frontal series. This peculiar
character is totally absent in T. dubia, in no specimen of which
is a frontal unsculptured space seen, although in respect to size
* Kitchin (1), p. 65. t Ibid., P 69.

The Invertebrate Fauna of the Uitenhac/e Series. 113

all the individuals examined considerably exceed the dimensions at
which this feature becomes noticeable in T. van. . . . Sharpe's type
is of larger dimensions than the known specimens of T. Jitbia.'

The extent to which an unsculptured space near the lower part of
the frontal margin may be present in T. ran is really somewhat
variable, but a few of the lowest ribs of the anterior series seem
always to fail to extend at their upper terminations to the valve
margin. In some individuals, two or three of these lower ribs may
be slightly bent down at their anterior ends in manner reminiscent
of the strong, angular bending at the front end of these ribs in
T. stowi, as well as in the Indian T. v-scripta. It may also be
noted that although the inclination of the frontal series of costae in
T. van is always as steep as in the specimen depicted in Sharpe's
figure, yet these ribs may in some instances attain a stronger
development and may be of a rather more robust character.

A shell possibly related to T. van is T. kiihni G. Miiller,* from
Neocomian strata at a locality 23 km. west-south-west of Mtshinga
in German East Africa. To judge from the description and figures,
this is distinguished by an angular rib arrangement very similar to
that exhibited by the members of the group of T. v-scripta in India,
and T. van and T. stowi in South Africa. T. kiihni seems to differ,
however, by its less equilateral form and the persistent ornamenta-
tion of its area. The figure of an imperfect specimen of T. kiihni
(fig. 8) shows a curved frontal profile and the ribs of the frontal
series obliquely crossing the growth-lines in manner that recalls
the same characters in T. van, though still more reminiscent
of T. rccurva Kitchin, from the Oornia beds. It is as yet not
possible to say, however, whether T. kiihni is more nearly related to
T. van or to the group of T. v-scripta, though it may be anticipated
that a study of the youthful characters will ultimately throw light on
this point.

Trigonia hetcroscidpta Stanton,t from the Belgrano beds (Lower
Cretaceous) of Patagonia, is very probably closely related to T. van,
but differs by the much shorter and more elevated triangular figure.
The first-formed ribs of the posterior series are vertical or backwardly
inclined, and not forwardly sloped as in T. van, while several
successive anterior ribs terminate abruptly on the flank of a single
vertical posterior rib. In the neanic stage, however, the plan of
sculpture as well as the form of the valve appears to agree closely
with that of T. ran.

* G. Miiller (1), p. 501, Taf. xxv., figs. 0-8.
f Stanton (3), p. 20, pi. iv., figs. 16-18.

114 Annals of tlic South African Museum.

E. A. Philippi* has figured some Chilian forms (for example,
T. arsinoe Philippi, T. foveata Philippi) which may be related to
T. ran; they are characterised by great posterior elongation, very
convex anterior profile, and two series of ribs on the flank meeting
at an angle. The general aspect, indeed, at once suggests the
probability of relationship with the two South African types, but the
Chilian shells are so unsatisfactorily figured, and their preservation
appears to have been so imperfect that further and critical com-
parison is not possible.

Whiteaves has drawn attention to the features wherein Trigonia
divers icostata Whiteaves,! from the Cretaceous of Queen Charlotte
Islands, bears resemblance to T. van. These are: the elongated and
posteriorly strongly produced figure of the shell, the convex anterior
profile and recurved umbonal region, and the general plan of
sculpture on the flank. In T. diver sicostata, however, the anterior
rilis are much coarser in character than in T. van, and are directed
almost parallel to the lower valve margin, thus forming a right angle
with the posterior ribs instead of an acute angle. A good distinctive
feature is seen in the area of the Canadian shell, which is coarsely
ornamented by strong longitudinal ribs, and this alone is sufficient
to indicate that the two forms are in no way nearly related, but that
the common characters of shape and broad plan of flank sculp-
ture have been quite independently attained. Whiteaves ascribes
T. diver sicostata to the section Scaphoidese, but all members of this
division bear evidence of a clavellate ancestry, and longitudinal
ornamenting ridges on the area are unknown amongst them. Such
ornaments are essentially characteristic of the Costatae, from which
T. dicerxirostata was quite probably derived, and rapid divergence
from the typical pattern of the section might well result in this
aberrant form, in manner analogous to that exhibited in some of the
modified Costatse of the marine Oomia beds in Cutch. T. van
shows no signs of a connection with the section Costatse at any
early growth-stage that can be studied, but the ornaments of the
area are transverse from the first.

The peculiarly ornamented Trigonia doroscJiini Eichwald,| from
the Neocornian of Tukusitnu Bay (Alaska), may finally be brought
into comparison. This appears to me, however, to be well separated
from the group of T. van (as well as from the group of T. v-scripta
in India) by the more truncated and less convex anterior margin and
by the manner in which the frontal ribs are directed at right angles

* K. A. Philippi (1), pis. 34, 35. f Whiteaves (1), p. 68, pi. x., fig. 1.

I Eichwald (2), p. 180, pi. xiii., figs 12-14; xiv., figs. 1-4.

The Invertebrate Fauna of the Uiteuhage Series. 115

to the frontal urn-gin. It seems doubtful from Eichwald's figures
whether a single form alone is represented, for it is difficult to
believe that the originals of pi. xiii., fig. 12, pi. xiv., fig. 1, and
pi. xiv., fig. 3 can belong to one species. Other points in which
these differ from T. van are the great extent of the frontal ribs, the
horizontal position of these, and the persistent transverse ornaments
on the area. Eichwald's description and figures leave the affinities
of T. doroschini obscure, but I think it improbable that it is related
to the group of T. vau.

TRIGONIA STOWI sp. nov.
Plate VI., figs. 4, 4a, 46, 5 ; VII., fig. 1.

Description. The shell is of very elongated form, moderately
inflated, anteriorly pointed, posteriorly very much produced, and
slightly gaping at the siphoual margin. The umbones are situated
at some point lying between one-quarter and one-third of the
shell's total length from the anterior extremity ; they are well
incurved and slightly recurved. The umbonal region is fairly
prominent and inflated. Posteriorly to the umbo, the very long
cardinal margin forms a straight or very gently concave outline and
passes posteriorly by a curve into the short, convex siphonal margin.
Anteriorly to the umbo, the valve margin slopes downwards with
straight profile to the projecting anterior extremity, where it forms a
sub-angular or sharply curved junction with the lower margin ; the
latter forms a gentle and evenly convex profile as it is traced between
the anterior and posterior extremities of the shell.

At no stage of growth is a marginal carina developed ; in the
youthful stage its place is represented by a blunt carinal fold which
passes with advancing growth into a still broader and less well
defined rounded fold of the valve. At the siphonal end of an adult
individual this loses definition and the valve becomes evenly convex.
In the youthful shell, until the valve attains a height of about
8 mm. measured from the umbonal apex, the ornamentation
consists of numerous, crowded linear ribs which are parallel to the
lower margin and extend from the frontal border across the flank ;
they pass over the carinal angle and across the area. Up to a
distance of between 5 mm. and 10 mm. from the umbonal apex these
ribs extend across the escutcheon also, and terminate either at the
cardinal margin or just before reaching it. Subsequently they cross
the area though not the escutcheon, until a distance of about 10 or
12 mm. from the umbonal apex is reached, after which they occupy
the flank only, terminating at the carinal angle, while the area subse-

116 Annals of the South African Museum.

quently remains smooth. In the space between 10 mm. and 20 mm.
from the umbonal apex, the very blunt ridge demarcating the area
from the escutcheon is ornamented by minute, transversely elongated
nodes or short raised lines, each measuring about one millimetre in
length. At a distance 'of about 10 mm. below the umbo the ribs on
the flank become bent down in the middle in angular form and
become more widely spaced. The transition from the straight to
the angularly bent flank-ribs is somewhat sudden. Three or four
bent ribs are successively produced, having a more delicate anterior
and more robust posterior limb, and an increasing acuteness of the
angle. Subsequently, an anterior and a posterior series of inclined
ribs may be spoken of, not strictly coinciding with each other in
number. The anterior ribs, numbering about 14-16 in the adult,
often cease to extend to the frontal margin after the shell has attained
half its adult dimensions ; their upper, anterior terminations are
situated on the flank at an increasing distance from the margin, and
forming acute angles with these, with upwardly directed apices, are
several short, steeply inclined, weak and rather broader ribs. Each
of these is only a few millimetres in length and might be considered
to represent the downwardly bent anterior termination of a rib of
the anterior series. The most forward portion of the flank is usually
devoid of sculpture and marked only by ridges and furrows of
accretion. The lowest ribs of the anterior series become in
varying degree irregularly nodose or broken into wavy lines of nodes
with loss of regularity near the lower valve margin. The ribs of the
posterior series have their upper termination at first close to the
carinal fold, and successively at a gradually increasing distance
from this, so that a narrow portion of the flank adjacent to the fold
is smooth. These ribs are not developed to the posterior end of the
flank in the adult, but there is here a smooth tract only marked by
lines of growth. The ribs of the posterior series, developed to the
number of about 14, are very steeply inclined, and all have a slight
forward slope. They are of rounded form, are about 3 mm. in
breadth, and have slightly narrower interspaces.

The area is relatively narrow, and is marked throughout the adult
period by a longitudinal linear depression. In the posterior half of
the area this marks off a narrow superior and broader inferior
portion. There is no defined inner carina, and the line of delicate
and imperfectly developed nodes which marks the inner limit of the
area at a distance of 10-20 mm. from the umbonal apex is a
transitory feature, and does not persist with subsequent growth.
The escutcheon is relatively very long and is of lanceolate form, and

The Invertclirati 1 Fauna of the Uitcnhagc Scries. 117

though imperfectly demarcated from the area, it is marked by its
excavated form and smooth surface. Bidges of growth which cross
the area are replaced by very delicate and scarcely perceptible lines
of accretion on the escutcheon. At a distance of 15 mm. from the
um bones the escutcheon in each valve is fully as broad as the area.
The ligament pit is relatively short and broad.

Dimensions. (1) (2) (3)

Greatest length 80 . 95 110mm.

Height, measured from the umbo 42 . 44 50 ,,

Greatest depth of a single valve ... 14 . 15 . 20 ,,

Occurrence. Marine Beds of Sunday's and Zwartkop's Eivers.
Specimens in the collection of the Geological Society of London are
labelled " Zwartkop Elver" (Eubidge) ; " McLoughlin's, bed No. 5 "
(Stow) ; " Above Modder Drift, No. 3 bed " (Stow).- In the British
Museum (Natural History) a fine specimen is labelled " Sundays
Elver, Pont." A specimen sent to me from the South African
Museum is believed to come from the Sunday's River cliffs. An
immature specimen, 15 mm. in length, and a portion of the flank of
an adult in the form of an external mould, resembling T. stowi, were
collected by Messrs. Rogers and Schwarz at Walton's Farm on the
Sunday's River, just below Dunbrodie (307). These may probably
represent T. stowi, though they might equally well be ascribed to
T. rait, so far as their characters allow of determination. Another
immature specimen, from the railway cutting between milestones
24A 24f on the Uitenhage-Graaff-Reinet railway (309) may also
perhaps belong to this form. Mr. Rogers collected T. stowi in a
small kloof, three miles up the left bank of Sunday's River (17h).
A hand-specimen containing one complete and two fragmentary
valves was also obtained by him from a cliff W. 20 S. from Comley's
house, right bank of Sunday's River (90h).

Remarks. This form shows some variation in regard to the shape
of the frontal profile and the position of the umbones in relation to
the anterior extremity. In some examples, the anterior sculpture is
more robust and more regular than in others. The thickened and
downwardly directed terminations of the anterior ribs are some-
times a very prominent feature, in other cases they are less
noticeable. In some individuals the anterior ribs are very
closely spaced, and towards the lower part of the adult valve may
be much broken up into nodes (as exemplified by specimen 17h).
Sometimes a small portion of the flank, most anteriorly situated, is

* See Stow's table of cliff-exposures at localities on " Upper Sundays Kiver " and
" Lower Sundays River " ; Stow (1), tig. 3.

9

118 Annals of the South African Museum.

smooth, in other cases the ribbing extends to the frontal margin
at all parts. Near the pallial margin of large individuals senile
characters may be seen in the cessation of sculpture and the
appearance of several strong furrows of accretion.

Two right valves in the specimen numbered 90h have a more
rounded and less sharply pointed anterior profile than any of the
other specimens studied. They also have the urn bones rather more
anteriorly placed. The other extreme, illustrating the less inequi-
lateral form of the valves and the more marked pointing of the
anterior profile, is shown by an individual from the collection of
the South African Museum. Characters quite intermediate between
these extremes are exhibited, for example, by a specimen numbered
12006 in the collection of the Geological Society of London. I
believe, therefore, that the material examined only suffices for the
establishment of a single species.

Stow appears to have believed this form _ to represent T. van
Sharpe. \Vhen dealing with the Triyonia of the Oomia beds of
Cutch, I made reference to T. stoivi as an undescribed form allied to
T. van.- 1 ' In addition to the specimens there referred to, I have
since had the opportunity of examining further material in the col-
lections of the Geological Society and the British Museum (Natural
History), and also the fine specimens sent to me from South Africa.
T. stowi is a very well characterised form, peculiar for its elongated
outline, its pointed anterior extremity and greatly produced posterior
region, its striking flank-sculpture and its siphonal gape. Although
in many points, and particularly in the sculpture of the adult, it very
strongly recalls the Oomia T. v-scripta Kitchin, yet its youthful
characters plainly reveal its close relationship to T. van Sharpe (see
above). Until the young shell has attained a height of about 10 mm.,
these two African forms are so alike that I have been unable to find
any feature by which they may with certainty be distinguished. In
the adult stage, however, T. stoivi has a more elongated outline and
is more strongly produced posteriorly ; its umbones are relatively
further removed from the anterior extremity ; the upper and lower
valve-margins converge towards the front to join in more or less
pointed form, and produce a frontal profile which stands in contrast
to the sweeping convex outline of the frontal margin in T. vau. The
angularly bent and swollen anterior portions of the ribs in the frontal
series, which produce such a peculiar pattern in the sculpture of the
anterior quarter of the flank in T. stoivi, are developed to a much

Kitchin (1), pp. 66, 74.

The Invertebrate Fauna of the Uitenhage Series. 119

less noticeable extent in T. van,, and the bending is absent in some
individuals.

When studied in connection with the shells of the group of
Trigonia v-scripta from the Oomia beds of Cutch, T. stoioi can only
be closely compared with T. v-scripta itself. Though, when adult,
it resembles this in most striking manner in the backward position
of the umbones, the posterior elongation and general form, and
especially in the character of the flank ornamentation, T. stoioi may
be readily distinguished by its relatively more elongated and poste-
riorly attenuated figure, and its anteriorly more acutely converging
upper and lower margins. T. v-scripta is a shorter shell relatively
to height, particularly in the late adult and fully grown state, while
its valves are somewhat flatter and less inflated. In the youthful
stage the two are widely different in character ; T. v-scripta then has
a very few coarse concentric flank-ribs, which, continuing across
the well-marked carinal angle, pass obliquely forwards over the area
in the form of attenuated thread-like raised lines, and terminate at a
delicate linear ridge which represents an inner carina. Similar
characters distinguish the young T. rccarva, another Oomia form.
This, like T. stoiui, has strong posterior elongation in the adult, and
a smooth tract on the posterior portion of the flank ; but it is well
contrasted by its much smaller size, its rounded convex frontal
profile, its more crowded and much less steeply sloped anterior ribs,
and its imperfectly developed, short posterior ribs.

T. hcterosculpta Stanton,* from the Belgrano beds (Lower Cre-
taceous) of Patagonia, is very probably a related form. Its youthful
characters are closely similar to those of T. stoiui, but the adult shell
is much shorter, more triangular in form, and much less elongated
posteriorly, w T hile there are marked differences in the details of
sculpture.

TKIGONIA CONOCARDIIFORMIS (Krauss).
Plate VII., figs. 2, 2, 25, 3, 4.

1843. Lyriodon conocardiiformis F. Krauss, Amtlicher Bericht liber

die zwanzigste Versammlung der Gesellsch. deutsch.

Naturforscher und Aerzte zu Mainz im Sept., 1842,

p. 130.
1850. Lyrodon conocardiiformis F. Krauss, Nov. Act. Acad. Cass.

Leop. -Carol. Nat. Cur., vol. xxii., pt. 2, p. 454, Tab. 49,

figs. la-Id.

* Stanton (3), p. 20, pi. iv., figs. 16-18.

120 Annals of the South African Museum.

1879. Trigoniaconocardiiformis J. Lycett, Mon. Brit. Foss. Trigoniae,
pp. 210, 211, and woodcuts; p. 230. (Palseontographical
Society.)

1903. Trigonia conocardiiformis C. Burckhardt, Beitriige zur Kennt-
niss der Jura- und Kreideformation der Cordillere. Palae-
ontographica, Band 1., Taf. xiii., figs. 3-5.

Supplementary Descriptive Note. Krauss gave an admirable
detailed description of this very characteristic shell, but I am able to
supplement this by some additional observations, based on the
examination of material more favoxirably preserved than that which
appears to have been at his disposal. There are one or two points,
also, in which the figures accompanying Krauss's description are
rather misleading.

With regard to the ribbing of the flank, it must be noted that this
does not always strictly conform with the simple plan illustrated in
the figures given by Krauss and Lycett. It may be remarked that
if the figures of the German author are imperfect, that given by
Lycett to illustrate the exterior of a left valve is still less character-
istic, although it was intended to supply the deficiencies of the
earlier illustrations.

Several specimens now examined are in such condition as to show
the form of the sculpture in early growth-stages, and two individuals,
in particular, have the ornaments of the nepionic and neanic stages
well preserved. One is an imperfect left valve which measures
9 mm. in length, and would be 7 mm. in height if uninjured at the
pallia! border ; it occurred in a hand-specimen, containing adult
T. conocardiiformis, T. van, and other typical forms, from the
Sunday's River. The other specimen is an immature individual
having both valves in place, imperfect posteriorly, but beautifully
preserved in the umbonal region. This was collected by Mr. Rogers
from a cliff on the right bank of Sunday's River on Commando
Kraal (104h). Other specimens, also, show the ribbing of the neanic
stage, though less perfectly. It is seen that until the valve attained
a height of about 6 mm., measured from the umbonal apex, the
flank was ornamented by simple concentric ribs, running parallel to
the lower margin. Close to the apex the crests of the ribs are
situated at a distance of about 0'5 mm. apart ; at about 5 mm.
below the apex the distance between the ribs has increased to 1 mm.
These concentric ribs extend to the frontal margin. Traced back-
wards, they pass over the flank to a definite carinal ridge, and
becoming more delicate in character, turn sharply forwards and
pass across the relatively narrow area. There is no inner carina

The lucertcbratc Fauna of tlic UitcnJiage Scries. 121

even at this early stage, but the ribs encroach slightly upon the
smooth escutcheon. This is clearly seen in the neanic stage, where
the escutcheon begins to increase rapidly in breadth. Very close to
the umbonal apex, however, it is difficult to observe traces of an
escutcheon, and here the attenuated and delicate linear ribs of the
area pass obliquely forward until almost reaching the cardinal
margin, when they die out. Where they disappear, very close to
the cardinal border, the ribs are extremely attenuated and crowded,
while directed to form an acute angle with the valve-border. The
longitudinal linear groove of the area which becomes so marked a
feature in the adult, is only faintly indicated at the close of the
neanic stage.

As the succeeding growth-stage is entered upon that is, when the
valve exceeds 8 mm. in height the carinal angle rapidly becomes
rounded and blunt, and soon takes the form of a broad fold. The
ribs of the flank are now prominent and robust, and are separated
by interspaces 3 mm. broad. They pass obliquely downwards when
traced forwards. There is some variation shown in the relation of
the anterior ribs to the succeeding posterior ones in the adult stage.
Sometimes the first four or five strong ribs have their posterior
terminations on the flank at successively greater distances from the
carinal fold. Immediately behind these, and forming a separate
series having an almost vertical arrangement, are the numerous and
much more delicate ribs of the posterior series. They have their
superior terminations immediately below the carinal fold. The sixth
or seventh rib of the anterior series is in continuity with the second
or third rib of the posterior series, forming the lower and more
strongly developed portion of it, this lower portion being directed to
form a very obtuse angle with the upper portion. Posteriorly to
this, the ribs are not divisible into two series. In other specimens,
however, the successively formed ribs have their upper terminations
alike close under the carinal fold of the valve, and there is no
tendency to division of the ribs into two series. There is also con-
siderable variation in the strength and number of the ribs. In some
specimens the ribs are more strongly developed and are rather less
numerous than in other individuals of equal size ; sometimes their
more strongly swollen character may be particularly noticeable on
the posterior half of the flank.

The area throughout the adiilt stage is devoid of ornamentation
except the well-marked longitudinal groove. The escutcheon is of
very elongated lanceolate form. It is not separated from the area
by any definite carinal ridge, but is well marked off for the most

122 Annals of the South African Museum.

part by being sunk in concave form. Towards the posterior end it
is scarcely demarcated.

Dimensions. Trigonia conocardiifonnis may attain very large
dimensions. A specimen presented by Atherstone to the Geological
Society of London has a maximum length of 164 mm. This
specimen, which is not quite complete at the lower margin under
the umbonal region, must originally have had a height (measured
at this part) of 95 mm. The ligament groove is 45 mm. long. A
specimen figured by Krauss is 84 mm. in length and 42 mm. in
height (at the umbonal region), while the depth of a single valve,
measured from the figure, is 24 mm. In several individuals
examined by me, the height is somewhat greater than this, in
relation to the length.

Occurrence. This form is found in the Marine Beds at various
localities on the Sunday's and Zw-artkop's Eivers. It was obtained
by Messrs. Rogers and Schwarz from the railway cutting between
milestones 24^-24-J on the line from Uitenhage to Graaff-Reinet
(297, 298) ; also at the white krantz on Wolve Kraal on the north
bank of Sunday's River. The same authors have mentioned the
occurrence of this shell in a conglomeratic bed at Plettenberg's
Bay.* Specimens in the collection of the South African Museum
are from the Sunday's River. In 1905, Mr. Rogers obtained
examples in the cliff on Commando Kraal, right bank of Sunday's
River (104h) ; in the highest beds on Zoet Geneugd, right bank of
Sunday's River (67h) ; on a bare slope W. 30 S. from the middle
of Barkly Bridge, on the farm Olifant's Kop, Sunday's River (21h) ;
and from the Nek S. 33 E. from Comley's house, right bank of
Sunday's River (86h).

Remarks. A very striking feature exhibited by this Trigonia is
the sudden transition of sculptural characters at the close of the
neanic stage. The abrupt manner in which the crowded concentric
ribs give place to coarse, widely spaced, inclined ribs, recalls the
analogous transition in Trigonia vau and T. stowi ; in these, the
youthful stage has similar though more delicate concentric orna-
ments, and these are replaced almost as abruptly by angularly bent,
coarse ribs. The youthful characters of T. conocardiifonnis seem
perhaps to suggest an ancestry similar to that from which T. vau
and its ally were derived. In the young T. conocardiifonnis,
however, the carinal ridge is more clearly defined, and in passing
across this the ribs are sharply bent to form an angle. The early
characters do not seem to indicate alliance with either the Scabrae

* Schwarz (1), pp. 53, 61 ; Kogers and Schwarz (1), p. 5 ; Rogers (1), p. 295.

The Invertebrate Fauna of the Uitenhage Series. 123

or the CostataB ; but in view of the remarkable manner in which
widely divergent types of Trigonia in the Oomia beds of ditch
show, by their neanic characters, their relationship to the Costatae,
it would not have been surprising to obtain evidence of a similar
descent in T. conocardiiformis. The young stages, however, exhibit
no trace of longitudinal ornamentation on the area, and it is difficult
to surmise its true derivation.

T. conocardiiformis, by reason of its peculiar characters of outline
and sculpture, has for long held a somewhat isolated position among
the representatives of the genus. Lycett at first expressed the
belief that it is an abnormal example of the Clavellatae, only
remotely related to other members of the section ; * but he after-
wards modified his opinion and associated this form with the
" crenulated examples of the Scabrte." t Definite indications of
such a relationship, as we have seen, are not to be recognised.

It is a point of great interest that on the South American
continent, an apparently very close ally of T. conocardiiformis has
been found to occur, and in association with two members of the
Pseudo-quadratae. The shells described by Burckhardt \ under
the name " Trigonia aff. conocardiiformis " were collected from
strata ascribed to the Lower Neocomian at Las Lajas (Argentine),
and they certainly bear a very remarkable resemblance to this
South African form. In general figure and outline the similarity
is very striking, while differences in the hinge apparatus of the left
valve are of a minor character. The principal points of distinction
are in the nature of the sculpture of the flank in the adult stage ;
regarding the sculptural plan in the youthful stage of the South
American shell, no information is as yet forthcoming. The tendency
in some few individuals of T. conocardiiformis to show a develop-
ment of two distinct series of ribs on the flank during the early
adult stage is illustrated in more complete and emphasised manner
during the whole adult period in the South American shell. Most
of its anterior ribs, though in continuity with ribs of the posterior
or vertical series, form an obtuse angle with these, and pass across
the flank towards the frontal border in a direction more nearly
horizontal than the downwardly directed anterior portions of the
ribs in T. conocardiiformis. The contrasted aspect of the sculpture
in the two forms is well illustrated in plate xiii. of Dr. Burckhardt's
monograph, where figures of African and South American individuals
are presented side by side.

* Lycett (3), p. 121 (1875). f Ibid., p. 210 (1879).

I Burckhardt (2), p. 72, pi. xiii., figs. 1, 2.

124 Annals of the South African Museum.

Another South American form which is probably also nearly
related, is Trigonia eximia E. A. Philippi," from the Tinguirica
valley in Chili. This represents a type of shell very closely com-
parable with T. conocardiiformis, but the complete differentiation of
the ribs into a posterior and an anterior series is a marked feature
of the adult stage. With regard to this character of the ribbing,
the form described by Burckhardt under the name Trigonia aff.
conocardiiformis may be regarded as illustrating a somewhat inter-
mediate type of sculpture between the two extremes, T. conocardii-
formis and T. eximia. Another probably allied shell has been
recorded by Haupt f from the Neocornian of Loteno, on the Eio
Neuquen, on the east slope of the Argentine Cordillera, under the
name Trigonia cf. eximia Philippi. This is said to differ from
Philippi's type chiefly in having the dividing line between the
anterior and posterior ribbing more obliquely directed. T. eximia
is referred by Haupt to the section Undulatse, but this is an obvious
error. These aberrant forms in South Africa and South America
have in common certain peculiar characters of shape and ornamenta-
tion by which they differ in marked manner from all divisions of
the genus known to occur in the Jurassic rocks. It is not improbable
that we are here dealing with representatives of several parallel
series developed from some common ancestral species or group of
species, showing rapid departure from the ancestral type along
similar lines of development, the successive phases being attained,
however, at an unequal rate. On the other hand, the possibility
that convergence is illustrated is not remote. In the above descrip-
tion of T. conocardiiformis, differences observable in the plan of
sculpture after the close of the neanic stage are ascribed to individual
variation. I believe this to be sufficient to account for such
differences, but the possibility is not excluded that in the Uitenhage
beds, two very closely similar forms, undergoing parallel develop-
ment, are present. It may be that those individuals which show
some division of the ribbing into two distinct sets forming an angle
with one another, at the beginning of the adult stage, illustrate a
series which has passed through a stage similar to that represented
in the adult T. eximia, and that this stage, with angularly disposed
costate ornamentation, has become suppressed by tachygenesis and
reduction. The entire absence of characters of sculpture which
approach those of T. eximia, in the other individuals from Cape
Colony, might probably be due to the same cause.

* E. A. Philippi (1), p. 76, pi. xxxiv., fig. 3.
t Haupt (1), p. 216.

The Invertebrate Fauna of the Uitenhagc Series. 125

TRIGONIA TATEI Neumayr.
1867. Trigonia cassiope E. Tate, Quart. Journ. Geol. Soc., vol. xxiii.,

p. 158.
1881. Trigonia tatei M. Neumayr, in E. Holub and M. Neumayr,

Denkschr. d. k. Akad. Wiss., Math. -Nat. CL, Band xliv.,

p. 275, pi. ii., fig. 3.

Occurrence. A very imperfect and ill-preserved fragment of a
costate Trigonia occurs in bard limestone with Pleiirotni/a baini
from Grass Eidge, three miles east-north-east of Uitenhage (335).
In all probability this represents T. tatei.

Bemarks. Although no complete or satisfactorily determinate
specimens of this well characterised form are included in the collec-
tion under examination, it may be useful to draw attention to one
or two points of interest in reference to the occurrence of this shell,
the only member of the section Costatae hitherto found in the
Uitenhage beds. In 1877, Lycett :;: cast doubt upon the correctness
of Tate's identification of this South African form with the European
Oolitic shells named T. cassiope by d'0rbigny,t and Neumayr sub-
sequently saw the necessity of applying a new name. T. tatei
certainly cannot be united with any other known costate form, and
although the general outline and nature of the ribbed flank has a
close parallel in several familiar European Costatae, yet the narrow
area and elongated escutcheon, the delicate and little-prominent
carime and inter-carinal ridges, and the relatively very delicate
beaded ornamentation of area and escutcheon, are very distinctive
features.

Two specimens preserved in the collection of the Geological
Society of London are labelled " Zwartkop Eiver " (H. Longlands)
and " McLoughlin's Best" (Major Eocke) respectively, and the
former specimen appears to have been the one upon which Tate's
identification was based. The length is relatively great compared
with the height ; the cardinal margin is elongated, while the siphonal
margin is short. The area is slightly convex in form, without a
marked median carina or groove ; the escutcheon is large, and
ornamented by raised lines of granules running parallel to the
ridges of the area. There are about 18 ribs on the flank of
the larger specimen. The elongated form, the very delicate carinae,
the fine intercarinal sculpture, the convex area, and the elongated
granular escutcheon, are all characters which at once recall the

* Lycett (3), p. 172. f d'Orbigny (4), vol i., p. 308.

126 Annals of the South African Museum.

same features in T. tennis, from the Oomia beds in Cutch.* This,
however, is strongly distinguished from T. tatei by the very crowded
and delicate costate ornamentation of the flank, and by the slight
convexity of the valve.

In view of the comparative rarity of the Costatge in the Cretaceous
rocks, the occurrence of so typical a member of the section in the
Uitenhage Series might appear at first sight to lend some support
to the view of those who have maintained that a part, at least, of
these beds must be ascribed to the Upper Jurassic. It is clear,
however, that in any attempt to uphold such a conclusion, less
weight can now be attached to the presence of Costatse than when
our knowledge of the distribution of this division of the Trigonice
was founded principally upon European occurrences. In Europe
itself, representatives of the section are sparsely present in the
Cretaceous, and these mostly show deviations which at once dis-
tinguish them from the typicalJurassic forms. Trigonia peninsularis
Coquand,t from the Aptian of Spain, exhibits a marked degeneration
of characters in its later growth-stages, while T. carinata Agassiz,|
from the Neocornian and Aptian, which may be brought into the
most intimate connection with the Costatae, shows equally far-
reaching modifications, although an examination of Lycett's figured
specimens and other well-preserved individuals fails to reveal any
characters which can be taken to justify Lycett's separation of this
form under a separate sectional heading. The typical features of
the section Costatse are exemplified, however, in a small Trigonia
described by E. Ascher from the Hauterivian of Silesia. ||

Although these European forms may be looked upon as lingering
representatives of a section which had passed its maximum develop-
ment, we must look further afield in order to complete the history of
this strongly characterised and long-lived division of the genus. In
the marine Oornia beds of Cutch, which may now be regarded as
of Lower Cretaceous rather than of Upper Jurassic age, aberrant
derivatives of the CostataB provide a striking feature in the molluscan
fauna ; but here also have been found two representatives, T. tenuis
Kitchin and T. parva Kitchin, which retain in perfect manner the
typical characters of the section. South America has also furnished
another unmodified Cretaceous costate form in T. anguste-costata,
described by Behrendsen *i from strata which he regards as Upper

* Kitchin (1), p. 35, pi. iii., tigs. 5, 6. f Coquand (1), p. 129, pi. xxiii., tig. 3.

+ Agassiz (1), p. 43, Tab. vii., tigs. 7-10 ; Lycett (3), p. 179, pi. xxxv., figs 3-6,
(1877). See also Collot(l).

|| Ascher (1), p. 159 [25], pi. xiii. [ii.], fig. 10. H Behrendsen (1), p. 6, pi. 3, fig. 7.

The Invertebrate Fauna of tlie Uitenhage Series. 127

Cretaceous, at Caryilauhue (Argentine) ; this is accompanied by
a typical member of the Cretaceous section Scabrse, T. transatlantica
Behr. T. anguste-costata is a small triangular shell, which in the
delicate character and close spacing of the flank-ribs recalls the
Indian T. parva Kitchin, from which, however, it is widely
distinguished by the very different outline and the delicate ornamen-
tation of the area. In the perfect development of these longitudinal
ridges of the area, T. anguste-costata retains in typical manner an
essential feature of the section, a differentiation of the valve-
sculpture which so strongly and constantly characterises all the
Costatse. With reference to the occurrence of the costate Trigonia
in Cretaceous rocks, Behrendsen cites, in addition to T. peninsu-
laris Coquand and T. carinata Agass., the following shells : T. longa
Ag., T. pennata Sow., T. cardissa Ag., and T. indica Stoliczka. This
list, however, is misleading, as may be ascertained from a careful
examination of these forms. T. longa and T. pennata certainly
cannot be included in the section Costatae, while it is doubtful
whether T. indica stands in very close connection with this division.
T. cardissa, on the other hand, is a typical representative of the
section, but it has been shown to be Jurassic," and not Cretaceous,
as at first suggested on insufficient grounds by Agassiz.

GENUS CAEDITA J. G. Bruguiere.

CARDITA NUCULOIDES Tate.
Plate VII., figs. 5, 5a, 5b.

1867. Cardita nuculoides E. Tate, Quart. Journ. Geol. Soc., vol.
xxiii., p. 162, pi. vii., figs. la-Id.

Occurrence. Tate's figured specimen, numbered 11028 in the
collection of the Geological Society, came from the Sunday's Eiver.
A specimen obtained by Mr. Eogers is from the left side of the
Coega Valley, half a mile down from the railway (466g).

Remarks. The specimen found by Mr. Eogers is a well-preserved
left valve which differs slightly in outline from Tate's figured type :
it is a little more drawn out obliquely, and the convex fold of the
valve which runs obliquely from the umbonal region to the postero-
ventral corner is rather more pronounced than in the original
type. Tate's specimen, however, although having both valves in
place, is not so perfectly preserved, and some allowance may also
be made for individual variation. The general form of the shell,

* Bigot (1), p. 292.

128 Annals of the South African Museum.

and the complete agreement in the sculpture, leave no doubt
that the specimen from Coega Valley must be identified with
C. nuculoides.

The form of the shell is that of a four-sided figure with
rounded angles. The anterior margin is short, the posterior margin,
on the other hand, very extensive. The umbonal region is relatively
weakly developed. The surface is beautifully ornamented by
numerous, minute, radial striae which cannot be seen with the
naked eye. The intercrossing of the striae with concentric growth-
lines results in a delicate cancellation.

GENUS ASTAETE J. Sowerby.

ASTABTE LONGLANDSIANA Tate.

1867. Astarte long land siana R. Tate, Quart. Journ. Geol. Soc.,
vol. xxiii., p. 158, pi. viii., figs. 5a, 5&.

Occurrence. Specimens in the collection of the Geological Society
are from the Zwartkop's River. M.r. Rogers obtained an immature
example of this form in the Coega Valley east of the railway,
one mile up the line from Coega station (477g).

Remarks. The immature specimen from the Coega Valley
measures only 19 mm. in height. It has both valves in position,
and the umbonal region of the right valve is perfectly preserved.
The umbo is acutely pointed and is ornamented by closely spaced
concentric ribs up to the apex. Tate says that the valves are
ornamented with " slightly elevated ridges of growth," but it would
be more correct to say that the ornamentation consists of well-raised
concentric ribs. The ribs show some little irregularity of spacing.
The surface between the rib-summits frequently shows finer
concentric lines, which appear here and there to have the regularity
of definite sculpture, though in places they resemble ordinary ridges
of growth. The escutcheon is narrow, steep-sided, and deeply
excavated, and the ribs of the flank terminate abruptly at the acute
margin which bounds the escutcheon.

SUB-GENUS ERIPHYLA F. Stoliczka ( ? W. M. Gabb).
ASTABTE (EBIPHYLA) HEBZOGI (Goldfuss).

1840. Cytherea licrzogii (Hausmann) A. Goldfuss, Petrefacta
Germanise, Band ii., Lief. 7, p. 239, Tab. cxlix., fig. 10.

The Invertebrate Fauna of the Uitenliage Series. 129

1843. Astarte capensis F. Krauss, Amtlicher Bericht iiber die

zwanzigste Versarnmlung der Gesellsch. deutsch.

Naturforscher und Aerzte zu Mainz im Sept., 1842,

p. 130.
1850. Astarte herzocjii F. Krauss, Nov. Act. Acad. Caes. Leop. -Carol.

Nat. Cur., vol. xxii., pt. 2, p. 447, Tab. 47, figs. 2a-2e.
1905. Astarte herzoyi A. W. Eogers, An Introduction to the

Geology of Cape Colony, p. 291, fig. 25 (1).

Occurrence. Specimens before me are from the uppermost
red bed in a kloof east-north-east of Red House 011 Zwartkop's
River (326, 327), and from a krantz near Picnic Bush, north-east
of Red House (328). Messrs. Rogers and Schwarz also found this
shell in the cutting on the road running from the farm Perseverance
on to the plateau, and leading to the Salt Pan. Stow obtained
specimens at McLoughlin's Bluff and between the Addo and Modder
Drifts, on the Sunday's River. Hertzog's collection, including this
shell, was obtained in the Sunday's River district about eighteen
miles from Enon. : ' : Krauss collected specimens on the left bank
of the Zwartkop's River, below Uitenhage.

Remarks. Krauss furnished such an accurate description of this
shell that it will only be necessary to supplement this by a discussion
of its narrower relationships. Astarte herzogi cannot be looked upon
as a typical representative of its genus, and it is by no means a
simple matter to assign it to its correct position among recognised
sub-generic groups. The principal characters to be borne in mind
in this connection are as follows. The shell has a lenticular form
and frequently an almost circular outline ; it is ornamented by
crowded concentric ribs and sulcations forming a well-developed
sculpture. The lunule is short, but is sharply bounded and of very
deep and concave form ; the escutcheon is very narrow and
elongated, almost entirely occupied by the long ligament space,
and very sharply marked off from the flank. In the right valve
there is a distinctly developed posterior lateral dental process,
situated behind the elongated external ligament, and an anterior
(sub-lunular) depression in the hinge-plate for the reception of an
anterior lateral process of the left valve. The pallia! margin of the
valve is strongly notched on its inner side ; the pallial line is
posteriorly rather weakly impressed.

We are therefore dealing with an Astarte in which lateral hinge
processes are definitely developed. The evidence for the presence or

* See Hausniann (1), p. 1459 (as Cytherea).

130 Annals of tJie South African Museum.

absence of a shallow pallial indentation, it must be admitted, is incon-
clusive in the material I have examined, but this, after all, is a com-
paratively minor point, and one of less importance than the nature
of the hinge. A fact to be noted is that shells which exhibit a
similarly developed hinge are chiefly characteristic of Cretaceous
rocks in other regions.

Stoliczka : ' : thought that Astarte herzogi belonged to Speyer's
genus Grotriania,^ but in reality it is widely enough removed from
this, which has a very different form and is characterised by a very
large and profoundly excavated lunule and escutcheon, and by the
absence of lateral teeth. Much closer agreement is shown to the
shells known as Eriphyla lenticularis (Goldfuss) \ and other forms
to which the name Eriplnjla has been applied. Unfortunately, the
employment of this name has been attended with some confusion.
The shell upon which Gabb founded the name (E. umbonata
Gabb) occurs in the Shasta-Chico Series of California, || and it
was thought to be allied to Astarte. The characters of the shell
were unfortunately not diagnosed with satisfactory precision,
and Stoliczka first stated in full detail the distinguishing
features of Cretaceous shells (E. lenticularis) which he believed
to be generically identical with Eripliyla umbonata Gabb.
Stoliczka, however, brought the genus into relationship with
Dosinia, though he adduced no convincing reason for such a view,
except the presence of a pallial sinus. His description of the
hinge characters shows that they agree with those of Astarte herzoyi,
but he mentioned the pallial sinus as a leading feature of his shells,
and made no reference to any crenulation of the margin. He says :
" It appears probable that some of the Jurassic Astarte (A. excavata
and others) belong to this genus, but a careful examination of the
hinge and of the pallial line, which is broad though very faint, and
also of its sinus, will be necessary." Again, he remarks: "The
hinge-teeth of Eripliyla closely approach those of Astarte, but these
have no distinct lateral teeth, nor a deep lunule or sinus."

Whether Stoliczka did right in ascribing the widely distributed
E. lenticularis to Gabb's imperfectly characterised genus is a
question which still remains unanswered. He, at any rate, used the
name in connection with definite and precise diagnostic characters,
and his reading has been accepted by Holzapfel,** who maintains

* Stoliczka (2), p. 286 (1871). t Speyer (1), p. 496, Taf. si., fig. 6.

I Goldfuss (1), Band ii., p. 228, Tab. cxlvi., fig. 16 (1837).

Gabb (1), p. 180, pi. 24, fig. 162. || See Diller and Stanton (1).

<[ Stoliczka (2), p. 156 (1870). ** Holzapfel (1), p. 196.

The Invertebrate Fauna of the Uitenhage Series. 131

that so long as it cannot be shown that Eripliyla Stoliczka differs in
essentials from Eripliyla Gabb, this name must be retained for shells
having the characters set forth by Stoliczka. Previously, however,
J. Bohrn : ' : had remarked upon the unsatisfactory nature of Stoliczka's
use of Gabb's name, in view of the great uncertainty respecting the
characters of the Californian shell, and he employs the name Dozyia
for Lucina lenticularis Goldf., a name applied by Bosquet in 1868 to
the same form. Meek t also doubted the correctness of Stoliczka's
view in identifying the European and Indian shell with Gabb's
genus, but his remarks on the subject only add further confusion and
help to show the futility of attempts to utilise Gabb's imperfect
description and figure in a generic characterisation.

From the foregoing it appears clear to me that in the absence of
further information regarding the Californian type, the published
account of Gabb's genus is totally insufficient to support the validity
of the name proposed by him ; the repeated attempts to utilise
Gabb's description and figure have proved so unavailing that the
name as based upon Californian material may be reasonably ignored.
In further support of this contention it is only necessary to mention
that Whitfield} has so conceived the meaning of Gabb's description
as to apply the name Eripliyla to a Cretaceous shell from Syria
which is characterised by triangularly elliptical outline, strongly
prominent umbonal region, a short internal ligament, and surface
sculpture resembling that of Cliione.

The question then arises, whether the name Eripliyla as defined
by Stolic/ka may be applied to E. lenticularis, in view of the fact
that Bosquet had previously proposed to call this Dozyia lenticularis.
It is true that this generic name was only published in a tabular
list, without description or discussion, but it was accompanied by a
reference to Goldf uss's type. I should have no hesitation in
following Dr. J. Bohrn and employing Bosquet's name on the
strength of the definite reference to Goldfuss, but unfortunately the
description and figure given in the " Petrefacta Germanise " deals
only with external characters, and no mention was made of those
features which are of generic or sub-generic value. These were first
elucidated by Stoliczka, and herein lies the justification for following
Prof. Holzapfel in accepting the name Eripliyla, as defined by
Stoliczka.

Eripliyla lenticularis has a shallow pallial sinus and its valve-
margins are not crenulated. It might therefore appear that, on the

* J. Bohm (1), p. 125. t Meek (2), p. 123.

\ Whitfield (1), p. 403, pi. vi., figs. 2-5. Dewalque (1), p. 368.

132 Annals of the South, African Museum.

strength of these points, Astarte lierzogi should be excluded from
this division of the genus, though in reality these characters are of
very small value and appear to be sufficiently inconstant to be dis-
carded as guides to the sub-generic grouping. The hinge-characters
above described, on the other hand, are remarkably constant in
several Cretaceous forms which differ in regard to the marginal
crenulation, and it appears that the posterior indentation of the
pallial line is only of specific value at the most, appearing occasion-
ally, as in a few other " integripalliate " genera. A shallow pallial
sinus is shown to be present in the figures of Aptian shells ascribed
by Pictet and Eenevier ::: to Astarte buclii F. Roem. and A. obovata
J. Sow., and in both of these the margin is crenulate. Astarte striata
J. de C. Sow., from the Blackdown Beds of England, shows very
close agreement with E. lenticularis, and, like it, has a shallow sinus
and a smooth margin. Then again, the same shell agrees closely in
the hinge and other features with Astaric lierzogi, though the latter
has a crenulated margin. This last character, indeed, is clearly one
of very small significance. In dealing with Astarte and its divisions,
von Zittel t has ignored the notching of the margin, while several
authorities on the living forms have abandoned this structure as even
of specific value. \ Eegarding the value of the distinctive characters
of Eriphyla, as here accepted, in establishing its claim to the rank of
a separate genus, opinions may be expected to differ ; I am not
convinced that a definite separation from Astarte is expedient, or
warranted by the features which distinguish Eriphyla from typical
members of that genus, and I therefore prefer for the present to
follow Zittel in employing the name Astarte in the broad sense, and
including Eripyhla as a sub-genus.

In tracing the relationships of A. licrzogi, it is somewhat surprising
to find that the closest agreement is exhibited with shells from the
Neocomian of Lincolnshire. These occur in the Claxby Ironstone
at Willingham and Benniworth Haven, and are most probably to be
assigned to a horizon comparable with a part of the zone of Belem-
uitcs lateralis or possibly the lowest part of the zone of Belemnites
jaculum at Speeton ; specimens are preserved in the Museum of
Practical Geology in London, and in the Sedgwick Museum at
Cambridge, and have been identified by Mr. Woods as Astarte lavis
(Phill.).:j The similarity to A. licrzogi is striking, but the following

* Pictet and Eenevier (1), pi. x., tig. 1; xi., fig. 1.
f Zittel (5), p. 65. J See Jeffreys (1), p. 309.

Pavlow and Lamplugh (1), p. 29 (of authors' copy); Pavlow (1), able,
p. 548. || Woods (3), vol. ii., p. 115 (1906).

The Invertebrate Fauna of the Uitenhage Scries. 133

points of distinction appear to be constant and to justify the definite
separation of these two forms. In the Lincolnshire specimens the
shell is as a rule less orbicular, and attains greater height in propor-
tion to length ; at the same time, the valve is rather more flattened
towards the inferior margin in fully grown individuals. The lunule
has greater relative length and is less deeply excavated and less
concave in form than in A. hcrzogi. The noticeable fact that the
sculpture in the Lincolnshire examples is less well denned than in
the African shell may perhaps be in some measure due to the
mode of preservation. The form and size of the lunule certainly
affords a good separating character, and as regards the outline
of the shell, the difference above mentioned seems to hold good
if specimens of average proportions be compared, though it must
be admitted that Astarte herzogi exhibits considerable shape
variation, and specimens with shorter and higher outline occur
not infrequently.

As already remarked, Astarte striata Sow.,* from the Blackdown
Beds, agrees in many respects, but it is a much more compressed and
flattened shell, has greater length in relation to height, and shows no
marginal crenulation.

Astarte beaumonti Leym.,f from the Neoconiian of the Aube, is
likewise characterised by a sharply demarcated and deeply excavated
lunule, and is ornamented by ridges and sulcations similar to those
of A. herzogi; it may probably agree, too, in the internal characters,
but it differs by its inequilateral form and oblique elongation.

Astarte biichi P. Eoemer I (Lower Cretaceous, Perte du Ehone)
seems to agree well in internal characters, though it is not clear
whether a posterior lateral tooth of the right valve is plainly
developed. The sub-lunular groove in the hinge of the right valve,
shown in Eoemer's figure, seems to indicate the presence of an
anterior lateral tooth in the left valve. The sharply cut, elongated
escutcheon and the well-sunk short lunule are similar, but Eoemer's
shell differs from A. herzogi by being considerably more inequilateral
in shape and also less incurved at the umbones, besides having much
less perfectly developed concentric surface sculpture.

Astarte samanni de Loriol, from the Portlandian of the north of
France and of England, || is more similar again, and may be closely

* J. de C. Sowerby (1), vol. vi., Tab. 520, fig. 1 (1826).

f Leymerie (2), p. 4, pi. 4, fig. 1.

\ F. Boemer (1), p. 20, fig. 4.

De Loriol and Pellat (1), p. 68, pi. vi., fig. 9.

|| Blake (l),p. 232, pi. x., fig. 5.

10

134 Annals of the South African Museum.

related. The ornamentation, though coarser, with broader and more
widely spaced ribs, is of essentially the same kind, and there is the
deeply cut lunule. It may be judged from de Loriol's figure of a
left valve that a posterior lateral hinge process is present in the right
valve. The shell, while appearing to have very closely similar
hinge characters, differs from A. Jierzogi by its more elongated form
and relatively less height. English specimens from the Portland
Sands of Swindon, ascribed to A. scemanni, have a denser costate
ornamentation than the example figured by de Loriol, and in this
they more closely resemble A. herzogi. Miss E. G. Skeat has drawn
attention to the fact that these English specimens, which are well
represented in the Sedgwick (formerly Woodwardian) Museum at
Cambridge, exhibit a shallow pallial sinus."

A shell described by G. Miiller under the name Eripliyla stuhl-
manni, from the Neocomian at a locality 35 km. west of Mtshinga in
German East Africa,! clearly belongs to the same sub-generic group.
In the characters of the hinge and in the presence of marginal
crenulation, it resembles A. herzogi, but differs by the absence of
the surface ridges and sulcations and by the more inequilateral form
and more prominent umbones. Eripliyla stuhlmanni is accompanied
by a Gervillia probably identical with G. dentata Krauss. Dr. Miiller
refers his shell to Eriphyla without comment, and was no doubt
influenced by the presence of the lateral teeth.

In the orbicular outline, the surface ornamentation and the type
of hinge, Astarte jugosa (Forbes) I from the Upper Cretaceous of the
Trichinopoly district (Utatur stage) approaches somewhat closely to
A. herzogi. The Indian shell was referred by Stoliczka to Speyer's
genus Grotriania, but this was an error of judgment. Its valves, it
is true, have a somewhat deepened lunule and escutcheon, but do not
compare in this respect with those of the true Grotriania, and, more-
over, have plainly developed lateral hinge-processes fitting into
opposing grooves. In this respect A. jugosa agrees with A. herzogi,
and may be classed in the same sub-division of Astarte. According
to the description and figures given by Stoliczka, it has a crenulated
valve-border and the pallial line is without a sinus. The shell
differs from A. herzogi by the greater compression of the valves, the
greater depth of the escutcheon and hinge-plate, and the more nearly
circular outline.

* Skeat and Madsen (1), p. 124.

t G. Muller (1), p. 553, Taf. xxi., figs. 3, 4 ; Taf. xxii., figs. 8-10.

I Forbes (2), p. 142, pi. xvii.,fig. 7.

Stoliczka (2), p. 289, pi. x., figs. 12-14.

The Invertebrate Fauna of the Uitenhage Series. 135

Eripliyla argentina Burckhardt,* from the Neocomian of Las
Lajas (Argentine), is also a similar shell, as regards the hinge, the
deep lunule and escutcheon, and the development of concentric orna-
ments. It differs, however, from A. lierzogi by the considerably more
elongated figure and the coarseness and spacing of the ornamenting
ridges. E. argentina, it may be observed, has a weak pallial sinus,
and Dr. Burckhardt remarks that the margin is probably crenulated.

ASTARTE (ERIPHYLA) PINCHINIANA Tate.
Plate VII., figs. 6, 6ft.

1867. Astarte pinchiniana R. Tate, Quart. Journ. Geol. Soc.,
vol. xxiii., p. 157, pi. ix., fig. 7.

Supplementary Descriptive Note. The shell has a sub-orbicular
outline, with rather acutely pointed, anteriorly directed umbones.
The lunule is sharply demarcated from the flank, and is of deeply
excavated form, bounded outwardly by a sharp ridge which is con-
cave in profile. The lunule is of relatively great length. The
escutcheon is narrow, deep, and sharply bounded, and is occupied
very largely by the ligament space.

The valve-surface is ornamented during the neanic stage by
numerous and closely crowded, very regular and delicate concentric
ribs, terminating posteriorly at the margin of the ligament-space.
This ornamentation becomes obscure at a distance of 3-5 mm. below
the umbo, and subsequently gives place to numerous delicate, raised
growth-lines, too irregular to be confounded with definite sculpture.
Stronger ridges and furrows of accretion appear in the late adult
stage.

The shell-substance is very thick in relation to the size of the
specimens. The interior of the valve-border is strongly crenulate.

Dimensions. (1) (2) (3)

Height, measured from the umbo 15 . 18 18 mm.

Length 15 1S'5 . 19

Depth of a single valve 3 - 5 .5 4 ,,

Number (3) is the specimen figured by Tate.

Occurrence. Collected by Miss Wilman at Coega River. The
specimens in the collection of the Geological Society in London are
from the Sunday's River.

Bemarks. Tate's description and figure of this elegant form must
appear sufficiently inadequate w 7 hen it is realised that a most dis-

* Burckhardt (2), p. 76, pi. xii., figs. 3-f>.

136 Annals of the South African Museum.

tinctive feature is the very elongated, sharply demarcated and deeply
excavated lunule, well preserved and exhibited in the specimen un-
satisfactorily figured by Tate. The widely erroneous statements of
shell-measurements, so numerous in that author's paper on the South
African fossils, are difficult to account for and may best be ignored.
Tate's comparison of A. pinchiniana with two English Oolitic forms,
A. excentrica Morr. and Lye. and A. puviila J. de C. Sow., is also
unfortunate, since a very much closer resemblance is shown to several
Cretaceous forms. A. excentrica'''' is much more triangular in outline
and has the lunule scarcely defined, while A. pumila f is a narrow,
convex, inequilateral shell widely different in type from the one we
are considering.

The question whether A. pinchiniana should be referred to the sub-
genus Eriphyla, as defined by Stoliczka, cannot be decided by an
examination of the material which has been at my disposal. In no
instance have I been able to ascertain the characters of the hinge, and
it is therefore uncertain whether lateral hinge-processes are present
in either valve. At the same time, the external characters of shape
and the presence of a deep and sharply defined lunule, suggest the
strong probability that the arrangement of hinge and lateral teeth is
the same as in Eripliyla, but it would, of course, be unwise to accept
close agreement in external features as justification for a definite
conclusion on this point. For the time being, a provisional reference
to Eripliyla may be permitted. This question might seem to be of
subsidiary interest were it not that the known characters of
A. pinchiniana give this shell so close a resemblance to members of a
principally Cretaceous group of forms, and as additional evidence for
the age of this fauna, such resemblance must be taken fully into
consideration. Remarks on the application of the name Eripliyla in
dealing with certain Cretaceous forms of Astarte will be found above
in the discussion concerning the relationships of Astarte (Eripliyla}
herzogi.

A. pinchiniana differs from immature examples of A. herzogi bythe
relatively more extended lunule and the disappearance of concentric
sculpture at the close of the neanic stage. It bears a great outward
resemblance to immature specimens of A. striata Sow.,:]: having a
similarly elongated lunule, with the marginal profile of concave form
in front of the umbo ; but in .4. pinchiniana the concentric surface

* Morris and Lycett (1), part iii., p. 83, pi. ix., fig. 8 (1855).
t J. de C. Sowevby (1), vol. v., Tab. 444, fig. 2 (1824).
I Ibid., vol. vi., p. 35, Tab. 520, fig. 1 (1826).

The Invertebrate Fauna of the Uitnilitige Series. 137

ridges are absent in the adult, and the interior of the valve-margin is
strongly crenulated.

Astarte rhodani Pict. and Camp.,* from the " Gault" of the Perte-
du-Khone, is of similar general type, but is truncated posteriorly.
Shells from the Gault of Cosne (Nievre) figured by de Loriolf under
the name A. rhodani have a much more closely similar, rounded
outline; they appear, however, to have a slightly shorter lunule, and
they are probably without marginal crenulation.

Astarte dupiniana d'Orb.J from the Albian of Prance, closely
resembles A. rhodani Pict., and like this, is abruptly truncated
posteriorly ; in this it offers a contrast to the more rounded profile of
A. pinchiniana. A. dupiniana possesses the character of marginal
crenulation, but differs from the African shell in having a less
strongly developed lunule and escutcheon.

The presence of well-developed sculpture in the neanic stage of
A. pinchiniana and its disappearance in the succeeding growth-
stages, indicates degeneration, so far as this character is concerned,
from a wholly sculptured ancestry.

GENUS ANTHONYA W. M. Gabb.

ANTHONYA LINEATA sp. nov.
Plate VII., figs. 7, 8.

Description. The shell is of slender, elongated form, much pro-
duced posteriorly. The valves are flattened and compressed, having
very slight convexity. The umboiial region is not strongly pro-
minent ; it is situated at a distance of about one-third of the shell's
total length from the anterior extremity, where also the shell has its
greatest height. The upper valve-margin, posteriorly to the umbo,
slopes down gradually, giving a very slightly concave outline. It has
an angular junction with the siphonal margin. The siphonal margin
is very short and shows a straight outline, directed slightly pos-
teriorly when traced down to its inferior termination. In front of
the umbo, the valve-margin is at first forwardly produced with a
downward slope and straight outline, passing then by a curve into
the convex anterior margin. This in turn passes by regular curve
into the elongated, slightly convex inferior margin. The siphonal

* Pictet and Campiche (1), 3 e Partie, p. 319 (1866) ; Pictet and Eoux (1), p. 437,
pi. 32, fig. 5 (1852).

t de Loriol (4), p. 94, pi. xii., figs. 1-7.

t d'Orbigny (3), p. 70, pi. 264, figs. 4-6 (1844).

138 Annals of the South African Museum.

margin marks the posterior limit of a weakly defined postero-
superior area of the valve-surface. This is only demarcated from the
flank by a weak and flattened fold of the surface which passes
obliquely backwards from the umbo to the postero-inferior angle of
the valve.

The ornamentation consists of very delicate, closely spaced,
rounded, concentric ridges, separated by impressed linear interspaces.
The ridges traverse the flank, parallel to the pallial border, then turn
sharply upwards to cross the area in a direction parallel with the
siphonal border. On the area the ornaments are slightly coarser
and less regular, and less strongly developed than on the flank.

An internal cast of a left valve shows traces of very weak rounded
ribbing, running parallel to the inferior border, but these markings
are confined to the middle part of the valve and terminate posteriorly
at the oblique, weak carinal ridge. In addition to the two short,
diverging, cardinal teeth, there are indications of a narrow, lath-like
ridge (represented on the cast by a narrow hollow) running parallel
with the upper valve-margin for a short distance in front of the
umbo, and another similar narrow depression extends for a longer
distance close to the valve-margin behind the umbo. Just above the
posterior part of the broadly oval anterior adductor impression,
is a small, well-marked, oval pedal muscle-scar. The somewhat
elongated posterior adductor impression is situated close to the upper
valve-border, half-way between the umbo and the posterior margin,
and just below the narrow, lath-like process which is represented on
the cast by a groove close to the valve-margin. The pallial line on
reaching its posterior extremity turns sharply forward with an
angular bend.

Dimensions. (1) (2)

Length 15 . 22mm.

Greatest height 8 . 12 ,,

Occurrence. Specimen (1), preserved as a mould of the external
surface of a left valve, was found on the left side of the Coega
Valley, half a mile down from the railway (461g). Specimen (2),
which is the cast of the interior of a left valve, came from a bare
slope W. 30 S. from the middle of Barkly Bridge, on the farm
Olifant's Kop, Sunday's Eiver (35h).

Remarks. There can be no reasonable doubt that the two
specimens above described, showing the exterior and interior
characters respectively, belong to one species. They are of interest
as representing a genus of shells which is elsewhere only known to
occur in rocks of Cretaceous age. These valves have a very delicately

The Invertebrate Fauna of the Uitenhage Series. 139

shaped, flattened form, and though their depth cannot be satisfac-
torily measured in either specimen, it could not have exceeded two
or three millimetres. In the example from Olifant's Kop, the pallia!
border does not lie in a median plane of symmetry, but is laterally
bowed, with a slight convexity towards the observer. This may
possibly have been an individual abnormally distorted in life, or the
distortion may perhaps have been brought about during the process
of fossilisation, but it was probably not a normal character.

Antlionya cultriformis Gabb, : ' : the type-species of the genus, from
the Cretaceous rocks of California, differs in being considerably
more elongated and attenuated posteriorly, and in having more
oblique posterior truncation. It also lacks the close linear orna-
ments which cover the whole surface of A. lineata, though it has a
similar type of sculpture in the umbonal region.

A. cantiana Woods,! from the Folkestone Beds of Folkestone, is
more inequilateral, with the umbonal region more elevated and
nearer to the anterior extremity. A species described by Mr. Woods,
though unnamed, from the Lower Greensand of Atherfield, is
distinguished by the posterior elongation and attenuation of its
shell, and by its smooth surface.

Anthony a cornmliana (d'Orb.) \ appears to approach more closely
in form, especially such a specimen as that figured by Miss Skeat ;
but .4. lineata is less inequilateral and is not so obliquely truncated
posteriorly.

GENUS TANCEEDIA J. Lycett.

TANCEEDIA SCHWARZI sp. nov.
Plate VII., figs. 9, 9a, 10.

Description. The shell has an elongated trigonal outline, well
produced anteriorly. The umbonal region is prominent, the margin
before and behind it falling away rapidly. The urnbo is situated
within the posterior half of the valve, though not distant from the
middle ; it is fairly strongly incurved. The upper margin in front of
the umbo forms a straight line in profile, and only curves on reaching
the anterior extremity, where it passes by a sudden and sharp bend
into the long, gently convex inferior margin. At the posterior
end of the short, straight hinge-line the shell-outline is obliquely

* Gabb (1), p. 182, pi. xxx., fig. 236.

t Woods (3), vol. ii., part 3, p. 130, pi. xix., figs. 4, 5 (1906).
t d'Orbigny (3), p. 74, pi. cclxiv., figs. 7-9 (1844), described as Crassatella ;
Skeat and Madsen (1), p. 178, pi. vi., fig. 13 (as Ptychomya).

140 Annals of the South African Museum.

truncated. The oblique posterior border has a sub-angular junction
with the inferior border. The greatest height occurs at the umbo.
The valves are weakly inflated, most strongly so posteriorly to the
middle, and are anteriorly compressed and flattened.

On the posterior side of the valve, a well-marked carinal ridge
extends obliquely backwards from the umbo to the posterior angle of
the shell, cutting off a narrow, flattened area from the flank. Near
the umbo, the surface of the area is directed at right angles to that
of the flank, but the angle becomes more obtuse when traced
towards the posterior end of the valve. The surface of the area is
slightly concave in the neighbourhood of the umbo, but becomes
flat posteriorly. The surface of the flank close to the antero-superior
margin curves over with a convex surface towards the margin.
The surface of the valves is marked only by occasional faint growth-
lines.

Immature specimens have a rather more pointed anterior outline
than an individual of larger dimensions.

Dimensions. (1) (2) (3)

Length 16 . 16 22mm.

Height, measured from the umbo 10 . 11 15 ,,

Greatest depth of a single valve 4 . 4 . 5 ,,

Occurrence. In the kloof east-north-east of Bed House, on the left
side of the Zwartkop's Eiver (324) ; this form is also found in the
Marine Beds of the Sunday's Eiver, where it occurs associated in
the same hand-specimen with Trigonia ran, Trigonia conocardii-
formis, Act&onina atherstoni, and other characteristic shells. A
specimen of this kind is from the collection of the South African
Museum. Mr. Rogers collected a fine example of this form from a
cliff W. 20 S. from Comley's house, right bank of Sunday's Eiver
(95h).

Remarks. The specimens examined are unfortunately so preserved
in hard matrix that it has been found impossible to investigate the
internal characters. The outward features of the shell, however, are
so well marked and so characteristic, that one can scarcely suggest
a doubt as to the generic position, unless, indeed, it should happen
that we are dealing with a striking instance of parallelism, of which
there is so far no evidence. It is with Jurassic forms that the most
striking similarities are shown.

This shell may be most aptly compared with familiar English
Oolitic forms. T. extensa Lycett,* from the Inferior Oolite, is very

* Lycett (1), pi. xi., tig. 9; Morris and Lycett (1), part iii., p. 93, pi. xiii., tig. 6
(1855).

The Invertebrate Fauna of the Uitenhage Series. 141

similar in outline, but differs in having the urnbones more centrally
placed, and the posterior carinal ridge less prominently developed
and less steeply inclined. The posterior area is narrower, and
this and the carinal ridge have greater relative length than in
T. scliwarzi. In the African shell the urnbonal angle is sharper
and the umbonal region more prominently projecting.

Tancrcdia brevis Lycett," from the Great Oolite, is also closely
similar in form, and may be distinguished principally by its slightly
greater convexity, its less prominently projecting umbonal region,
and its more pointed anterior outline.

Again, the resemblance shown by T. sclucarzi to T. angnlata
Lycett,! from the Great Oolite, is very close, particularly in the
position of the unibones, the long antero-superior slope of the outline,
and the inclination of the carinal ridge ; in T. angulata, however,
the outline is rather more obtuse at the posterior angle, and the
aspect of posterior truncation is more marked, while the anterior
outline has a rather more pointed form. The resemblance of T.
schiuarzi to these Jurassic types is indeed so striking as necessarily
to arrest attention, and it is clear that such a form, if known before,
would have been seized upon by those who believed the Uitenhage
fauna to be of Oolitic age. How little reliance should be placed on
the evidence of a single occurrence such as this is well shown by a
study of the associated types, which can only be taken to indicate the
Lower Cretaceous age of this fauna. Although the genus Tancrcdia
reached its greatest development during Jurassic times, T. americana
Meek and Hayden I is known from Cretaceous strata on the Upper
Missouri and on Cache La Poudre River, in Colorado. The generic
position of T. americana has not been disputed, so far as I am aware,
but it is a much larger shell, not closely comparable with the one
we are discussing.

A shell from the Upper Aptian of Spain, described by Coquand
as Tellina gibba, bears considerable resemblance to some forms of
Tancredia, and may very well be a representative of this genus,
though Stoliczka suggested that it belongs to Gray's Tellinella.
Coquand himself stated that it differs from other fossil Tellina by
its elongated form and particularly by the strong carinal ridge.
Speaking generally, this shell shares the main outward characters of

* Morris and Lycett (1), part iii., p. 92, pi. xiii., fig. 8 (1855).
f Lycett (2), 341, pi. xiv., fig. 5 ; Morris and Lycett (1), part iii., p. 94, pi. xiii.,
fig. 9 (1855).

\ Meek (2), p. 142, pi. 38, fig. 1.

Coquand (1), p. 101, pi. viii., figs. 9, 10.

142 Annals of the South African Museum.

Tancredia schwarzi, from which it differs by its greater elongation
and relatively reduced height; it has a more obtuse unibonal angle,
and the umbonal region is consequently considerably less prominent.
Although there is also some resemblance to shells which have been
ascribed to the genus Palceomya Zittel and Goubert,* Palceomya
deshayesi Zittel and Goub., from the Corallian, upon which the
genus was established, is a more elongated shell and is more
inequilateral and less definitely carinated posteriorly. Its umbonal
angle is also very much more obtuse. Palceoinya autissiodorensis
(Cott.) de Loriol,f from the Portlandian of the Yonne, while more
equilateral and more sharply carinated than the last, is also
relatively much more elongated than the African shell, and has
much less sloping upper outlines, with inconspicuous umbonal
region and obtuse umbonal angle. It may be noted that Miss E. G.
Skeat ;[ has referred this Portlandian shell to the genus Tancredia,
and expressed the belief that these supposed separate genera may
have to be united ; she points out that they appear to agree very
closely both in external and internal characters.

GENUS THETIEONIA R Stoliczka.
THETIRONIA PAPYKACEA (Sharpe).
Plate VII., figs. 11, lla.

1856. Ceromya papyracea D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,
vol. vii., p. 195, pi. xxii., fig. 8.

Supplementary Description. The shell is of rounded sub-quadrate
or oblong outline, with the length slightly greater than the height ;
it is strongly inflated in the umbonal half. The umbones are
strongly incurved and are anteriorly directed ; they are situated at a
distance of less than one-third of the shell's total length from the
anterior margin. The hinge-line is only very slightly curved and has
a scarcely perceptible downward slope when traced backwards from
the umbo, passing posteriorly by a somewhat abrupt curve into the
elongated posterior margin; this margin, also, is only very slightly
convex in profile and gives the shell an aspect of vertical posterior
truncation. The lower border forms a gently convex profile. There
is no keel on the posterior side of the valve.

The surface of the very thin shell is ornamented by numerous,

* Zittel and Goubert (1), pi. viii., figs. 6-8.

f de Loriol and Cotteau (1), p. 510, pi. v., figs. 12-14.

I Skeat and Madsen (1), p. 129.

The Invertebrate Fauna of the Uitenhage Series. 143

delicate, radial, linear striae, which at the lower margin of an adult,
and especially in the posterior half of the valve, may he separated by
spaces exceeding a millimetre in breadth. The granular ornaments
which occur on the striae are delicate though prominent, and widely
spaced (nearly 4 mm. apart on the same radial line) towards the
lower margin in the posterior half of the valve, but less conspicuous
and much more closely spaced in the anterior half. ::: The radial
striae leave their impression on the cast when the thin shell becomes
removed.

The well-impressed line on the posterior side of a cast, passes far
up and has its angular apex situated close to the umbonal apex. Its
anterior limb becomes weakly impressed when traced downwards
from the angle for a distance of 7 mm. or 8 mm., and appears to
dwindle and disappear at its lower end instead of bending forwards
as a well-defined line. On the anterior side of the cast there is
a single radial linear impression, less clearly defined than those of
the posterior side ; it appears most clearly marked at a distance
of about 6 mm. from the umbonal apex, and dies out on entering
the lower half of the valve.

Dimensions. (1) (2) (3)

Length 22 . 25 . 47 mm.

Height measured from the umbo 19 30 ,,

Greatest depth of a single valve ... . 10 11 ,,

Number (3) is the specimen figured by Sharpe.

Occurrence. Collected in the railway cutting between milestones
24|-24f on the line from Uitenhage to Graaff-Reinet, about three
miles from Uitenhage (315) ; also found by Miss M. Wilman at
Coega. The locality given by Sharpe is " Zwartkop River."

Remarks. Sharpe referred this shell with some doubt to the
genus Ceromya, and he evidently had not the opportunity of
observing the characters of the cast, which would have set at rest
all doubts as to a generic determination. In addition to the well-
impressed angular line on the posterior side of the umbonal region
of the cast, the thin shell and the surface markings afford additional
indications of generic position. In the fine specimen collected by
Bain and figured by Sharpe, the shell is so preserved that the delicate
ornaments of the surface are obscure, and their presence only becomes
clearly evident when the specimen is very carefully examined in a
good light under slight magnification. The specimen from Coega,

* It may, perhaps, be more correct to speak of some of these markings as puncta-
tions, each of which has a well-raised circular rim ; they represent the bases of very
short spines.

144 Annals of the South African Museum.

figured here, is so well preserved that these surface characters are
plainly visible to the naked eye.

The occurrence of a representative of this exclusively Cretaceous
genus has considerable interest when regarded as additional evidence
for the age of this molluscan fauna. It is perhaps fortunate that the
shell before us, when well preserved, is sufficiently characterised by
its own distinctive form to preclude confusion with European mem-
bers of the genus, especially as the nomenclature of these has for so
long been in an unsatisfactory state. The European forms appear
to show considerable variation, and are often preserved merely as
casts, while the delicacy of the shell renders it liable to distortion,
and true characters of similarity or difference are, therefore, often
difficult to establish. In like manner, immature or ill-preserved
specimens of Th. papyracea may seem to differ somewhat widely
from the large individual figured by Sharpe, and may appear to
approach some of the European forms ; hence, a brief comparison
with certain of these may perhaps be not without value.

The shells from the Blackdown Beds described under the name
Corbula Iceviyata by Sowerby* seem to represent the same form as
that afterwards named Thetis major, \ while some shells from the
Lower Greensand of Atherfield (Isle of Wight) cannot always be
satisfactorily distinguished from small specimens of the Blackdown
form. Others from the Lower Greensand in the Isle of Wight,
named Thetis minor by the same author, \ are frequently preserved
as casts (the specimens from Shanklin), but appear to be usually
well distinguished from those above-mentioned by their greater con-
vexity, their more prominent umbones, and frequently straighter
and longer hinge-line. Some individuals, however, are very difficult
to separate, and Eoemer j united these two forms under the name
Thetis soicerbyi. Forbes, again, divided Boemer's Th. sowerbyi into
varieties minor and major ; |] but these are both regarded by Mr.
Woods as synonymous with J. de C. Sowerby's Thetis minor.
Mr. Woods also considers that d'Orbigny's Tlietis Icevigata is
identical with Sowerby's Th. minor. Despite some difficulty in
the comparison of certain individuals, we may on the whole satis-
factorily distinguish between Thetironia lavigata (J. Sow.) which
includes Th. major (J. de C. Sow.) and Th. minor (J. de C. Sow.),
which includes Th. Icerigata (d'Orb.)

* J. Sowerby (1), vol. iii., Tab. 209, figs. 1, 2 (1818).

f J. de C. Sowerby (1), vol. vi., Tab. 513, figs. 1-4 (1826).

+ Ibid., vol. vi., Tab. 513, figs. 5, 6 (1826).

F. A. Roemer (2), p. 72 (1841). || Forbes (1), 242.

The Invertebrate Fauna of the Uitenhage Series. 145

Thetironia papyracea is distinguished from Th. Icevigata by its
greater inflation, its anteriorly situated umbones, its posterior trunca-
tion, and the absence of strong radial striae on the posterior side in
the neighbourhood of the cardinal margin. It is widely separated
from Th. minor by its inequilateral form and long, truncated posterior
margin, and apparently also by the obscure lower termination of the
anterior limb of the angular line impressed on the cast.

The Cenomanian shell ascribed by d'Orbigny to Thetis major,
which was renamed Thetis rotomagensis by Pictet and Campiche, is
much more equilateral, and appears to have the umbonal region less
prominent and less inflated than in Th. papyracea, and if d'Orbigny's
figure be correct, the angular line on the cast does not extend so far
up towards the umbonal apex. The anterior limb of the line is also
bent forwards at its lower end and passes in well-defined manner
across the flank.

Thetironia renevicri (de Loriol), : ' ; from the Hauterivian of Sainte-
Croix and Mont Saleve, has a general outline and degree of convexity
somewhat approaching that of Th. papyracea, but the Swiss shell
has the limbs of the angular line of the cast much more widely
diverging, and the anterior limb forwardly bent and passing relatively
closely to the anterior valve-margin when traced downwards ; it
is also more equilateral, with much greater height in relation to
length.

Thetironia genevensis (Pict. and Roux),t which occurs in the
" Gault " of the Perte-du-Khone, Cosne, Sainte-Croix, and other
places, is distinguished by its more equilateral and more circular
outline, and by the form of the angular line on the cast, though
it approaches Th. papyracea in its inflation and in the prominence
of the umbonal region. Thetironia prestensis (Pictet and Cam-
piche) I (Aptian) lacks the elongation and posterior truncation that
characterises Th. papyracea, and its linear markings on the cast
follow a widely different course.

Thetironia sanctce-crucis (Pict. and Camp.) is still further removed
from the African shell, not only by its outline, but especially by
the great relative breadth and shallowness of the posterior sinus on
the cast.

A shell which much more closely approaches Th. papyracea in the
general form and outline and in the position of the umbones is

* de Loriol (1), p. 65, pi. ix., fig. 11.

t Pictet and Koux (1), p. 420, pi. xxx., fig. 2 (1852).

\ Pictet and Campiche (1), 3" Partie, p. 205, pi. cxii., fig. 6 (1865).

146 Annals of the South African Museum.

Th. caucasica (Eichwald),* from " Greensand " in Daghestan, said
to be of Gault horizon. When compared with other representatives
of the genus, this, like the African form, is of relatively elongated
outline ; but in Th. caucasica the shell is slightly more equilateral
and the posterior border is not so abruptly truncated as in Th.
papyracea, but is more convex in profile, while the course followed
by the anterior limb of the angular line on the cast is a different
one.

Although the true systematic position of the genus Thetironia
cannot yet be said to be established beyond doubt, it is clear that
the angular linear marking on the internal casts in no degree coincides
with the course of the pallial line. The genus has therefore been
erroneously held to be related to members of the Veneridse, but
although its suggested affinity to the Lucinidae seems to accord
better with what is known of its characters,! yet the evidence for
such relationship is not of a satisfactory character. Mr. H. Woods,
who has examined some material which is very favourably preserved
for the purpose of comparative study, has found points of analogy
between TJtctironia and Protocardia, both in the characters of the
hinge and the ornamentation ; but after drawing attention to some
features which distinguish these genera, and carefully weighing the
available evidence, Mr. Woods concludes that "although Thetironia
resembles the Cardiidae in several respects, yet therpoints of difference
are too great to allow of its being included in that family." J

THETIRONIA OBLONGA sp. nov.
Plate VII., figs. 12, 12a, 126.

Desertion. The shell is of elongated form, with the length con-
siderably greater than the height. The umbones are situated a short
distance anteriorly to the middle of the shell and are fairly well
raised and prominent and are strongly incurved. The cardinal
margin is very long and is only slightly curved, falling very gently
when traced forwards from the umbo and still more gradually
posteriorly to the umbones. The greatest height of the shell is at
the umbonal part. The height near the posterior end of the shell is
greater than in the anterior quarter. In lateral aspect, the cardinal
margin is seen to pass by a curve into the posterior margin, which

* Eichwald (1), p. 709, pi. xxvi., fig. 7; Anthula (1), p. 90, Taf. iv., fig. 6.

t See remarks on this point by Miss E. G. Skeat ; Skeat and Madsen (1), p. 177.

t Woods (3), vol. ii., part 4, p. 167 (1907).

The Invertebrate Fauna of the Uitenhage Series. 147

at once falls very steeply with very gently convex outline. This
passes in turn by a curve (broader than that of the postero-dorsal
outline) into the inferior margin, which has an almost straight out-
line. The short frontal border has an evenly convex outline. The
shell is greatly inflated below the umbonal region, but the valves are
compressed in the neighbourhood of the posterior border.

The outer surface of the very thin shell-wall, so far as it has been
observed, appears to show only the faintest traces of radial striae,
but delicate radial linear markings are to be seen on the surface of a
cast of the interior. The punctate ornaments of the surface may be
separated from one another, on the same radial line, by a space of
at least 3 mm. in the posterior part of the shell, near the pallial
margin. They are less strongly developed and are more closely
spaced in the anterior part of the valves.

The well-marked angular linear impression on the posterior side
of the umbonal region of the internal cast, has its apex extending
almost to the umbonal apex, so that the apical part of the angular
line is hidden from view by the incurvation of the contiguous
umbones, in a specimen in which the valves are united. The line
forms a very acute angle. The posterior limb is relatively weakly
impressed, but the anterior limb is well incised and has a very slight
backward inclination when traced away from the umbonal apex. In
the specimen here described, it only extends for about 5 mm. and
then ceases abruptly, while the posterior limb extends for at least
8 mm. and dwindles away.

Dimensions.

Length 25 mm.

Height, measured from the unibo 17 ,,

Greatest depth of a single valve 9 ,,

Occurrence. Collected at a kloof, S. 5 W. from Comley's house,
on the right bank of Sunday's River (83h).

Remarks . Only one specimen of this form has been found, but it
is well characterised by its long cardinal and pallial margins and
the relatively small height in comparison with length, as well as by
the peculiar form of the angular linear impression on the cast in the
umbonal region. The specimen is preserved in the form of an
internal cast with portions of the shell adhering, and it is not in the
least distorted. So far as I am aware, there is no described species
of Thetironia with which this new form can be brought into close
comparison.

148 Annals of the South African Museum.

GENUS TEAPEZIUM Megerle von Miihlfeldt.

TRAPEZIUM? TATEI sp. nov.
Plate VII., figs. 13, 13a.

Description. The shell is well elevated in figure, with the umbonal
region rising prominently, and well removed from the anterior
extremity. The cardinal margin is almost straight posteriorly to the
umbo, and slopes back to form a rounded obtuse angle with the
straight, very steeply falling posterior border. In front of the umbo
the margin falls rapidly, with slightly convex outline, to the sharply
convex anterior border. The inferior margin gives a broadly convex
outline, and has a sharply angular junction with the posterior
border. The greatest height occurs at the umbo, the greatest con-
vexity at about the middle of the valve.

On the posterior side of the valve a sharp carinal ridge passes
in steeply inclined direction from just behind the umbonal apex to
the postero-inferior angle of the valve-margin. This carina marks
off a very well-defined, flattened posterior area, the surface' of which
is inclined at a sharp angle to that of the remainder of the valve.
The valve-surface is devoid of sculpture, but is marked by numerous
delicate lines of growth.

Dimensions. (1) (2)

Length 9-5 . 14 mm.

Height 8 . 12 ,,

Depth of a single valve . 4 ,,

Occurrence. Found on the left side of the Coega Valley, half a
mile down from the railway (452g). An imperfect left valve,
apparently of the same species, occurring in the same hand-specimen
with Solecurtus sjj. and Meretrix uitenhagensis (310), was found at
Grass Ridge, three miles east-north-east of Uitenhage.

Remarks. Unfortunately there is only a single perfect specimen,
a left valve, available for description, but this is very well preserved,
although the interior characters cannot be studied.

Another shell of still more doubtful generic position, with which
this form is comparable in some respects, is Corbula ? rockiana
Tate. :;: The general habit is very similar, but a very careful com-
parison with Tate's type-specimen (number 11023 in the collection of
the Geological Society) seems to preclude any idea of uniting these
two forms. Tate's species is less elongated, more elevated in figure,
and considerably more inflated, particularly in the umbonal region,

* Tate (1), p. 159, pi. viii., tig. 8.

The Invertebrate Fauna of the Uitenhage Series. 149

and its carina falls more steeply and forms a line more nearly
straight. In Trapezium ? tatei, the carina follows a more oblique
direction and forms a more curved line when the valve is viewed
in lateral aspect. In Corbula ? rockiana the umbonal region is
broader and more massive, and the incurvation more pronounced.
It may be remarked that Tate's figure of Corbula ? rockiana is not
accurately drawn. The figure gives the idea that there is a depres-
sion or concavity on the surface of the valve, whereas this is really
not the case in the specimen itself.

A shell of apparently very similar type, so far as external
characters go, has been described by G. Mtiller* from the Lower
Cretaceous of German East Africa as Mactra stromeri. Another
comparable form when large specimens are selected is Mactra ? angu-
lata J. de C. Sowerby, from the Blackdown Beds of England.! This,
however, is less produced in front and the umbonal region is
not so prominently developed and is less incurved. The urnbo in
Trapezium ? tatei is more anteriorly placed and more forwardly
directed.

Owing to the scanty material available for study, and the fact that
the nature of the interior is at present unknown, the generic position
of this shell cannot be satisfactorily settled, although the provisional
assignment to Trapezium may, perhaps, not prove incorrect. It
seems possible, however, that this may be a member of the group,
typified by Cypricardia bathonica d'Orb.,} of the Great Oolite, for
which Fischer has proposed the name Pseudotrapezium, although
there can be no certainty on this point.

GENUS CYPEINA Lamarck.
CyrEiNA EUGULOSA Sharpe.

1856. Cyprina rugulosa D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

vol. vii., p. 195, pi. xxii., fig. 9.

Occurrence. This characteristic form was obtained from the green
sandy beds, crowded with the remains of oysters, in the cliff below
the old school-house at Dunbrodie, Sunday's Eiver (325). Sharpe
recorded it from " Sunday Eiver, in greenish shelly grit," and Stow
mentions its occurrence above the Modder Drift on the Sunday's

* G. Miiller (1), p. 563, pi. xxv., fig. 15.
t J. de C. Sowerby (2), p. 341, pi. xvi., fig. 9.
\ Morris and Lycett (1), part ii., p. 75, pi. vii., fig. 8 (1853).
Fischer (1), fasc. xi., p. 1075 (1887i.

11

150 Annals of the South African Museum.

Eiver, in association with the characteristic Trigonice, of the Marine
Beds. In 1905 Mr. Rogers found this species in the cliff on Buck
Kraal, Sunday's Eiver (128h).

Remarks. Cyprina rugulosa, in its inflated character, the position
of the umbones, and the wrinkled surface, bears no slight resem-
blance to C. regularis d'0rb., :;: from the lower Gault of Europe ; it
differs chiefly in the manner in which the upper margin slopes away
posteriorly into the posterior margin. The European form has
greater relative height at its posterior end, with an accompanying
aspect of truncation.

Some individuals of C. rugulosa, in which the shell attains a rather
greater height in proportion to length than in the majority of
specimens I have examined, approach somewhat closely in form to
a Cyprina recorded by Dacqu6 from strata of supposed Aptian age
in Somaliland ; f but C. rugulosa is less strongly inflated, and
most examples are further distinguished by their rough surface
markings.

CYPEINA BOECHERDSI Tate.

1867. Cyprina borcherdsi E. Tate, Quart. Journ. Geol. Soc., vol. xxiii.,
p. 161, pi. viii., fig. 2.

Occurrence. Found in the railway cutting between milestones
24i_24f on the line from Uitenhage to Graaff-Eeinet, about three
miles from Uitenhage (313). The specimens described by Tate were
also from the Zwartkop's Eiver.

A specimen obtained by Mr. Eogers from the left side of the Coega
Valley, half a mile down from the railway (467g), must also be
referred to the same species.

Remarks. Tate's type-specimen is unfortunately inaccurately
figured, and is represented as considerably more elongated than it
really is. It measures 42 mm. in greatest length and 31 mm. in
height, while the corresponding measurements of the illustration in
Tate's paper are 47 mm. and 27 mm. respectively. The greatest
depth (convexity) of this type-specimen (a single valve) is about
10 mm. A specimen obtained by Messrs. Eogers and Schwarz is an
immature individual measuring 17 mm. in height ; it is imperfect
anteriorly, and consequently has the aspect of being more equilateral
than Tate's shell, though a careful comparison leaves no doubt about
its identity.

* d'Orbigny (3), p. 100, pi. 272, figs. 3-6 (1844); Pictet and Campiche (1),
3 e Partie, p. 224, pi. cxv., figs. 1, 2 (1865).
t Dacque (1), p. 16, Taf. ii., fig. 9.

The Invertebrate Fauna of the Uitenhage Series. 151

I have been unable to ascertain the nature of the hinge, and must
regard the generic determination of this form as provisional. A
shell having very similar outward form is Cyprina sioindonensis,
described by the late Prof. Blake ::: from the Portlandian Swindon
Sands of England.

GENUS MEEETEIX Lamarck (sensu laid).

MERETKIX UITENHAGENSIS sp. nov.
Plate VII., figs. 14, 14a; VIII., figs. 1, la.

Description. The shell is of somewhat variable ovate outline,
with the umbones situated at about one-third (or slightly less) of the
shell's total length from the anterior extremity. The shell-substance
is relatively thin and the aspect of the valves considerably com-
pressed and flattened, particularly in the lower half of the individual.
The cardinal margin slopes down only very gently when traced back
from the umbo, giving a slightly convex outline, and passes by a
curve into the posterior border which is evenly convex in outline,
or most sharply curved towards the lower part. This border
usually has considerably greater extent than the frontal border,
which is more sharply curved and limited in height by the some-
what rapid convergence of the upper and lower borders in front
of the umbo. The inferior border gives an evenly convex out-
line. The urnbones are little-prominent and gently incurved;
they share in the relatively compressed character of the shell.
The greatest height of the valve occurs a little posteriorly to
the umbo.

The surface is covered with very minute and delicate, concentric,
raised linear ornaments, separated by narrow, thread-like striae ; in
the lower half of the valve there are about eight of these raised
lines within the space of a millimetre. The lines are not all of
equal strength, nor are the interspaces equal in breadth, yet
they have a much greater aspect of regularity than that shown
by mere striae and ridges of growth. Behind the umbones, the
flank of each valve passes over into the well-sunk ligament space
without carination, but forming a rounded, blunt, pillow-like mar-
gin which, when observed in profile, conceals the ligament. The
ligament (preserved in some specimens) in an individual measur-
ing 14 mm. in length, extends back from the umbones for a distance

of 5 mm.

* Blake (1), p. 232, pi. x., fig. 2.

152 Annals of the South African Museum.

Dimensions.- (1) (2) (3) (4)

Length 14 . 16 . 17 20mm.

Height, measured at the

middle of the valve 11 . 12 . 13 16

Greatest depth of a

single valve 3 . 3'5 . 3'5 4 ,,

Occurrence. Found in the kloof east-north-east of Eed House, on
the left side of the Zwartkop's Eiver (324), and at Grass Eidge, three
miles east-north-east of Uitenhage (310) ; also in the railway cutting
between milestones 24|-24f on the line from Uitenhage to Graaff-
Eeinet, about three miles from Uitenhage (316). A specimen sent
from the South African Museum is from the Sunday's Eiver,
occurring with HolcostepJianus cf. atherstoni (Sharpe). Mr. Eogers
obtained specimens of this form, in 1905, from the left side of
Coega Valley, half a mile down from the railway (453g, 454g) ;
from a bare slope, W. 30 S. from the middle of Barkly Bridge,
on the farm Olifant's Kop, Sunday's Eiver (24h, 26h, 28h-30h) ;
and from the highest beds in the kloof behind Colchester, left bank
of Sunday's Eiver (493g, 498g).

Remarks. Although this elegant shell shows some variation in
outline and in the degree of inflation, yet the individuals here brought
together agree on the whole very closely and are well characterised.
The chief features of the shell are the generally rounded and soft
outline, the inconspicuous, rounded umbonal region, and the very
compressed form of the valves.

In outline, the shell resembles some specimens of Venus
orbignyana Forbes," from the Lower Greensand of the Isle of
Wight, which differs, however, by the considerably stronger inflation
and the more marked anterior excavation under the umbones.
Meretrix uitenhagensis has a less steeply sloping upper margin
posteriorly to the umbones, and the umbones are less prominent.
The minute concentric linear striee are developed also in Venus
orbignyana, though more faintly marked than in the African shell.

Meretrix parva (Sow.),t also from the Lower Greensand of
England, is more circular in outline, more nearly equilateral, and
considerably more inflated in form.

Meretrix brongniarti (Leymerie)J is a larger and more massive
shell, and even if compared at the same dimensions is seen to be more
elongated and posteriorly attenuated in outline, and more equilateral.

* Forbes (1), p. 240, pi. ii., fig. 5.

t J. de C. Sowerby (1), vol. vi., Tab. 518, figs. 4-6 (1826).

{ Leymerie (2), p. 5, pi. v., fig. 7; pi. vii., fig. 1.

The Invertebrate Fauna of the Uitenhage Series. 153

Meretrix labadyei (d'Archiac),* from the Tourtia of Tournay, has
an almost identical outline, but its valves are much more convex.
The same character distinguishes several other Cretaceous forms
ascribed to Cytherea or Venus, which, when compared with our shell,
are seen to have a very similar outline.

In the shape of the shell, the inconspicuous umbones, and the
compressed form, resemblance is shown to Tapes picteti de Loriol,t
from the Gault of Cosne (Nievre), which, however, is not so elongated
relatively to height, and moreover, has a coarser concentric orna-
mentation. A similar resemblance is shown to Tapes patagonica
Stanton,J from the Belgrano beds (Lower Cretaceous) of Patagonia,
but the points of distinction are plainly seen in the less elongated
figure, the greater inflation, and the coarser concentric ornamenta-
tion of the Patagonian shell.

A shell from supposed Lower Cretaceous strata in the Cameroons
(left bank of Mungo Eiver), described by von Koenen as Cytherea
wohltmanni, differs from Meretrix uitenhayensis in the greater
inflation, the more inequilateral form, and the prominence of the
umbonal region.

A word may be added regarding the generic position of this form.
In no specimen has the interior been seen, so that a precise generic
determination cannot really be made with certainty ; but a com-
parison with other species in which the hinge-teeth are known,
justifies a provisional reference to Meretrix, if this name be applied
in the broad sense in which the name Cytherea has for long been
used, with reference to Cretaceous forms. The tendency of modern
work is to set closer and closer limits to the application of long-
established generic names amongst lamellibranchs, as in other
classes of Mollusca. It is highly probable, when evidence of
internal characters can be obtained, that an extension of this
principle to Cretaceous forms, on the lines carried out in the classifi-
cation of recent and Tertiary species, may eventually show the in-
applicability of the name Meretrix (equivalent to Cytherea as
commonly used) to such a form as the one here described. From
practical considerations, however, it will often be necessary, as in
the present case, to continue to utilise in a broad sense a name
which, though perhaps technically wrong, conveys as definite a
meaning as the available evidence for the time being allows.

* d'Archiac (1), p. 303, pi. xiv., fig. 7. t ^ e Loriol (4), p. 64, pi. vii., fig. 21.

I Stanton (3), p. 23, pi. iv., figs. 12, 13.

von Koenen (1), p. 36, Taf. iv., tigs. 6, 8, 9 ; since shown to be of Upper
Cretaceous age, see Solger (1).

154 Annals of the South African Museum.

GENUS PSAMMOBIA Lamarck.

PSAMMOBIA ATHERSTONI Sharpe.

1856. Psammobia atherstoni D. Sharpe, Trans. Geol. Soc. Lond.,
ser. 2, vol. vii., p. 196, pi. xxii., fig. 11.

This was found in the cliff below the old school-house at Dun-
brodie, on the right bank of Sunday's Eiver (321, 325). A specimen
from this locality, submitted to me, agrees very closely with the
largest individual figured by Sharpe. The Dunbrodie specimen
measures 25 mm. in length and 15 mm. in height ; it is a left valve,
having the characteristic compressed form and flattened flank and
very inconspicuous urobo. Sharpe records this shell from the
" Sunday River near Enon, in a grit sometimes full of the casts of
the shell." At Dunbrodie Psammobia atherstoni is associated with
Actceonina atherstoni, Turbo atherstoni, Pccten cottaldinus, and other
forms. Messrs. Rogers and Schwarz mentioned the occurrence of
Psammobia atherstoni in the Wood Bed series of the Bezuidenhout's
River below Blue Cliff,* but this name was probably applied to
specimens which I have referred to the genus Unio.

The generic determination of this species must be regarded as
provisional. No specimens showing satisfactorily the internal
characters have been available for study, and true relationship with
Psammobia (or Gari Schumacher, if this be regarded to have
synonymic value) still remains to be proved. The propriety of
considering the name Gari to possess the same significance as that
which has been ascribed to Psammobia (sensu lato) is perhaps still
open to question. The shells we are dealing with in the present
instance are smooth and without radial markings, and this fact,
together with the slight doubt concerning even the broader generic
relationships, seems to justify the retention, for the time being, of
Lamarck's more familiar and more widely accepted name. It
appears reasonable, under the circumstances, to use as a provisional
measure a nomenclature which clearly indicates the supposed rela-
tionships, though it may perhaps be technically erroneous. To
hazard a "correction" of nomenclature on an insecure basis of
imperfect knowledge is a step for which it would probably be more
difficult to find justification.

* Eogers and Schwarz (1), p. 13.

The Invertebrate Fauna of the Uitenhage Series. 155

GENUS SOLECUETUS H. D. de Blainville.

SOLECURTUS Sp.

Text-figure 1.

Description of a Single Specimen. The specimen is a right valve,
slightly imperfect at the siphonal margin. The umbo is very incon-
spicuous, and is situated slightly in front of the middle of the shell.
The upper margin slopes down very gently in front and behind the
umbo. The frontal margin is short and has a rounded convex
profile ; towards this margin the upper and lower valve-borders very
gently converge. Towards the siphonal margin the valve has a
slightly greater height than in the anterior half. The surface is
marked by lines of accretion, but no trace of ornamentation has
been observed. The inflation of the valve is very slight.

FIG. 1. Solccurtits sp., nat. size. Survey Collection.

Dimensions.

Length (if restored) 31 mm.

Height at the umbo 12 ,,

Depth of the valve, about 3 ,,

Length of siphonal margin 9 , ,

Length of frontal margin 6 , ,

Occurrence. Found at Grass Eidge, 3 miles east-north-east of
Uitenhage (310).

Remarks. The single specimen examined is not quite complete
at the posterior end, and, unfortunately, the surface is so preserved
as to leave no traces of faint linear sculpture, if such markings were
ever present. By following the growth-lines it is possible to arrive
at the original form of the posterior border. In shape the shell
agrees very closely with Solecurtus warburtoni Forbes,* from the
Aptian of Atherfield (Isle of Wight) ; the umbo occupies a similar
position, and the valve is anteriorly rounded, with diminished height
towards the frontal border, while posteriorly the height is greater.
The only clearly observable difference is that, in specimens of similar
dimensions, the measurement from the umbo vertically to the
inferior margin is relatively rather smaller in the English shell.

* Forbes (1), p. 237, pi. ii., fig. 1.

156 Annals of the South African Museum.

This also has very faint and delicate radial, linear ornamentation on
the anterior part of the flank, but the African shell may originally
have borne similar minute sculpture, and on this point nothing
further can at present be said.

Similar forms were described by d'Orbigny, under the name
Solen, from the Lower Cretaceous (S. robinaldinus) ' and from the
Chalk (S. cequalis).} Gueranger has figured a shell from the Ceno-
manian of the Sarthe under the name Solecurtus (equalis,l to which
our specimen appears to bear a close resemblance, so far as com-
parison is possible.

GENUS MACTEA Linn.

MACTRA ? DUBIA sp. nov.

Plate VIII., figs. 2, 2a, 3, 3a.

Description. The shell has greater length than height, and the
umbones are situated at rather more than one-third of the shell's
total length from the anterior extremity. The inflation is moderate,
and most strong just above the middle of the valves. The umbones
are somewhat weakly developed and are not very prominent. The
cardinal margin and the posterior margin form together a curved
outline which passes down, posteriorly very steeply, to a marked
angular junction with the lower border. Prom the umbo a well-
marked carina passes obliquely across the posterior part of the
valve, down to the postero-inferior angle of the valve-margin. The
carina marks off a flattened or very slightly concave postero-superior
area which occupies less than one-quarter of the total valve-surface.
In front of the umbo the valve-margin slopes down to form a some-
what sharply curved outline in front, the foremost point of the
anterior margin falling well within the lower half of the shell. The
long pallial margin shows a broadly convex outline, sometimes
slightly flattened towards the posterior end. The greatest height of
the valve occurs at the umbonal part. The surface is smooth, with
numerous delicate growth-lines, and shows closely spaced radial
rows of very minute punctae, only visible under a lens.

Dimensions. (1) (2)

Length 15 . 15mm.

Height, measured from the umbo 12 . 12 ,,

Greatest depth of a single valve 4 . 3 '5 ,,

* d'Orbigny (3), p. 320, pi. 350, figs. 1, 2 (1845).
t d'Orbigny (3), p. 321, pi. 350, figs. 5-7 (1845).
J Gueranger (1), pi. xv., fig. 3.

The Invertebrate Fauna of tJie Uitenhage Series. 157

Occurrence. Found in the cliff on Buck Kraal, Sunday's Eiver

Remarks. The doubtful reference of this form to Mactra is, it
must be admitted, unsatisfactory. The generic position, in fact, is
very uncertain, for the characters of the hinge are unknown and the
ligament space is relatively extensive. The general aspect of the
shell does not accord well with either Cyprina or Meretrix. The
specimens share with Mactra angulata J. de C. Sow. (Blackdown
Beds),* the sub-angular junction of the posterior and inferior margins
and the presence of a denned ridge running obliquely from the
umbo down to the base of the posterior margin. In M. angulata,
however, the shell is more triangular and less ovate in outline, and
the umbonal region is more prominent and tumid, less anteriorly
placed, and less strongly directed forwards.

Mactra warrenana Meek and Hayden (Cretaceous of Dakota) i is
in some degree comparable, but differs by its more trigonal form,
more prominent umbonal region, and the presence of a defined
lunule of relatively large size.

GENUS PLEUEOMYA L. Agassiz.

PLEUROMYA BAINI (Sharpe).

Plate VIII., figs. 4, 4a.

1856. Myacites? bainii D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,
vol. vii., p. 195, pi. xxii., fig. 7.

Occurrence. The locality given by Sharpe was merely " Sunday
Eiver." Specimens were collected by Messrs. Eogers and Schwarz
at Grass Eidge, 3 miles east-north-east of Uitenhage (317, 318,
334, 335), from an outcrop of nodular limestone, the highest fossili-
ferous outcrop of this locality, where many characteristic fossils of
the Sunday's Eiver Beds were found. J P. baini was also found to
occur commonly in the railway cutting between milestones 24^-24f
on the line from Uitenhage to Graaff-Eeinet. Mr. Eogers obtained
a very well-preserved specimen from the highest beds in the kloof
behind Colchester, Sunday's Eiver Valley (495g).

Remarks. There is some variation in the form of the shell, and
the inferior margin, though never showing a strongly curved outline,
is not always so straight as depicted in Sharpe's figure. When well

* J. de C. Sowerby (2), p. 341, pi. xvi., fig. 9.
f Meek (2), p. 208, pi. xxx., fig. 7.
| Eogers and Schwarz (1), p. 9.

158 Annals of the South African Museum.

preserved the surface is seen to be covered with closely spaced
radial lines of delicate granules.

Although possessing no highly distinctive external features, this
shell retains a certain characteristic aspect, while approaching some-
what closely to some European representatives of the genus. It
may usually be easily distinguished from the larger Pleuromya
lutraria (Krauss),* another characteristic shell of the Sunday's
Eiver Beds, with which it is associated. If mature specimens be
compared, P. lutraria is distinguished by its much larger dimen-
sions. If small specimens of this be brought into comparison with
mature examples of P. baini of equal size, there is a considerable
general similarity between them ; but P. lutraria, when young, has
a definite ornamentation of rounded concentric ribs, whereas P. baini
is a smooth shell, with the surface only marked by lines of growth
and minute granules. The umbones of P. lutraria are rather more
prominent, and are perhaps sitiiated a little nearer to the anterior
extremity, while the margin in front of the umbo slopes down more
suddenly than in P. baini.

Pleuromya neocomiensis (d'Orb.) \ is a similarly elongated form,
but is distinguished by marked anterior truncation. P. rostrata
(d'Orb.) {' is more produced posteriorly and has a more curved
inferior outline. Some individuals of P. baini closely resemble a
Pleuromya from the Lower Greensand of the Isle of Wight which
has been considered, apparently erroneously, to represent d'Orbigny's
P. neocomiensis, and is much more likely to be identical with
P. scliroderi (Wollemann), from the Neocomian of North Germany.
To judge from Dr. Wollemann's description and figure, P. scliroderi
approaches very closely indeed to P. baini.

Another shell which shows considerable resemblance to P. baini
occurs in the Neocomian sandstone of the Teutoburger Wald, and
was ascribed by Weerth, || though perhaps erroneously, to d'Orbigny's
Panopcea neocomiensis ; the general similarity is great, but the shell
figured by Weerth has an inferior margin presenting a rather more
curved outline.

The shell from the Neocomian of North Germany, described by
F. A. Koemer 1; as Pleuromya solenoides, is an elongated form with

* Krauss (2), p. 447, Tab. xlvii., fig. 1.

t d'Orbigny (3), p. 329, pi. 353, figs. 3-8 (1845).

I d'Orbigny (3), p. 333, pi. 355, figs. 3, 4 (1845).

Wollemann (1), p. 126, Taf. v., fig. 7 (as Panoptca).

|| Weerth (1), p. 37, Taf. viii., fig. 7.

[ F. A. Eoemer (3), p. 330, pi. xli., figs. 20 21.

The Invertebrate Fauna of the Uitenhage Series. 159

straight upper margin and little-curved lower margin. P. baini
differs from this in being more equilateral and less produced and
truncated posteriorly.

As regards the generic position of this shell, although the hinge-
characters have not been described and I have been unable to inves-
tigate them, there is no reason to doubt that we are dealing with a
typical Pleuromya, to which genus belong also, in all probability, the
majority of the Neocomian forms to which the name Panop&a was
formerly applied. There is no reason to suppose that the hinge-
characters of P. baini differ essentially from those of P. lutraria,
with which it is associated ; and although it was the nature of the
hinge that led Krauss to propose for the latter form the separate
generic name Anoplomya, it appears plain from Terquem's : ' : detailed
studies of the genus Pleuromya that the name proposed by Krauss
must be regarded as a synonym a view already adopted by Zittel.f

GENUS GONIOMYA J. L. E. Agassiz.

GONIOMYA Sp.

This genus is represented by a single specimen of a small right
valve. It is unfortunately imperfect, with a large part of the shell
substance removed, and it is embedded in a very hard matrix which
cannot be removed without further injury to the delicate shell.

Description. The valve has little convexity and is posteriorly well
produced. The urnbo is rather pointed and prominent and shows
a weak fold of the valve-surface extending for a short distance on
its posterior side. The shell-substance is very thin and delicate.
The ribbed ornaments of the surface are developed already close
to the umbonal apex, where the anterior and posterior ribs are very
delicate and closely spaced and are steeply inclined to form the
V-pattern. The angle of the V is very acute, and the successive
angles formed by the junction of the ribs of the two series are
situated at first just below the umbonal apex, and then below one
another on the flank on a slightly oblique line posteriorly inclined,
so that the lowest angles of the sculpture are situated more back-
wardly than those above. The most backwardly situated ribs,
which do not contribute to the angular ornamentation, are pos-
teriorly inclined when traced down from their upper terminations.
Posteriorly to the umbo there is a broad smooth area devoid of
sculpture on the upper part of the valve, but this is not sharply
demarcated from the flank.

* Terquem (1) ; Terquem (2). t Zittel (5), p. 125.

160 Annals of the South African Museum.

The anterior ribs are narrower and perhaps rather more pro-
minent than the posterior ribs. Delicate and fairly regular lines
of growth cross the surface of ribs and interspaces alike. The
surface of the shell, where well preserved, is seen to be covered
by very numerous and delicate, radially disposed lines of minute
granules.

Dimensions. The anterior part of the specimen is hidden in the
matrix, but to judge from the course taken by the anterior growth-
lines, the length of the valve may be fairly accurately estimated as
22 mm. The umbo is situated at about one-third of the total length
from the anterior extremity, assuming that the above length-measure-
ment is correct. The height of the valve at the umbonal region is
13 mm. The height of the siphonal border is 8 mm.

Occurrence. Collected by Mr. Eogers from the highest beds in
the kloof behind Colchester, Sunday's Eiver Valley (489g).

Remarks. This is the first specimen of Goniomya recorded from
the Uitenhage Series. The condition of the single valve available
for comparison scarcely justifies a specific determination, and the
question of relationships cannot be usefully discussed without better
material for study but the specimen probably represents a new
species.

It is interesting to find the genus Goniomya represented here, in
strata which yield the peculiar Trigonice of the group of T. vau
which so strongly simulate the genus Goniomya both in the
arrangement of the costate ornaments and in the general habit.
Though the specimen above described at once recalls these
Trigonice, it may be readily distinguished from them by the very
thin shell-wall and the presence of surface granulation, as well as
by the details of ornamentation. In Trigonia vau Sharpe and its
allies, the neanic stage is ornamented by concentric ribs passing
across the flank and area, and the inclination of the ribs with
resulting angular pattern is not produced until the early adult
stage. In this Goniomya, the costse formed an anterior and a pos-
terior series, steeply inclined, already in the neanic stage, at a very
much shorter distance from the umbonal apex than in the Trigonice,
mentioned.

GENUS THRACIA W. E. Leach.

THEACIA sp.
Plate VIII., fig. 5.

A single specimen, which must be ascribed to this genus, is
scarcely so well preserved or so strongly characterised as to allow

TJie Invertebrate Fauna of the Uitenhage Series. 161

of a thorough comparison with known forms, or to warrant the use
of a new specific name. The posterior end is slightly broken away
so that the exact outline of the shell here cannot be clearly seen.
The length is considerably greater than the height ; the valves are
of a compressed form, most convex in the uppermost third, more
flattened below. Theumbones are little conspicuous and are directed
very slightly backwardly ; they are situated just posteriorly to the
middle of the shell. The upper margin slopes down very gently in
front of the umbo, with an almost straight profile which passes into
the evenly convex outline of the frontal margin. The posterior com-
pressed area is well defined, especially in the neighbourhood of the
umbo.

Dimensions.

Length 40 mm.

Height, measured from the umbo 29 ,,

Depth of one valve 8 ,,

Occurrence. In the railway cutting between milestones 24-| 24f
on the line between Uitenhage and Graaff-Keinet, about three miles
from Uitenhage (331).

Remarks. Comparison with English specimens of Thracia pliil-
lipsi Boem.,* from the Speeton Clay, shows that these differ widely
from the shell before us by their much more inflated and prominent
umbonal region and more steeply sloping antero-superior margin.
In general form and outline the African shell shows close simi-
larity to Thracia subtruncata Meek, from the Cretaceous strata
of Sucia Islands (British Columbia).! The outline also some-
what nearly resembles that of Thracia gracilis Meek and
Hayden,} from Cretaceous sandstone at the mouth of the Judith
River (on the Missouri), but the latter is rather more elongated
posteriorly.

Another shell which may be brought into close comparison occurs
in the Aptian of Spain (Obon and Utrillas) and was described by
Coquand as Periploma lorieri. To judge by Coquand's figure, this
differs from the Uitenhage form by the steeper inclination of the
carinal ridge and the more nearly parallel direction of the upper and
lower margins anteriorly to the umbo.

* Mya depressa Phillips, (1), Tab. 2, fig. 8 (non Sow.); Thracia phillipsiF. A.
Boemer, (2), p. 74, Taf. 10, fig. 1 (1841).
t Whiteaves (2), p. 140, pi. 17, fig. 7.
J Meek (2), p. 224, pi. 39, fig. 6.
Coquand (1), p. 100, pi. ix., figs. 5, 6.

162 Annals of the South African Museum.

Thracia robinaldina (d'Orbigny) :;: is distinguished by its more
oblong and less ovate outline, and its more conspicuous umbones.
The African shell compares again more closely with the figure of a
specimen from the Lower Cretaceous (Boiling Downs Formation)
of Queensland, ascribed by Etheridge with some doubt to Thracia
primula Hudleston, and figured under the generic name Corimya.}
Less similarity is shown to the specimen originally named Thracia
primula, I preserved in the British Museum (Natural History) ; this
has greater relative height anteriorly to the umbo, a more curved
inferior margin, more strongly compressed valves, a less inflated
umbonal region, and less definitely developed posterior carination.

GENUS GASTEOCH^NA L. Spengler.
GASTROCH^NA DOMINICALIS Sharpe.

1856. Gastrochc&na dominicalis D. Sharpe, Trans. Geol. Soc. Lond.,
ser. 2, vol. vii., p. 198, pi. xxiii., fig. 4.

Sharpe described under this name a shell which had bored into
a fragment of wood, found on the Sunday's Eiver " near Enon."
He also mentions the occurrence of Gastrochcsna boring into
fragments of wood and Triyonia-she\l, also from the Sunday's
Eiver.

Fragments of fossil wood bored by Gastrocluena were collected by
Messrs. Eogers and Schwarz from an oyster bed at the base of the
cliff below the old school-house at Dunbrodie, Sunday's Eiver (336).
The shells are concealed within the short calcareous tubes which
line their cavities, and some of these crypts measure about 6 mm.
in length. There is no reason to doubt that these specimens repre-
sent the same form as that described by Sharpe. This shell was
also found boring into lignite in strata of the " Wood Bed " series
of the Bezuidenhout's Eiver below Blue Cliff station.

* d'Orbigny (3), p. 380, pi. 372, figs. 1, 2 (1845).

t Jack and Etheridge (1), p. 481, pi. 28, fig. 11.

} Hudleston (1), p. 245, pi. ix., fig. 7.

Sharpe spoke of this fragment as " bone," but this was corrected on p. 228
of Sharpe's paper and by Prof. T. Rupert Jones in an editorial footnote to Tate's
paper in 1867, Tate (1) , p. 155.

The Invertebrate Fauna of the Uitenhage Series. 163

CLASS GASTEROPODA.

GENUS PATELLA Linnteus.
PATELLA CAPERATA Tate.

1867. Patella caperata B. Tate, Quart. Journ. Geol. Soc., vol. xxiii.,
p. 152, pi. vii., fig. 1.

Occurrence. A single specimen, on Exocjyra imbricata Krauss,
from the collection of the South African Museum, was obtained at
Coega. Tate's figured specimen, in the collection of the Geological
Society (No. 11,003), is labelled " Prince Alfred's Eest " (Sunday's
Eiver).

Remarks. The specimen from Coega is more strongly elevated
in form than Tate's original type. The apex in both is excentric
in position, and as a result of this, the outline from the apex to the
margin on the shorter side of the shell falls more steeply than that
on the opposite side. Annular markings and growth-lines are more
marked on Tate's specimen, because its surface has suffered less from
weathering than that of the other individual.

The figured type of this species measures 28 mm. in longest
diameter at the base, and the measurement at right angles to
this, across the shortest diameter, is 22 mm. The height is 10 mm.
The true dimensions of the Coega specimen cannot be measured
satisfactorily because the shell has so suffered from weathering that
its original margins are not preserved. There are indications that
a firmly established station had been taken up, on the Exogyra. In
one place, on the surface of the Exogyra, in the position which the
serrated margin of the Patella formerly occupied, there are indenta-
tions which correspond with the terminations of the costse on the
Patella.

GENUS PLEUBOTOMABIA J. L. M. Defrance.
PLEUROTOMARIA sp.

Description. A large, ill-preserved and much-weathered specimen
consists of at least five whorls. The body- whorl is flattened on its
under side and its outer margin is obtusely angular ; the surface of
the whorl above this margin is very slightly convex. The suture
falls at the bluntly angular margin of the whorls, so that the general
figure of the shell in profile is that of a cone with little excavation at
the sutural depressions and inconspicuous convexity of the outline

164 Annals of the South African Museum.

between these. The mouth aperture is of transversely oval form.
The surface is so far weathered that no sculpture can be seen except
on the base, where traces of numerous raised longitudinal (spiral)
ornamenting ridges, crossed by ridges of growth, may be detected.
The spiral angle is about 85.
Dimensions.

Height 90 mm.

Greatest width of base 100 ,,

Height of aperture 44 , ,

Approximate breadth of aperture 55 ,,

Occurrence. Collected by Miss Wilman at Coega.
Remarks. This single specimen is the first representative of its
genus yielded by the Uitenhage beds, and it is a matter for regret
that the preservation is so indifferent that a more precise description
of characters is impossible. We must await the addition of further
material before the nature of the sculpture can be ascertained and
the nearer relationships made clearer ; in the meantime it would be
unjustifiable to propose a specific name. In general figure the shell
is not unlike Pleurotomaria (Leptomaria) tithonia Zittel,* but the
spiral angle is rather wider than in that form, and if viewed in
aperture-aspect it is seen that the whorl is deeper in section in the
Uitenhage shell, and the mouth much less transversely elongated.

The form of the whorls is not unlike that of Pleurotomaria
tardensis Stanton,f from the Belgrano beds (Lower Cretaceous) of
Patagonia, which is also similar by reason of its large dimen-
sions ; but in the Coega specimen the outer margin of the body-
whorl is less broadly rounded and the spiral angle is rather
smaller.

GENUS TUEBO Linnaeus.

TURBO ATHEBSTONI Sharpe.

Plate VIII., figs. 6, 6a, 7, la, 7b.

1856. Turbo atherstoni D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

vol. vii., p. 200, pi. xxviii., figs. 15, 16.
1856. Turbo baini D. Sharpe, Ibid., p. 200, pi. xxviii., figs. 17, 18.

Supplementary Descriptive Note. The shell consists of about six
whorls, the first four of which have a smooth, flat, or very slightly
concave upper surface, sloping down very gently from the spiral
suture. The outer side of the whorl is truncated, with a sharp and

* Zittel (4), p. 337, Taf. 50, figs. 5, 6.
t Stanton (3), p. 29, pi. vii., figs. 1, 2.

The Invertebrate Fauna of the Uitenhage Scries. 165

prominently projecting longitudinal (spiral) keel limiting the trun-
cated band above, and also a similar one below. Under the lower
marginal keel the surface of the whorl is gently convex in profile and
slopes rapidly in. Just below the lower marginal keel there is a
narrow, deep spiral sulcus bounded below by a delicate thread-like
longitudinal keel. Below this there are four weaker raised spiral
lines which are already developed in the third whorl, if not before.
Until the fourth whorl is reached, the slightly concave outer band
between the prominent marginal keels is smooth, but before this
whorl is completed, in some individuals at its commencement, a
weaker central keel appears.

The spiral angle varies somewhat, and averages about 70. The
last two whorls expand rapidly. The spiral suture is situated on the
lower marginal keel of the whorls. The mouth has an angular
outer lip, corresponding with the form of the whorls. The outer
margin of the aperture is thin and sharp ; under the columella the
margin is slightly thickened.
Dimensions.

) Height of a specimen with four complete whorls 5 mm.

I Diameter of the fourth whorl 4 ,,

( Height of a specimen w r ith five (or six ?) whorls 14 ,,

\ Diameter of the body-whorl 9 ,,

Occurrence. Found in the cliff below the old school-house at
Dunbrodie on the Sunday's Eiver (351), where it is abundant.
Specimens in the collection of the Geological Society of London are
from "the lowest strata of the Zwartkop crag," and "greenish grit
with Ostrece from Sunday River," as recorded by Sharpe. This form
was obtained by Mr. Rogers in 1905 in a cliff W. 20 S. from Comley's
house, right bank of Sunday's River (95h).

Remarks. It is clear from a careful examination of the specimens
in the collection of the Geological Society, together with the new
materials supplied to me, that Sharpe was in error when he separated
the shells described by him as Turbo atlicrstoni and T. baini. The
differences which caused him to make the separation are in reality
due to changes in the nature of the ornamentation during successive
stages of growth ; but the specimens at his disposal were very small
and the materials so comparatively unsatisfactory that the mistake
is not surprising. All specimens having the two marginal keels
only, and consequently agreeing with the type of T. atherstoni, are
very small individuals which have not reached the stage in which, by
the intercalation of a central marginal keel and the more prominent
development of the spiral ornaments on the lower part of the whorl,

12

166 Annals of the South African Museum.

the features of T. baini are exhibited. The change in characters is
found to be very constant, although some slight variation may be
shown as regards the exact period, in relation to the precise number
of whorls developed, when the additional keel makes its appearance.
In this connection also, differences of preservation prove very decep-
tive. As the adult stage becomes completed, it is noticeable how
relatively rapidly, by the greater expansion of the whorls, the shell
gains in length when compared with the proportional dimensions of
the young individual.

This well-characterised form seems to resemble very closely Turbo
reedi W. Keeping," from the Lower Cretaceous of England, and may
be more aptly compared with this than with any other European
shell with which I am acquainted. The type of ornamentation is
essentially similar as regards major features, and while Keeping
gave no account of the variation of sculptural characters when
traced through successive stages, it seems probable that the central
marginal keel became added between the two prominent keels during
the course of individual growth, just as in T. atherstoni. The
principal points by which the English shell differs, are seen in the
greater number of spiral ribs on the lower part of the body-whorl
and in the very delicate spiral striation and resulting granulation of
the surface.

A somewhat analogous type of ornamentation is exhibited by a
shell from the Upper Neocomian of Utrillas (Teruel), described as a
Trochus, under the name T. maestrei, by de Verneuil and Loriere. t
This, however, shows only two marginal spiral keels, while the base
of the body-whorl is ornamented by several strong, well-spaced
keels. Further points of distinction are the fine spiral lines which
ornament the whorls, and the wider spiral angle.

A form which perhaps merits mention by way of comparison
occurs in the Upper Jurassic Aitcella-beds of Novaya Zemlya, and
was ascribed by Tullberg to Minister's Turbo capitaneus.\ This has
ornaments similar to those of T. atherstoni, but there is less overlap
of the whorls, the upper and lower marginal keels are less
prominent and of similar strength to the central one, and the
outline of the whorl is more rounded and less angular. There are
apparently other differences, so far as it is possible to make com-
parison with the figure of the relatively large specimen figured by
Tullberg.

* Keeping (1), p. 97, pi. iii., fig. 13.

t de Verneuil and Loriere (1), p. 23, pi. ii., fig. 9.

i Tullberg (1), p. 9, pi. ii., fig. 1.

The Invertebrate Fauna of the Uitenhage Series. 167

There is much closer resemblance, again, to Turbo bitropistus
Ascher, from the Grodischter Schichten (of Hauterivian age) in
Silesia.* This, however, has a relatively narrower band between the
two sharp marginal keels, and moreover, the surface of the shell is
ornamented with numerous and delicate spiral lines.

TURBO ROGEBSI sp. nov.
Plate VIII., figs. 8, 8, 8b.

Description. The shell consists of six whorls, possibly more.
The whorls are flattened above, and the upper surface slopes very
gently away from the spiral suture, terminating abruptly at a
prominent marginal keel which marks the upper limit of the outer
truncated face of the whorl. This truncated portion is straight in
profile in the earlier whorls, but becomes gently but distinctly
convex in the fourth or fifth whorls ; it is bounded below by a
prominent marginal spiral keel similar to the one above. Already in
the third whorl, a central spiral keel makes its appearance, and this
soon grows in strength, so that in the fourth whorl it is almost, if
not quite, as prominent as the two pre-existing marginal keels, and
subsequently it remains quite as conspicuous as these. At the com-
pletion of the fourth whorl, or perhaps a little before this, two
additional well-marked, thread-like, raised linear ornaments become
intercalated, one above and one below the central marginal keel.
The lower part of the body-whorl shows a rounded, convex surface
ornamented by about five delicate spiral keels, of which the upper-
most is the strongest and is separated by a narrow sulcus from the
lowest marginal keel.

The spiral angle is about 75. The overlap of the whorls is such
that the suture coincides with the lowest keel of the marginal area.
In the earlier whorls the outwardly truncated form of the whorls
gives a step-like profile with vertical outer face ; in the later growth-
stages (about the fifth whorl) the upper surface of the whorls becomes
a little more convex and less abruptly demarcated from the marginal
face, while the latter becomes more convex and merges gradually
into the lower part of the whorl, so that the marginal truncation
becomes successively obscure and lost and the whorl attains an
aspect of even convexity. The mouth is then almost circular in
outline, with thin outer lip not showing angularity of form. In the
fifth whorl an indistinct and weakly developed sulcation appears on
the upper surface of the whorl, immediately above the upper marginal
keel and running parallel to it.

* Ascher (1), p. 139 [5], pi. xii. [i.], figs, la-lc.

168 Annals of the South African Museum.

Dimensions.

Height of specimen with five whorls ... 11 mm.
Diameter of the fifth whorl 7 ,,

Occurrence. In the cliff below the old school-house at Dunbrodie,
on the right bank of Sunday's Eiver (282).

Remarks. This shell, represented in the collection by only two
specimens, appears to be so closely related to Turbo atherstoni that
at first I felt some hesitation in definitely separating it. A careful
comparison with the specimens of T. atJterstoni from Dunbrodie and
those in the museum of the Geological Society shows, however, that
it possesses distinctive characters which appear in no specimen of
T. atherstoni that I have examined ; the points of difference are as
follows. In T. atherstoni the upper and lower marginal keels are
very prominent and the outer area of the whorl concave in profile ;
the marginal keels in T. rogersi are less pronounced and the outer
truncated area flat and vertical in profile in the earlier whorls. The
central keel of T. rogersi makes its appearance much earlier and
gains in strength so that it rapidly becomes as prominent as the two
original peripheral keels ; this equality is not attained in T. ather-
stoni. In T. rogersi two additional raised ornamenting lines appear
on the peripheral area, while larger specimens of T. atherstoni, illus-
trating a more advanced stage of growth, have shown no similar
development. Another feature of distinction is seen in the manner
in which the whorls of T. rogersi lose their angularity of form with
advancing growth and present an even convexity of surface not seen
in the largest specimens of T. atherstoni. It is impossible to speak
with certainty of any differences in the spiral angle, but the com-
parison of additional specimens, when these are forthcoming, may
possibly show that the angle of T. rogersi rather exceeds that of
T. atherstoni.

A Turbo described by Zittel from the Stramberg beds under the name
Turbo eryx d'Orb., var. major'''- is of similar type to T. rogersi, but
differs in the more expanded form and wider spiral angle, and in the
details of the spiral linear ornaments.

TURBO MINUTULUS sp. nov.
Plate VIII., figs. 9, 9a, 9b.

Description. The shell consists of at least four whorls, and these
are flattened above and sloping gently away from the suture, but
vertically truncated at the periphery. The truncated marginal band

* Zittel (4), p. 321, pi. xlviii., fig. 10.

The Invertebrate Fauna of the Uitenhage Series. 169

is bounded above and below by a prominent longitudinal (spiral)
keel ; these two principal keels are of equal strength, and while the
lower one is smooth (or nearly so), the upper one is bluntly nodular.
Each of the rounded nodes of this keel is prolonged in wedge shape
downwards into the marginal band, but the apices of the wedges die
out before they reach the lower keel. At the end of the fourth
whorl the wedge-like ornaments appear to be losing the well-defined
character they possessed at the beginning of this whorl and before,
while early in the fourth whorl an intermediate (central) peripheral
keel makes its appearance, though it is less strongly developed than
the two pre-existing marginal keels.

Below T the marginal band the surface of the body- whorl has an
evenly rounded outline ; it is ornamented by three prominent spiral
keels, one situated near the lower marginal keel and almost as
strongly developed as this, the other two, rather weaker, below and
in close proximity to one another. The spiral angle is about 70.
The spiral suture is immediately under the lower marginal keel.
The mouth aperture has equal breadth and height. The outer lip is
sharp and thin and slightly angular, in accordance with the form of
the whorl. The inner lip is slightly thickened.

Dimensions.

Height of specimen with four complete whorls 8 mm.

Greatest diameter of the last whorl 6 , ,

Height of aperture 4 ,,

Occurrence. Cliff below the old school-house at Dunbrodie, on the
right bank of Sunday's Eiver (305, 351).

Remarks. This form, though apparently a near ally of Turbo
atherstoni and T. rogersi, is readily separable from these by the
regular nodose ornaments on the upper marginal keel and peripheral
zone below this, as well as by the stronger development and smaller
number of the spiral keels on the under surface of the whorl, below
the lower marginal keel. It seems probable that this shell would
attain dimensions as large as those reached by T. atherstoni, and the
appearance of a central marginal keel in the last whorl of the largest
individual examined very strongly recalls the increase of the spiral
ornaments in the associated forms mentioned.

TUKBO sp.

An imperfect specimen, consisting of a body- whorl deprived of the
spire, may be provisionally referred to this genus. The greatest
breadth across the base is 8 mm., and the diameter of the mouth
aperture which is nearly circular, measures 4 mm. The surface is

170 Annals of the South African Museum.

ornamented by rounded, shallow spiral grooves, and blunt spiral
ridges having little prominence. These ornaments are so disposed
that above the peripheral area a weakly marked ridge forms the
outer limit of a narrow shoulder which is without spiral sculpture,
while on the lower part of the peripheral area is a spiral groove,
slightly broader than the three remaining grooves which lie below
it. The surface is crossed transversely by numerous and well-
marked ridges and furrows of growth, which are so regularly
developed as almost to appear to constitute part of the sculpture.

Occurrence. This was found, in association with the three forms
previously described, in the cliff below the old school-house at
Dunbrodie, Sunday's Eiver (351).

Remarks. This imperfectly preserved specimen is quite distinct
from the other forms assigned to the genus Turbo, described above,
but it will be well to await the collection of further material before
applying a specific name. It can hardly even be said that the generic
position is established beyond doubt. Although the rounded form
of the whorl and the impressed spiral grooves might suggest at first
sight that we are dealing with a fragment of an immature Neritopsis ?
turbinata Sharpe, :1: a comparison with Sharpe's specimens shows
that this is not the case. The spiral ornaments lack the regularity,
and the grooves are broader, less sharply incised and less regularly
spaced than those of Sharpe's type. A complete comparison, it is
true, is difficult, since the examples of Neritop&is 1 tiirbinata pre-
served in the Geological Society's museum are all much larger than
the fragmentary specimen here described, and the involution of
Sharpe's shell is such that it is not possible to make satisfactory
comparison at the same stage of growth ; but the observable diffe-
rences in the ornamentation are in themselves sufficiently significant.

Considerable resemblance is shown to a shell from the Neocomian
of German East Africa described by G. Miiller as Delpliinula afri-
cana,} such similarity, in fact, as to suggest that we are dealing
with a closely allied form. The shape and proportions of the body-
whorl appear alike, and in each case there is the strong spiral groove
on the lower part of the peripheral area. In the figure of DelpUinula
africana a spiral keel is seen, situated between this groove and the
keel which defines the shoulder above. In the Dunbrodie specimen,
which is of smaller dimensions than Miiller's type, the beginnings of
a similar keel, though weakly developed, are becoming apparent in
the most advanced portion of the whorl. As regards other charac-

* Sharpe (1), p. 198, pi. xxiii., fig. 5.
f G. Miiller (1), p. 557, Taf. xix., fig. 11.

The Invertebrate Fauna of tJie Uitenhage Series. 171

ters, it is not easy to make a detailed comparison, and it must suffice
to have drawn attention to the apparently close resemblance between
these shells. In his employment of the generic name Delphinula
for forms such as these, Miiller follows the usage of Stoliczka.
Fischer has restricted the use of the name to living types, but it has
been applied to certain Turbo-like shells of Jurassic and Cretaceous
age by several leading authorities." In the case of the single speci-
men here dealt with, while it may be considered most expedient in
the meantime to use the name Turbo as one of convenience, it
appears not improbable that better preserved material may even-
tually show that a separation is necessary.

In the form of the whorl and the ornamentation great similarity
is also shown to the figure of a specimen from the Aptian of Sainte-
Croix, identified by Pictet and Campiche, perhaps wrongly, with
Turbo mnnitus Forbes, t The African specimen is much smaller
than this Swiss individual, but so far as a comparison with the figure
is possible, the general agreement is very close. The specimen
depicted by Pictet and Campiche in fig. 1 of the same plate has a
taller spire and less expanded whorls than the original of fig. 3, and
agrees much more closely with the typical T. munitus from the
English Lower Greensand, and correspondingly less closely with
this specimen from South Africa, apart from the differences in the
ornamentation.

GENUS NATICA Lamarck.

NATICA UITENHAGENSIS sp. nov.

Plate VIII., figs. 11, lla; ? figs. 10, lOa.

Description. The shell consists of at least five whorls. The spire
is very short, the body-whorl overlapping rather more than one-half
of the preceding whorl and expanding relatively rapidly ; the body-
whorl occupies rather more than two-thirds of the whole height of
the shell. The spiral suture is somewhat deeply impressed though
not definitely channelled ; the upper part of each whorl, adjacent to
the suture, is slightly flattened to form a narrow rounded ledge, as
seen in profile, the outer limit of which is not sharply defined, but
forms a curved outline passing down into the rather flattened upper
half of the whorl. This flattening of the whorl above the middle
zone is most marked in the body-whorl of a well-grown individual,
and even a slight depression of the surface here may be developed

* See remarks by Huclleston and Wilson (1), p. 20 ; also Stoliczka (1), p. 368

(1868).
t Pictet and Campiche (1), 2* Partie, p. 480, pi. Ixxxiv., figs. 3-3c (1863).

172 Annals of the South African Museum.

during the fifth whorl. At the middle zone, and below, the whorl is
evenly convex in outline.

The shell-wall is thick. The surface of the whorls is marked by
numerous distinct, transverse growth-lines, which, however, do not
form an obtrusive feature. The spiral angle is about 90. The
inner lip of the aperture is considerably thickened and forms a raised
callus which bounds a well-impressed though closed umbilical slit.
Just below this, and adjacent to the inner lip, the surface of the
whorl presents a narrow, slightly flattened space. The height of the
aperture is much greater than the breadth.

Dimensions. (1) (2)

Height of shell 15 . 30mm.

Height of body- whorl at the aperture 11 . 24 ,,
Greatest width at aperture 8 . 12 ,,

Occurrence. Two well-preserved specimens in the collection of
the Geological Society of London are labelled " Zwartkop, Dr.
Atherstone, 1876." A specimen from the railway cutting between
milestones 24^ 24f on the railway from Uitenhage to Graaff-Eeinet,
about three miles from Uitenhage (350), is referred with some doubt
to the same species.

Remarks. The specimen from the railway cutting (350) may per-
haps be identical with Atherstone's two individuals, upon which this
species is founded. It differs from them by its rather lower spire
and slightly wider spiral angle, and by the more complete overlapping
of the whorls. These differences may, however, be due to individual
variation.

Natica atherstoni Sharpe, :;: another Uitenhage shell, is distin-
guished by its much more slender and elongated figure and narrow
apical angle (75).

Compared at similar dimensions, Natica Icevigata (Desh.) Leym.,f
a Lower Cretaceous form, has much less overlap of the whorls, and
consequently a higher spire and narrower spiral angle. N. dupini
(Desh.) Leym.4 on the other hand, has a shorter spire and wider
spiral angle, with wider and more inflated whorls and less elongated
mouth aperture. The shell figured by d'Orbigny as N. dupini
shows much greater similarity to N. uitenliagensis, but this too has
a wider spiral angle and a broader aperture as well as a distinct
umbilical opening.

* Sharpe (1), p. 200, pi. xxviii., fig. 22.

f Leymerie (2), p. 13, pi. 16, fig. 10.

J Leymerie (2), p. 13, pi. 16, fig. 7.

d'Orbigny (2), p. 158, pi. 173, figs. 5, 6 (1843).

The Invertebrate Fauna of the Uitenhage Series. 173

In form and outline and in the shape of the aperture great resem-
blance is shown to the shell from the Stramberg beds figured by
Zittel :;: as Natica (Ampullina) elegans Sow. ; this differs, however,
by the slightly less overlap of the whorls and by the thinner shell ;
the inner lip is considerably less thickened than in the African form.
It seems doubtful whether the Stramberg shell is identical with the
English Portland Natica to which Sowerby gave the name IV. elegans,
and although Sowerby's figure t much resembles fig. 23a in plate 45
of Zittel's work, yet typical specimens of the Portland shell show a
higher spire. Natica uitenhagensis is well distinguished from the
English N. elegans by the much thicker shell and the lower spire.
The shells from the " Portlandien superieur " of Boulogne, ascribed
to Natica elegans Sow. by de Loriol and Pellat, { whose determina-
tion was accepted by Hudleston and Wilson, are also slightly
higher in the spire, and the aperture is broader and more rounded
below.

The form described by Stanton as Lunatia constricta, from the
Belgrano beds of Patagonia, j| shows great similarity to N. uiten-
hagensis, but differs apparently by the more strongly impressed
sutures, the slightly less overlap of the whorls, and by the marked
transverse furrows.

NATICA ROGERS: sp. nov.
Plate VIII., figs. 12, 12a, 13, 13a.

Description. The shell is small, and consists of four whorls in
the specimens examined. The body-whorl overlaps a little less than
one-half of the preceding whorl, and occupies about two-thirds of
the whole height of the shell. The spiral suture is well impressed
or slightly channelled. The whorl-surface traced downwards from
the suture presents an evenly convex surface.

The shell-wall is rather thick. The surface is marked by numerous
transverse lines of growth and there is also a tendency to produce
coarser, rounded, transverse ridges and furrows. The spiral angle
is about 90. The mouth aperture is broadly rounded below and
narrowed and somewhat pointed above. The height of the aper-
ture does not greatly exceed the breadth. The outer lip is sharp.
Below, the margin of the aperture is thickened. On the inner side

* Zittel (4), p. 289, Taf. 45, fig. 23.

f J. de C. Sowerby (2), p. 347, pi. xxiii., fig. 3.

{ de Loriol and Pellat (1), p. 27, pi. iii., figs. 13-15.

Hudleston and Wilson (1), p. 81.

|| Stanton (3), p. 31, pi. vi., figs. 10, 11.

174 Annals of the South African Museum.

of the aperture is a well-developed callus. There is a slightly
developed, short, closed umbilical slit.

Dimensions. (1) (2) (3)

Height of the shell 11 . 12 . 13mm.

Height of the body-whorl at the aperture 8 . 8 . 9 ,,
Greatest width of aperture 6 . 5 - 5 . 6 ,,

Occurrence. Found by Mr. Eogers in the cliff on Buck Kraal,
Sunday's River (136h).

Remarks. This form is distinguished from small specimens of
Natica uitenliagensis sp, nov. by the rather more expanded and less
elevated figure of the shell, and the broader and more rounded form
of the aperture. A specimen from the neighbourhood of Uitenhage
(350), ascribed above with some doubt to N. uitenhagensis , is com-
parable in point of size with individuals of N. rogersi. It differs
from these by its broader apical angle, shorter spire, narrower and
more elevated figure, and relatively higher body-whorl and aperture.

NATICA '? MIRIFICA sp. nov.
Plate VIII., figs. 14, 14a.

Description. The shell consists of at least six whorls, and has a
somewhat elongated turbinate outline with the body-whorl well pro-
duced and narrowed below. The body-whorl overlaps about half
the preceding whorl, and occupies rather less than two-thirds of the
total height of the shell. The spiral suture is well impressed, and
below it the whorl-surface bulges out in the form of a convex spiral
fold. Immediately below this rounded swelling of the whorl-surface
is a depression of the surface forming a concave spiral band. This
impressed zone is less well developed in the earlier whorls, more
pronounced in character in the later ones. At the middle zone of
the whorl the surface is again convex, and below this the whorl-
surface slopes inwards with less marked convexity. The concave
and convex zones in the upper half of the whorl merge into one
another without the production of any angularities of the surface
or of the outer lip of the aperture.

The shell-wall is moderately thick. The surface is smooth and
only marked by transverse growth-lines. The spiral angle is about
70. The aperture is elongated, its height being about twice as
great as its maximum breadth. It is angular at its upper extremity
and has a narrowed, rounded outline below. The inner lip is
thickened and a callus is developed. There is a short, narrow,
but closed umbilical slit.

The Invertebrate Fauna of the Uitenhage Series. 175

Dimensions.

Height of shell 27mm.

Height of aperture 17 ,,

Greatest width of aperture 10 ,,

Occurrence. Collected from the cliff on Buck Kraal, Sunday's
Eiver (137h). A fragment of a specimen, probably belonging to the
same form, was obtained from the cliff W. 20 S. from Comley's
house, right bank of Sunday's Eiver.

Re-marks. This is a rather peculiar shell, and I am unable to
state with certainty its true generic position. In spite of the
elongated form, the relatively narrow spiral angle, the narrowed
aperture, and the body-whorl produced and narrowed below, the
characters of the shell appear on the whole to conform with those
of naticoid type, while the close and smooth texture and lustrous
appearance of the surface, when well preserved, seem to favour
alliance with some division of the Naticce. A striking feature of
the shell is the longitudinal (spiral) depression of the surface of the
whorls in their upper part. This character in less emphasised form
is not unknown in Natica, though I am not aware that any described
species exhibits it in such a marked degree as in the shells under
discussion. The well-known Natica bulbiformis from the Gosau
beds, figured by J. de C. Sowerby,* has a distinct depression of the
surface corresponding in position with the stronger sulcation hi this
African form. N. bulbiformis also has a relatively tall spire, but it
is otherwise well distinguished by its more cylindrical whorls, the
deep channelling at the suture and the more oblique direction of the
mouth in relation to the long axis of the shell. Natica angulata
from the same beds,! first figured by Sowerby, also shows in some
degree a corresponding depression in the surface of the whorls.

There is a very close resemblance between Natica ? mirifica and
a Cretaceous gasteropod from the steppes of Astrakhan, described
and figured by B. Eehbinder under the generic name Odostomopsis.\
The general form of the shell and the undulating outline of the
whorl-surface is strikingly similar, but generic identity seems
excluded by the absence of any fold on the columella in the
African form. It may be questioned whether the specimens
depicted in Eehbinder's figures 12 and 13 are identical with the
originals of any of the other figures given by him under the same

* In Sedgwick and Murchison (1), pi. xxxviii., fig. 13 ; see also Zekeli (1), p. 45,
Taf. viii., fig. 2.

t Zekeli (1), p. 46, Taf. viii., fig. 4.

\ Eehbinder (1), p. 139, pi. ii., figs. 12, 13.

176 Annals of the South African Museum.

specific name, and it is very doubtful whether they are correctly
referred to Odostomopsis abeihensis (Blanck.), but it is certain that
the Uitenhage form here ascribed with doubt to Natica cannot be
brought into close comparison with Whitfield's genus."

GENUS ACT^ONINA A. d'Orbigny.

ACTJEONINA ATHERSTONI (Sharpe).

Plate VIII., figs. 15, 15a, 16, 16a, 166.

1856. Actceon atherstoni D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,
vol. vii., p. 200, pi. xxviii., fig. 19.

Supplementary Descriptive Note. The shell consists of about six
whorls, in the last of which there is a tendency to develop a steeply
sloping and slightly convex shoulder below the suture, demarcated
from the flattened central portion of the whorl by an ill-defined,
blunt spiral ridge. The previous whorl also shows these characters,
though in less marked degree, but the earlier-formed whorls have
a more evenly rounded outline, most convex near the suture, more
flattened below. The whorls are ornamented over their whole
surface by numerous delicate, impressed, linear, spiral striae which
are crossed by more conspicuous furrows and rounded ridges of
accretion, most noticeable in the last whorl. The spiral angle
is about 45.
Dimensions.

Height of a specimen with six whorls 19 mm.

Greatest diameter of the last whorl 9 ,,

Height of the aperture 10

Greatest width of the aperture 4*5 ,,

Occurrence. Collected at Grass Eidge, three miles east-north-east
of Uitenhage (333), and from a clay-pit in the lower part of the
Marine Beds on the left bank of the Zwartkop's River near Eawson
Bridge (343) ; also at Dunbrodie, Sunday's River (283, 284). Speci-
mens from the South African Museum are from the Sunday's River.
The record of occurrence given by Sharpe was " the lowest strata of
the Zwartkop crag." Mr. Rogers collected specimens of this form
in 1905 at the bare slope W. 30 S. from the middle of Barkly Bridge,
on the farm Olifant's Kop, Sunday's River (24h) ; from the highest
beds in a kloof behind Colchester, Sunday's River (499g) ; and from
a cliff W. 20 S. from Comley's house, right bank of Sunday's River
(92h). From this last locality was also obtained a specimen (95h)

* Whitfield (1), p. 425.

The Invertebrate Fauna of the Uitcnhacje Scries. 177

which may possibly belong to this species, though it possesses certain
distinguishing characters which are noted below.

Remarks. Sharpe's figured type, in the collection of the Geological
Society of London, is an unsatisfactorily preserved immature speci-
men with the shell partially removed. The smallest example sent
to me from South Africa measures 6'5 mm. in length, and is con-
siderably larger than the figured type, but a careful comparison
convinces me that these and the larger shells, one of which attains
a length of 19 mm., represent the same form. All the specimens
submitted to me have their whorls covered throughout with the
delicate spiral striae, and it was probably the imperfect preservation
of his material which led Sharpe to believe that these markings are
confined to the lower part. Eelatively to the size of the whorls, the
spiral ornamentation is strongest in youthful and early adult stages,
weakest in the last whorl. A single specimen collected by Mr.
Eogers (95h) differs from the majority of the individuals examined
by its taller spire, narrower spiral angle and less inflated body-
whorl : the degree of overlap in the whorls is also rather less,
and the body-whorl occupies less of the total height of the shell
(see plate viii., fig. 17). This may possibly represent a distinct
species, but it bears so close a general resemblance to A. atJierstoni
that it is difficult to arrive at a decision on this point, especially as
some variation is observable in the other individuals.

The examination of well-preserved adult specimens shows that
this species possesses a mouth of oval form, rounded below, with
sharp outer lip and a columella quite devoid of folds. It therefore
falls within d'Orbigny's genus Actaonina as restricted by Meek, :;:
under which name it was correctly quoted by Messrs. Rogers and
Schwarz in their report on the survey of 1900.

There is some resemblance between Actceonina atherstoni and the
Belgian shells ascribed by Briart and Cornet t to the Blackdown
species named Phasianella striata by J. de C. Sowerby, and
described under d'Orbigny's name Phasianella soiverbyi.\ This
has a shorter spire than the African form, but to judge from the
figures given by the Belgian authors, the slender outline of the shell
and the shape of the mouth are very similar, although the two forms
are perhaps not even generically identical, and the striae are coarser
in the Belgian shell. In another Blackdown species, described by
Sowerby under the name Phasianella formosa,^ the shape of the

* Meek (1), p. 91. f Briart and Cornet (1), p. 35, pi. iii., figs. 20, 21.

t d'Orbigny (4), vol. ii., p. 152.

J. de C. Sowerby (2), p. 343, pi. xviii., fig. 14.

178 Annals of the South African Museum.

whorls and the mouth closely approaches that of Actceonina ather-
stoni, but the spire of the English shell is relatively shorter, and the
spiral striae are confined to the lower part of the whorls.

A specimen from the Upper Cretaceous beds of Umkwelane Hill,
Natal, has been described as a variety of Actaonina atherstoni by
R. Etheridge, jun. (var. umkwelanensis).--' To judge from the figure
of this supposed variety, there is a general resemblance to Actaonina
atherstoni, although the spiral striae are depicted as of a coarser
character than in the Uitenhage form. Mr. Etheridge's remarks
on his new variety do not suffice to enable a satisfactory com-
parison to be made, but I am inclined to doubt the likelihood of
near relationship between these forms.

Actaonina haugi Ascher, from the Grodischter Schichten,! the
equivalent of the Hauterivian in Silesia, is of similar general type
to A. atherstoni, and is ornamented over the whole surface of the
whorls by minute spiral striae. It is more slender and elongated,
however, and more cylindrical in form, and has a narrower spiral
angle and less convex whorls than A. atherstoni. The specimen
numbered 95h, mentioned above, approaches closely to A. haugi in
the form of the spire, but has a more convex and less cylindrical
body-whorl.

GENUS LIMN/EA Lamarck.
LIMN.EA BEMOTA sp. nov.
Plate VIII., figs. 18, 18a.

Description of a Single Specimen. The specimen, which is im-
perfect at the apical end, consists of nearly four whorls, and is
wound in the form of a tall spire, having a somewhat pupoid figure.
The outline of each whorl slopes down very steeply from the spiral
suture, giving the whorl a gently and evenly convex profile. The
volutions are relatively high between the sutures. The body-whorl
overlaps about one-third of the previous whorl, and occupies nearly
one-half of the total height of the shell. The rather steeply inclined
spiral suture is not deeply impressed.

The spiral angle is about 30. The shell-wall is thin, and its outer
surface is marked by numerous wrinkles and furrows of accretion.
The mouth aperture is in the form of an acute angle above, and is
of rounded oval outline below. Its height is about twice as great as

* Etheridge (1), p. 87, pi. ii., fig. 38.

t Ascher (1), p. 152 [18], Taf. xii. [i.], fig. 12o-c.

The Invertebrate Fauna of the Uitenhagc Scries. 179

its breadth. The outer lip is thin and the inner lip only furnished
with a very thin callus-coating.

Dimensions.

Height of shell 33 mm.

Height of body- whorl at the aperture ... 17
Greatest width of the aperture 8'5 ,,

Occurrence. The specimen was collected by Mr. Eogers from the
cliff on Buck Kraal, Sunday's Eiver (138h).

Remarks. This is the only fresh-water form included among the
fossils found in the cliff-section on Buck Kraal, and it seems most
probable that it represents a merely stray occurrence in marine
strata, and does not indicate the presence of a fresh- water bed. The
specimen may have become washed down into marine surroundings
by flood-waters, but its fresh appearance suggests that it could not
have been subjected for long to the action of currents before becoming
embedded.

The resemblance shown by this specimen to Limncea longiscata
Brongn., * which occurs so abundantly in the Headon Beds of the
Hampshire basin and in the Paris basin, is really very striking.
Although selected specimens of L. longiscata may approach very
closely in all characters to the form with which we are dealing, yet
this may be well distinguished from most specimens of the Tertiary
shell by its more slender and elongated figure. The spiral angle of
L. longiscata is frequently wider, and the body-whorl is almost always
more tumid than in Limn&a remota.

CLASS CEPHALOPODA.

GENUS PHYLLOCEEAS E. Suess.
PHYLLOCERAS ROGEKSI sp. nov.

Plate VIII., figs. 19, 19o-c.

Description of a Single Specimen. The shell is very involute, with
minute umbilicus. The umbilical wall is not abruptly marked off
from the flank. The whorls have a laterally compressed form, and
the flanks show a slightly convex surface. The height of the whorl,
measured from the umbilical rim, is greater than the breadth seen in
cross-section. The greatest breadth of the whorl-section occurs at

* Edwards (1), p. 85, pi. xii., figs. 3<i-7i ; Deshayes (1), p. 92, pi. xi., figs. 3, 4
(1825).

180 Annals of the South African Museum.

about the middle of the flank, and from here the whorl narrows very
gradually towards the periphery, which forms a relatively broad,
flattened arch.

The surface is ornamented with closely set, delicate, linear trans-
verse ribs, which have a very slight forward inclination when traced
from the umbilicus towards the periphery. On a portion of the last
whorl which measures 9 mm. from the umbilical centre to the
periphery, there are 12 ribs crowded into a space of 2 mm. at the
middle of the flank. There are no constrictions.

The specimen is imperfect, and is wholly septate. The course of
the lobe-line is relatively complex in its details. The siphonal
(external) lobe is narrow and very deep, and the small siphonal
saddle at its termination narrows somewhat abruptly to a sharply
pointed summit. The siphonal lobe is as deep as the first lateral
lobe.

Dimensions.

Greatest diameter, if completed, about 30 mm.

Greatest breadth of the last whorl in cross-section 16 ,,
Height of the last whorl at the centre, in section... 13 ,,
Occurrence. Found in the kloof behind Colchester, Sunday's
River Valley, in the middle beds (3h).

Remarks. This specimen, so far as I am aware, is the only repre-
sentative of the genus Phylloceras which has been found in the
Uitenhage Series. It is, unfortunately, not a complete individual ;
the body- chamber is absent, and a portion of the last whorl pre-
served is also broken away. The surface ornaments are only pre-
served in a few small patches on the early part of the last whorl,
where the outer surface has escaped destruction. In the remaining
portions of the surface which are exposed, sufficient of the shell
substance is present to prevent the septal suture-line from being
seen. It has, therefore, been necessary to etch away part of the
shell with acid in order to obtain a view of the suture-line, but owing
to the mode of preservation of the cast, the picture obtained is not
so satisfactory or complete as might be desired.

It has been possible, however, to trace the course of the lobe-line
so far as concerns the siphonal lobe, the first lateral saddle and the
first lateral lobe. The most striking feature is the depth of the
siphonal lobe, which touches a radius which is reached by the first
lateral lobe. In other respects the lobe-line, so far as observed, very
much resembles that given by d'Orbigny in the case of his Ammonites
picturatus,* from the Lower Neocomian of France ; although both
* d'Orbigny (1), p. 178, pi. liv., figs. 4-6 (1841).

The Invertebrate Fauna of the Uitenhayc Series. 181

suture-lines are so complex, the agreement, in fact, is remarkably
close. Phylloceras picturatum has somewhat less inflated whorls
than the African form, and its surface is said to be entirely smooth.

In the depth of the siphonal and first lateral lobes of the septal
suture, there is agreement with Phylloceras semisulcatum (d'0rb.) :|:
but the suture of that form is much less complex in its details.

There is considerable similarity of form to a specimen from the
Wernsdorfer Schichten (Barremian of Silesia) described by Uhlig
under the name Phylloceras cf. guettardi Easp.t The whorl-section
of this shows that the flanks converge slightly towards the periphery,
which is narrowly arched. In the African specimen the flanks are
not so markedly convergent, but are more nearly parallel, and they
are perhaps a little less convex. The peripheral area is broader, and
in the cross-section of the whorl a greater breadth is maintained up
to the peripheral part. There are no noteworthy points of agree-
ment in the course of the septal suture of these two forms. In the
shell described by Uhlig, which is said to have a suture identical
with that figured by d'Orbigny for Phylloceras guettarcli,\ the
siphonal and first lateral lobes are of unequal length, and the suture
is considerably less complex in its details.

The specimen here described appeal's to represent a new species,
but it will be necessary at some future time to supplement the
present imperfect account by additional notes, and by the illustra-
tion of the septal suture of a more favourably preserved individual.

GENUS BOCHIANITES P. Lory.

BOCHIANITES GLABEB Sp. nOV.

Plate VIII., figs. 20, 21.

Description. The shell has a delicately attenuated form, and
increases very slowly in diameter. Near the delicately pointed
posterior end the shell is cylindrical, but becomes slightly com-
pressed laterally, with corresponding elliptical section, when traced
towards the anterior end. The surface is devoid of ornamentation,
and is marked by numerous minute lines of growth which show
a slight anterior arching on the siphonal side, and when traced
across the flank of the shell are directed somewhat posteriorly ; they
follow a straight course (or have an exceedingly gentle and broad

* d'Orbigny (1), pi. liii., fig. 6 (1841).
t Uhlig (1), p. 182, Taf. iv., fig. 9.
{ d'Orbigny (1), pi. liii., fig. 3 (1841).

13

182 Annals of the South African Museum.

anterior arching) when passing across the anti-siphonal side. In
addition to these lines there are very faint and indistinct shallow
depressions of the surface, of varying breadth, which follow a course
similar to that of the growth-lines.

It has only been possible to follow completely the course of the
septal line at an early stage of growth, where the shell has a
diameter no greater than 2 mm., and, as might be anticipated, the
suture is here of a comparatively simple character. While the main
lobes and saddles have come to full development as regards their
general course, the minor indentations of the suture have only in
small degree made their appearance. The siphonal, anti-siphonal,
and lateral lobes are narrow and deep, and have almost equal
elongation. At this comparatively early stage the trifid character
of the lateral lobe is already making its appearance. Both the
siphonal and lateral saddles are divided into two limbs by a narrow
and shallow subsidiary lobe. Each limb of the saddle is in turn
divided by a shallow, pointed indentation. The lateral saddle is
slightly broader than the siphonal saddle. The small secondary
saddle at the base of the siphonal lobe is at this stage weakly
developed and has a blunt summit.

Dimensions.
Length of a specimen with body-chamber, but incomplete

at initial and anterior ends 50 mm.

Diameter at the posterior end 2 ,,

Greater diameter at the anterior end 5 ,,

Occurrence. Found on the road below the railway cutting, one
mile from Eawson Bridge on the main line, up side (277, 338).
Crushed specimens obtained from the clay-pit on the left bank of
Zwartkop's Eiver near Rawson Bridge (278, 339, 344), referred by
Messrs. Rogers and Schwarz with some doubt to Baculites, may
probably represent the same form.

Remarks. The adoption of the name Bochianites, proposed by
P. Lory" for Baculites neocomiensis d'Orb. and its allies, seems to be
well justified when a comparison is made between the septal sutures
of the forms to which the name is applied and those of the true
Baculites as exemplified, for instance, by B. bacidoides (Mant.) or
B. bohemicus Fritsch and Schloenbach. Whether we are to regard

* Lory (1), p. 133, footnote ; Lory (2), p. 129, footnote. [I have not been able to
consult the latter work quoted, but give the citation on the authority of von
Koenen (2), p. 397, and G. Boehm (3), p. 26. The papers Lory (1) and Lory (2)
have identical title and date of year, but I have not ascertained which was issued
first ; one is presumably a reprint of the other, except in pagination.]

The Invertebrate Fauna of the Uitenhage Series. 183

Bochianites as a separate genus or as a sub-genus of Baculites might,
perhaps, be considered doubtful, and on this point G. Boehm, who
adopts the name," has refrained from expressing an opinion. Sarasin
and Schondelmayer ascribe a generic value to the name, and con-
sider that a comparison of the septal sutures indicates that
Bochianites and Baculites have originated separately ; t the dif-
ferences revealed by such a comparison are, in fact, so striking and
so constant that with present knowledge it appears to be a reasonable
and expedient course to regard these forms as generically distinct.
In Bochianites there is a siphonal lobe, an anti-siphonal lobe, and
on either side one main lateral lobe. These are relatively narrow
and of almost equal depth. An important distinctive feature is the
trifid termination of the lateral lobe. On either side there are two
relatively broad saddles, each divided fairly symmetrically by a
narrow subsidiary lobe. There are thus typically only four main
lobes and four saddles, though in some forms (for example B. undu-
latus von Koenen) the lateral saddle is so deeply divided that it
might be almost regarded as forming two saddles separated by a
lobe less deep than the lateral lobe. To obtain the septal suture
of Baculites, we must imagine the subsidiary lobe of the lateral
saddle to be so deepened as to constitute an additional lateral lobe ;
at the same time the anti-siphonal lobe becomes very much less
deepened, while the first lateral lobe should have a paired instead
of a trifid termination. These relations are very well revealed by
a comparison of the suture-line in Bochianites neocomiensis J and
Bochianites waterinyi with that in Baculites baculoides \\ and
Baculites bohemicus.^.

There can be no doubt that the Uitenhage specimens must be
referred to Bochianites, as clearly indicated by the course of the
lobe-line. One of the chief distinctive characters of the form here
dealt with appears to be the absence of surface ornaments, and even
allowing for imperfect preservation, it may be regarded as certain
that the available material would have shown definite traces of
ribbing if such had existed. Specimens from the clay-pit near
Eawson Bridge, which I think may represent the same species, are
much crushed, but the shell- substance is in great part preserved and
retains its nacreous lustre. In these there is no trace of ribbing,
and the shell appears to have been a smooth one.

* G. Boehm (3), p. 26. f Sarasin and Schondelmayer (1), p. 177.

I d'Orbigny (1), p. 560, pi. 138, fig. 4 (1842).

G. Boehm (3), p. 26, fig. 3. || d'Orbigny (1), p. 562, pi. 138, fig. 11 (1842).

IT Jahn (1), pi. viii., fig. 7 ; Woods (1), p. 76, pi. ii., fig. 10.

184 Annals of tlic South African Museum.

A form with which it might seem possible to bring Bochianites
glaber into close comparison is J5. oosteri Sarasin and Schondel-
mayer, :;: from the Swiss Neocomian. This is also devoid of
ornamentation so far as has been ascertained ; but it appears to be
well distinguished by the great breadth of the lateral saddle in
relation to the siphonal saddle, and by the deeply divided form of
the lateral saddle. This division is carried so far, indeed, that it
may be said that two lateral saddles are developed. If B. glaber
possessed in maturity a septal suture having the relations of lobes
and saddles similar to those exhibited in B. oosteri, it would be right
to expect that some signs of it would be evident even at an early
stage. Traces of the suture of B. glaber where the shell has attained
a diameter of 4 mm. are still of a very simple character, and show
no promise of such sub-division at a later stage of growth as that
which characterises the mature B. oosteri. Moreover, the septa of
B. glaber are much more closely approximated than those of the
Swiss shell, in which the length of the chambers is a special
feature.

Bochianites undulatus von Koenen,t from the lower Aptian of
North Germany, is well distinguished from B. glaber by the well-
developed ornamentation and the deeply divided lateral saddle, as
well as by other features.

Baculites rotundas Reuss,} from the Planermergel of Bohemia, is
also a smooth, slowly tapering form, but without particulars of the
septal line it is impossible to make a close comparison. B. rotundus
may be a true baculite, and Pictet has even suggested that its real
position may be in the genus Hamites.

GENUS HOLCOSTEPHANUS M. Neumayr (scnsu stricto}.

The great majority of the ammonites included in the collections
under examination belong to that section of the original genus
Holcostephanus exemplified by d'Orbigny's Ammonites astierianus,
the form which was regarded by Neumayr to be the best known
typical species of his genus. To this species and some allied forms
Pavlow gave the name Astieria, when recognising that the wide
application of the name Holcostephanus was not in accordance with

* Sarasin and Schondelmayer (1), part 2, p. 179, pi. xxiv. , figs. 3, 4, and text-
figure 6.

t von Koenen (2), p. 398, pi. liii., figs. 11, 13, 14.
J Reuss (1), part i., p. 24, pi. vii., fig. 4 (1845).
Neumayr (1), p. 922.

The, Invertebrate Fauna of the Uitenhage Series. 185

the results and requirements of modern work, and that a division
into a number of genera or sub-genera had become necessary." If
we were to ascribe to the name Holcostephamis the broad signification
allowed by Neumayr, Zittel, and others, it would certainly be con-
venient to use the name Astieria to denote one of several sub-genera,
were this procedure not contrary to the recommendations of the
International Zoological Congress.! It will be more correct, as
Lemoine has pointed out, to restrict the name Holcostephanus to
that portion of the genus, as originally conceived, which is typified
by H. astierianus, and apply it in place of Astieria. I

The separation of Spiticeras from Pavlow's Astieria, as proposed
by V. Uhlig and F. Suess, appears to be a well-founded step in the
recognition of further divisions within the broadly conceived genus
Holcostephanus. Holcostephanus, sensu stricto (= Astieria Pavlow,
emend. V. Uhlig and P. Suess) thus becomes a group having very
narrow limits, and some division of opinion is likely to exist con-
cerning the question of its generic or sub-generic value. This at
present must remain a matter for individual judgment, and in view
of our rapidly growing knowledge and the other causes which con-
tribute to the present instability of ammonite-classification, it is a
question of comparatively subordinate importance. For the purposes
of the present work the name will be employed in a generic sense,
as the equivalent of the restricted Astieria of Uhlig and F. Suess.

It is unfortunate that the specimens of Holcosteplianus obtained
from the rocks of the Uitenhage Series are usually in a state of
preservation unfavourable for exhaustive study. Thus, in no case
has it been possible to ascertain the true course of the lobe-line, while
evidence regarding the form of the body-chamber and mouth aperture
is likewise still scanty. We must also look to further collecting to
furnish materials for a study of individual development in the various
forms represented, and it is clear that there is much to be added in
order to supplement the following imperfect account of this interesting
assemblage. There can be no doubt, however, that the characters of
shape and ornament displayed by the available specimens suffice to
enable us to read aright the relationships of these forms, though
perhaps not always with the precision that might be desired.

It may be pointed out that in the diagnosis of Astieria given in
Uhlig's monograph on the Fauna of the Spiti Shales a sentence

* Pavlow and Lamplugh (1), pp. 471, 491 (113, 133 of authors' copy).
f Regies Internationales de la Nomenclature Zoologique adoptees par les Congres
Internationaux de Zoologie ; articles 9, 25, 29. Paris, 1905.

J Lemoine (1), p. 181. Uhlig (4), p. 86.

186 Annals of the South African Muscitm.

occurs which does not seem to express quite accurately the relations

most frequently exhibited in the secondary ribbing. It is stated that,

starting from the tubercles at the margin of the umbilical wall, there

are groups of secondary ribs " which bifurcate higher up and admit

of intercalary ribs." It would perhaps have been more correct to

say that bifurcation of the secondary ribs on the flank (away from

the tubercles) occurs rather as an exception than as a rule, since it

is certainly either absent or very rare in the majority of known

" Astieria." In typical specimens of H. astierianus (d'Orb.), as

figured by d'Orbigny himself," no bifurcation in the costas of the

rib-bundles after they leave the tubercles is apparent, but the finely

ribbed forms (d'Orbigny's plate 28, fig. 4), which have been named

H. sayni by Kilian and Astieria scissa by Baumberger, show the

division of the ribs here and there on the flank. In all the South

African representatives of Holcosteplianus the groups of secondary

ribs proceed from the compressed umbilical tubercles and pass over

the periphery to the tubercles of the other side without undergoing

any division. It may be noted, however, that the ribs forming a

group on one flank may become so disposed when traced to the

opposite flank that they do not all reunite at one tubercle ; thus, of

three ribs departing from a tubercle on one side, two may unite

again at a tubercle on the opposite side of the whorl, while one may

either pass to an adjacent tubercle or may terminate in the space

between two tubercles at the umbilical rim.

There is one other point in Uhlig's diagnosis of Astieria which
seems to call for emendation. It is stated that the ornamentation
" consists of a varying number of primary costee which start from the
suture, and at the passage from the umbilical wall into the sides
swell up into tubercles." A specimen obtained by Messrs. Eogers
and Schwarz near Uitenhage, and described below (H. uitenhagensis
sp. nov.), is so characterised that it can only be ranged among the
"Astieria" ; but in this example the ornamentation commences with
the marginal tubercles, and no primary umbilical costae are developed
in the last whorl. It seems therefore necessary to say that although
primary umbilical ribs are present in the majority of the Holcostepliani,
yet they do not form an essential part of the sculpture in all growth-
stages. It may be that in the specimen referred to, umbilical ribs
were developed in the earlier growth-stages represented by the inner
whorls, but these, owing to unfavourable preservation, unfortunately
cannot be studied. In this specimen, which is of considerable size,
the ultimate whorl probably represents a stage approaching, or in,
* d'Orbigny (1), p. 115, pi. 28, fig. 1 (1840).

TJie Invertebrate Fauna of the Uitenhage Series. 187

the final phase of individual growth, and it is only in this whorl
that the umbilical wall has been satisfactorily exposed and freed
from the matrix.

The predominance of Holcostephani in the cephalopod-fauna of
the Uitenhage beds is no less striking than the manner in which
certain of these lend themselves to close comparison with European
forms. There is apparently near relationship to types well distributed
in the Valanginian and Hauterivian of the European continent and
more sparingly represented in England, and in this fact we may
recognise the most reliable evidence for a correlation of the
Uitenhage Marine Beds with European standards. It will be
well, for the present, to regard with the greatest caution the ap-
parently peculiar geographical distribution of Holcostephamts, since
so little is yet known of any cephalopod-faunas, in tropical or sub-
tropical latitudes, which may be brought into correlation with that
of the Uitenhage Series. Some remarks on this subject have already
been made in the previous section of this memoir.

I have to acknowledge my indebtedness to Mr. G. C. Crick for
kindly assisting me to compare some of the specimens described
below with representatives of Holcostephanus preserved in the British
Museum (Natural History).

HOLCOSTEPHANUS ATHERSTONI (Sharpe).

1856. Ammonites atherstoni D. Sharpe, Trans. Geol. Soc. Lond.,

ser. 2, vol. vii., p. 196, pi. xxiii., fig. 1.
1882. Olcostephanus atherstoni M. Neurnayr, in E. Holub and

M. Neumayr, Denkschr. d. k. Akad. Wiss., Math. -Nat.

01., Band xliv., p. 272.
1892. Olcostephanus (Astierm) atherstoni A. P. Pavlow (partim], in

A. P. Pavlow and G. W. Lamplugh, Bull. Soc. Imp. Nat.

Mosc., annee 1891, Nouv. Ser., tome v., p. 495 (p. 137 of

authors' copy). (Probably not pi. xvii., fig. 14.)
Occurrence. Collected by Miss M. Wilman at Coega. A specimen
from the South African Museum is probably from the Sunday's
Eiver. The type-specimen in the Geological Society's collection in
London is from the Sunday's River (registered 10975). Messrs.
Eogers and Schwarz have recorded this form from the Zwartkop's
Eiver valley in the neighbourhood of Uitenhage. :;: Mr. Eogers has
also noted its occurrence at several localities in the valleys of the
Coega and Sunday's Eivers. |

* Rogers and Schwarz (1), p. 9. t Rogers (2).

188 Annals of the South African Museum.

Remarks. Some characteristic points in H. atlierstoni are the
strong, backwardly directed umbilical ribs, the steep umbilical wall,
the relatively narrow and profound umbilical cavity, and the con-
siderable breadth and inflation of the whorls. The involution until
the shell has attained large dimensions is such that the umbilical
wall of a given whorl falls upon the umbilical tubercles of the
previous whorl, wholly concealing the secondary ribbing. In the
large type-specimen of H. atherstoni, measuring 140 mm. in greatest
diameter, the last -formed whorl is beginning to show a little less
involution, and its most advanced part leaves the tubercles of the
previous whorl wholly exposed, and also the commencement of the
rib-bundles of the flank.

It has been stated by several authors that H. atlierstoni occurs in
the Valanginian and Hauterivian of Europe, and since such an
occurrence would be of high interest, both from a stratigraphical
and paloeontological point of view, I have taken some pains to test
the value of these identifications. The result has been to convince
me that none of the determinations can be upheld with certainty.

Pavlow considered Holcostephanus psilostomus Neum. and Uhlig, *
from the Neocomian of North Germany, to be identical with
H. atherstoni, or at least that the two forms represent no more than
varieties of a single species. Uhlig, on the other hand, believes this
view to be erroneous,! and points out that H. atlierstoni has an
almost semicircular cross-section and greater breadth. There can
be no doubt that, though closely comparable, these types are not
identical. Another allied North German form, from the zone of
Hoplites radiatus (Lower Hauterivian) has been described and
figured by von Koenen J under the name Astieria aff. psilostoma.
In this, the rate of increase in the breadth of the whorls is not so
rapid as in H. atlierstoni, and the ribbing is a little coarser. The
pronounced development of constrictions is another distinguishing

feature.

Another European form which bears considerable resemblance to
H. atherstoni has been described by von Koenen as Astieria convoluta
from the Lower Hauterivian of North Germany. This appears to
have similarly inflated whorls and a narrow umbilicus, but the
number of umbilical tubercles is fewer than in H. atherstoni at a
comparable stage of growth, and the ribs are coarser and less
numerous.

* Neumayr and Uhlig (1), p. 149, Taf. xxxii., fig. 2.

t Uhlig (4), p. 132. + von Koenen (2), p. 151, Taf. liv., fig. 2.

von Koenen (2), p. 146, Taf. xxxix., figs. 4n, 46.

The Invertebrate Fauna of the Uitenliage Scries. 189

Holcostcplianus multiplicatus Neum. and Uhlig (non Eoemer) *
from the Upper Valanginian of North Germany, which has been
renamed Astieria ventricosa by von Koenen, f approaches H. cither-
stoni in the tine character of the ribbing, but is well separated by the
much higher whorl-section and the conspicuous development of con-
strictions, as well as by other characters.

Certain forms occurring in the south-east of France and in the
Maritime Alps, ascribed by Kilian to H. atlierstom,\ seem to have
been so named in the belief that H. multiplicatus Neum. and Uhlig
(= Astieria ventricosa yon Koenen) is identical with H. atlierstoni.
Baumberger has also spoken of the " Zone of Holcosteplianus multi-
plicatus Neum. and Uhlig, non Eomer = H. atlierstoni Sharpe " in
the Lower Hauterivian of the Swiss Jura. Early in the present
year, however, he has taken pains to show that the earlier identifi-
cation of H. multiplicatus with H. atlierstoni was erroneous. ||

When we come to discuss the correctness of Baurnberger's view
that a group of forms represented richly at the base of the Haute-
rivian (the so-called Astieria Zone) in the Swiss Jura is really
inseparable from Holcosteplianus atlierstoni, the difficulty of forming
a final judgment becomes very great. Baumberger discusses the
question in great detail, *I and his conclusion is put forward judicially
and not with any claim to be the final word in the matter. Some,
at least, of the specimens figured by Baumberger bear a strikingly
close resemblance to H. atlierstoni, and it appears safe to conclude
that the relationship is a very close one. It appears possible that
more than a single species is represented in the Swiss material
brought together by Baumberger under Sharpe's specific name, but
I am not in the position to form a definite judgment on this point.
Of the larger specimens figured by Baumberger, which from their
size are most fitly comparable with the large type-specimen from
South Africa, that depicted in plate xxiv., fig. 2, seems to agree
most closely in the fineness of the ribbing. The specimen
shown in plate xxiii., fig. la, exhibits more clearly the
characters of the umbilical part of the shell, and allowing that
this is an internal cast, the figure might well have been drawn
from a South African specimen. The apertural view (fig. 15), how-
ever, unless the peripheral part is much crushed down, shows that

* Neumayr and Uhlig (1), p. 150, Taf. xxxiii., fig. 2.
f von Koenen (2), p. 144.

J Kilian (1), pp. 716, 726, 735; Kilian (4), p. 865; Kilian and Leenhardt (1),
pp. 973, 979, 981 ; Matte (1), p. 151.

Baumberger (1), 2 er Theil, p. 3. ]| Baumberger (1), 4 er Theil, p. 40.

M Baumberger (1), 4 er Theil, pp. 39-47 (Astieria atlierstoni).

190 Annals of the South African Museum.

the whorl is not nearly so highly arched in section as in H. atherstoni.
In this character, the specimen shown by Baumberger in his text-
figure 115 agrees more closely. The whorl-section shown in fig. 114,
on the other hand, is not quite so highly arched in proportion, and
seems to agree better with that shown by the South African
specimens described below as H. cf. atherstoni. As regards the
development of constrictions, these, if truly developed, are so in-
conspicuous in South African specimens that they can scarcely
be detected, and they never form a feature like that shown in
Baumberger's text-figure 116.

It must be admitted that much remains to be known about the
true H. atherstoni. As yet, we are without information regarding
the limits of individual variation, the characters of shape and
sculpture in successive growth-stages from early youth onwards,
and the course of the septal suture throughout the individual de-
velopment. This information can only be obtained by the exami-
nation of an extensive suite of specimens, and in the meantime it is
not possible to form a decided opinion on the question of identity
with the Swiss; specimens dealt with by Baumberger. It seems
necessary, with the present imperfect data for comparison, to regard
that question as still awaiting settlement. At the same time, we
may presume the relationship between these Swiss and South
African forms to have been so intimate as to furnish further highly
suggestive evidence for a correlation between the Uitenhage beds
and the strata situated about the base of the Hauterivian in
Europe.

It may be noted that Baumberger gives reasons for considering
that the form from the Hauterivian of the Maritime Alps ascribed
by Kilian * to H. atherstoni, is not identical with the supposed
H. atherstoni from, the Swiss Jura, above discussed.

It has been stated by Sayn and Roman that Holcostephanus
atherstoni occurs in the Hauterivian inferieur at localities in
Languedoc.f In answer to an inquiry concerning this French form,
M. Eoman very courteously sent me three specimens from the Lower
Hauterivian of Saturargues (Herault) which had been determined by
M. Kilian. Two of the specimens were named H. atherstoni, and
the third, a very finely ribbed example, somewhat flattened laterally
by crushing, bore the name H. sayni Kilian. After a careful exami-
nation of these three specimens I was unable to satisfy myself that
they represented two separate species. All three are very finely

* Kilian (4), p. 865, pi. Ivii., fig. 1.

t Sayn and Eoman (1), pp. 623, 625, 629, 632, 638, 639.

The Invertebrate Fauna oj the Uitcnhage Scries. 191

ribbed, and though the ribs seemed most crowded in the specimen
referred by M. Kilian to H. sayni, yet the differences in this respect
did not appear to me to be at all considerable, and other points of
distinction, such as the more discoidal form, were probably to be
accounted for by distortion through crushing. A comparison of
these specimens with Sharpe's type, and other examples of
H. atJierstoni, shows that in H. atherstoni the shell is much more
inflated and less discoidal : the whorl is broader in section and less
highly arched. The umbilicus is relatively much narrower and
more profound. The umbilical wall falls more abruptly and steeply
into the umbilical cavity. The involution is greater. In the French
specimens the umbilicus is of a broader and shallower type, and the
involution is such that the tubercles, together with part of the ribbing
of the flank, are visible in the penultimate whorl. The involution,
in fact, is less than in the last whorl of the considerably larger type-
specimen of H. atherstoni. The primary (umbilical) ribs are also
much more weakly developed than in H. atJierstoni. The ribbing of
the flank and periphery of H. atJierstoni is of a somewhat coarser
type, and the ribs are rather more prominent and are less closely
crowded together than those of the French specimens. At a selected
stage of growth comparable in the two forms, a space of 20 mm.
along the periphery includes 9 ribs in H. atherstoni and 13 in the
French shells. In H. sayni an occasional bifurcation of a rib, or
the intercalation of a shorter rib between two longer ones, may be
observed to occur on the flank at a varying but usually considerable
distance away from the tubercles. In H. atherstoni, the ribs of the
flank all have their commencement either at, or in the near vicinity
of, the tubercles, and bifurcation of the ribs has not been observed
on any part of the flank.

It is clear, therefore, that these French specimens are not very
closely comparable with H. atherstoni, and I think it highly probable
that all three represent H. sayni Kilian," a finely ribbed species very
nearly allied to H. astierianus (d'Orb.).

A specimen from the Neocomian of Speeton (zone of Belemnites
jaculum), preserved in the Scarborough Museum, was figured by
Pavlow (Bull. Soc. Imp. Nat. Mosc., 1892, pi. xvii., fig. 14) and con-
sidered by him to be identical with H. atherstoni. Through the
kindness of Mr. H. Ascough Chapman and Mr. J. A. Hargreaves, of
Scarborough, I have been able to compare this Yorkshire individual
with Sharpe's type and other examples of H. atherstoni from South

* Kilian and Leenhardt (1), p. 976 ; Sarasin and Schondelmayer (1), part 1, p. 38,
pi. iv., figs. 2, 3 ; d'Orbigny (1), pi. 28, fig. 4.

192 Annals of the Soutli African Museum.

Africa. The specimen is preserved unfavourably for an exhaustive
comparison, but certainly bears a very close resemblance to H.
atherstoni, particularly in the inflation of the whorls and the general
character of the ribbing. Some readily apparent differences may at
the same time be noticed. At a comparable stage of growth the
Speeton form is distinctly more closely and more delicately ribbed ;
umbilical ribs are hardly denned, and certainly have not attained the
development to be observed at a similar stage in H. atherstoni. A
further point is that in all specimens of H. atherstoni the flattened
umbilical marginal tubercles have either a slight backward direction
when traced towards the flank, or their disposition is strictly radial ;
in the Yorkshire example, on the other hand, the tubercles have a
strong forward inclination. These differences cannot be overlooked,
and they strongly suggest that the Speeton form may not be
identical with that from South Africa ; but it must be admitted that
the two approximate very closely to one another. It may suffice to
have drawn attention to these points of difference while refraining
from any final expression of opinion concerning the intimate relation-
ship of these forms a question which can only be ultimately settled
by the comparison of further material. At the same time, I am
inclined to think that the Speeton form will not be found to be
identical with H. atherstoni.

Other finely ribbed specimens from the lower part of the zone of
Belemnites jaculum at Speeton, which have been referred by Prof.
Pavlow and Mr. C. G. Danford to Holcostcphanus astierianus (d'Orb.),
are also in some respects comparable with H. atherstoni ; but they
are usually of small size, and a satisfactory comparison with the
much larger available examples of H. atherstoni is difficult. One
such individual, in which the development of constrictions does not
form a feature, has been figured by Mr. Danford under the name
Olcostephanus (Astieria} astieri* It is laterally compressed and
relatively highly arched, and the umbilical ribs are very weakly
developed and are not backwardly inclined. The secondary ribbing,
which is of a very crowded and delicate character, has a stronger
forward inclination than that of H. atherstoni. The tubercles also
have a forward inclination, which is not the case in the African
species.

Karakasch has described a Holcostephanus from the Lower
Cretaceous of Biassala (Crimea) under the name Holcostephanus
(Astieria} cf. atherstoni Sharpe.f It appears, from the figure given

* Danford (1), pi. xi., figs. 7, la.
f Karakasch (1), p. 103, pi. L, fig. 3.

The Invertebrate Fauna of the Uitenhagc Series. 193

by Karakasch, to differ from the African species by the relatively
wider umbilicus and less degree of involution, but to resemble it in
possessing backwardly inclined umbilical ribs. Kilian has described
an ammonite from the Hauterivian of the neighbourhood of Escrag-
nolles (Maritime Alps) as Holcostephanus gueblmrdi* and states that
this corresponds to H. atherstoni Karakasch, but not to the type of
Sharpe. In the specimen figured by Kilian the ribs of the flank are
strictly radial in direction, so that there is no close agreement with
H. atherstoni.

Another form which may be somewhat nearly related to H.
atherstoni is II. schenki (Oppel), from the Spiti Shales in Tibet,! but
this is less densely ribbed and is further distinguished by the
more flattened arch of the whorl-section, and by the presence of
strongly marked constrictions.

An inflated form from the so-called " Gault " on the Island of
Chasik and from Tukusitnu Bay (Alaska), figured by Eichwald as
" Ammonites astierianus d'Orb. aff." and compared by him with
Eoemer's Ammonites multiplicatus, may perhaps be related to H.
atherstoni ; but according to Eichwald's insufficient figure the
primary ribs are almost exactly direct (radial) in their course. A
complete comparison with this unsatisfactory figure is impossible,
since a side view only is given. |

HOLCOSTEPHANUS cf. ATHERSTONI (Sharpe).

A single specimen from the Sunday's River, in the collection of the
South African Museum, agrees very closely with Sharpe's type in all
points except one the inflation of the shell. The whorl-section has
rather greater breadth in relation to its height, and the peripheral
area presents a more broadly rounded surface and is less highly
arched in sectional view than that of H. atherstoni. The shell
accordingly retains a rather more globose aspect, the flank of the
whorl is less extensive and less flattened and becomes more insensibly
merged in the broad peripheral area, while the umbilical cavity is
correspondingly rather more profound. The dimensions of the
specimen, which comprises a considerable part of the body-
chamber, are as follows :

Greatest diameter 130 mm.

Greatest breadth of the last whorl in cross-section

(estimated) 95 , ,

* Kilian (4), p. 866, pi. Ivii., figs. 2a, 1b.

t Oppel (2), p. 286, Tab. 81, figs. 4-4c; Uhlig (4), p. 130, pi. xviii., figs. 2a-2c.

I Eichwald (2), pi. viii., fig. 13.

194 Annals of tlie South African Museum.

Greatest height of the last whorl at the centre, in section 35 mm.
Greatest diameter of the umbilicus, measured from the

umbilical rim between the tubercles 48 ,,

In the type-specimen of H. atherstoni the maximum diameter is
140 mm., the greatest breadth of the last whorl 90 mm., and the
height of the last whorl in section at its anterior part 42 mm. It is
thus seen that the height of the whorl-section in relation to its
breadth is somewhat different. In the more inflated specimen, when
the breadth of the whorl is 86 mm. its height at the centre is 32 mm. ;
in H. atherstoni (type-specimen) I have estimated that with a
breadth of 86 mm. the height is 40 mm. The two left-hand
specimens, depicted in greatly reduced size, in a text-figure pub-
lished by Drs. Hatch and Corstorphine,* may perhaps represent this
inflated form.

Mr. G. C. Crick has shown me a specimen in the British Museum
(registered 46534) which closely agrees with the one here discussed.
It is from the Sunday's Eiver (from A. G. Bain's collection), and has
at some time been determined as Olcostephanus atherstoni. It has
an inflated form and breadth of whorl-section which seem to dis-
tinguish it from H. atherstoni, as exemplified by Sharpe's figured
type, in the manner described above in the case of the other
specimen. The greatest diameter of this individual in the British
Museum is 145 mm. The greatest breadth of the ultimate whorl
would, when complete, have been almost 100 mm., while the height
of this whorl at the centre at its anterior end would have slightly ex-
ceeded 35 mm. In the breadth of the umbilicus and in all other
characters there is the closest agreement with the specimen from the
South African Museum. This would seem to indicate that fixed and
reliable characters are exhibited which may suffice to separate this
more globose shell from the true H. atherstoni, and it might perhaps
be possible to establish the constancy of these distinctive features
were a suite of specimens of undoubted H. atherstoni, at a suitable
stage of growth, available for comparison. Unfortunately this is not
the case, and the only specimen of H. atherstoni that I have seen
which illustrates a comparable stage is Sharpe's figured type. Com-
parison with this has revealed the slight shape differences above
noted, but beyond this there is entire agreement ; the characteristic
ornamentation, the degree of involution and the width of the
umbilicus are the same. There is as yet no material for a
comparison of the septal sutures. I therefore do not yet feel
justified in proposing a new specific name for the specimens here
* Hatch and Corstorphine (1), p. 243, fig. 65 (as Olcostephanus bainii).

The Invertebrate Fauna of the Uitenhage Series. 195

dealt with, and whatever may ultimately become known regarding
their exact relationship to H. athcrstoni, which they so closely
resemble, it is certain that so far as present evidence goes, this
relationship must be presumed to be very intimate.

HOLCOSTEPHANUS WILMAN.E Sp. nOV.

Plate IX., figs. 1, la.

Description of a Single Specimen. The shell is laterally much com-
pressed and is discoidal in form ; the flattened flanks of the ultimate
whorl are relatively extensive, while the peripheral area is narrow and
presents a somewhat flatly convex surface. The whorl is narrow and
highly arched in section, and the height of the whorl-section,
measured at the middle, is almost equal to the breadth, which is
greatest at the umbilical margin. The whorl, however, is almost
parallel-sided, so that the breadth of its section taken at the middle
of the flank is little less than at the umbilical tubercles. The in-
volution is such that about two-thirds of the preceding whorl are
embraced.

The umbilicus is relatively shallow, and occupies one-third of the
total diameter. The umbilical rim is not abruptly defined, but the
surface of the flank curves down somewhat gradually to the umbilical
wall, which falls very steeply to the spiral suture. Primary umbilical
ribs commencing at the suture slope back very obliquely, and, quickly
gathering strength, swell up at the umbilical rim to form strong,
elongated, much-compressed tubercles, present to the number of
eighteen in the ultimate whorl. The secondary ribs proceed from the
tubercles chiefly in groups of three, but there are occasionally four ribs
in a group in the ultimate whorl, and frequently a rib is inserted so
as to terminate between two tubercles. The ribs at first swing
forwards from the tubercle, and bend back at the middle of the flank
to take an almost direct course across the periphery. The ribs are
prominent in character, and on the peripheral area in the anterior
part of the last whorl their crests are 4 mm. apart.

Immediately behind the oral margin is a strong constriction, and a
portion of a lateral auricular extension is preserved.

Dimensions.

Greatest diameter 90 mm.

Greatest breadth of the last whorl in cross-section 30 ,,

Height of the last whorl at the centre, in section 28 ,,

Greatest diameter of the umbilicus, measured from the

umbilical rim between the tubercles 30

196 Annals of the South African Museum.

Occurrence. The specimen is in the collection of the Geological
Society of London (registered 10975A), and the label affixed to it
bears the following record of locality and collector : " Aasvogel
Krantz above Modder Drift, Sunday's River, S. Africa. Dr.
Atherstone, F.G.S., 1876."

Remarks. The individual above described is almost complete,
comprising the body-chamber, but the exact length of this cannot be
ascertained, nor is the shell so preserved that the lobe-line is any-
where visible. The specimen has undergone some distortion from
crushing, but making every allowance for this, it is clear that we are
dealing with a form well distinguished by its high whorl-section and
extensive flattened flanks. In the discoidal form, the relatively
shallow umbilicus, and the degree of involution, we are reminded of
H. asticrianus (d'Orb.),* but H. wilmance is readily separated from
this by its very strong, oblique umbilical ribs and tubercles, and by
the inclination and the coarseness of the secondary costse.

This appears to be the specimen mentioned by Pavlow t as a
compressed variety of H. atherstoni, but it was recognised that
the distinctive characters are sufficient to warrant a new specific
name (op. cit. p. 492). As pointed out by Pavlow, the specimen
is distinguished from the typical H. atherstoni by the presence
of rather fewer umbilical ribs and by the more strongly developed
secondary costse. In addition, H. atherstoni is a considerably
more strongly inflated shell, and has a relatively narrower and
much more profound umbilicus. The points of distinction are in
fact so great that the only course open is to definitely separate
the two forms.

There is much stronger resemblance to H. psilostomus Neum. and
Uhlig,:J from the Neocomian of North Germany. The ribbing is of
very similar character, but in H. psilostomus the umbilical ribs are
rather more numerous, the breadth of the umbilicus is somewhat
greater relatively to that of the flank, and the shell, although much
more laterally compressed than H. atherstoni, is still considerably
more inflated and less discoidal than in H. wilmance. I believe that
the presence of the above characters of distinction must be con-
sidered sufficient to warrant the provisional separation of these two
forms, but there can be little doubt that the relationship is intimate,
and there is no other known Holcostephanus with which H. ivilmance
can be brought into such close comparison.

* d'Orbigny (1), p. 115, pi. 28, fig. 1 (1840).

f Pavlow and Lamplugh (1), p. 496 (p. 138 of authors' copy).

\ Neumayr and Uhlig (1), p. 149, Taf. xxxii., fig. 2.

The Invertebrate Fauna of the. Uitenhage Series. 197

A shell from the Lower Hauterivian of North Germany described
by von Koenen :;: under the name Astieria aff. psilostoma also shows
great similarity to H. wilmancs, but may be readily distinguished by
the wider umbilicus, the strongly developed constrictions, and the
more inflated and less discoidal form of the whorls.

Pavlow thought that small specimens from the Neocomian of
Spain, figured by Nickles \ as Holcostephanus hispanicits, might
possibly belong to this form ; but though a thorough comparison
with these figures is difficult, and the character of the ribbing
appears to be very similar, it seems to me most probable that the
much greater inflation of the Spanish shell would be found sufficient
to separate it. If the specimen figured by Nickles in fig. 11
represents the same form as that depicted in fig. 3, then it is clear
that H. wilmancB is a much more discoidal shell. The small
specimen represented by Nickles in fig. 10 is only drawn in side
view, but it appears to possess a prominence of the umbilical rim
and a depth of the umbilical cavity which may be taken to indicate
the somewhat strongly inflated character of the shell. Pavlow draws
attention to the existence of two specimens which he identifies with
the South African form above described ; the one is from Aigles
(Basses-Alpes), preserved in the Pictet collection in the museum at
Geneva, and the other, from the Shasta group of California, is in the
geological collection of the University of Moscow.

Holcostephanus tonsbergensis (Weerth),} from the Neocomian of
the Teutoburger Wald, may possibly also be related, so far as can be
judged from a comparison with Weerth's figure 4.

HOLCOSTEPHANUS BAINI (Sharpe).

1856. Ammonites bainii D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

vol. vii., p. 197, pi. xxiii., fig. 2.
1882. Olcostephanus baini M. Neumayr, in E. Holub and M.

Neumayr, Denkschr. d. k. Akad. Wiss., Math-Nat. Cl.,

Band xliv., p. 272.

Although no specimen undoubtedly referable to this form is
included in the collection under examination, a few remarks under
this heading may be useful as a preface to the account of an
individual, to be described below, which I believe to be very closely
related to H. baini.

* von Koenen (2), p. 151, Taf. liv., fig. 2.
t Nickles (1), pi. ix., figs. 3 and 10.
I Weerth (1), p. 16, Taf. iv., fig. 4.
14

198 Annals of the South African Museum.

Occurrence. H. baini has been found at localities on the Zwart-
kop's and Sunday's Eivers, and its occurrence in the Zwartkop's
River Valley below Uitenhage has been noted by Messrs. Rogers and
Schwarz.* Specimens in the collection of the Geological Society
(Sharpe's type, registered 10976A) and in the British Museum
(registered 52052) are from the Sunday's River.

Remarks. Neumayr thought H. baini to be most closely related
to H. schenki (Oppel),f from the Spiti Shales of Tibet, and there is
indeed a great resemblance between the two in all the more impor-
tant features ; but H. baini is more closely ribbed, and as mentioned
by Uhlig, ]: in regard to the fineness of the ribbing, H. schenki
occupies a position just midway between H. baini and H. atherstoni.
Neumayr and Uhlig also thought close relationship to exist between
H. baini and H. keyserlingi Neum. and Uhl. from the Neocomian of
North Germany, but the manner of division of the ribs is really a
strong distinguishing feature. In H. baini none of the ribs bifurcate
after leaving the nodes. The relationship of H. baini to H. diptyclms
(Keys.) and H. polyptyclms (Keys.) | from Petschora-Land is also no
doubt much more remote than Neumayr supposed, and these forms,
as well as H. keyserlingi, clearly do not belong to the same division
of the Holcostephani (sensu lato) as H. baini and H. schenki.
Nikitin,*: in fact,, has strongly criticised Neumayr's suggestion of
relationship between H. schenki and these two ammonites from
Petschora-Land, and Pavlow has also referred to this matter.**

Remarks on the points of resemblance between H. baini and
H. rogersi sp. nov. will be found appended to the description of the
latter given in these pages.

Neumayr ft dealt fully with the question of a suggested resem-
blance between H. baini and a Jurassic ammonite from Madagascar
ascribed to J. Sowerby's Ammonites herveyi,H and showed that these
really have no significant characters in common. The points of
resemblance are in fact so slender as not to merit further discussion
here ; but it may be remarked that Neumayr probably misrepre-
sented the case when he wrote as follows : " Newton thinks to have
found close relationship between Macroceplialites herveyi from
Madagascar and Olcostephanus baini Sharpe from the Uitenhage

* Kogers and Schwarz (1), p. 7.

t Oppel (2), p. 286, Tab. 81, figs. 4a-c ; Uhlig (4), p. 130, pi. xviii., figs. 2a-c.

I Uhlig (4), p. 132. Neumayr and Uhlig (1), p. 156.

|| Keyserling (1), p. 327, pis. xx., figs. 4, 5 ; xxi., figs. 1-3 ; xxii., figs. 9, 10.

IF Nikitin (1), p. 122.

** Pavlow and Lamplugh (1), p. 488 (p. 130 of authors' copy).

ft Neumayr (5), p. 6. }} Newton (1), p. 334.

The Invertebrate Fauna of the Uitenhage Series. 199

Formation." In the description of the Madagascar fossils, the
opinion that these two forms are truly related was not expressed,
and it is not necessary to infer that such a view was actually
entertained.

Kilian :;c has suggested that H. baini may represent an immature
stage of H. athcrstoni (Sharpe), but this is certainly not the case.
H. atherstoni at a comparable growth-stage is much more finely
ribbed, to mention only one point of distinction.

With regard to the type-specimen of Ammonites baini Sharpe,
preserved in the collection of the Geological Society, it is perhaps
well to note that in the late Prof. J. F. Blake's published list of the
types and figured specimens in the Society's museum, reference is
accidentally made to a wrong specimen.! The individual figured by
Sharpe was unfortunately overlooked by Mr. C. D. Sherborn when
preparing a manuscript catalogue of the collection some years ago.
The specimen erroneously noted as the type (registered 10976) was
presented by Atherstone and bears the date 1876 ; J I consider it to
represent a hitherto undescribed form, and it is dealt with in these
pages under the name Holcosteplianus modderensis.

HOLCOSTEPHANUS cf. BAINI (Sharpe).
Plate IX, fig. 2 ; X., fig. 1.

A single specimen, from the collection in the South African
Museum, agrees so closely with Sharpe's Ammonites baini that
it has seemed questionable whether it should not be considered
identical.

Description. The specimen, which is considerably larger than
known examples of H. baini, consists of at least four whorls, and
the form of these as well as the degree of overlap in the earlier
whorls is the same as in H. baini. Some of the shell substance is
retained and the specimen is partly preserved as a cast, but it
exhibits in very imperfect manner the course of the septal sutures,
so that these cannot be satisfactorily copied for illustration. The
body-chamber is not preserved. In the interior whorls the umbilical
wall of each whorl falls upon the umbilical tubercles of the pre-
ceding whorl, but in the latter half of the last whorl the degree of
involution becomes slightly reduced, so that the nodes of the
preceding whorl are completely exposed and the beginnings of the
Hank ribs proceeding from them can be just discerned. At the same

* Kilian (4), p. 865. t Blake (2), p. 59.

\ Sharpe's figure of H. baini was published in 1856.

200 Annals of the Sonth African Museum.

time, the slope of the umbilical wall of the last whorl becomes
slightly less steep.

There are sixteen strong umbilical marginal tubercles in the last
whorl preserved, and from these the ribs of the flank and periphery
proceed most frequently in groups of three, though occasionally only
two ribs are given off from the nodes. Sometimes a rib may be
situated between two nodes at one of its extremities, while springing
from a node at its opposite end. These relations of the ribs, how-
ever, are the same in H. baini. The ribs as they depart from the
tubercles to cross over the periphery are not forwardly inclined, but
pass almost straight across the whorl. In the most advanced
portion of the specimen the crests of the ribs upon the peripheral
area are situated at a distance of 6 mm. apart. Well-marked
constrictions are present as in H. baini.

Dimensions.

Greatest diameter of specimen 92 mm .

Greatest breadth of last whorl in cross-section 53 ,,

Greatest height of last whorl at the centre (section) 24 ,,

Greatest diameter of the umbilicus, measured from the

umbilical rim between the tubercles 45 ,,

Occurrence. This specimen is believed to come from the Sunday's
Eiver.

Remarks. There can be no doubt that the form here represented
is very closely related to H. baini, and I was for some time
undecided whether definitely to identify the two. Allowing for the
difference in dimensions between this specimen and the examples of
H. baini that I have seen, which are smaller, the only apparently
valid character of distinction is in the disposition of the ribs. In
H. baini the ribs proceeding over the whorl from the nodes have a
well-marked forward throw, but such an inclination is not seen in
the specimen above described. Mr. G. C. Crick, who has kindly
examined the specimen, considers that this is a significant point of
distinction, but admits that in other respects the two forms are very
closely similar. With regard to the inclination of the ribs as a
constant and reliable character, it may be remarked that in the
British Museum collection a specimen of H. baini (registered 52052)
which strikingly resembles Sharpe's type in all other points, agrees
with it also in showing the same inclination of the ribs. It therefore
appears evident that in this character of the ribbing we have a
distinctive feature for separation, whatever this alone may be w r orth.
There is unfortunately as yet only a single specimen available for
comparison, and since in other respects it shows such close agree-

The Invertebrate Fauna of the Uitenkage Series. 201

meat with H. baini, I am unable to arrive at a definite opinion as to
the nearness of its relationship to that form, and feel hardly justified
in proposing a new specific name. It is to be hoped that the acquisi-
tion of additional material may enable a more thorough comparison to
be made and lead to a clearer knowledge of the actual relationships.

The specimen has much general resemblance to H. rogersi sp.
nov., but is more coarsely ribbed. In H. rogersi, too, the ribbing
shows a still greater departure from that of H. baini, in that the
secondary costae as they proceed from the tubercles are not truly
direct in their course across the periphery, but have a very sli ht
backward throw.

HOLCOSTEPHANUS KOGERSI Sp. 11OV.

Plate IX., fig. 3 ; X., fig. 2.

Description of a Single Specimen. The shell is involute to the
umbilical tubercules, and the ultimate whorl is broad in section
and broadly rounded on the peripheral area. The umbilical wall
falls very steeply and the umbilical rim is bluntly rounded. The
primary ribs, commencing above the spiral suture, are narrow,
and rather weakly developed. They have a slight backward slope
as they are traced up to the marginal nodes. There are about
sixteen of these laterally compressed, not very prominently
developed tubercles in the ultimate whorl. From these the ribs
of the flank proceed mostly in groups of three, though there
are occasionally only two ribs terminating at a node, or a rib-ending
may now and then fall between two nodes. These ribs as they pass
from the tubercles over the peripheral area are very slightly reclined,
that is, have a backward slope in relation to a radius. In the
anterior portion of the ultimate whorl the crests of the ribs on the
peripheral area are separated from one another by a space of 3 mm.

Two strongly developed constrictions are visible on the ultimate
whorl, and they are situated opposite to one another. The specimen
is entirely septate, but so preserved that the figure of the lobe-line
cannot be traced.

Dimensions.

Greatest diameter 63 mm.

Greatest breadth of the last whorl in cross-section 40 ,,

Height of the last whorl at the centre, in section 16 ,,

Greatest diameter of the umbilicus, measured from the

umbilical rim between the tubercles 25 ,,

Occurrence. The specimen, in the collection of the South African
Museum, is from the Sunday's Eiver.

202 Annals of the South African Museum.

Remarks. Although I have only seen a single specimen, and
the state of preservation of this is not so good as might be desired,
I have no hesitation in pronouncing it to represent a hitherto
undescribed form. It is apparent that though distinctively charac-
terised, particularly by the feature of reclined costation, this
ammonite may be brought into closest relationship with H. baini
(Sharpe) and H. schenki (Oppel), and particularly with the latter.
H. baini has the whorl relatively narrower and more highly arched
in section, while its ribbing is noticeably coarser. On the peri-
pheral part of the whorl at a comparable stage there are seven ribs
in H. rogersi within a space which includes only six ribs in
H. baini. Moreover, the ribs in H. baini have a marked forward
inclination as they depart from the nodes.

Eeference to the published descriptions and figures of H. schenki,"
from Shangra, east of Puling (Tibet), seemed to show a very close
resemblance between the two forms, but I have been able to
supplement this by an actual comparison with Oppel's type-
specimen in the State Palaeontological Collection at Munich, and
when making this comparison I was fortunate in having the kind
assistance of Prof. J. F. Pompeckj. H. schenki has rather more
numerous umbilical tubercles and is slightly more densely ribbed
than H. rogersi. In H. schenki the umbilicus is relatively a little
wider and in cross-section the whorl is rather wider between the
umbilical rims, in relation to the height. The most notable point
of distinction, however, is in the course taken by the secondary
ribs as they pass from the tubercles over the periphery. In
H. schenki they have a forward inclination, while in H. rogersi they
are slightly reclined.

There is considerable resemblance between H. rogersi and H.
guebhardi Kilian, i from the Hauterivian of the neighbourhood of
Escragnolles in the Maritime Alps, but in H. guebhardi the ribbing
of the flank is strictly radial in direction.

HOLCOSTEPHANUS MODDEKENSIS Sp. nOV

Plate X., figs 3, 3a.

1892. Astieria schenki A. P. Pavlow (partim), Bull. Soc. Imp. Nat.

Mosc., annee 1891, Nouv. Ser., tome v., p. 493 (p. 135

of authors' copy).

Description of a Single Specimen. The shell has rapidly expanding
whorls, and is involute to the umbilical marginal tubercles in the

* Oppel (2), p.