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AUNEN ASS

OF THE

SOUTH AFRICAN MUSEUM

VOLUME VII.

ANNALS

OF THE

SOUTH AFRICAN MUSEUM

VOLUME VII.

DESCRIPTIONS or tue PALHONTOLOGICAL MATERIAL

COLLECTED sy tae OFFICERS or tHe GEOLOGICAL SURVEY

oF CAPE COLONY anp OTHERS.

PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

AND

THE GEOLOGICAL COMMISSION or tHe COLONY or THE

CAPE OF GOOD HOPE

BY West, Newman & Co., LONDON

1913.

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TRUSTEES OF THE SOUTH AFRICAN MUSEUM.

The Right Hon. Jonn Xavier Merriman, M.L.A.

Tuomas Murr, C.M.G., LL.D., M.A., F.R.S., F.R.S.E., Superintendent-General

of Education.

JoHN Wituism Jaccer, M.L.A., F.R.Stat.S.

SCIENTIFIC STAFF OF THE SOUTH AFRICAN

MUSEUM.

Louis ALBERT PrértncuEy, D.Sc., F.Z.S., F.E.S., Director.

Wiuu1amM Frepertck Purcewt, B.A., Ph.D., Keeper of Land Invertebrates, except

Insects. /

ArrHur Witttam Rocers, D.Se., F.G.S., Keeper of the Geological and Minera-

logical Collections.

Rosert Broom, D.Se., M.D., Keeper of the Paleontological Collections.

EK. P. Putnuirs, M.A., F.L.S., Assistant in Charge of the Herbarium.

K. H. Barnarp, B.A., Assistant in Charge of Fish and Marine Invertebrate

Collections.

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

LIST OF CONTRIBUTORS.

C. W. ANDREWs.

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

Uitenhage Beds of Cape Colony ......... HS a Gin GoUuDOmS ab OOOO

R. Broom.

F.

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

Notice of Some New South African Fossil Amphibians and Reptiles ..

On-a/large Extinct Species Of Buvalte acne o<seciee deme cee eee nee

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

On the Shoulder Girdle of Gynognathus 12.2. ..s.ecnosr see eeenieneee

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

I YE OO: 5% wie Gye se cuslersieks giao Seti aistne ciase Ginie Ateiata. lain © ape RE RTE es

On the Dinosaurs of the Stormberg, South Africa ............eceeeeee

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

MUTED S. ete crete cle iche ays ernie et sieisyens eileen roneiertercieriiiciereereiicioets

Ona Species of Tylosaurus from the Upper Cretaceous Beds of Pondoland

On a New Type of Cynodont from the Stormberg ..............-e000-

On Some Points in the Structure of the Dicynodont Skull ............

OntthesManustandePes ol Pan erasais cei-reie elelerelersieieieicie cieicicineinie siete

On a New Mesosaurian Reptile (Noteosaurus africanus)........ 0.00000.

AURevisionlof the: Reptilesiofithe Karrool ca. acetic ce eerie:

L. Kireuin.

The Invertebrate Fauna and Paleontological Relations of the Uitenhage

DELIOS sic. svarole olrsesole. snevel cena) opstote otonete oie: aes ey avsnereleteasvs re ieyenetere ieee toner:

W. D. Lane.

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

Camp Bultalowivertecrietetrereretiiierrs scl lorie toler ttc ttciratcrer-

H. Woops.

Echinoidea, Brachiopoda, and Lamellibranchia from the Upper Creta-

ceous Limestone of Need’s Camp, Buffalo River ................6-

PAGE

309

LIST OF PLATES.

PLATE

|

|

.

|

|

Filisparsa ramosa, d’Orbigny.

Filisparsa fragilis, Marsson.

Diastopora compressa (Goldfuss).

Idmonea virgula, dOrbigny.

Crisina cenomana, d’Orbigny.

Crisina excavata, d’Orbigny.

Crisina marginata, d’Orbigny.

Tervia dorsata (von Hagenow).

Tervia gibbera, Gregory.

Entalophora virgula (von Hagenovw).

Entalophora conjugata (von Reuss).

Kntalophora echinata (Romer).

Spiropora verticillata (Goldfuss).

Entalophora madreporacea (Goldfuss).

Coptosoma capense, sp. nov.

Serpula cf. concava (J. Sow.).

Pecten (Syncyclonema) orbicularis, J. Sow.

Pecten (Camptonectes) cottaldinus, d’Orbigny

Pecten (Camptonectes) projectus, V'ate.

Pecten (Chlamys) cf. subacutus, Lam.

Lima (Acesta) obliquissima, Tate.

Lima (Mantellum) neglecta, Tate.

Perna atherstoni, Sharpe.

Mytilus uitenhagensis, sp. noy.

Modiola baini, Sharpe.

Nucula uitenhagensis, sp. nov.

Grammatodon jonesi (Tate).

Unio uitenhagensis, sp. nov.

Trigonia ventricosa (Krauss).

Trigonia kraussi, sp. nov.

Trigonia rogersi, sp. nov.

Trigonia rogersi, sp. nov.

Trigonia holubi, sp. noy.

Trigonia herzogi, sp. nov.

Trigonia rogersi, sp. nov.

Trigonia vau, Sharpe.

Trigonia stowi, sp. noy.

vill List of Plates.

PLATE

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.

VII.

NV

Meretrix uitenhagensis, sp. nov.

Mactra? dubia, sp. noy.

Pleuromya baini, Sharpe.

| Thracia sp.

|

Turbo atherstoni, Sharpe.

Turbo rogersi, sp. nov.

Turbo minutulus, sp. nov.

VIII. Natica uitenhagensis, sp. nov.

Natica rogersi, sp. nov.

Natica? mirifica, sp. nov.

Actzonina atherstoni (Sharpe).

| Limnea remota, sp. nov.

Phylloceras rogersi, sp. nov.

Bochianites glaber, sp. nov.

Meyeria schwarzi, sp. noy.

Holeostephanus wilmane, sp. nov.

Holecostephanus cf. baini (Sharpe).

Holcostephanus rogersi, sp. nov.

Meyeria schwayzi, sp. nov.

U

|

Holeostephanus cf. baini (Sharpe).

Holcostephanus rogersi, sp. nov.

Holcostephanus modderensis, sp. nov.

Meyeria schwarzi, sp. nov.

XI. Holcostephanus uitenhagensis, sp. nov.

Hybodus africanus, n. sp.

Ccelacanthus africanus, Broom.

Ceratodus ornatus, n. sp.

Hydropessum kannemeyeri, g. et sp. nov.

Helichthys draperi, S.-Voodward.

| Helichthys browni, g. et sp. nov.

Oxygnathus browni, n. sp.

| Cleithrolepis minor, n. sp.

. Pholidophorus browni, n. sp.

XIV. Gryponyx africanus, g. et sp. nov.

( Gryponyx africanus, g. et sp. nov.

| Massospondylus carinatus, Owen.

‘| Massospondylus harriesi, sp. nov.

| Aitonyx palustris, g. et sp. nov.

List of New Generic Names. 1X

PLATE

XVI. Massospondylus harriesi, sp. nov.

Massospondylus harriesi, sp. nov.

XVII.) Aitonyx palustris, g. et sp. nov.

. Geranosaurus atavus, g. et sp. nov.

XVIII. Plesiosaurus capensis, sp. nov.

XIX.— ( Pr ; ;

XXI, | Propappus rogersi, sp. nov.

-xr1_ ( Lylosaurus capensis, sp. nov.

XXIL. ; ; ;

( 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 GENERIC NAMES

INTRODUCED IN THIS VOLUME.

PAGE

PULOMY Kawi.) Yen MIN OSAULIA) sO LOOM y araleicls)scleveveiele re «olor eieis)o clove siels\ else BOC DOE 304

aura Oy (Cynodontia) BLOOM | er airs eve eye siete a cieole ele eve sel 1« © elle) sie s)e) ese 272

IBICCASAULU Selo Oe LST OOMMUmEctay evarerereietrenelen ietslcieielercicl sieves ollelcloielorerocciele 11 lsaieiere 276

Geranosaurus; n..¢. (Dinosauria); Broom “Vase... s 2.66 ie cee eee nesses 306

Cryponyx, ne 2. (Dinosauria), Broome nrirciis| <6 aie lsc elocie ese ao «cle elsialeie 294

Gyposaurus, n> g> (Dinosauria); Broom) <3. 5 ~ 2ccie os oe ewe ee lel nse 293

Heleophilus, n. g. (Mesosauria), Broom ......... 2... cece cece creo cece cone 277

Helichthys, n. g. (Actinopterygii), Broom. ... 2... 52.2... 000cee st cess ewees 254

Hydropessum, n. g. (Actinopterygii), Broom .......... 2022 ceeeceeeeeeecs 266

Noteosaurus, n. g. (Proganosauria), Broom ........ 0... cece rece eee ee eeee 308

Tritheledon, n. g. (Cynodontia), Broom............ccccsceeeceres pG00G0C 334

INDEX OF GENERA AND

SPECIES.

——>———_

A. PAGE PAGE

Bocutanires, Lory........ ovonon 181

ACANTHODISCUS, (WiUtlig 20 «cies si «le 207 borcherdsi (Cyprina), Tate ...... 150

Acusta, H. d A. Adams ........ 71 | bronni (Astarte), Krauss ........ 26

AcTtonina, @Orbigny ......6.4. 176 | bronni (Seebachia),............ 43, 47

acutus (Heleophilus), Broom .... 277 | browni (Euskelesaurus), Hualey.. 292

ANLUROSAURUS, Owen ......00%- 363 browni (Helichthys), Broom...... 254

/ALURosUCHUS, Broom .......... 364 browni (Massospondylus), Seeley 293

AMMORWOS JRO oa 5acnaoon Kc 304, 364 ~browni (Oxygnathus), Broom . 259

africanus (Belemnites), J'ate .... 27 browni (Pholidophorus), Broom .. 267

africanus (Capitosaurus), Broom.. 271 Busauis, Hamilton-Smith........ 279

africanus (Coelacanthus), Broom.. 253 |

africanus (Gryponyx), Broom .... 294

africanus (Hybodus), Broom .... 252 C.

africanus (Noteosaurus), Broom.. 358

ALOPECODON, Broom .....sesccee 363

ALOPOSAURUS, Broom ...eeseee . 3863

AUMMMONITES Gysveieisasie'sie eteus.s cictel 25

angulosa (Idmonea), d’Orbigny .. 6

ANTHODON, Owen weer eeots cect 362

IAN THHONIWAN MG LDU St ieiererenerelersiersrerniete BY,

antipodum (Berenicea), Tate .... 33

ARCHHOSUCHUS, Broom ....+..0+- 062

ARCTOGNATHUS, Broom ........<. 363

ARCTOSUCHUS: (BROONUs oislelelele cieierele 363

ARNOGNATHUS, Broom ........-. 363

AsTARTE, J. Sowerby ........ 26, 128

atavus (Geranosaurus), Broom 306

atherstoni (Acteonina), Sharpe 37, 176

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 (Holecostephanus), Sharpe

27, 28, 31, 197

baini (Modiola), Sharpe ........

eee (Mytilus), Sharpe.......... 33

baini (Pareiasaurus), Seeley...... 353

baini (Pleuromya), Sharpe ...... 167

IBAURIA, SB TOOTUs merieeienieeiee ~. 212, 364

BELEMNITES, Lamarck .......... 210

BERENICEA, Lamarch..........<- 33

calypso (Idmonea), @’Orbigny .. 5

Campronrecres, Meek .......... 65

capense (Coptosoma), Woods .... 13

capensis (Ceratodus), S.-Woodward 253

| capensis (Equus), Broom ........ 281

| capensis (Gyposaurus), Broom .. 293

capensis (Orinosaurus), Lydekker 292

capensis (Plesiosaurus), Andrews 309

capensis (Semionotus), S.- Woodward 262

| capensis (Tylosaurus), Broom .... 332

caperata (Patella), Yate ........ 163

Capitosaurus, Milnster ........ 271

CARDITA, BrUguiene | lereiieie rere 127

carinatus (Massospondylus), Owen 291

CARYOPHYLLIA, Stokes ........¢. Ti

cassiope (Trigonia), Z'ate 32, 125

cenomana (Crisina), d’Orbigny .. 5

CERATODUS PA GASSI2 \ selene 253

CHELYPosauRUsS, Broom ...... bo ahos}

CHDAM YS) BOLtEN) > «sewlereerics 69

CISTECEPHALUS, Owen ......+.+- 363

| CLEITHROLEPIS, Heerton ........ 264

C@LACANTHUS, Agussiz .......... 253

85 |

complicata (Crassatella), Tate .. 26, 27

compressa (Diastopora), Goldfuss 3

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

conjugata (Entalophora), v. Reuss 9

conocardiiformiis (Trigonia), Krauss

23, 27, 35, 42, 119

cottaldinus (Pecten), d’Orbigny

37, 48, 65

| CRASSATELLA, Lamarck ........ 26

| Crioceras, @Orbigny ........6. 27

Index of Genera and Species.

PAGE

CRISINARGONDIGNI!... esse oss s 4

Cryprocynopon, Seeley ........ 363

CYNOCHAMPSA, Owen ..........-. 363

CYNoDRACO, Owen ......... sadcd. ett

CynoGNaTHus, Seeley ........ 283, 363

cynops (Bauria), Broom ........- 272

CyNosucHuS, Owen..... Sher ecote. bess 363

CXPRINAS LEQMATCK seo cies vic cccee 149

D.

decurrens (Tervia), Pocta ...... U

DELPHINOGNATHUs, Seeley ...... 362

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

DrapEMopoN, Seeley ...........- 364

IDTAALURODON, BROOM we <cls cece. 363

Drasropora, Lamouroux ........ 2

DictyoryGE, Egertoit.......ee.0- 262

IDTGYNODON, O1ven aeons sss 339, 363

disticha (Idmonea), Michelin .... 5

dominicalis (Gastrochena), Sharpe 162

dominicalis (Pholadomya), Sharpe

dorsata (Tervia), v. Hagenow ....

draperi (Helichthys), S.-Woodward 257

dubia (Mactra?) Kitchin ........ 156

E.

KiccasavRus, Broom .......-- 276, 362

echinata (Entalophora), Rémer 9

IDWOADOERE TROON comoonooou0doK 363

EINDOTHIODON, Owen .....0..00.. 363

FENNTALOPHORA, Lamouroux ...... 8

IHOSUCHUSS Vatson eeicce sect eens 364

HI QUUS IStNTUCUS) aeleieneeie oreie ole oie Bo eel

ErtpHostoma, Broom ......- 363

ERIPHYDA, Stoliczka ..........>- 128

ErytHrosucuus, Broom.......... 364

Kunorosaurus, Seeley ......... . 364

Kurarxerta, Broom (ined.) ...... 366

KuskeLesaurus, Hualey...... 292, 364

excavata (Crisina), d’Orbigny.... 6

JaPOERAIS MG) céocuacapoducauc 16, 77

extoni (Cleithrolepis), S.-Woodward 264

F.

Finisparsa, d@’Orbigny .......... 1

formosa (Dictyopyge), Broom .... 262

fragilis (Filisparsa), Marsson .... 2

fungatus (Hamites), Brongniart.. 25

G.

GALEcHIRUS, Broom .......... 55 6B

GAMMORSHB Oona cece s «aeciee 362

GaLErPus, Broom ......... Son00 — Gly)

|

|

33 |

7

XI

PAGE

GALESAURUS, Owen .....-.....-- 363

GASTROCHENA, Spengler ..... 162

GERANOSAURUS, Broom ...... 306, 364

gibbera (Tervia), Gregory .... 7

glaber (Bochianites), Kitchin .. ‘38, 181

GuaNnosucHus, Broom...... a Aeisis.s 363

goldfussi (Trigonia), Tate ...... 32

GomPHoGNatuHus, Seeley .......- 364

Goniomya, J. L. Agassiz ........ 159

Gorconops, OLWAD coecoocdcooode 363

Gramraronox, Meek & Hayden ws 88

GROTRIANIA, Speyer ..... SOeuOCC 26

(Carvaronpec, JERI coodeoudce 294, 364

GYPOSAURUS, Broom ........ 293, 364

18,

Jaana, J2ANts GoesoodbonoeDoUC 25

harriesi (Massospondylus), Broom 299

HELEOPHILUS, Broom........0-0- 277

HELEOSAURUS, Broom .......- s 364

IIELEOSUCHUS, Broont: o.oo... .c oc 364

HeuicutuHys, Broom ...... apetevete 254

herzogi (Astarte), Goldfuss.. 26, 47,128

herzogi (Lyrodon), Goldfuss .... 25

herzogi (Trigonia), Goldfuss 34, 40, 101

HoxcostrerHanus, Neumayr...... 184

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

Horranorarsus, Seeley ......-- 293

FLOWISTA, 15 7;OOT0N werctclelcl ace cletch eve A 364

FAlY@NASUCHUS, Broom ....6...0- 363

TLVBODUSs AGQSS1Z) «yee eletse recreate 252

Hyproprssum, Broom .......-.- 266

Tr

IctrDOGNATHUS, Broom .......... 363

Icripoprsis, Broom ..... Snole el arenes 363

JETT OSUCHUSS B?OOM a.) o1eicie) leletetor= 363

IpmoneEa, Lamouroux.........26: +

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

imbricata (Placunopsis), Tate 32

intermedius (Hamites), J. Sowerby 25

J.

jonesi (Grammatodon), Tate 48, 88

jonesiana (Ostrea), Tate ........ 31

K.

kannemeyeri (Ceratodus), Seeley.. 253

_ kannemeyeri (Hydropessum), Broom 266

| kannemeyeri (‘Trematosaurus),

BROOD: oeoanooo oogdDo C040 ; 270

| Karoomys, Broom ......+-.++0+: 364

kolbei (Dicynodon), Broom ...... 338

|

Xll

PAGE

kraussi (Cucullea), Tate ........ 43

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

L.

LacazELuA, M.-Chalmas ..... Sica melD

Lima, Brugwiére ...... oe oo. efi

TIMNawA, Lamarck .......-sss« 178

lineata (Anthonya), Kitchin 137

longlandsiana (Astarte), Tate .. 128

IGYCORHINUS; BOOM \.4 alee cele 363

Tivcosaumus,, Oven ms eeiciielcicde- 363

LV COSUCHUS POON Mee eine ce icmare 363

TAVRODONI re aye oto.sie. 5 0 Stee kce's ev ercions 25

LystTROSAURUS, Cope ...........- 363

M.

Macrra, Linneus . - L566

madreporacea (Entalophora), Gold-

UBS) wee asks, lel wes aue tere avers ore aoe 10

MantELuuM, Bolten ......+.0.0- 74

marginata (Crisina), d@’Orbigny .. 6

Massosponpyuus, Owen .. 291, 293, 364

MELINODON, Broom ....c..ccoes 364

MmREDRIX. 5 QILanClomrciterettete miele 151

MESOSAURUS, Gervais.......-00- 364

MeEsosucuus, Watson ........0. 364

Wibopundy WECM Saococscococ00K6 212

MicroGomeHopon, Seeley ........ 364

minor (Cleithrolepis), S.-Woodward 266

minutulus (Turbo), Kitchin 168

mirifica (Natica?), Kitchin ...... 174

modderensis (Holcostephanus)

did (Op arearn po ht rD SoG OE 202

Moproua, Lamarck .......... ; 85

MoscHoers, Broom ......-.ccee-- 362

Weaniquisy JG CAE) nooncgou90e- 82

ING

INUNCOING ILMB soaocooucoande 171

neglecta (Lima), Z'iate ........ 37, 74

INELTHHAl OD) TOULGMeeerre sere 17

Norrosaurus, Broom........ 358, 364

NorocHampsa, Broom........ 307, 364

INUCULA, Lamarchiae eect 87

nuculoides (Cardita), Tate ...... 127

NYTHOSAURUS, Owen .........+-- 363

O.

obliquissima (Lima), Tate ...... 71

oblonga (Thetironia), Kitchin . 146

orbicularis (Pecten), J. Sowerby... 65

Ontnosaurus, Lydekker ........ 292

ornatus (Ceratodus), Broom...... 253

Index of Genera and Species.

PAGE

OSTREA, Linn@us <<. ..2s0-..0.- iy 7/7

| OstREA (Exocyra), 7’. Say...... 16, 77

OUDENODON, Owen .....2.2-- ae 338

OxyGNnaTHuS, Egerton .......... 209

12

PALAcRoDOoN, Broom ..... sieve 364

IPATTGUAN AGED 70 ONUMETeeinateleelcioe: 364

palustris (Autonyx), Broom ...... 304

papyracea (Thetironia), Sharpe .. 142

PARDOSUCHUS se 1OONtUteerieeierereteler 363

PAREIASAURUS, Owen ........ 353, 362

INGO ID BQN 3565 0000000000 163

IPinlepion, WLP gacoganoan5bo506 65

PrctEn (NritHEa), Droult ...... 17

PELOSUCHUS, Broom seacnceccces 362

PERNA, (BiUguUce4nes veercdeltleleiletere WG 03

PHOLIDOPHORUS, Agassiz ........ 267

PHYLLOCERAS, SwesS &....000 s+ 0s 179

pinchiniana (Astarte), Tate .. 33, 135

pinchiniana (Serpula), Tate ...... 64

INNA LGUNILEUS reels) elelsiereie'e sia 06

Puacunopsis, Morr. & Lycett. . 32

PLEsIosAuRuUsS, Conybeare ........ 309

| SPER UROMYA, A GUSStz teiimcerecle eis 3 Lo”

PLEUROTOMARIA, Defrance ...... 163

priscus (Bubalis), Broom ........ 279

priscus (Kecasaurus), Broom .... 276

PRISTERODON, Hurley .........- 363

PRISTEROGNATHUS, Seeley ........ 363

PROCOLOPHON, Owen ........ 3 362

PropicyNopon, Broom ....... neo GlaB

projectus (Pecten), Tate........ 48, 66

Propaprus, Seeley .......... 322, 02

PROTEROSUCHUS, Broom ......0- 364

PSAMMOBIA. EQManchseiiee cto as 154

PTYCHOMYA, AQQSS1Z <.000002..6 26,210

R.

ramosa (Filisparsa), d’Orbigny .. 2

IREMONDIA, G@b0.. oniceicee ince 26

remota (Limnea), Ivitchin .... 178

yiconoi (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

8.

SAUROSTERNON, Hualey ........ 362

SCALOPOSAURUS, Owen .......0-. 363

Index of Genera and Species. xl

PAGE

ScaPpanopon, Broom ..... ..... 363

schenki (Astieria), Pavlow ...... 202

schwarzi (Meyeria), Kitchin .... 212

schwarzi (Tancredia), Kitchin. . 38, 139

ScyLacoenatuus, Broom (ined.) .. 366

Scynacops, Broom (ined.) ...... 366

ScyLacosaurus, Broom .......... 363

ScymnoenatuHus, Broom ..... .. 3093

Scymnoruinvs, Broom (ined.) .... 366

ScymNosauRus, Broom .........- 363

Srrpacuia, Neumayr............ 26

SEMIONOTUS, Agassiz ............ 262

SERPULA, Linn@us ...........6. 63

SESAMODON, Broom ............ 364

simplex (Reptotubigera), de Loriol 4

skirtopodus (Hortalotarsus), Seeley 293

SoLEcurtTts, de Blainville ...... 155

spinosissimum (Crioceras), Newmayr

27, 31

Sprropora, Lamouroux.......... 10

stowi (Trigonia), Kitchin .. 41, 42,115

subacutus ef. (Pecten), Lamarck.. 09

subanceps (Ammonites), Tate.... 27

subjurensis (Placunopsis), Tate .. 32

SyncycLonemMaA, Meek............ 65

Ty

WAMIBORRIAG SCCLEY) ic aieieeies ee ose) CO

SAN COREDEN, 57 Cettin vaery. «ste nie ciel 139

TAOGNATHUS, Broom ,....-2...... 363

TAPINOCEPHALUS, Owen.......... 362

tatei (Trapezium ?), Kitchin .... 148

tatei (Trigonia), Newmayr ...... 27,34,

38, 40, 125

AUATROPS Di OONUEN sete icieices ite 362

tenuis (Helichthys), Broom...... 258

{ui DUUMED ec baceoocnK0e56N 6

THAMNASTR@A, Lesauvage........ 62

Tuecripea (LacazeELua), W.-Chalmas 15

THECODONTOSAURUS, Broom ......- 364

THELEGNATHUS, Broom.......... 362

PAGE

THeETIRONIA, Stoliczka .......... 142

THRAGCTAS Deachs. a.cac<-s 22.4 L6O

TIGRISUCHUS, Owen ........ 362, 363

TITANOSUCHUS, Owen ......... 363

TrapEezium, v. Miihlfeldt ........ 148

TremaTosacrus, Braun.......... 270

TRIBOLODON, Seeley ...........- 364

TRIGONIA, Brugwiére...........- 91

TrrracHopon, Seeley .......... 364

TRITHELEDON, Broom ........ 334, 364

TROCHOSAURUS, Broom ........0. 363

ToORBONINNECUS) Wao scien ieee 164

TynosAuRuS; Marsh = jase <6 332

U.

| uitenhagensis (Holcostephanus),

TRGECRUID stereos iieea rarest 206

| uitenhagensis (Meretrix), MKitchin 151

| uitenhagensis (Mytilus), Avtchin.. 82

uitenhagensis (Natica), Kitchin .. 171

uitenhagensis (Nucula), Kitchin.. 87

uitenhagensis (Unio), Kitchin.... 89

Wixi to) WYATT cogq aoe Sau0de 6a46 89

V.

29, 41, 42,

48, 49, 110

vau (Trigonia), Sharpe ....

| ventricosa (Trigonia), Krauss 27, 29,

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verticillata (Spiropora), Goldfuss 10

virgula (Entalophora), v. Hagenow 8

virgula (Idmonea), d’Orbigny .... 4

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(1)

1.—Polyzoa and Anthozoa from the Upper Cretaceous Limestone of

Need’s Camp, Buffalo River—By W. D. Lane, 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

Danian of Faxoe, 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.

DIASTOPORIDA.

Genus FILISPARSA, d’Orbigny.

Gregory * places this genus among the Diastoporide because of

its resemblance to Proboscina. He says it may be regarded as

“a group of Proboscine with an erect habit’; and goes on to show

how some species of Pil’sparsa correspond with some of Proboscina.

But he also points out that Pergens{ placed the genus in the

Idmoneide. The nature of the zoarium and distribution of the

zoccia are such that the genus occupies an intermediate place

* J. W. Gregory, B.M. Cat. Cret. Bryozoa, vol. i., 1899, p. 67.

+ HE. Pergens, Rev. de Bry. du Cret. fig. par d’Orb., Bull de la Soc. Belge de

Geol., iii., 1890, Mem., p. 339.

I

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.

FInisPpARsSA RAMOSA, d’Orbigny.

Plate I.,' fig. 1.

1853. Filisparsa ramosa, d’Orbigny, Bry. Crét., p. 819, pl. 756

figs. 18-22.

Zoarium of erect, flattened, solid branches, in one plane. Zocecia

only opening on one face of the zoarium. Apertures tend to arrange-

ment in Y-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. Zocecia immersed.

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

figured.

Distribution. Turonian and Senonian.

FILISPARSA FRAGILIS, Marsson.

Plate I., fig. 2.

1887. Filisparsa fragilis, Marsson, Bry. Riig.; Pal. Abh., vol. iv.,

p. 36, pl. iii., fig. 8.

Zoarium of erect, cylindrical, or slightly flattened solid branches,

in one plane. Zoccia 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. Zocoecia immersed,

but the boundaries between them are more or less apparent as

ridges. One fragment out of seven is figured.

Distribution. Senonian.

Genus DIASTOPORA, Lamouroux.

Diastopora is here used, as in the British Museum Catalogue, for

Diastoporidee with an erect uni-, bi-, or even multi-laminar zoarium,

consisting of fronds or of very flattened branches.

Polyzoa and Anthozoa from Upper Cretaceous Limestone. 3

DIASTOPORA COMPRESSA (Goldfuss).

Plate I., fig. 3.

1827. Ceriopora conpressa, Goldfuss, Petref. Germ., p. 37, pl. xi.,

fig. 4.

1851. Ditaxia convpressa, von Hagenow, Bry. Maastr. kr., p. 50,

pleiv.; 1g5 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.

conupressa from Mesenterrpora Hyselyi, de Loriol (Urg. Land.: Mém.

Soc. hely. Sci. Nat., vol. xxiii., 1869, p. 40, pl. iii., 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 pl. 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 Oberfliche 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 zowcia

at that level. It is possible that in M. Hyselyi the direction of the

zocecia 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 Campicheana (Bry. Crét.,

1853-4, p. 800, pl. 784, figs. 6 and 8) also resembles Diastopora

compressa and Mesenteripora Hyselyt. 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. Crét. fig. par

d’Orbigny, 1890, pl. xi., fig. 7), showing it to be a form quite distinct

from D. convpressa and M. Hyselyt.

Gregory places as synonyms of Diastopora compressa the two

forms figured by d’Orbigny as Mesenteripora compressa and M.

neoconensis (Bry. Crét., 1853-4, pl. 756, figs. 10-13 and 7-9); but

both these forms have emergent zoccia.

Among the material described the only specimen referable to this

Species is a small fragment of the base of a frond. The earliest

zocecia, 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.

IDMONEIDA.

Genus IDMONEA, Lamouroux.

IDMONEA VIRGULA, d’Orbigny.

Plate I., fig. 4.

1851. Idmonea virgula, d’Orbigny, Bry. Crét., pl. 631, figs. 15-17.

1863. Leptotubigera simplex, de Loriol, Invert. Mt. Saléve, p. 127

pl. 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’Orbigny’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

Idmoneidie 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 Leticrisina, 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 Idmoneide,

in which the zoarium is erect, and consists of solid branches, which

are subeylindrical or moderately compressed; the reverse face is

well developed, but does not overlie an “axial rod” as in Retecava ;

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.

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

Polyzoa and Anthozoa from Upper Cretaceous Limestone. 5

CRISINA CENOMANA, d’Orbigny.

Plate I., fig. 5.

1851. Crisina cenomana, d’Orbigny, Bry. Crét., p. 732, pl. 614,

figs. 1-5.

1845. Idmonea disticha, Michelin, Icon. Zooph., p. 204, pl. lii.,

fig. 18.

1853. Idmonea calypso, d’Orbigny, Bry. Crét., p. 733, pl. 747,

figs. 10-14.

1890. Idmonea cenomana, Pergens, Revision des Bry. Crét. fig. par

d’Orbigny, p. 344, pl. xii., fig. 3.

Zoarium subcylindrical, and the reverse face is convex or flat;

there is no keel on the obverse face ; the zocecia 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 Retecava 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 f. Abbotti (Gabb and Horn), (Journ.

Acad. Nat. Sci. Philadelphia, ser. ii., vol. iv., 1860, p. 404, pl. Ixix.,

figs. 45-47), but in this form there is a ridge down the obverse face

of each zocecium.

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 zoccial boundaries are hardly

visible and the reverse face is convex. In the second (pl. xlv., fig. 4),

resembling d’Orbigny’s figure of Idmonea calypso, the zocecial 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 JI. calypso, Pergens says,

‘“‘quelquefois les branches sont aplaties a la partie inférieure et

forment ainsi une variété décrite comme J. 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, Pergens, Revision des Bry. Crét. 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. Idmonea excavata, d’Orbigny, Bry. Crét., p. 742, pl. 749,

figs. 11-15.

Zoarium subtriangular with a flat 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. Crét., 1853, p. 744, pl. 750, figs. 1-5), and Idmonea

lata, d’Orbigny (Bry. Crét., 1853, p. 734, pl. 748, figs. 6-10) resemble

this form, but have only three apertures in a lateral series. Jdmonea

marginata, d’Orbigny (Bry. Crét., 1853, p. 744, pl. 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. Crét., p. 744, pl. 749,

figs. 20-23.

1853. Idmonea angulosa, d’Orbigny, Bry. Crét., p. 735, pl. 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 C. excavata (d’Orbigny),

but has fewer apertures in a lateral series, and the zoccia are

immersed.

Distribution. Turonian and Senonian.

Genus TERVIA, Jullien.

Tdmoneide 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 7’. dorsata (von Hagenow).

TERVIA DORSATA (von Hagenow).

Plate I., fig. 8.

1851. Idmonea dorsata, von. Hagenow, Bry. Maastyr. kr., p. 31, pl. 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 ’’’; zocecia 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. korye. Hory.; Ceska. Ak. Fr.

Jos. Pratze, sect. 2, p. 18, pl. 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 zocecia

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 1:°5 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 + to throw up branches of the genus

Crisina. The general zocecial characters are those of Tervia decurrens

(Pocta), in which species it is provisionally placed.

Distribution. Cenomanian.

ENTALOPHORIDA.

Genus ENTALOPHORA, Lamouroux.

ENTALOPHORA VIRGULA (von Hagenow).

Plate I., fig. 10.

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

1840, p. 646.

1851. Pustulipora virgula, von Hagenow, Bry. Maastr. kreide.,

Delis aplel., gs a3.

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 zowcia

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

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

Polyzoa and Anthozoa from Upper Cretaceous Lumestone. 9

are immersed. Entalophora Pergensi, Gregory,* which the above

definition would also include, has thinner branches and the apertures

still further apart than in HL. virgula.

This is the commonest species of Cretaceous Entalophora and has

been found in most of the localities whence Cretaceous Polyzoa have

been recorded; but only one specimen has been found among the

material here described.

Distribution. Neocomian—Danian.

ENTALOPHORA CONJUGATA, von Reuss.

Plate I., fig. 11.

1872-3. Entalophora conjugata, von Reuss, Bry. unt. Quad. Palzon-

togr:, volo xx., part: 1., p. 117, pl. 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 zocecia are immersed and

their boundaries may be faintly marked. This form is very like

EH. geminata + (von Hagenow), but the zocecia are shorter, the boun-

daries of the zocecia are less clearly marked, and the apertures occur

in groups of three instead of two as in H. geminata. A single

specimen of this species occurs among the material here described.

Distribution. Cenomanian.

ENTALOPHORA ECHINATA (Romer).

Plate I., fig. 12.

1840. Pustulopora echinata, Romer, Verst. Nordd. kr., p. 22, pl. v.,

fig. 23.

1846. Pustulopora echinata, von. Reuss., Verst. Bohm kr., p. 64,

pl. 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 zocecia are immersed, with their boundaries more or less apparent.

This species is very nearly allied to H. 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, pl. x.,

NE By Oy Lc

+ Pustulipora geminata, von Hagenow, Bry. Maas. kr., 1851, pl. i., fig. 11, p. 20,

10 Annals of the South African Musewm.

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 MADREPORACEA (Goldfuss).

Plate I., fig. 14.

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

figs. 12a and 5b.

1851. Pustulipora madreporacea, von, Hagenow, Bry. Maastr. kr.,

Des pla, fe: ts.

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 Kast London material are doubtfully referred to this

species. All are very short fragments of stout branches, and, in so

far as the zocecial characters can be determined, resemble EH. madre-

poracea more closely than any other form.

Distribution. Cenomanian to Danian.

Genus SPIROPORA, Lamouroux.

Spiropora should probably be united with Hntalophora, for in

many species of the latter genus the apertures are arranged

spirally. Spiropora, however, may be defined as resembling

Entalophora, but the apertures are arranged in whorls, or, if in

spirals, these are very much looser than in Hntalophora.

SPIROPORA VERTICILLATA (Goldfuss).

Plate I., fig. 13.

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

plexi

1851. Cricopora verticillata, von Hagenow, Bry. Maastr. kr., p. 20,

plo ie pee a2:

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 zocecia are immersed, but the

Polyzoa and Anthozoa from Upper Cretaceous Limestone. ee

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 Caryophyllia, but the

species cannot be determined owing to their encrusted condition.

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2.—LHchinoidea, Brachiopoda, and Lamellibranchia from the Upper

Cretaceous Limestone of Need’s Camp, Buffalo River—By

Henry Woops, M.A., University Lecturer in Palaozoology,

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 Perna were found in the hard erystalline 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.

HCHINOIDEA.

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 interambulacra. 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. Hach 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.

Fia. 1.—Coptosoma capense, 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 Czdaris, and a portion of the test of an

irregular form which may perhaps belong to Casszdulus.

Echinoidea, Brachiopoda, and Lamellibranchia 1d

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.

Fic. 2.—Thecidea (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, F, 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 Thecidea vermicularis (Schlotheim),* but is not

so deep as in 7’. longirostre, Bosquet.t

* For figures and references see Geinitz, Das Elbthalgeb. in Sachsen (Paleonto-

graphica, vol. xx., pt. 1, 1872), p. 162, pl. xxxv., figs. 35-38. Bosquet, Brach.

Terr. Crét. Supér. de Limbourg (1859), p. 26, pl. iii., figs. 4-11. Posselt, Brach.

danske Kridtformat. (Danmarks geol. Underség., N" 6, 1894), p. 53, pl. iii.,

figs. 23-25.

+ Bosquet, op. cit., p. 33, pl. iv., figs. 4-6

16 Annals of the South African Museunt.

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.

LAMELLIBRANCHIA.

OsTREA, sp.

A small specimen of part of a left valve is probably an example

of O. vesicularis, Lamarck.* That species occurs in the Ariyalur

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.

Fic. 4.—Perna, sp. Left valve. In-

ternal cast of part near the umbo

Fic. 3.—Ostrea (Exogyra), sp. Right and of part of the ligamental area.

valve. Natural size. Natural size.

OstREA (ExoGyra), sp.

Text-figure 3.

There are two right valves which resemble O. (Hxogyra) Langloisi,

Coquand, from the Senonian (Santonian) of Algeria and Tunis.{

* Coquand, Monogr. du Genre Ostrea (1869), p. 35, pl. xiii., figs. 2-10. Stoliczka,

Cret. Fauna S. India, vol. iii. (1871), p. 465, pl. xlii., figs. 2-4, pl. xliii., fig. 1, pl.

xlv., figs. 7-12. Peron, Brachiopodes, etc., de la Tunisie (1893), p. 175. Wanner,

Paleontographica, vol. xxx., pt. 2 (1902), p. 119, pl. xvii., figs. 10-12.

} Peron, op. cit., p. 146, pl. 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.

PERNA, 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.

Fic. 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 Ariyalir Group (Senonian) of Southern India,

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

2

18 Annals of the South African Museum.

and also P. cereviciana, Petho,* from the Senonian of Cereviié

(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

Fic. 6.—Perna, sp. Internal cast. A considerable portion of the posterior part

of the shell appears to be missing. x 4.

the flattened part of the shell extending ventrally from the umbo is

similar in both species.

* Die Kreide- (Hypersenon-) Fauna d. Peterwardein. Gebirges (Paleonto-

eraphica, lii., 1906), p. 234, pl. xvii.

:

Echinoidea, Brachiopoda, and Lamellibranchia. 19

NOTE ON THE LOCALITY.

These fossils were obtained by Mr. J. W. Woods, of Hast London,

from Need’s Camp, on the Buffalo River, 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. RoGeErs.

* 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, R.E., to be 1,144 feet above sea-level.

bo

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13.

14.

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( 20)

EXPLANATION OR -PiA hd:

Potyzoa AND ECHINOIDEA FROM THE Upprr Creracrous oF NEED’s Camp,

Burrato River, Cape Conony.

. Filisparsa ramosa, d’Orbigny. x 12. Obverse face of part of a zoarium,

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

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

(p. 408).

. 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).

. Idmonea virgula, VOrbigny. x 12. Obverse face of zoarium (p. 410).

. Crisina cenomana, d’Orbigny. x 12. Obverse face of a specimen resembling

d’Orbigny’s figure of Idmonea calypso, showing strongly-marked zocecial

boundaries (p. 411).

. Crisina excavata (VOrbigny). 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).

. Crisina marginata (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).

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

(p. 413).

. Tervia gibbera, 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).

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

Entalophora conjugata (von Reuss). x12. Part ofazoarium. 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).

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

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

Entalophora madraporacea (Goldfuss). x12. Part of a worn zoarium

(p. 416).

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

Ann.S. Afr Mus. Vol. VII. Pas

Lee ae

HO)

T.A.Brock del. West, Newman collo.

Need’s Camp Fossils.

(21 )

3.—The Invertebrate Fauna and Paleontological Relations of the

Uitenhage Series.—By F. L. Kircntn, M.A., Ph.D., F.G.S.

CONTENTS.

PAGE

I. INTRODUCTION 3 oa a a Ses ans 21

II. THe AGE OF THE FAUNA.

(a) Summary of Previous Work ... oe as ae 25

(b) Additional Evidence for Age ... fe aes ade 30

III. Comparison witH ExtTrA-HuROPEAN FAUNAS,

(a) Possible Traces of a Related Fauna wm the South-

west of Madagascar <2 39

(b) Comparison with the Fauna ¥ the Oonn Chests 4h im

Cutch oe 3 39

(c) Traces of a Related ae im ‘the Gee Die

and in Hazara (N.W. Himalayas) $i Sea 46

(d) Neoconian in German Hast Africa ... Sa see 46

(e) Neocomian in South America... ose a af 49

IV. Tue DistRIBUTION OF THE UITENHAGE FAUNA IN RELA-

TION TO SOME THEORETICAL QUESTIONS ... bie’ ol

V. Descriptions, CriticAL Norss, AND Lists oF FossIts 60

VI.-Summary oF RESULTS _... ois ie st. Sa PH,

VII. List or WORKS CITED ... Bar ws ar “3, Doo

I.—INTRODUCTION.

THE marine invertebrate fossils collected in 1900 by Mr. A. W.

Rogers and Mr. E. H. L. Schwarz from the Uitenhage Series, and

forwarded to me by Mr. Rogers on behalf of the Geological Commis-

sion, were found at localities near Uitenhage and below this on the

Zwartkop’s River, and in the valley of the Sunday’s River near and

below Dunbrodie. Further materials have been sent from the South

3

22 Annals of the South African Musewm.

African Museum, principally comprising specimens presented by

R. Rubidge 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. Rogers and Schwarz,

some Shells of the genus Unio were collected in the Bezuidenhout’s

Valley below Blue Cliff.

In 1905 Mr. Rogers devoted some time to a further examination

of the Uitenhage beds in the valleys of the Sunday’s River and

Coega River, 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 Neumayr, 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. Rogers in 1905 well

bear this out, for they include a larger proportion of new species

than that shown by the collections made in 1900.+

In the following pages I have discussed the present state of our

knowledge regarding the paleontological 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. ]

+ Rogers and Schwarz (1).

} For an account of the survey of 1905, see Rogers (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 Reports 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 River 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

molluses 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. Rogers 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 River Beds

are partly replaced by conglomerate resembling that of the Enon

Beds, but here yielding Trigonia conocardiformis, 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} 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 which 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 Rivers. It appears highly probable, however, from the

paleontological evidence alone, that no very considerable extent of

time is represented by the whole 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. 58, 61; Rogers and Schwarz (1), p. 5; Rogers (1), pp. 282-

296. See also Rogers (2), pp. 13, 15.

+ Stow (1).

24 Annals of the 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. +

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 Huropean 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-Huropean 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. R. 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 my 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 Thammnastrea 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.

+ 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 River 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 7. dedalea 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.}

The examination of a small collection of lamellibranchs obtained

by F’. Krauss in 1839 from strata exposed on the Zwartkop’s River,

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 molluses, obtained by

R. Rubidge on the Sunday’s River, 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‘) referred

the Uitenhage fossils with doubt to the Lias, basing this opinion

upon the prevalence of a supposed Liassic form, ‘‘ Gryphea incurva.”’

The shell mistaken for this, however, was Hxogyra imbricata, pre-

viously described by Krauss, which, as we shall see, bears only a

superficial resemblance to the Gryphea mentioned, and is in reality

closely comparable with certain Lower Cretaceous forms of Hxogyra.

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 Rivers. The forms described,

principally Mullusea, led Sharpe to the conclusion that they most

nearly resembled Huropean species of the Middle and Lower

Oolites; he compared his Ammonites atherstont with A. macro-

cephalus and A. herveyi, while believing Ammonites baina to be

related to A. huwmphresianus and other Lower Oolitic forms.

* Hausmann (1), p. 1457.

+ Goldfuss (1), Band II., p. 202, pl. 137, fig. 5 (1837); p. 239, pl. 149, fig. 10

(1840).

t Krauss (1), pp. 129, 130. § Krauss (2).

|| Portlock (1). {| Bain (1).

26 Annals of the South African Museum.

In 1867, a paper by Ralph 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 Trigonie 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 palzontological types which are analogous to, or

identical with, those distributed in successive zones in Hurope.

In his monograph on the Cretaceous lamellibranchs of Southern

India, Stoliczkat made some reference to Uitenhage forms. He

evidently believed Tate’s Crassatella complicata to belong to the

genus Ptychonya, 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

Remondia 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 7’. twber-

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 Kemondia.

A. bron 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 Remondia.

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

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

{ 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

Trigonie, Lycett} referred briefly to the Trigonie 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 TJ. goldfusst, 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,} 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

(7. tater 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 atherstont

(Sharpe) ; Holcostephanus baini (Sharpe); Crioceras spinosissumum

(Hausm.) Neumayr; Trigonia ventricosa (Krauss); Trigonta cono-

carduformis (Krauss) ; Ptychomya complicata (Tate)§; and Hxogyra

imbricata Krauss. He suggested that Tate’s Ammonites subanceps,

which was thought by Tate to resemble the Jurassic A. anceps Rein.,

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. canaliculatus Schloth. and 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). + Lycett (3), p. 230 (1879).

{ Holub and Neumayr (1). § Misquoted ‘‘ implicata’’ by Neumayr.

|| Neumayr (4).

28 Annals of the South African Musewm

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 Holcostephanus,

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 | has since confirmed this view, after studying South

African specimens, and has placed Holcostephanus atherstoni and

H, baint in his generic or sub-generic division Asteria, which

comprises numerous Neocomian types; he even goes so far as to

identify Holcostephanus 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 haying 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 Trigonia-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).

+ Pavlow and Lamplugh (1), pp. 492-497 (134-139 in authors’ copy).

{| Neumayr and Ublig (1), p. 149, pl. xxxii., fig. 2.

§ Rogers and Schwarz (1), p. 17.

|| Rogers (1), pp. 281-318; Rogers (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,+ but Andrew Wyley,

in 1859, placed the Enon Beds so low as the New Red Sandstone,

and correlated the Sunday’s River 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.f’ In 1878

W. T. Blanford, basing his view upon a study of Tate’s and Stow’s

papers, concluded that the beds containing Hamutes 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. vauw were still

higher.***

In 1880 Griesbach classed the Uitenhage beds as Jurassic,}} while

the same view was adopted by T. Rupert Jones in 1884 t{ and by

Moulle in the following year.s§ Even so recently as 1897, Futterer

has accepted Sharpe’s and Tate’s conclusions.||||_ On the other hand,

Gurich,‘\’ Schenck,*** and Molengraaff +++ have referred these beds

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

} 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.

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

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

Moulle (1), p. 216. \||| Futterer (1), p. 625.

{11 Giirich (1). *** Schenck (1), p. 231.

ttt 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 R. B. Newton published a useful

summary of previous work on the Cretaceous conchology of South

Africa,t and furnished a complete list of the known Mollusca from

the Uitenhage beds, which he classed as Neocomian. In their text-

books de Lapparent? and Kayser § have followed Holub and Neu-

mayr in assigning a Neocomian age. Passarge, in his work on the

Kalahari,|| has tabulated the Uitenhage Series with the Upper

Jurassic and Lower Cretaceous. Lemoine believes it to be of

Lower Cretaceous age.

(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 Holcostephanus, 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-

stont (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 Nicklés 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 R. Tate t} gave a tabular list

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

{ de Lapparent (1), p. 1267. § Kayser (1), p. 444.

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

** Pavlow and Lamplugh (1), pp. 492, 496, (134 and 138 of authors’ copy).

tt Tate (1), p. 166.

The Invertebrate Fauna of the Uitenhage Series. 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 atherstoni and H. baini, which were

erroneously thought to be allied to well-known Oolitic forms, there

is Tate’s Ammonites swhanceps, a single specimen, regarding which

that author wrote: ‘‘ It is doubtful whether this be not a mere variety

of A. anceps Reinecke, 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. botelle Kilian. These were thought by Pavlow to have

been wrongly identified, and are united by him under the name

Hoplites heteroptychus, 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

Belemmites lateralis Phill. above the “ Rjasan-Horizont”’ in Russia

(Mostja River).* Hoplites subanceps 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 O. costata Sow.

from the Oolites; but it is in reality a true Hxogyra quite compar-

able with shells of the type of H. subplicata Roem.,} from the

Hilsconglomerat of North Germany.

Pecten 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. A. Roemer (1), Nachtrag, p. 25, pl. xviii., fig. 17; F. A. Roemer (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. jwrensis

Roem., may perhaps be an Anomia, and in some respects closely

resembles A. pseudoradiata d’Orb.,* from the Aptian. Placunopsis

embricata Tate is compared in the table with P. substriata Lyc.,

from the Oolites. P. semistriata (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. zmbricata, 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 Neumayyr.

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. Tvrigonia goldfussi Tate is an immature

shell, and in all probability illustrates the youthful stage of one of

the large and robustly ornamented Pseudo-quadratee which form

such a striking feature in the Uitenhage fauna. It certainly does

not represent either Trigonia goldfussi Agassiz or T. painer Lye.

(=T. goldfusst Morr. and Lyc.), 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 } repeatedly used the name “ 7’. goldfussi’’ in his

paper, but I have been unable to ascertain to which of the Trigonie

he referred.

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

* @Orbigny (4), p. 84; Woods (3), vol.i., p. 27, pl. v., figs. 1-3 (1899).

+ @Orbigny (3), p. 754, pl. 489, figs. 1-3 (1848).

{ Stow (1), pp. 497-514.

The Invertebrate Fauna of the Uitenhage Series. 33

Pinna atherstont Sharpe and P. sharpet 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.

Mytilus baint Sharpe was compared with M. sowerbianus 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. sowerbzana (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.

Pholadomya domimcalis 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 pinchiniana 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 Hriphyla and is certainly most closely com-

parable with Cretaceous forms.

Berenicea 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 Serpule 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. quwinquangularis Goldf. t

(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 Renevier (1), p. 60, pl. vi., fig. 7, 1855 ; Moesch (1), p. 93, pl. xxxiii.,

fig. 2. t Gregory (1), p. 112.

} Goldfuss (1), Band i., p. 230, pl. 68, fig. 8 (1831).

34 Annals of the South African Museum.

it is not distinguishable from the English Lower Cretaceous Serpule

that have been variously referred to Serpula filifornuis J. de C. Sow,

and S. plexus J. de C. Sow. We thus see that in the case of the

two Trigonie and the two Serpuleé 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 T’rigonia, 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. krausst, and 1’. rogers, all possessing the characters of the section

Scabree—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 7. tate, a typical member of the

section Costate, but although this division of the Trigonieé has its

maximum development in the Oolitic rocks, it is also represented in

strata of Lower Cretaceous and even later age.

Trigoma pemnsularis Coquand,* from the Aptian of Spain, cer-

tainly shows characters of degeneration not shared by T. tatez, but

the Oomia strata of Cutch have yielded two normal representatives

of the Costatze, which, together with some degenerate derivatives of

this section, are accompanied by Trigoni@ which exhibit a decidedly

Cretaceous aspect. Another unmodified costate form, 7. anguste-

costata Behr. + occurs in the Argentine Republic 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 herzogs (Goldf.), another very characteristic Uitenhage

form, was believed by Lycett to belong to the Quadrate, which, so

far as known, are exclusively Cretaceous. Steinmann,§$ however,

pointed out that Trigonia herzogi differs in several important par-

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

+ Behrendsen (1), p. 6, pl. 3, fig. 7.

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

§ Steinmann (2).

The Invertebrate Fauna of the Uitenhage Series. 30

ticulars from all known Quadratz, and found that the special features

which distinguish this Uitenhage shell from the Jurassic Clavellate

on the one hand and the Cretaceous Quadrate 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 Trigonie, which were the only examples known to exhibit these

peculiar distinguishing characters. Trigoima newquensis Burckhardt, *

which occurs with J. transitoria in Neocomian rocks at Las Lajas

on the Rio Agrio (Argentine), falls under the same heading, while

T. mamillata + from the Oomia strata in Cutch must also be referred

to this group. Still another form, described in these pages (7’. holubz),

is clearly referable to the same category. Although the five members

of the Pseudo-quadrate known to us are all similarly differentiated

from the typical Clavellatz, 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 Quadrate,

and though definite indications on this point fail us, it is quite

probable that the Quadratz and Pseudo-quadratze represent parallel

and independent groups of derivatives arising from Jurassic Clavel-

late. It is therefore evident that the Trigoni@ of the group Pseudo-

quadratz cannot yet be considered in themselves to furnish quite

such reliable guidance to geological age as the true Quadratz, but at

the same time their general aspect, by comparison with the Quadrate,

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 Quadratz differ from the

Clavellatze are also exemplified in the Pseudo-quadrate. Lycett

laid great emphasis on the presence or absence of sculpture on the

escutcheon in distinguishing between the Jurassic Clavellatz on the

one hand, and the Cretaceous Clavellate and Quadratz on the other,

and in regard to this feature all the Pseudo-quadratz agree with the

Quadrate 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 conocarduformis (Krauss) was thought by Lycett to fall

* Burckhardt (2), p. 74, Taf. xiv., figs. 4-6.

¢ Kitchin (1), p. 100, pl. ix., figs. 8, 9; pl. x., figs. 1-3.

{ Remarks on this subject will be found to follow the description and the dis-

cussion of the relationships of Trigonia holubi, in these pages.

36 Annals of the South African Museum.

within the section Scabrz, 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 Clavellatz

in a manner somewhat analogous to that which distinguishes the

Scabree, 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 Trigonia-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 Trigonie@ 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 widely 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. Solecwrtws 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 Pecten

which can only be identified with the Cretaceous P. orbicularis

J. Sow. and P. cottaldinus d’Orb., while a third form may be most

aptly brought into comparison with P. swbacutus 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. deecks

Frech; G. aviculoides Sow.), has a close counterpart in G‘. anceps

Desh. and G. swblanceolata d’Orb., in the Lower Cretaceous of Europe.

A form closely similar, and perhaps identical, occurs also in the

Neocomian of German Hast 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

* Frech (1). + Frech (1), pp. 612, 613.

The Invertebrate Fauna of the Uitenhage Series. 37

and characteristic Uitenhage shell is Hxogyra 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. Reference 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. wilmane 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 paleontological 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 River) associated with plant-bearing beds, and

are in the lower part of the Uitenhage Series ; according to Messrs.

Rogers and Schwarz they may be classed with the so-called ‘‘ Wood

Bed” series. Amongst the fossils collected here are Act@onina

atherstom (Sharpe), Cyprina rugulosa Sharpe, and a Pecten which

I have identified as Pecten cottaldinus dOrb. While Acteonina

atherstont is found also in the highest part of the Marine Beds

on the Zwartkop’s River and with the characteristic Trigoni@ on the

Sunday’s River, Stow recorded the occurrence of Cyprina rugulosa

with similar associates. Although the Wood Bed series was not

seen in the Zwartkop’s River Valley below Uitenhage, the lowest

part of the Marine Beds in the Zwartkop’s River section, exposed

in a clay-pit near Rawson Bridge, yielded Acteonina atherstone and

4

38 Annals of the South African Musewm.

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 Rawson

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 River Valley, yielded no single form which suggests

stronger affinities to Upper Jurassic than to Lower Cretaceous

types; but those very forms, such as Tvrigonia tatei Neum. or

Tancredia schwarzi sp. noy., 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 River district in 1905, Mr. Rogers

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

Trigome) were observed to occur, together with layers of lignite,

at the very base of the Sunday’s River Beds, where Mr. Rogers 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.

Rogers 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. Rogers believes that from the same cause his own

records are likely to prove faulty, and that exhaustive collecting

would 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 paleontological stage.

Allowing, then, that the cephalopods, supported in no small degree

* For the published account of the 1905 survey, consult Rogers (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 (= Astierra 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-Huropean regions which, in greater or

less degree, bear the imprint of a similar facies.

III.—COMPARISON WITH EXTRA-EUROPEAN FAUNAS.

(a) Possible Traces of a Related Fauna im the South-west of

Madagascar.—Douvillé 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.t Mention is made of a large

Trigonia, said to be analogous to some of the Trigonie of the Oomia

beds in Cutch and of the Uitenhage Series, and with this is asso-

ciated a shell referred by Douvillé to the genus Pycnodonta, and said

by him to be closely comparable with Hxogyra imbricata Krauss

(also referred to Pycnodonta by Prof. Douvillé). On the evidence of

these fossils, the strata containing them are considered by Douvillé

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.

(b) 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.

+ Douvillé (2), p. 8388; Douvillé (4), p. 215; Lemoine (1), p. 176.

40 Annals of the South African Musewm.

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.

Stoliczka+ first directed attention to the great similarity between

a Trigonia collected by Wynne during the geological survey of

Cutch,* and the South African 7. ventricosa (Krauss), and the identity

of these was afterwards confirmed when the Oomia shell was definitely

referred to J. ventricosa by Waagen,$ Feistmantel,|| and W. T.

Blanford.‘/ 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, 7. mamuillata is comparable

with 7’. herzogi and the South American 7’. transitoria in the manner

in which it exhibits characters somewhat intermediate between those

of the Clavellatea and the Quadrate, but it bears a still closer

resemblance to 7. holubi sp. nov. Owing to their large and massive

shells, and their well-characterised sculptural plan, the members of

this group of Trigonie 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 Costatz with members of the Scabra, both in the Oomia

and the Uitenhage strata—a fact to which I have already referred.

T. parva accompanies 7’. ventricosa in Cutch, and although T” tens,

another costate form, has not been recorded from any of the

localities where 7’. ventricosa is found, it occurs in beds at least not

older than those from which the remaining Oomia Trigoni@ have

been obtained. In the Uitenhage Series we have 7’. tatez, which,

although not closely comparable with T. tenwis, 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). + Stoliczka (2), p, 315 (1871).

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

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

“| 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

Costate 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 Trigonie 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

Oomia beds and the group of JZ. vaw in the Uitenhage Formation.

The similarity of one of the Oomia T'rigonie to T. vaw Sharpe seems

to have been first pointed out by Feistmantel,* and W. T. Blanford t

shortly afterwards suggested that a shell from the Oomia Group,

which was ascribed by Feistmantel to the genus Goniomya, might in

reality represent the African 7. vau. Which of three Oomia forms

was here and elsewhere} referred to in this connection is uncertain,

but it is possibly the one which I have described under the name 7’.

v-scripta,§ although the less well-known 7’. dubia in reality bears a

much closer resemblance to JT. vauw, particularly in the outline.

When dealing with the Oomia Trigonie I pointed out that 7’.

v-scripta, when adult, more closely approaches an undescribed

Uitenhage form which is associated with 7. vaw. This is T. stowz,

described in these pages, and it is certainly closely allied to 7’. vaw ;

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 7’. v-scripta are

also practically the same as in 7’. recwrva, 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 Trigoni@, to distinguish

between the ‘‘ Group of 7. vaw’’ and the ‘‘ Group of 7’. 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. + W. T. Blanford (1), p. 118.

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

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

42 Annals of the South African Musewm.

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 homao-

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

Trigonié, 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 carinz 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 umbones relatively far from

the anterior extremity. In the Oomia beds these points are illus-

trated in varying degree in certain degenerate derivatives of

Costate, and in the group of TY. v-scripta; in the Uitenhage beds

they are exemplified in 7’. vau, T. stowi, T. rogersi, and T’. conocardic-

formis. In both Trigonia vauw 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 7’. stowz 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

The Invertebrate Fauna of the Uitenhage 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 Hxogyra occurring in Oomia beds at the Trummo

River is certainly not distinguishable from individuals of H. 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 herzogi

Krauss, though it is not identical with it; there are fragments of a

large, coarsely-ribbed Cucwllea which, so far as can be seen, shares

all the distinguishing features of the strongly characterised Cucullea

krausst 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 Seebachia 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 :—

Uitenhage Beds. Oomia Beds.

Exogyra imbricata Hxogyra inbricata

Cucullea kraussi Cucullea kraussi

Gervillia dentata Gervillia dentata ?

Astarte herzogi Astarte sp., near herzogi

Trigonia ventricosa Trigonia ventricosa

Trigonia holubt

Trigonia herzogi | (Pseudo-quadrate) Trigonia mamillata

Trigome of the vau Trigome 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.

ike 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-

branehs 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 T'rigonie to corroborate

fully his view concerning the Portlandian age.* But a detailed

study of the Trigonie has shown that this belief was not well founded,

and the relationship of any of the Oomia forms to the Portlandian

Gibbose is at the best a matter for conjecture. I have provisionally

referred one of these Trigoni@ to the group Gibbose 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 7’. spisszcostata to the Gibbose 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 Trigone

which exhibit characters simulating those of the Portlandian Gibbose

have been shown to be allied to the section Costatze, and nothing

quite comparable with these diversely modified derivatives is known

in Jurassic rocks, though the late adult stage of 7. peninsularis Cogq.,

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 Fawna of the Urtenhage 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. pulchra, both members of the

section Scabre. Though claiming corroboration of his views from

Trigonie 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 we 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 Paleontological Collection of the State at Munich shows that

the Indian form is rather thicker and more inyolute, 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 the South African Musewm.

nothing in the fauna of the Oomia 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

Neocomian in Europe.

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

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 Tvrigonia smeei, a characteristic fossil of the

Oomia Group in Cutch, though it seems possible that this may be

7’. crassa, an Oomia form of similar type.t The small collection

of fossils from near Coconada was examined and named by Stoliczka,

and it comprises Znoceramus 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) Neoconuan in German East Africaa—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 Trigonieg. 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 beyschlagi G. Miller, which here

accompanies 7’. ventricosa in great abundance, appears to have no

counterpart in the Uitenhage beds, but it clearly belongs to the same

category of modified Costate as 7’. smeez|| and ZT. crassa‘ from the

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

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

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

§ G. Miiller (1). || J. de C. Sowerby (8), pl. lxi., fig. 5.

“| Kitchin (1), p. 44; pl. iv., figs. 4—6, pl. v., figs. 1-3.

The Invertebrate Fauna of the Uitenhage Series. 47

Oomia beds in Cutch; the resemblance to T. crassa is indeed very

striking, though I have elsewhere stated reasons why these two

forms cannot be considered identical.* But 7. beyschlagi is certainly

either closely related to 7’. 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

smeei 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 beyschlagit with Neumayr’s excellent correspond-

ing illustration of Seebachia.§ A glance at these figures proves

beyond question that Miller was right in assigning his shell to the

genus T'rigonia, 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,

30 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 (Ptychonya hauche-

cornet Mill.) which, though distinct from P. convplicata (Tate), may

possibly be nearly related to it. TZrigonia bornhardti Miller, from

the same locality, is only so far comparable with the Uitenhage-

Oomia Trigonie, 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 T’rigonia-beds compares very closely with the shells

from Ntandi described by Miiller under the name Hriphyla stuht-

mannt, 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. + Ublig (8).

+ G. Miller (1), pl. xix., fig. 3.

§ Holub and Neumaypyr (1), pl. ii., fig. 40.

48 Annals of the South African Musewm.

Still greater interest attaches to a Trigonia which strongly recalls

the peculiar types belonging to the group of 7. v-scripta in the

Oomia Trigonia beds and the Uitenhage group of 7. vaw. This

is T. kiihni Mill., 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. Miller has also brought it into comparison. Trigonia kiihna, ib

is true, seems to be distinguished from 7. vaw 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 7’. 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 J. v-scripta or to T. vaw and

T. stowt 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 Roem., which was found with Gervillia dentata, and Arca

witenhagensis Miller, which accompanies Trigonia beyschlagz. They

may be brought into near comparison with Pecten projectus Tate and

Arca 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 described 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 Hast

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 Trigonia, 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 Series. 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 Zrigoni@ from the Belgrano beds, T. sub-

ventricosa Stanton, a member of the section Scabre, and T’.. hetero-

sculpta Stanton, believed by Stanton to be allied to 7. vaw Sharpe.

Trigonta subventricosa certainly approaches very closely to 7’. ven-

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 7’. kraussi sp. nov., which occurs with 7’. 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 Scabre, 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 TZ. scabricola Lycett and 7’. alzformis

Park. are typical members in Europe.

The ornamentation of 7. heterosculpta is of such a kind as to

recall instantly the conspicuous shells of the groups of 7. v-scripta

and 7. vau. The principal points of resemblance are thé 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. 7’. heterosculpta

is certainly not to be brought into comparison with the European

Undulatz, but its whole aspect suggests a close connection with the

group of 7. vau. Its near relationship to 7. vaw and 7’. stow? 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 7. v-scripta is exactly what we might expect to

find, having regard to the geographical relationships. The adult

* Hatcher (1). + 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 Oomia 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 Hxogyra which

has much in common with Exogyra imbricata Krauss; and to judge

from the description of O. 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 molluscan 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 + 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) ; }

these beds yield another member of the Pseudo-quadrate (7’.

neuquensis Burckhardt) and a Trigonia very closely resembling the

South African 7’. conocardiiformis,§ as well as an Hriphyla in some

respects comparable with the South African Astarte (Hryphyla)

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

p. 297. + 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 “ T'rigonia cf. aliformis Park.” from supposed Upper

Neocomian beds at the Arroyo Pequenco may possibly be identical

with the Patagonian 7. suwbventricosa. It is accompanied by an

oyster which Behrendsen identified with Hxogyra couloni, and found

to agree well with the figures of Ostrea couloni cited by Bayle and

Coquand+ from the Neocomian at Arqueros in Chili. The Chilian

shells ascribed to O. cowloni bear a considerable resemblance to the

Uitenhage Hxogyra imbricata Krauss, with which Coquand even later

identified them,} and accompanying this oyster in Chili is Trigoma

delafosset Bayle and Coquand, the resemblance of which to 7’. ven-

tricosa has been elsewhere remarked upon.$ Associated with these

forms is a Crioceras identified by Bayle and Coquand with the

European C. duvali Lév., thus recalling the analogous association

of Trigonia ventricosa, Crioceras spinosissimum and Exogyra imbri-

cata in South Africa. Of the above-mentioned Mollusca, Trigonia

delafosset, T. transitoria, T. newquensis, and T. cf. conocardiformis

suggest very strongly their relationship to Uitenhage and Oomia

forms. It should be noted also that Philippi has described Trigonie

from localities in Chili, which share the characters of peculiar

sculpture and siphonal elongation shown by the group of T. vaw

Sharpe, while 7. eximua R. A. Philippi, from the Tinguirica valley

in Chili, appears to be closely related to 7. conocardiiformis (Krauss)

and Burckhardt’s 7. ef. conocardiformis.

IV.—THE DISTRIBUTION OF THE UITENHAGE FAUNA

IN RELATION TO SOME THEORETICAL QUESTIONS.

We may now briefly consider in what measure the correlation

of the Uitenhage Neocomian fauna with the similar assemblages in

German Hast 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 Neumayvr’s theory of distvri-

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

paleontological character, it may be of advantage briefly to review

the evidence now available. Moreover, it is advisable to glance at

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

+ 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 paleeogeographic

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 paleontological 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 Range as possible indications of an

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

+ Stow (1), p. 546; H. F. Blanford (1), pp. 534-540; Medlicott and Blanford

(1) pp xxine Icy lixcxit 297,

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

The Invertebrate Fawna of the Uitenhage Series. 53

encroachment of the southern development into the northern

region.* With reference to the same subject, Mr. R. 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.”’ +

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.§ From 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 possibile reference to its state of completeness during

any part of the Cretaceous period. Dr. F. Kossmat ‘i 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.

t Newton (1), p. 333. § Neumayr (5).

|| W. 'T. Blanford (4). “| 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; + 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} 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.

+ Since these sentences were written (in 1903), Lemoine has expressed the same

opinion ; see Lemoine (1), p. 391.

+ Pavlow and Lamplugh (1), p. 493 (p. 135 of authors’ separate copy).

§ Uhlig (4), p. 132.

The Invertebrate Fauna of the Uitenhage Series. 5)

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 paleontological 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 East 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. Miiller’s suggestion + to the effect that

Stow’s record of Uitenhage lamellibranchs from the Zambesi,{

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. + G. Miiller (1), p. 571.

t 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 River Isakondry, east of Tullear.* Professor Douvillé

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. +

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. Douvillé 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 Trigonie 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. Douvillé 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 River

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 réle 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. + Douvillé (3), p. 435.

t Douvillé (2), p. 388; Douvillé (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 palzontological and

stratigraphical point of view, between the deposits of India and

of the south of Africa (fauna with dicynodont reptiles; Glossopteres

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.{ 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; Douvillé (4), p. 215; Boule (5);

Boule and Thevenin (1), p. 59; Lemoine (1), p. 232.

+ Haug (1), p. 397. t Haug (1), p. 398.

§ Boule and Thevenin (1), p. 59. See also Woods (4), p. 348.

|| Kilian (2); Kilian (3).

‘| Choffat (1); Choffat (2); Choffat (3); Lemoine (1), p. 396.

58 Annals of the South African Musewm.

although the known evidence seems to be on the whole destructive

of Neumayvr’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 Huropean’’ 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. + 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 Astieria) 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. Boehm (2); Tornquist (1), p. 285; Ortmann (1); Stanton (2)

Burckhardt (3), p. 179. + Burckhardt (3), p. 183-185.

60 Annals of the South African Musewm.

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 OF

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 Holcostephanus bain, H. modderensis, and

H. wilmane, 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 Series. 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’’ 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.*

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.’’ JI am 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.

+ Rogers and Schwarz (1); also Rogers (1), pp. 281-292.

and ‘ neanic,’

62 Annals of the South African Musewm.

In the case of the specimens from the collection of the South African

Museum labelled ‘‘Sunday’s River,’’ no more detailed account of

locality is available, but these are all typical Mollusca of the Marine

Beds or Sunday’s River Beds. Of all the fossils examined, the only

forms which indicate fresh-water conditions are the Unio from the

Wood Beds of the Bezuidenhouts River, below Blue Cliff station, and

a single specimen of Linnea from the section at Buck Kraal, Sunday’s

River. 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 paleontological

grounds to suppose that this small marine assemblage, characterised

by the prevalence of Gasteropoda, is materially older than the fauna

yielded by the Trigonia-bearing Marine Beds.

The collection obtained by Mr. Rogers from the Marine Beds of

the Coega River and Sunday’s River 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. Rogers should be consulted.* This work adds

materially to our knowledge of the Sunday’s River 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. Rogers, 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. Rogers in a patch of marine

Uitenhage beds at Knysna Estuary + only represent the product of

a hasty search, and it is believed that careful collecting at this new

locality may yield good results.

Crass ANTHOZOA.

Genus THAMNASTRAA } Lesauvage.

THAMNASTRA Sp.

There are several specimens of a T’hamnastrea from the locality

between milestones 244 and 242 on the railway between Uitenhage

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

{ 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-Reinet (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 Thammnastrea.

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.

Crass ANNELIDA.

GENus SERPULA Linnezus.

Serpuna cf. concava (J. Sowerby).

Plate IT., 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. 348, pl. xviii., fig. 10.

», 1884. Vermicularia concava R. 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 ................4. 9 mm.

Greatest external diameter of the tube at the close

Ole THERCOU EM STACK 26.2 ces nee 0s sc cedeumienwsieselesone Sis

Occurrence.—A single specimen was found between milestones

64 Annals of the South African Museum.

245-243 on the railway between Uitenhage and Graaff-Reinet (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. damesi Noetling, + 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) pinchinana R. Tate, Quart. Journ. Geol.

Soc., vol. xxiii., p. 163, pl. ix., figs. 10a—c.

The large cylindrical tubes of this form are sometimes seen

attached to massive lamellibranch-shells such as Hxogyra imbricata

Krauss, or one of the pseudo-quadrate Trigoni@. 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), pl. 6, fig. 5.

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

The Invertebrate Fauna of the Urtenhage Series. 65

Crass LAMELLIBRANCHIATA.

Genus PECTEN O. F. Miiller.*

Sus-Genus SYNCYCLONEMA F. B. Meek.

PrEcTEN (SYNCYCLONEMA) ORBICULARIS J. Sowerby.

Plate II., figs. 2, 3.

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

Britain, vol. i1., p. 193, tab. 186.

1902. Pecten (Syncyclonema) orbicularis H. Woods, Mon. Cret.

Lamell. Engl., vol. i., part iv., p. 145, pl. xxvii., and

text-fig. 1. (Palezeontographical 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. Oneof 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 pl. 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. Rogers in the Coega River Valley. Length 13 mm.;

height 15 mm.

Locality—Sunday’s River (279, 280). Coega Valley, on the left

side of the valley, half a mile down from the railway (453g).

Sus-ceEnus CAMPTONECTES F. B. Meek.

PECTEN (CAMPTONECTES) COTTALDINUS d’Orbigny.

Plate IT., fig. 4.

1847. Pecten cottaldinus A. d’Orbigny, Paléont. Frang., Terr. Crét.

vol. iii., p. 590, pl. 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, pl. xxix., figs. 1-3.

(Palzontographical Society.)

A specimen collected by Messrs. Rogers 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

warkings 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 River (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. Miiller from the Neocomian of German East Africa.

PECTEN (CAMPTONECTES) PROJECTUS Tate.

Plate II., figs. 5, 5a.

1867. Pecten projectus R. Tate, Quart. Journ. Geol. Soc., vol. xxiii.,

p. 165; pl. ax. 128.16.

Supplementary 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 striz, 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 Uitenhage 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 hydrothecz 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 strie 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 ; i) : PAN)

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

Occurrence.—Railway cutting between milestones 241-243 on the

Uitenhage to Graaff-Reinet railway (297, 350); also obtained by Miss

M. Wilman at Coega River. Tate says: ‘Collected by Dr. Rubidge

in a soft, yellowish-grey, sandy limestone at the Zwartkop River

Heights, and at Prince Alfred’s Rest; in both localities it is

frequent.”

Remarks.—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 stri,

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 Musewm.

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 strize

with a punctate structure, but no encroachment of this sculpture into

the interspaces between the linear stria. Pecten suprajurensis

Buvignier,| an Upper Jurassic form, approaches much more closely

to P. projectus 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 buchi Roem.,} 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 strie.

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 stria, 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 Reuss, yet it appears from the descriptions

that punctate linear strize with smooth interspaces characterise these

forms.

Nilsson’s P. virgatus, of Upper Chalk age from Sweden,t+t+ is

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

+ Buvignier (1), p. 24, pl. xix., figs. 21-23.

¢ F. A. Roemer (1), pl. xiii., fig. 8 (1836), and Nachtrag, p. 27; de Loriol, Royer

and Tombeck (1), p. 389, pl. xxii., figs. 12, 13.

§ F. A. Roemer (1), Nachtrag, p. 27; d’Orbigny (3), p. 592, pl. 482, figs. 4-7

(1847); Woods (3), vol. i., p. 157, pl. xxix., figs. 4-6 (1902).

|| See also figures given by de Loriol under the name P. arzierensis ; Loriol (3),

pl. iv., figs. 3-5. ‘| Geinitz (1), p. 16, Taf. iii., fig. 13.

** Geinitz (2), Theili., p. 193, Taf. 43, fig. 15 (1872); Theil ii., Taf. 10, fig. 1,

(1872) ; Reuss (1), part ii., p. 28, Taf. 39, fig. 6 (as P. divaricatus).

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

The Invertebrate Fauna of the Uitenhage Series. 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 striz 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 Neocomian 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) pueyrrydonensis 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 kamerunensis 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.

Sus-cEnus CHLAMYS J. F. Bolten.

PrECTEN (CuHuLAmys) cf. susacutus 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, pl. 2, figs. 28, 33. + Zittel (2), p. 109, pl. xvii., fig. 8.

t Whiteaves (3), p. 242, pl. 32, fig. 4. § G. Miller (1), p. 550, Taf. 24, fig. 7,

|| Stanton (3), p. 12, pl. iv., fig. 1.

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

6

70 Annals of the South African Musewm.

Cf. 1847. Pecten subacutus A. d’Orbigny, Paléont. Frang., Terr. Crét.,

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

», 1872. Pecten subacutus H. B. Geinitz, Das Elbthalgeb. in

Sachsen, part i., p. 195, pl. xliv., fig. 5 (Paleeonto-

graphica, vol. xx.).

,», 1902. Pecten (Chlamys) subacutus H. Woods, Mon. Cret. Lamell.

Bmegl.; vol. is part) iv., p. 169s pk xxxi., fess 7-9:

(Paleeontographical Society.)

Descriptive Note.—A single specimen sent from the collection of

the South African Museum seems to approach very closely to P. sub-

acutus. 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 2mm. 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 85 mm.

Occurrence.— Marine Beds of Sunday’s River (304).

Remarks.—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 urgonensis de Loriol,* common in the Urgonian of Grand

Saléve, 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, pl. C, figs. 25, 26.

The Invertebrate Fauna of the Uitenhage Series. 71

Genus LIMA J.G. Bruguiére.

Sus-cenus ACESTA H. and A. Adams.

Lima (ACESTA) OBLIQUISSIMA Tate.

Plate IT., fig. 7.

1867. Lima obliquissima R. Tate, Quart. Journ. Geol. Soc., vol. xxiii.,

p: 196, pl. ix., fig: 3.

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 mm. from the umbo, they may be separated by interspaces

2mm. 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

(aoa ERs] emma Aacaoh cath atom ts ORG Aer ay erase eae ae 40 ,,

Greatest depth of a single valve ..................00. Ome

Occurrence.—Railway cutting between milestones 244-243 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 Placuwnopsis 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. LL. 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 Acesta, *

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.s

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, pl. vii., fig. 13.”

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

pl. iii., fig. 5.

{| H.and A. Adams (1), p. 558. § E. Philippi (1), p. 680.

The Invertebrate Fauna of the Uitenhage Series. 73

sculpture of the flank, it is true, recalls that of some Plagiostome,

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,} and latterly

by Philippi, while regarded by Fischer as only representing a

section of Lima.t This last author merely quotes the living type

species, L. excavata. 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.,|; 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 wndata (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. + Zittel (5), p. 26 (1881).

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

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

“| Leymerie (2), p. 10, pl. 8, fig. 8; d’Orbigny (3), p. 528, pl. 414, figs. 9-12

(1847). ** Matheron (1), p. 182, pl. 29, figs. 3, 4

74 Annals of the South African Musewm.

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 Roemer’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 ; Z. longa is further distinguished by the possession of

great height measurement in proportion to its length.

There is closer agreement, again, between ZL. obliquissima and

L. tenuwitesta Whitfield, | from Upper Cretaceous rocks in Syria.

The two are practically identical in outline and the relative length

of the superior border, but L. tenwitesta has the linear striz of the

surface more numerous and closely crowded at a given distance

from the umbonal apex.

Sus-cENus MANTELLUM J. F. Bolten.

Lima (MANTELLUM) NEGLECTA Tate.

Plate II., figs. 8, 8a.

1867. Lima neglecta R. Tate, Quart. Journ. Geol. Soc., vol. xxiii,

p. 156, pl. vii., figs. 4a, 40.

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, pl. v., fig. 6.

+ F. A. Roemer (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).

+ 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 LD. (Mantellwm)

neglecta and L. parallela J. de C. Sow.;* but in Lima neglecta

the oblique elongation is greater and the posterior ribs are more

crowded and delicate. Lima (Mantellum) gaultina Woods + is more

closely similar to Z. 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 PERNA J. G. Bruguiére.

PERNA ATHERSTONI Sharpe.

Plate II., fig. 9.

1856. Perna atherstont D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

vol. vil., p. 193, pl. xxii., figs. 4, 4a.

Occurrence.—This shell was found at Dunbrodie (Sunday’s River),

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.”

fiemarks.—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, pl. v., figs. 14, 15 (1904).

t Woods (8), vol. ii., part 1, p. 31, pl. v., figs. 16-20 (1904).

76 Annals of the South African Museum.

variation. The convex fold of the valve, passing back from the

umbo, 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 arounded 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 umbo, is less inflated

anteriorly, and is more obliquely elongated. There are further

points of difference which it is unnecessary to recount.

Genus PINNA Linneus.

PINNA ATHERSTONI Sharpe.

1856. Pinna atherstoni D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

WOl. vile) ps 193; plixxu shes Wy

Occurrence.—Railway cutting between milestones 244-243 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 notedby Messrs. Rogers and Schwarz that

this shell, associated with Holcostephanus atherstont, 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. hartmanim, as figured by Goldfuss,} 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), pl. ii., fig. 5.

+ Goldfuss (1), Band ii., p. 164, Tab. exxvii., fig. 3b (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 OSTREA Linneus.

OSTREA sp.

Specimens of Ostrea (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-

chena 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 EXOGYRA T. Say.

EXoGYRA IMBRICATA Krauss.

1843. Hxogyra imbricata F. Krauss, Amtlicher Bericht tiber die

zwanzigste Versammlung der Gesellsch. deutsch.

Naturforscher und Aerzte zu Mainz im Sept., 1842,

pe L298;

1850. Exogyra imbricata F. Krauss, Noy. Act. Acad. Cees. Leop.-

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

2a-2d.

1856. Gryphea imbricata D. Sharpe, Trans. Geol. Soc. Lond.,

Sera) vol. vil, p. 197, pl. xxiii, figs: 3a, 3d.

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

78 Annals of the South African Musewm.

1867. Ostrea imbricata R. Tate, Quart. Journ. Geol. Soe., vol. xxiil.,

p. 164.

1904. Pycnodonta imbricata H. Douvillé, Bull. Soc. Géol. 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 River

valleys. It was met with at almost all the exposures in the

Zwartkop’s valley visited by Messrs. Rogers and Schwarz. HEx-

amples sent to me are from the railway cutting between milestones

244-243 on the railway from Uitenhage to Graaff-Reinet (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 River (300, 301, 302, 303).

Riemarks.—A striking character of the late adult and senile

condition of H. 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 lamin, 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

Ostreidz, 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 Hxogyra and Gryphea. 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 Hxogyra and Gryphea.” 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 Ostreide. +

He believes that Gryphea and Exogyra ‘are probably aberrant

members of the Ostreide, not typical forms in the line of evolution

of the group.”

* Leymerie (la). See also Peron’s remarks on these genera, Peron (1),

pp. 107-9. + Jackson (1), sections v. and vii.

The Invertebrate Fauna of the Urtenhage Series. 7

It is highly probable that the shells classed as Gryphea do not

represent a homogenetic group, but are polyphyletic in origin, in-

cluding repeated offshoots from an ostrean stock, While, in the

progress from Ostrea to Gryphea, there is diminished duration of

the attached habit, so that the typical Gryphea 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 Gryphea 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

Gryphea 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 ostreiform

aspect. Haxogyra 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

Gryphea 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 Cucullea 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 Hxogyra, and present the strongest contrast to

* Compare also figure of Gryphea alligata from the Corallian of NELUSSHE

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 Gryphea 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 H. imbricata often simulates that of

a true Gryphea 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 Hxogyra 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 EH. subsinwata. Leymerie first regarded all

these as varieties of Hxogyra sinuata,t and included forms to which

numerous specific names had previously been applied. He subse-

quently separated H. swbsiiwata as a species distinct from H. sinuata,

and used several varietal designations of both. Regarding the

value of the nomenclature employed by this author or by Coquand}

in dealing with the forms comprised within this perplexing group,

Iam 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 Hxogyra coulont 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 LH. imbricata

and believed it to be nearly related to ‘“ Gryphea couloni Defr.,”’

from which, however, he found it to differ in the following points:

“Tt is larger, more thick-shelled, longer and narrower; has a keel

narrower at the end (though always rounded), more concave laminz

lying much higher one upon the other, and a stronger umbo bent

further over the opercular valve, than in Gryphea coulom Defy.”

Some of the shells figured by d’Orbigny as Ostrea couloni§ show

no slight resemblance to selected individuals of H. imbricata, but are

narrower than the generality of the Uitenhage specimens, and the

imbricating lamelle 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). + Leymerie (1).

+ Coquand (2). § d’Orbigny (3), pl. 467, figs. 1-3 (1848).

The Invertebrate Fauna of the Uitenhage Series. 81

into comparison. The shells figured by Coquand* as Ostrea couloni

include narrow forms, more elongated and more decidedly carinated

than H. 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,t though itself very variable, is

typically widely separated from EH. 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 H. imbricata. Thus, Leymerie’s EH. sinwata, var.

elongata {| can be fairly closely matched, while extreme individuals

of H. wmbricata, in their narrow outline and very arcuate growth,

nearly resemble that author’s figure of H. swhsinuwata, 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,” from the Lower Cretaceous Gio beds

of Patagonia, may, in selected individuals, somewhat closely

approach H. 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 H. wmbricata is probably somewhat remote.

The shell from the Neocomian cf Arqueros in Chili described by

Bayle and Coquand as Ostrea coulon,** and afterwards brought into

association with a heterogeneous company under the collective name

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

figs. 1-6, 22.

+ J. Sowerby (1), vol. iv., Tab. 336 (1822). [As Gryphea. |

{ Leymerie (2), p. 17, pl. 12, fig. 2.

§ Ibid., pl. 12, figs. 6, 7.

|| Bayle (1), pl. exl., figs. 3-5.

{ Stanton (3), p. 11, pl. i., figs. 1, 2; pl. ii., figs. 1, 2.

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

82 Annals of the South African Musewm.

“ Ostrea aquilina”’ by Coquand,* closely resembles many specimens

of EH. wmbricata 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. Remes has considered

both the Chilian shell and Exogyra imbricata Krauss to be identical

with Sowerby’s Hxogyra sinuata,t but this is certainly an error,

The same author, however, has figured an Hxogyra from Stramberg, t

ascribed by him to Hxogyra subsinuata Leym., to which selected

individuals of H. 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

Douvillé § to H. ambricata Krauss, but this was an erroneous identi-

fication, as already pointed out by G. Miiller.|| The resemblance

of the small specimens figured by Douvillé to selected young

individuals of H. zmbricata 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

Douvillé’s types, if these represent the adult stage. The variation

in EH. 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 Douvillé to bear

a close resemblance to Hxogyra wmbricata Krauss.‘

Genus MYTILUS Linneus.

MytTILUS UITENHAGENSIS sp. nov.

Plate ©f., figs. 10; dy ila:

Description.—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. { Remes (1), p. 216.

t Remes (1), pl. xxi. (iv.), figs. 2a, 2b.

§ Douvillé (1), p. 230, pl. xii., figs. 8, 9. || G. Miiller (1), pp. 569, 570.

{| Douvillé (2), p. 388; Douvillé (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 ................0c.ceeeeseeeee 37 mm.

encthvok the cardinal margins... vos seqe) es eccaascaesaccoseeeeee Gi

Greatest breadth of flank between the carinal ridge and the

POSTEKIO“ Val Ve=mMaAnsin owe .seese ccna ctwesseccetee seeessateees 1345

Greatest width of the antero-ventral area in one valve ...... tne

Occurrence.—Found at Dunbrodie, Sunday’s River (319). Speci-

mens collected by Atherstone and preserved in the British Museum

(Natural History) are labelled ‘‘Cuylers.” Also collected by Mr.

Rogers from the lowest beds on Zoet Geneugd, right bank of

Sunday’s River (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 River

below Uitenhage, in association with Trigonia herzogi, T. conocardit-

fornus, 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 Mytilus. 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 lanceo-

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.)} from the Lower Greensand and Upper Greensand and

Blackdown Beds of England, WM. witenhagensis 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. witen-

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. witenhagensis 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,t may or may not be identical with the European form.

To judge from the description and figures, it shows considerable

similarity to MW. witenhagensis, 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 (8), vol. i., p. 110 (1900).

+ J. de C. Sowerby (1), vol. v., p. 55, Tab. 439, fig. 2 (1823); H. Woods (8),

vol. i., p. 110, pl. xviii., figs. 13-15, pl. xix., figs, 1-11 (1900).

t Whiteaves (3), p. 236, pl. 31, figs. 7, 7a.

The Invertebrate Fauna of the Uitenhage Series. 85

by J. Miiller,* and united by Holzapfel+ under the name Septifer

tequlatus, 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. witenhagensis 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 hinge-line is relatively longer than in M. witenhagensis, 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 MW. witenhagensis is Mytilus

triangularis J. Bohm,$ from the Trigonia-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 MM. triangularis, too, the flank between the carina

and the posterior margin is slightly narrower, while Dr. Bohm’s

figure 11 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.{! from the Lias of Avioth, Breux.

This closely resembles M. triangularis, from the Syrian Cretaceous,

in most outward characters, and differs from M. witenhagensis

chiefly in those points which distinguish MM. triangularis also,

though exhibiting a slight convexity of the antero-ventral area not

developed in these Cretaceous forms.

' Genus MODIOLA Lamarck.

MoptioLa BAINI Sharpe.

Plate IT., figs. 12, 12a.

1856. Modiola bain D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

Vol. vil. p. 193; pl. 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. + Holzapfel (1), p. 218, Taf. xxv., figs. 1-9.

t d’Archiac (1), p. 307, pl. xv., fig. 3. § J. Bohm (2), p. 218, Taf. vii., fig. 11.

|| For remarks on the age of this sandstone see Rehbinder (1), p. 87.

| Buvignier (1), p. 24, pl. xvi., figs. 35-37.

U

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 River valley, one mile

north-east of Rawson Bridge (281). Recorded by Sharpe from the

Sunday’s River at ‘C. Roe’s drift,’’ and by Stow from above Modder

Drift and from M’Loughlin’s Bluff, on Sunday’s River.

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 }

* d’Orbigny (4), vol. i., pp. 282, 312; J. Sowerby (1), vol. iii., Tab. 248, fig. 1 (1819).

+ Thurmann and Etallon (1), p. 223, pl. xxix., fig. 8 (1862); Loriol and Pellat

(2), p. 156 (312), pl. xviii., figs. 19, 20.

{ See also Loriol, Royer, and Tombeck (1), p. 348, pl. xix., figs. 10, 11.

The Invertebrate Fauna of the Uitenhage 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. medus

(d’Orbigny), an allied shell from the Kimeridge.

A shell of very similar type to M. baini is M. flagelltfera (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

Guéranger | as Mytilus sarthensis. It may be clearly seen from

that author’s photographic illustration that this differs from M. bain

by the relatively greater disparity in the number of large and small

ribs; in the French shell there are three small ribs to each of the

major ribs.

Modiola rubidger (Tate),} from the Uitenhage Marine Beds, also

bears a general resemblance to M. bauw. In M. rwbidger, 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. bani; 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. rubidger

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, pl. xvi.; fig. 9; Woods (3), vel. i., p. 99, pl. xvii., figs. 1, 2

(1900).

+ Guéranger (1), p. 17, pl. xxiii., fig. 1. t Tate (1), p. 157, pl. 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 .—:

Lien gil ssc nae eset eae noch haw tamer 17 mm

Greatest height. .65...saseeseee eee 11s aes

Greatest depth of a single valve ......... OE:

Occurrence.—Found by Mr. Rogers in a wash-out 100 feet above

Coega station, at a point one mile north of Coega Hotel (441).

temarks.—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 GRAMMATODON F. B. Meek and F. V. Hayden.

GRAMMATODON JONESI (Tate).

Plate II., figs. 14, 14a.

1867. Arca (Cucull@a?) jonesi R. Tate, Quart. Journ. Geol. Soc.,

vol. xxiti.,) ps l6L.plax. igo,

Occurrence.—Collected by Miss M. Wilman at Coega. This shell

is also recorded by Messrs. Rogers and Schwarz as occurring in the

highest of three beds of sandstone on the Grass Ridge road near

Uitenhage.t| The specimens described by Tate occurred “in a

* Woods (3), vol. i., p. 22, pl. iv., figs. 2, 3, 4 (1899).

+ Rogers and Schwarz (1), 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 River.”

Dimensions.— (1) (2) (3) (4)

Ibengtht etn esse scene 13 : 18 ; 21 ; 22 mm.

Hiei ght ae tasseisioceass 9. ff “ieheemea tutes 5 > cag: 3)

Depth of asingle valve 4 : — ‘ 6 , =

No. (2) is Tate’s figured specimen.

Remarks.—This form is closely similar in general type to G. secwris

(Leym.) * (Lower Cretaceous) and G. carinatus (Sow.) + (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 Arcide. In the use of the name

Grammatodon I follow Mr. Woods.

Compared with G. secwris, 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. jonest. 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

Kast Africa, which were correlated by G. Miiller with the Uitenhage

Series. Miller described and figured this under the name Arca

witenhagensis,§ and drew attention to the apparent relationship to

Arca jonest Tate, A. witenhagensis, which is associated with

Trigonia ventricosa (Krauss) and 7’. beyschlagi Miller, differs from

Grammatodon jonesi by the fewer number and much greater breadth

of its ribs.

Genus UNIO A. J. Retzius.

UNIO UITENHAGENSIS sp. nov.

Plate IT., figs. 15, 15a.

Description.—The shell is of elongated oval outline, posteriorly

produced, and laterally compressed in the postero-inferior region of

* Leymerie (2), p. 6, pl. vii., figs. 6, 7; Woods (3), vol. i., part 1, p. 44, pl. vii.,

figs. 14, 15, pl. viii., figs. 1, 2 (1899).

+ J. Sowerby (1), vol.i., p. 96, Tab. 44, lower figure (1813) ; Woods (8), vol. i.,

p. 45, pl. viii., figs. 3-8 (1899). t Woods (2).

§ G. Miller (1), p. 542, pl. 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. s2.--¢-- 4052-00 37 : 43 mm,

Greatest height)<5,...20).0¢scce-c--2 Ot ‘ Bsa

Greatest depth of a single valve 7 : 854,

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 River 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. Rogers 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 porrectus J. de C. Sow.,+ 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.

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

The Invertebrate Fauna of the Uitenhage 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) | 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

Germary, 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 witenhagensis. Unio planws Roem.|| 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 TRIGONIA J. G. Bruguiére.

TRIGONIA VENTRICOSA (Krauss).

Plate III., figs. 1, la.

1843. Lyriodon ventricosa F. Krauss, Amtlicher Bericht tiber 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.

Cees. Leop.-Carol. Nat. Cur., vol. xxii., pt. 2, p. 456,

Tab. 49, figs. 2c-2f, (exclude figs. 2a, 20).

1871. Trigonia ventricosa F. Stoliezka, Cretaceous Fauna of

Southern India, vol. ii., The Pelecypoda; p. 315,

pl. xv., figs. 9. 9a. (Palzeontologia Indica.)

1875. Trigonia ventricosa J. Lycett, Mon. Brit. Foss. Trigonie,

p. 119, and text-figures. (Palewontographical Society.)

1877. Trigonia ventricosa O. Feistmantel, Foss. Flora of the

Gondwana System, vol. i., pt. 3, p. 164. (Paleontologia

Indica.)

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

+ J. de C. Sowerby (2), p. 346, pl. xxi., fig. 17.

{ 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; Dunkerand 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. (Paleeontologia Indica.)

1893. Trigona ventricosa W. T. Blanford, in Medlicott and Blan-

ford, Manual of the Geology of India (2nd _ edition

revised and partly rewritten by R. D. Oldham), p. 225,

with figure.

1900. Trigonia- ventricosa G. Miller, Verstein. des Jura und der

Kreide, Deutsch-Ost-Afrika, Band vii., p. 5438, Taf. xix.,

figs. 4, 5.

1903. Trigona ventricosa F. L. Kitchin, Jurassic Fauna of Cutch,

vol. iii, pt. 2, The Lamellibranchiata; p. 104, pl. x.,

figs. 4-8. (Paleontologia 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 244-243 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. + Krauss found it

in the left bank of Zwartkop’s River below Uitenhage, associated

with Trigonia herzogi and T. conocardifornus.

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 beyschlagi,

which shows strong resemblance to TJ. crassa Kitchin, from the

Oomia marine beds in Cutch.

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

The Invertebrate Fauna of the Uitenhage Series. 93

In India, 7. 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 Scabre, 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 J. 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 JZ’. 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 7. tuberculifera.

A comparison of 7. ventricosa with T. krausst sp. noy. will be

found in the remarks appended to the description of the latter.

T. subventricosa 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. delafosset Bayle and Coquand,** from the Neocomian of Chil,

also shows a general agreement with TJ. ventricosa, and may

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

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

t Parkinson (1), p. 176, pl. xii., fig. 9; Lycett (3), p. 116, pl. xxv., figs. 3-6

(1875). § Lycett (3), p. 130, pl. xxvii., figs. 4, 5.

|| Stoliczka (2), pl. xv., fig. 10 (1871). {| Stanton (8), p. 18, pl. iv., figs. 19, 20

** Bayle and Coquand (1), p. 37, pl. viii., fig. 27; R. A. Philippi (1), p. 85,

pl. 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 TJ. 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, TJ. nepos

Paulcke, from the Neocomian of Chili, shows to some extent a

similarity to J’. 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 7. nepos and

T. delafosset into close relationship; yet he perceives in 7’. nepos an

exceedingly close resemblance to JT. baylei Dollfus, + 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. baylet and other

typical Jurassic Scaphoidex 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 7’. alifornis 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 Scaphoidez, the laterally compressed form,

the weak umbonal incurvation, the nodose carinze and delicately

sculptured area, no less than the relations of frontal to lateral

coste, 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 Scabre.

In the published description of the Trigoni@ 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.

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

The Invertebrate Fauna of the Uttenhage Serves. 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 KRAUSSI sp. Noy.

Plate IIT., figs. 2, 2a.

1850. Lyrodon ventricosus F. Krauss (partum), Nov. Act. Acad. Cees.

Leop.-Carol. Nat. Cur., vol. xxii., pt. 2, Tab. 49, figs. 2a,

2b (exclude figs. 2c—2f).

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 Musewm.

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) /- secs mee-es-ecst-2ceoss- se ances 75 mm

Height measured from the umbonal region......... Sosa

Greatest depth of a single valve................ssssee0 mo) oy

The Invertebrate Fauna of the Uitenhage Series. 97

Occurrence.—Messrs. Rogers and Schwarz record the occurrence

of this shell (“the large variety of JT. ventricosa’’) * in the railway

cutting between milestones 244-242 on the line from Uitenhage to

Graaff-Reinet, 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 River below Uitenhage.

Remarks.—This characteristic Trigonia, which belongs to the

section Scabre, may be said to exhibit in very exaggerated manner

some of the most striking features which render 7. 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. alifornis Park., and was even considered

identical with the latter by L. von Buch.{ TZ. kraussz 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 J. ventricosa

and the apparently closely allied 7. swhventricosa which occurs in

the Belgrano beds of Patagonia.

T. krausst 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 TZ’.

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 ZT’. ventricosa have been obtained by various

collectors, and the Uitenhage 7rigonie@ 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 Trzgone of the Oomia beds in Cutch the representa-

tives of J’. ventricosa, met with in abundance, agree with the smaller

typical form in South Africa.

T’. krawssi has approximately double the dimensions of 7’. 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

* Rogers and Schwarz (1), p. 10. + von Buch (1), p. 23.

98 Annals of the South African Museum.

width. In the adult 7. krausst, 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 7’. ventricosa, the anterior

ribs on the flank of 7. krausst are already much more robust and

more widely spaced than in 7’. 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 7. krawssi 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. krawssz, from the fact that it exhibits

all the late adult characters of 7. ventricosa while yet in the early

adult stage itself; the late adult characters of 7. kraussi are never

reached in 7. 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 TJ. 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 J. 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 7. ventricosa and T. krausst 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 TY. 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 JZ’. ventricosa. Moreover, no less than six

authors have figured the smaller shells under the name given

by Krauss.

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

The Invertebrate Fauna of the Uitenhage Serves. 99

TRIGONIA ROGERSI Sp. Nov.

Plate TIL., figs. 3, 83a; IV., fig. 1; V., fig. 2.

Description.—The shell is of somewhat elongated form; it is

anteriorly high and is posteriorly more siender 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 cost 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 cost 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 the 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 umbonal 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. Ata 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

84mm. 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 Uitenhage Serves. 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)

hength (estimaitied)) <25.....2.6.+.00- 78 ; 84 : 135 mm.

GreatestiWerghh s.tsicceceeseecdee ee 53 : 58 : G2;

Greatest poe of a single valve 19 23 5 32: 4,

Obtained by Mr. Rogers got a bare slope W. 30S.

from the middle of Barkly Bridge, on the farm Olifant’s Kop,

Sunday’s River (20h) ; also from Coega Valley, east of the railway,

one mile up the line from Coega station (472g).

Remarks.—This is an elongated representative of the section

Scabre, 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 Trigoni@ from the Uitenhage beds and from the

Oomia 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 ZT’. conocardiifornis (Uitenhage Series) and in members of the

group of 7’. v-scripta and certain degenerate Costatee (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 siphonal 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 7’. stow2.

In the neanic and early adult stages, 7’. rogersz bears a consider-

able resemblance to the same stages in 7’. 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 herzogiti (Hausmann) A. Goldfuss, Petrefacta

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

fig. 6.

1850. Lyrodon herzogii F. Krauss, Nov. Act. Acad. Cas, 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 fiir Mine-

ralogie, Band i,)p..220) Mat. vil, figs, 12 latex

figs. 1,2.

Occurrence.—This is an abundant and characteristic form in the

Sunday’s River Beds. It was recorded by Hausmann 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 Rivers. A fine specimen

sent to me from the South African Museum is from the Sunday’s

River (289). Messrs. Rogers and Schwarz found T. herzogi 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 2441-243 on the Graaff-

Reinet 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 7. 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 (4742); the cliff on Buck Kraal, Sunday’s River

(116h, 120h, 122h); and the cliffs on Zoet Geneugd, Sunday’s

River (99h).

LRemarks.—Trigonia herzogi is well known from the striking figure

given by Goldfuss, illustrating a fine specimen which was included

in a small collection of Uitenhage molluses 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 7. herzogi

with T. transitoria Steinm. in the group Pseudo-quadrate, the

characters of which are briefly discussed below in the remarks

appended to the description of Trigonia holubi. Steinmann has set

forth in great detail the differences that distinguish 7. herzogi from

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

The Invertebrate Fauna of the Uitenhage Series. 103

T. transitoria, which occurs in Neocomian strata in Bolivia, Chili,

and the Argentine Republic. 7’. transitoria shows a closer approxi-

mation of characters to the Clavellate; 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 TZ. transitoria the rows of nodes representing

the marginal and median carinee become joined by transverse rounded

ridges at an earlier growth-stage than in 7’. 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 7. transitoria, while ornamented at

an early stage in the African shell; in 7’. transitoria the escutcheon,

generally speaking, is more sparsely sculptured.

I have elsewhere pointed out the features of distinction between

T. herzogt and 7’. mamillata, from the Oomia beds of Cutch.*

T. mamillata shares the principal characters of the Pseudo-quadrate,

but is a shorter shell than 7. herzogi, and is vertically truncated

in front. In 7. mamullata 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. herzogi are backwardly directed and are not curved. On the

area of 7’. 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 7. 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. herzogt, 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 7’. 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 + represent J. herzogi,

but very greatly reduced in size.

TRIGONIA HOLUBI sp. nov.

Plate IV., figs. 2, 2a.

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. + 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 Uttenhage 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 Fo AO) 3) 20 2) l32inam:

Greatest height ...... 75 ; 78 : 87 ; eo)

Depth of asingle valve 18 ‘ 20 22 3 DiS sy,

Occurrence.—Specimens in the collection of the South African

Museum came from the Sunday’s River (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

LT. holwbt.

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 Musewm.

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 Trigonia allied to

T. herzogi; he gave the following description of specimens which

were presented by J. 8. Bowerbank to the Geological Society of

London: ‘‘ Trigonia sp.(?). This variety (or possibly distinct

species) is closely allied to Trigonia herzogi, 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. holwbi clearly cannot be united with either the Clavellatz or

the Quadratz, but its characters are those of the somewhat inter-

mediate kind which led Steinmann to establish the group Pseudo-

* Sharpe (1), p. 202.

The Invertebrate Fauna of the Uitenhage Series. 107

quadrate, to receive Trigonia herzogi (Goldf.), the well-known

Uitenhage form, and TJ. transitoria Steinmann, from Lower

Cretaceous strata in Chili, Bolivia, and the Argentine Republic.*

T. neuquensis Burckhardt,+ which is found associated with T. transi-

toria in the Neocomian Trigonia-beds of Las Lajas (Rio Agrio,

Argentine Republic), also falls within this division, and another

example of the same group is 7. mamillata, which occurs in the

Oomia Trigonia-beds near Goonaree in Cutch.} These five members

of the Pseudo-quadrate show similar deviation from the characters of

the two sections with which they may best be compared. Like the

Quadrate, 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 Clavellate, with which, both as regards

the youthful ornamentation and the adult form, it has much in

common. In the Quadratz the area is usually divided by its longi-

tudinal groove into a broader, superior, and narrower inferior

portion ; in the Pseudo-quadratz 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 Quadratze

but absent in the Pseudo-quadrate. 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-quadratie,

and may be almost twice as broad as the upper one.

T. holubt is most closely comparable with the Indian 7. mamillata,

to which it shows a striking similarity. In 7. holwbi, 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. mamuillata the phase in which

irregular transverse costz crossing the area form a dominant feature

is reached at a much earlier stage of growth than in 7. holubv.

Thus, in 7. manullata, strongly developed costz appear on the area

soon after the neanic stage is passed and long before half the adult

dimensions are attained. In 7’. holwbi, 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). + Burckhardt (2), p. 74, Taf. xiv., figs. 4-6.

{ Kitchin (1)}p) 100, pl. ix., figs. "8,95 pl. x. figs: 1-3:

108 Annals of the South 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 7. mamillata.

From Trigonia herzogi (Goldf.),* T. holubi 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. herzogi are straight, while those of 7. holubi are curved. The

backward slope of the posterior flank ribs in 7’. herzogi is another

point of distinction.

There is a marked general similarity to Tvrigonia transitorva

Steinmann,+ from South America, but the ornamentation differs

considerably. In TY. transitoria the tubercles of the costz on the

flank are placed closely against one another through all adult stages

instead of being openly spaced as in the ribs of T. holubi. 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 1s unnecessary to

enumerate.

T. neuquensis Burckhardt is at once readily distinguished from

T. holubt 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 7. holubi. In T. newguensis, 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 7’. holubt.

Note on the Development of the Pseudo-quadrate Trigonie.

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. holubi 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).

+ Steinmann (1), p. 260, Taf. ili., fig. 3; Steinmann (2), p. 221, Taf. vii.,

figs. 3, 4, Taf. viii., figs. 1-8; Burckhardt (1), p. 21, pl. xxv., figs. 1-8;

Burckhardt (2), p. 73, Taf. xiv., figs. 1, 2.

The Invertebrate Fauna of the Uitenhage Series. 109

of the tubercles of the flank coste—they become crowded closely

together on a given rib—is a very striking feature. In 7. mamullata

from Cutch, these characters are also coincident, but they are

acquired at a much earlier pevicd of growth. The members of the

Pseudo-quadratw, 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, 7. mamuillata

is in advance of T. holubt. In T. holubi, 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 7. mamillata, this trituberculate

stage is passed over rapidly and is superseded in very early adult

life. In 7. herzogz 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 7. holwbi. A point of great

interest is that in 7. mamillata and T. herzogi, 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 TZ. holubi, 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-quadratze 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

molluses and brachiopods give great probability to the truth of this

idea. We must suppose all these forms to have been descended from

true Clavellatz, just as in the case of the Quadratee of Europe, and

it is a very striking circumstance that the characters which mark

the Pseudo-quadrate 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 Quadrats# and Pseudo-quadrate,

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

from the ancestral clavellate characters) may be supposed to have

been in some degree independent.

110 Amnals of the South African Museum.

Confining our attention to the Pseudo-quadrate, 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-quadrate 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 Quadrate in Europe, and of

the comparable though independent Pseudo-quadratee of the south,

took place, so far as we know, invariably in Neocomian times.

TRIGONIA VAU Sharpe.

Plate VI., figs. 1, ta, 2, 2a, 20, 3.

1856. Trigonia vaw D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

vol. wil, ps LO4 ple exis tices:

? 1867. Trigoma vau R. Tate, Quart. Journ. Geol. Soc., vol. xxiil.,

pl vil, fe.6:

Occurrence.—The record of occurrence given by Sharpe was

“Sunday River, in greenish-grey grit, with fragments of wood

and shells; and in a shelly grit at the Zwartkop River.” Speci-

The Invertebrate Fauna of the Uitenhage Series. ia

mens preserved in the collection of the Geological Society of

London are labelled ‘Sunday River’? (Atherstone and Bain),

“Prince Alfred’s Rest’? (Rubidge), and “ Zwartkop River”

(Rubidge). Specimens sent to me from the South African

Museum are from the Sunday’s River. Messrs. Rogers and

Schwarz record 7. vaw from the following localities: in the neigh-

bourhood of Uitenhage, ascending the Grass Ridge road, in the

uppermost of three beds of sandstone exposed in the dry water-

courses, and in the railway cutting between milestones 244-24? on

the Graaff-Reinet line; on Sunday’s River, in the upper beds exposed

in a white krantz on Wolve Kraal, on the north bank of the river.

In 1905 Mr. Rogers collected specimens 300 yards below Addo

Drift (Tunbridge’s), left bank of Sunday’s River (40h, 41h, 42h).

An immature specimen (46h) from the same locality, may also

belong to Trigonia vau.

Remarks.—A large specimen of T. vaw 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 J. vaw from

T’. stow? appear up to the present to have escaped observation, and

it is probable that the shells cited under the name J. vaw by Stow *

are in reality to be ascribed to.7. stowi sp. nov. I have had the

opportunity of examining and comparing a considerable number

of excellently preserved specimens of these two Trigonie, and find

that the characters which distinguish them from one another are

very constant. Several fully-grown shells of 7. vaw 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. vaw 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.

Rogers and Schwarz at Walton’s Farm, just below Dunbrodie on

the Sunday’s River (307), and another individual measuring 17 mm.

in length, collected between milestones 244-242 on the Graaff-Reinet

railway (309). There is the same doubt concerning an immature

* Stow (1), pp. 499-505.

(1), ¥

112 Annals of the South African Musewm.

specimen from Addo Drift (46h). For the same reason it is uncer-

tain whether the young individual figured by Tate really represents

the true 7. vau. 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 TJ. vaw and its associate

when in the youthful stage. These are the members of the

Trigonia v-scripta group in the marine Oomia beds of Cutch,*

and a comparison of their youthful characters and those of 7. vaw

and 7’. stowt is given in the remarks which are appended to the

description of the latter. J. stowi when adult approaches closely

in many particulars to 7. v-scripta itself, but the adult 7. vaw 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 JT. dubia is not known, but it appears

reasonable to assume that it bears closer relationship to the other

members of the group of T. v-scripta than to T. vaw. In the adult

stages, too, points of distinction between T. dubia and T. vaw are

not wanting. These have been set forth in my account of the

characters of T. dubiat+ as follows: “In T. vau, the ribs of

the anterior series, though narrow and crowded as in ZT. 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 7. dubia, on the other hand, these anterior

coste 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 JT. vaw have reached

half their full dimensions, the costz 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 costae of the frontal series. This peculiar

character is totally absent in 7. dubia, in no specimen of which

is a frontal unsculptured space seen, although in respect to size

* Kitchin (1), p. 65. { Ibid., p. 69.

The Invertebrate Fauna of the Uitenhage Series. 113

all the individuals examined considerably exceed the dimensions at

which this feature becomes noticeable in T. vau. . . . Sharpe’s type

is of larger dimensions than the known specimens of T. dubia.”

The extent to which an unsculptured space near the lower part of

the frontal margin may be present in J. vaw 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 7. v-scripta. It may also be

noted that although the inclination of the frontal series of coste in

T. vau 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 7. vau 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 7. v-scrzpta in India,

and J’. vaw and T. stowz in South Africa. TZ’. kiihna seems to differ,

however, by its less equilateral form and the persistent ornamenta-

tion of its area. The figure of an imperfect specimen of 7. kiihnz

(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 7. vauw, though still. more reminiscent

of 7. recurva Kitchin, from the Oomia beds. It is as yet not

possible to say, however, whether 7’. kiihnz is more nearly related to

T. vau 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 heterosculpta Stanton,} from the Belgrano beds (Lower

Cretaceous) of Patagonia, is very probably closely related to 7’. vau,

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 JZ. vau, 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 7. vaw.

* G. Miller (1), p. 561, Taf. xxv., figs. 6-8.

+ Stanton (3), p. 20, pl. iv., figs. 16-18.

114 Annals of the South African Musewm.

R. A. Philippi* has figured some Chilian forms (for example,

T. arsinoe Philippi, 7’. foveata Philippi) which may be related to

T. vaw; 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

diversicostata Whiteaves,+ from the Cretaceous of Queen Charlotte

Islands, bears resemblance to 7’. vaw. 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 7. diversicostata, however, the anterior

ribs are much coarser in character than in 7’. vaw, 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. diversicostata to the section Scaphoidesw, 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 Costatee, from which

T. diversicostata 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 Costatz of the marine Oomia beds in Cutch. TJ. vaw

shows no signs of a connection with the section Costatz at any

early growth-stage that can be studied, but the ornaments of the

area are transverse from the first.

The peculiarly ornamented Trigomia doroschint HKichwald,{ from

the Neocomian of Tukusitnu Bay (Alaska), may finally be brought

into comparison. This appears to me, however, to be well separated

from the group of 7’. vaw (as well as from the group of 7’. 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

* R. A. Philippi (1), pls. 34, 35. + Whiteaves (1), p. 68, pl. x., fig. 1.

+ Hichwald (2), p. 180, pl. xiii., figs 12-14; xiv., figs. 1-4.

The Invertebrate Fauna of the Urtenhage Series. 115

to the frontal margin. It seems doubtful from Hichwald’s figures

whether a single form alone is represented, for it is difficult to

believe that the originals of pl. xiii., fig. 12, pl. xiv., fig. 1, and

pl. xiv., fig. 3 can belong to one species. Other points in which

these differ from 7’. vaw are the great extent of the frontal ribs, the

horizontal position of these, and the persistent transverse ornaments

on the area. Hichwald’s description and figures leave the affinities

of T. doroschini obscure, but I think it improbable that it is related

to the group of 7’. vau.

TRIGONIA STOWL sp. nov.

Plate VI., figs. 4, 4a, 40,5; VII., fig. 1.

Description.—The shell is of very elongated form, moderately

inflated, anteriorly pointed, posteriorly very much produced, and

slightly gaping at the siphonal 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

iower 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 Musewm.

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. Ata 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 Invertebrate Fauna of the Urtenhage Serves. uy

though imperfectly demarcated from the area, it is marked by its

excavated form and smooth surface. Ridges 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

umbones 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) lene pling. .sccoc...2-2-citcene 80 ; 95 0) ELOr mame

Height, measured from the umbo 42 : 44 ‘ i510) ee

Greatest depth of a single valve... 14 : 15 F DOs iss

Occurrence.—Marine Beds of Sunday’s and Zwartkop’s Rivers.

Specimens in the collection of the Geological Society of London are

labelled “ Zwartkop River’ (Rubidge) ; ‘‘ 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

River, 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 7’. stow?z, were

collected by Messrs. Rogers and Schwarz at Walton’s Farm on the

Sunday’s River, just below Dunbrodie (307). These may probably

represent 7’. stow2, though they might equally well be ascribed to

T. vaw so far as their characters allow of determination. Another

immature specimen, from the railway cutting between milestones

241-243 on the Uitenhage—Graaff-Reinet railway (309) may also

perhaps belong to this form. Mr. Rogers collected JZ’. stowz 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 8. 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 River” and

‘* Lower Sundays River”; Stow (1), fig. 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 umbones 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. vau

Sharpe. When dealing with the Trigone of the Oomia beds of

Cutch, I made reference to 7’. stow? as an undescribed form allied to

T. vau.* 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. stowt 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 7’. v-scripta WKitchin, yet its youthful

characters plainly reveal its close relationship to 7. vaw 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, 7’. stowi 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 7’. stow7, are developed to a much

* Kitchin (1), pp. 66, 74.

The Invertebrate Fauna of the Uitenhage Series. 119

less noticeable extent in 7. vaw, 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, 7. stow? can only

be closely compared with 7. 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, 7. stow? may

be readily distinguished by its relatively more elongated and poste-

riorly attenuated figure, and its anteriorly more acutely converging

upper and lower margins. TT. 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; 7. 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 7. recwrva, another Oomia form.

This, like 7. stow7, 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.

TL’. heterosculpta 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 7’. stow, but the adult shell

is much shorter, more triangular in form, and much less elongated

posteriorly, while there are marked differences in the details of

sculpture.

TRIGONIA CONOCARDIIFORMIS (Krauss).

Plate VII., figs. 2, 2a, 20, 3, 4.

1843. Lyriodon conocardiiformis F. Krauss, Amtlicher Bericht iiber

die zwanzigste Versammlung der Gesellsch. deutsch.

Naturforscher und Aerzte zu Mainz im Sept., 1842,

p. 180.

1850. Lyrodon conocardiiformis F. Krauss, Nov. Act. Acad. Ces.

Leop.-Carol. Nat. Cur., vol. xxii., pt. 2, p. 454, Tab. 49,

figs. la—ld.

* Stanton (3), p. 20, pl. iv., figs. 16-18.

120 Annals of the South African Museum.

1879. Trigonia conocardiiformis J. Lycett, Mon. Brit. Foss. Trigonie,

pp. 210, 211, and woodcuts; p. 230. (Palseontographical

Society.)

1903. Trigonia conocardiiformis C. Burckhardt, Beitrage zur Kennt-

niss der Jura- und Kreideformation der Cordillere. Palex-

ontographica, Band 1., Taf. xii., 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 favourably 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

pallial border; it occurred in a hand-specimen, containing adult

T’. conocardiuformis, T. vau, and other typical forms, from the

Sunday’s River. The other specimen is an immature individual

haying 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° 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 Invertebrate Fauna of the Uitenhage Serves. 121

even at this early stage, but the ribs encroach slightly upon the

smooth eseutcheon. 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 sueceeding 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 adult 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.

Dinensions.—Trigoma conocardiufornus 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 Zwartkop’s Rivers. It was obtained

by Messrs. Rogers and Schwarz from the railway cutting between

milestones 244-242 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 HE. 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 vaw and T. stow; 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 7. conocardiiformis seem

perhaps to suggest an ancestry similar to that from which 7. vau

and its ally were derived. In the young IT. conocarduformis,

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 Scabrze

* Schwarz (1), pp. 53, 61; Rogers and Schwarz (1), p. 5; Rogers (1), p. 295.

The Invertebrate Fauna of the Uitenhage Series. 123

or the Costate; but in view of the remarkable manner in which

widely divergent types of Trigonia in the Oomia beds of Cutch

show, by their neanic characters, their relationship to the Costate,

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. lLycett at first expressed the

belief that it is an abnormal example of the Clavellatz, 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 Scabre.’ + 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 7’. conocardiiformis has

been found to occur, and in association with two members of the

Pseudo-quadrate. The shells described by Burckhardt} under

the name ‘ Zrigonia aff. conocarduformis’’ 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 7’. 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. conocardiifornis. 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). + Ibid., p. 210 (1879).

t Burckhardt (2), p. 72, pl: xiii., figs. 1, 2.

124 Annals of the South African Musewm.

Another South American form which is probably also nearly

related, is Trigonia eximia R. A. Philippi,* from the Tinguirica

valley in Chili, This represents a type of shell very closely com-

parable with 7. 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 Tvrigonia aff.

conocardufornmis may be regarded as illustrating a somewhat inter-

mediate type of sculpture between the two extremes, 7’. conocardi-

formis and T. exumia. Another probably allied shell has been

recorded by Haupt} from the Neocomian of Loteno, on the Rio

Neuquen, on the east slope of the Argentine Cordillera, under the

name T'rigonia cf. exinva 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. 1’. exama

is referred by Haupt to the section Undulatie, 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 7. 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 7’. 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 7’. eximia, in the other individuals from Cape

Colony, might probably be due to the same cause.

* R. A. Philippi (1), p. 76, pl. xxxiv., fig. 3.

+ Haupt (1), p. 216.

The Invertebrate Fauna of the Uitenhage Series. 125

TRIGONIA TATEI Neumayr.

1867. Trigonia cassiope R, 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, pl. ii., fig. 3.

Occurrence.—A very imperfect and ill-preserved fragment of a

costate Trigonia occurs in hard limestone with Plewromya baina

from Grass Ridge, three miles east-north-east of Uitenhage (330).

In all probability this represents 7. tatez.

Remarks.—Although no complete or satisfactorily determinable

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 Costate 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 7. cassiope by d’Orbigny,t and Neumayr sub-

sequently saw the necessity of applying a new name. 1’. tated

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 Costatx, yet the narrow

area and elongated escutcheon, the delicate and little-prominent

carine 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 River” (H. Longlands)

and ‘“McLoughlin’s Rest’’ (Major Rocke) 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 carine,

the fine intercarinal sculpture, the convex area, and the elongated

eranular escutcheon, are all characters which at once recall the

* Lycett (3), p. 172. + d’Orbigny (4), voli., p. 308.

126 Annals of the South African Museum.

same features in 7. tenuis, from the Oomia beds in Cutch.* This,

however, is strongly distinguished from 7’. tate? 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 Costatz 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 Costate than when

our knowledge of the distribution of this division of the Trigoni@

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 typical Jurassic forms. Trigonia peninsularis

Coquand,+ from the Aptian of Spain, exhibits a marked degeneration

of characters in its later growth-stages, while 7. carinata Agassiz, {

from the Neocomian and Aptian, which may be brought into the

most intimate connection with the Costate, 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 Costate 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 Oomia beds of Cutch, which may now be regarded as

of Lower Cretaceous rather than of Upper Jurassic age, aberrant

derivatives of the Costate provide a striking feature in the molluscan

fauna; but here also have been found two representatives, 7’. tenwis

Kitchin and 7. 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 *' from strata which he regards as Upper

* Kitchin (1), p. 35, pl. iii., figs. 5, 6. + Coquand (1), p. 129, pl. xxiii., fig. 3.

t Agassiz (1), p. 43, Tab. vii., figs. 7-10; Lycett (3), p. 179, pl. xxxv., figs 3-6,

(1877). § See also Collot(1).

|| Ascher (1), p. 159 [25], pl. xiii. [ii], fig.10. [| Behrendsen (1), p. 6, pl. 3, fig. 7.

The Invertebrate Fauna of the Urtenhage Series. 127

Cretaceous, at Caryilauhue (Argentine); this is accompanied by

a typical member of the Cretaceous section Scabree, 7’. transatlantica

Behr. TZ. anguste-costata is a small triangular shell, which in the

delicate character and close spacing of the flank-ribs recalls the

Indian JT. 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, 7. 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

Costatz. With reference to the occurrence of the costate Trigonie

in Cretaceous rocks, Behrendsen cites, in addition to JT. peninsu-

laris Coquand and T. carinata Agass., the following shells: 7. 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. TJ. longa and T. pennata certainly

cannot be included in the section Costatze, while it is doubtful

whether 7’. 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 CARDITA J. G. Bruguiére.

CARDITA NUCULOIDES Tate.

Plate VII., figs. 5, 5a, 5d.

1867. Cardita nuculoides R. Tate, Quart. Journ. Geol. Soc., vol.

Xxiil., p. 162, pl. vii., figs. 7a—7d.

Occurrence.—Tate’s figured specimen, numbered 11028 in the

collection of the Geological Society, came from the Sunday’s River.

A specimen obtained by Mr. Rogers is from the left side of the

Coega Valley, half a mile down from the railway (466g).

ftemarks.—The specimen found by Mr. Rogers 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 Musewm.

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 striz which cannot be seen with the

naked eye. The intercrossing of the strie with concentric growth-

lines results in a delicate cancellation.

Genus ASTARTE J. Sowerby.

ASTARTE LONGLANDSIANA Tate.

1867. Astarte longlandsiana R. Tate, Quart. Journ. Geol. Soc.,

vol. xxiii., p. 158, pl. viii., figs. 5a, 50.

Occurrence.—Specimens in the collection of the Geological Society

are from the Zwartkop’s River. Mr. 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.

Susp-aENus ERIPHYLA F. Stoliczka (? W. M. Gabb).

ASTARTE (ERIPHYLA) HERZOGI (Goldfuss).

1840. Cytherea herzogii (Hausmann) A. Goldfuss, Petrefacta

Germanie, Band ii., Lief. 7, p. 239, Tab. exlix., fig. 10.

The Invertebrate Fauna of the Uitenhage Series. 129

1843. Astarte capensis F. Krauss, Amtlicher Bericht itber die

zwanzigste Versammlung der Gesellsch. deutsch.

Naturforscher und Aerzte zu Mainz im Sept., 1842,

p. 150.

1850. Astarte herzogw F. Krauss, Nov. Act. Acad. Ces. Leop.-Carol.

Nat. Cur., vol. xxii., pt. 2, p. 447, Tab. 47, figs. 2a—2e.

1905. Astarte herzogi A. W. Rogers, 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 on 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 Hnon.* Krauss collected specimens on the left bank

of the Zwartkop’s River, below Uitenhage.

Riemarks.—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 meansa

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 pallial 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 Hausmann (1), p. 1459 (as Cytherea).

130 Annals of the South African Musewm.

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.

Stoliezka* 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 Hriphyla hag been applied. Unfortunately, the

employment of this name has been attended with some confusion.

The shell upon which Gabb founded the name (HZ. wmbonata

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 (H. lenticularis) which he believed

to be generically identical with Hriphyla wmbonata Gabb.

Stolicezka, however, brought the genus into relationship with

Dosinia, though be 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 herzog,

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:

“Tt 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 Hriphyla 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). + Speyer (1), p. 496, Taf. xi., fig. 6.

t Goldfuss (1), Band ii., p. 228, Tab. exlvi., fig. 16 (1837).

§ Gabb (1), p. 180, pl. 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 Eriphyla Stoliczka differs in

essentials from Hriphyla Gabb, this name must be retained for shells

having the characters set forth by Stoliczka. Previously, however,

J. Bohm * 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 t has so conceived the meaning of Gabb’s description

as to apply the name Hriphyla 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 Chione.

The question then arises, whether the name Hriphyla as defined

by Stoliczka may be applied to ZL. lenticularis, in view of the fact

that Bosquet had previously proposed to eall 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 Goldfuss’s type. I should have no hesitation in

following Dr. J. Bohm and employing Bosquet’s name on the

strength of the definite reference to Goldfuss, but unfortunately the

description and figure given in the “ Petrefacta Germaniz ’’ 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 Hviphyla, as defined by

Stoliczka.

Hriphyla 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. + Meek (2), p. 123.

+ Whitfield (1), p. 403, pl. vi., figs. 2-5. § Dewalque (1), p. 368.

132 Annals of the South African Museum.

strength of these points, Astarte herzogi 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 Renevier* to Astarte buchi F. Roem. and A. obovata

J. Sow., and in both of these the marginis crenulate. Astarte striata

J. de C. Sow., from the Blackdown Beds of England, shows very

close agreement with H. 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 Astarte herzogi, 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 +} has ignored the notching of the margin, while several

authorities on the living forms have abandoned this structure as even

of specific value.} Regarding the value of the distinctive characters

of Hriphyla, 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 Hriphyla 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 EHripyhla as a sub-genus.

In tracing the relationships of A. herzogi, 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-

nites 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 levis

(Phill.).|| The similarity to A. herzogi is striking, but the following

* Pictet and Renevier (1), pl. x., fig. 1; xi., fig. 1.

+ Zittel (5), p. 65. + 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 Series. 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. herzogi. The noticeable fact that the

sculpture in the Lincolnshire examples is less well defined 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 beawmonti Leym.,+ from the Neocomian of the Aube, is

likewise characterised by a sharply demarcated and deeply excavated

lunule, and is ornamented by ridges and suleations 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 buchi F. Roemer{ (Lower Cretaceous, Perte du Rhone)

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 Roemer’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 Roemer’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 semanni 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).

+ Leymerie (2), p. 4, pl. 4, fig. 1.

t F. Roemer (1), p. 20, fig. 4.

§ De Loriol and Pellat (1), p. 68, pl. vi., fig. 9.

|| Blake (1), p. 232, pl. x., fig. 5.

10

134 Annals of the South African Musewm.

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. herzogi by its more elongated form

and relatively less height. English specimens from the Portland

Sands of Swindon, ascribed to A. semanniz, 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 Hriphyla stuwhl-

mann, 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. LHriphyla stwhlmanni is accompanied

by a Gervillia probably identical with G.dentata Krauss. Dr. Miller

refers his shell to Hriphyla 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) } 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 Grotriama, and, more-

over, have plainly developed lateral hinge-processes fitting into

opposing grooves. In this respect A. jugosa agrees with A. herzogt,

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.

+ G. Muller (1), p. 553, Taf. xxi., figs. 3,4; Taf. xxii., figs. 8-10.

+ Forbes (2), p. 142, pl. xvii., fig. 7.

§ Stoliczka (2), p. 289, pl. x., figs. 12-14.

The Invertebrate Fauna of the Uitenhage Series. 135

Eriphyla argentina Burekhardt,* 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. herzogz by the considerably more

elongated figure and the coarseness and spacing of the ornamenting

ridges. H. argentina, it may be observed, has a weak pallial sinus,

and Dr. Burckhardt remarks that the margin is probably crenulated.

ASTARTE (HRIPHYLA) PINCHINIANA Tate.

Plate VIL., figs. 6, 6a.

1867. Astarte pinchiniana R. Tate, Quart. Journ. Geol. Soc.,

VOlpxxite go. LO, plaix. fe 0.

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.

IGM Slag whens cia daseicecieSaa aor uelnsin viens 15 : aUS5) a IS) Ss

Depth of a single valve ............ SOLE 5 fo 59

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.

Remarks.—Tate’s description and figure of this elegant form must

appear sufficiently inadequate when it is realised that a most dis-

* Burckhardt (2), p. 76, pl. xii., figs. 3-6.

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 Lyc. and A. pumila 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 + 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 Hriphyla, 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 Hriphyla, 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 Hriphyla 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 Hriphyla in

dealing with certain Cretaceous forms of Astarte will be found above

in the discussion concerning the relationships of Astarte (Eriphyla)

herzogt.

A. pinchiniana ditters from immature examples of A. herzogi by the

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 A. pinchiniana the concentric surface

* Morris and Lycett (1), part iii., p. 83, pl. ix., fig. 8 (1855).

+ J. de C. Sowerby (1), vol. v., Tab. 444, fig. 2 (1824),

t Ibid., vol. vi., p. 35, Tab. 520, fig. 1 (1826).

The Invertebrate Fauna of the Uitenhage Serves. 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-Rhone, is of similar general type, but is truncated posteriorly.

Shells from the Gault of Cosne (Niévre) figured by de Loriol{ under

the name 4d. 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 dupiiana d’Orb.,t from the Albian of France, 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. noy.

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 umbonal 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), 8° Partie, p. 319 (1866); Pictet and Roux (1), p. 437,

pl. 32, fig. 5 (1852).

+ de Loriol (4), p. 94, pl. xii., figs. 1-7.

{ d@Orbigny (3), p. 70, pl. 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)

Tenet ata uanceeage eee eerie 15 ‘ 22 mm.

Greatest height, (...5252sasegss tees 8 : Ife

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 8. from the middle of Barkly Bridge, on the farm

Olifant’s Kop, Sunday’s River (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 pallial

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.

Anthonya cultrifornis 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.

Anthonya cornueliana (d’Orb.) | appears to approach more closely

in form, especially such a specimen as that figured by Miss Skeat ;

but A. lineata is less inequilateral and is not so obliquely truncated

posteriorly.

Genus TANCREDIA J. Lycett.

TANCREDIA 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 umbo 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, pl. xxx., fig. 236.

+ Woods (3), vol. ii., part 3, p. 130, pl. xix., figs. 4, 5 (1906).

t @Orbigny (3), p. 74, pl. cclxiv., figs. 7-9 (1844), described as Crassatella ;

Skeat and Madsen (1), p. 178, pl. vi., fig. 13 (as Ptychomya).

140 Annals of the South African Musewm.

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) (35)

HE ere lipas,. eetnckhdssme. at ese da eae 16 , 16 , 22 mm.

Height, measured from the umbo 10 : ia : 1 Roieee

Greatest depth of a single valve 4 : t ; Oss

Occurrence.—In the kloof east-north-east of Red House, on the left

side of the Zwartkop’s River (324); this form is also found in the

Marine Beds of the Sunday’s River, where it occurs associated in

the same hand-specimen with Trigonia vau, Trigonia conocardii-

fornuis, Acteonina 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 River

(95h).

Itemarks.—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. 1’. extensa Lycett,* from the Inferior Oolite, is very

* Lycett (1), pl. xi., fig. 9; Morris and Lycett (1), part iii., p. 93, pl. xiii., fig. 6

(1855).

The Invertebrate Fauna of the Uitenhage Series. 141

similar in outline, but differs in having the umbones 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. schwarz. In the African shell the umbonal angle is sharper

and the umbonal region more prominently projecting.

Tancredia 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 TZ. schwarz: to T. angulata

Lycett,{ from the Great Oolite, is very close, particularly in the

position of the umbones, 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 7.

schwarzi 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 Tancredia

reached its greatest development during Jurassic times, 7’. americana

Meek and Hayden { is known from Cretaceous strata on the Upper

Missouri and on Cache La Poudre River, in Colorado. The generic

position of 7’. americana has not been disputed, so far as I am aware,

but itis 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 Telline 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, pl. xiii., fig. 8 (1855).

+ Lycett (2), 341, pl. xiv., fig. 5; Morris and Lycett (1), part iii., p. 94, pl. xiii.,

fig. 9 (1855).

t Meek (2), p. 142, pl. 38, fig. 1.

§ Coquand (1), p. 101, pl. viii., figs. 9, 10. m

142 Annals of the South African Musewm.

Tancredia schwarzi, from which it differs by its greater elongation

and relatively reduced height; it has a more obtuse umbonal 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 Paleomya Zittel and Goubert,* Paleomya

deshayest 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. Palezomya autissiodorensis

(Cott.) de Loriol,t 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 THETIRONIA F. Stoliczka.

THETIRONIA PAPYRACEA (Sharpe).

Plate VII., figs. 11, lla.

1856. Ceromya papyracea D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

Voli vil, p. 195, spl. xxi, fies 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), pl. viii., figs. 6-8.

+ de Loriol and Cotteau (1), p. 510, pl. v., figs. 12-14.

t Skeat and Madsen (i), p. 129.

The Inwertebrate Fauna of the Uitenhage Series. 143

delicate, radial, linear stria, which at the lower margin of an adult,

and especially in the posterior half of the valve, may be separated by

spaces exceeding a millimetre in breadth. The granular ornaments

which occur on the striz# 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

strize 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)

Memes... ncns-nseriseneemsoero nese ess 22 25 : 47 mm.

Height measured from the umbo 19 é 22. : 30° |,

Greatest depth of a single valve.. 8 : 10 1A a

Number (3) is the specimen figured by SHarte!

Occurrence.—Collected in the railway cutting between milestones

241-943 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 markingsas 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 Kuropean 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 lable 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 levigata by Sowerby* seem to represent the same form as

that afterwards named Thetis major,t 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 nuinor 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 Roemer§ united these two forms under the name

Thetis sowerbyt. Forbes, again, divided Roemer’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 Thetis levigata 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 levigata (J. Sow.)—which

includes Th. major (J. de C. Sow.)—and Th. minor (J. de C. Sow.),

which includes Th. levigata (d’ Orb.)

* J. Sowerby (1), vol. iii., Tab. 209, figs. 1, 2 (1818).

+ 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

Thetiroma papyracea is distinguished from Th. levigata by its

greater inflation, its anteriorly situated umbones, its posterior trunca-

tion, and the absence of strong radial strie 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

weross the flank.

Thetironia reneviert (de Loriol),* from the Hauterivian of Sainte-

Croix and Mont Saléve, 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),+ which occurs in the

“Gault” of the Perte-du-Rhone, 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) | (Aptian) lacks the elongation and posterior truncation that

characterises Th. papyracea, and its linear markings on the cast

follow a widely different course.

Thetiroma sancte-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 7’. papyracea in the

general form and outline and in the position of the umbones is

* de Loriol (1), p. 65, pl. ix., fig. 11.

+ Pictet and Roux (1), p. 420, pl. xxx., fig. 2 (1852).

t Pictet and Campiche (1), 3° Partie, p. 205, pl. exii., fig. 6 (1865).

146 Annals of the South African Museum.

Th. caucasica (Kichwald),* 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 Zh. 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 Thetirona

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 ine. The genus has therefore been

erroneously held to be related to members of the Veneride, but

although its suggested affinity to the Lucinide 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 Thetironia 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 T'hetironia

resembles the Cardiidz in several respects, yet thejpoints of difference

are too great to allow of its being included in that family.” }

THETIRONIA OBLONGA Sp. Nov.

Plate VII., figs. 12, 12a, 120.

Description.—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

* Hichwald (1), p. 709, pl. xxvi., fig. 7; Anthula (1), p. 90, Taf. iv., fig. 6.

+ See remarks on this point by Miss E. G. Skeat; Skeat and Madsen (1), p. 177.

{ 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 strie,

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 .—

Wie ne bigs ceeetee aes eee oe see ce aroun este aia 25 mm.

Height, measured from the umbo......... dye

Greatest depth of a single valve ......... 9)

Occurrence.—Collected at a kloof, 8. 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 Musewm.

Genus TRAPEZIUM Megerle von Mihlfeldt.

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 shghtly convex outline, to the sharply

convex anterior border. The inferior margin gives a broadly convex

outhne, 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)

Lionel paneer one osmseeneaeace 9°5 ‘ 14 mm.

Height :..seccacsogstennes.cssonsets 8 : WAY

Depth of a single valve ......... — ee

Occurrence.—Found on the left side of the rece 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 sp. and Meretrix witenhagensis (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, pl. viii., fig. 8.

The Invertebrate Fawna of the Uitenhage Series. 149

and its carina falls more steeply and forms a line more nearly

straight. In Trapeziwm ? 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. Miiller* from the Lower

Cretaceous of German Hast Africa as Mactra stromert. 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 umbo in

Trapezium ? tatet 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 Trapeziwm 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 CYPRINA Lamarck.

CyYPRINA RUGULOSA Sharpe.

1856. Cyprina rugulosa D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

vOlenviie palo, pls xxi, tis, 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 River (325). Sharpe

recorded it from ‘Sunday River, in greenish shelly grit,’ and Stow

mentions its occurrence above the Modder Drift on the Sunday’s

* G. Miiller (1), p. 563, pl. xxv., fig. 15.

+ J. de C. Sowerby (2), p. 341, pl. xvi., fig. 9.

t Morris and Lycett (1), part ii., p. 75, pl. vii., fig. 8 (1853).

§ Fischer (1), fase. xi., p. 1075 (1887).

11

150 Annals of the South African Museum.

River, in association with the characteristic Trigoni@ of the Marine

Beds. In 1905 Mr. Rogers found this species in the cliff on Buck

Kraal, Sunday’s River (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’Orb.,* 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 Dacqué from strata of supposed Aptian age

in Somaliland;{+ but C. rugulosa is less strongly inflated, and

most examples are further distinguished by their rough surface

markings.

CYPRINA BORCHERDSI Tate.

1867. Cyprina borcherdsi R. Tate, Quart. Journ. Geol. Soe., vol. xxiii.,

p. L61, plicvill.; fie. 2.

Occurrence.—Found in the railway cutting between milestones

941-943 on the line from Uitenhage to Graaff-Reinet, about three

miles from Uitenhage (313). The specimens described by Tate were

also from the Zwartkop’s River.

A specimen obtained by Mr. Rogers 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. Rogers 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.

* @Orbigny (3), p. 100, pl. 272, figs. 3-6 (1844); Pictet and Campiche (1),

3e Partie, p. 224, pl. exv., figs. 1, 2 (1865).

+ Dacqué (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 swindonensis,

described by the late Prof. Blake * from the Portlandian Swindon

Sands of England.

Genus MERETRIX Lamarck (sensw lato).

MERETRIX UITENHAGENSIS sp. nov.

Plate VIL., figs. 14, 14a; VIII., figs. 1, 1a.

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 umbones 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 striz ; 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 stris 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, pl. x., fig. 2.

152 Annals of the South African Musewm.

Dimensions.— (1) (2) (3) (4)

Mengthy ace. ssdecaatase: 14 ; 16 : ky ; 20 mm.

Height, measured at the

middle of the valve 11 ; 12 : 13 : Gwaes

Greatest depth of a

single valve basse. 3 : 3 15 ae 3°Oune Anas

Occurrence.—Found in the kloof east-north-east of Red House, on

the left side of the Zwartkop’s River (324), and at Grass Ridge, three

miles east-north-east of Uitenhage (310) ; also in the railway cutting

between milestones 244-242 on the line from Uitenhage to Graaff-

Reinet, about three miles from Uitenhage (316). A specimen sent

from the South African Museum is from the Sunday’s River,

occurring with Holcostephanus cf. atherstoni (Sharpe). Mr. Rogers

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 River (24h, 26h, 28h—80h) ;

and from the highest beds in the kloof behind Colchester, left bank

of Sunday’s River (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 witenhagensis has a less steeply sloping upper margin

posteriorly to the umbones, and the umbones are less prominent.

The minute concentric linear striz 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)} 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, pl. ii., fig. 5.

+ J. de C. Sowerby (1), vol. vi., Tab. 518, figs. 4-6 (1826).

+ Leymerie (2), p. 5, pl.v., fig. 7; pl. 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 pictetc de Loriol,+

from the Gault of Cosne (Niévre), which, however, is not so elongated

relatively to height, and moreover, has a coarser concentric orna-

mentation. A similar resemblance is shown to Tapes patagontca

Stanton,{ 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 River), described by von Koenen§ as Cytherea

wohltmanm, differs from Meretrix witenhagensis 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.

* dArchiac (1), p. 303, pl. xiv., fig. 7. + de Loriol (4), p. 64, pl. vii., fig. 21.

t Stanton (3), p. 23, pl. iv., figs. 12, 13.

§ von Koenen (1), p. 36, Taf. iv., figs. 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. Psanumobia atherstoni D.. Sharpe, Trans. Geol. Soc. Lond.,

ser.2; vol. vile p. 196, piixxit.; fig: 11.

This was found in the cliff below the old school-house at Dun-

brodie, on the right bank of Sunday’s River (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 umbo. 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

Acteonina atherstoni, Turbo atherstoni, Pecten 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 Unvo.

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.

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

The Invertebrate Fauna of the Uitenhage Series. 155

Genus SOLECURTUS 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.

Fie, 1.—Solecurtus sp., nat. size. Survey Collection.

Dimensions.—

length) (if restored) .c20-..--es.-c.8e--s- 31 mm.

Height at the umbos:.2-25....20--...+63 1D nae

Depth of the valve, about ............ eae

Length of siphonal margin............ eas

Length of frontal margin ............ ay

Occurrence.—Found at Grass Ridge, 3 miles east-north-east of

Uitenhage (310).

femarks.—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.

*Horbes|(1) ps Zar, ple iis, fe. 1

156 Annals of the South African Musewm.

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. @qualis).| Guéranger has figured a shell from the Ceno-

manian of the Sarthe under the name Solecurtus equalis,} to which

our specimen appears to bear a close resemblance, so far as com-

parison is possible.

Genus MACTRA Linn.

Macrra? DUBIA sp. nov.

Plate VIIL., 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. From 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 puncte, only visible under a lens.

Dimensions.— (1) (2)

Ihengthy.. oscc.sassacseusemereeer sneer 15 ; 15 mm.

Height, measured from the umbo 12 : 1s

Greatest depth of a single valve 4 : 3°5,,

* d@Orbigny (3), p. 320, pl. 350, figs. 1, 2 (1845).

+ @Orbigny (3), p. 321, pl. 350, figs. 5-7 (1845).

+ Guéranger (1), pl. xv., fig. 3.

The Invertebrate Fauna of the Uitenhage Series. 157

Occurrence.—Found in the cliff on Buck Kraal, Sunday’s River

(141h).

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 defined 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) + 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 PLEUROMYA L. Agassiz.

PLEUROMYA BAINI (Sharpe).

Plate VIIL., figs. 4, 4a.

1856. Myacites ? bainit D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

VOl ovals LOO, pl exxil., fe 17.

Occurrence.—The locality given by Sharpe was merely ‘“ Sunday

River.’’ Specimens were collected by Messrs. Rogers and Schwarz

at Grass Ridge, 3 miles east-north-east of Uitenhage (317, 318,

334, 335), from an outcrop of nodular limestone, the highest fossil1-

ferous outcrop of this locality, where many characteristic fossils of

the Sunday’s River Beds were found.} P. baini was also found to

occur commonly in the railway cutting between milestones 243-243

on the line from Uitenhage to Graaff-Reinet. Mr. Rogers obtained

a very well-preserved specimen from the highest beds in the klootf

behind Colchester, Sunday’s River 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, pl. xvi., fig. 9.

+ Meek (2), p. 208, pl. xxx., fig. 7.

+ Rogers 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 Plewromya

lutraria (Krauss),* another characteristic shell of the Sunday’s

River 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. bane

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 situated a little nearer to the anterior

extremity, while the margin in front of the umbo slopes down more

suddenly than in P. bainz.

Pleuwromya 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

Plewromya 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. schréderi (Wollemann),§ from the Neocomian of North Germany.

To judge from Dr. Wollemann’s description and figure, P. schréderi

approaches very closely indeed to P. bain.

Another shell which shows considerable resemblance to P. baina

occurs in the Neocomian sandstone of the Teutoburger Wald, and

was ascribed by Weerth,|| though perhaps erroneously, to d’Orbigny’s

Panopea 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. Roemer { as Plewromya solenoides, is an elongated form with

* Krauss (2), p. 447, Tab. xlvii., fig. 1.

+ d’Orbigny (3), p. 329, pl. 353, figs. 3-8 (1845).

+ d’Orbigny (3), p. 333, pl. 355, figs. 3, 4 (1845).

§ Wollemann (1), p. 126, Taf. v., fig. 7 (as Panopea).

|| Weerth (1), p. 37, Taf. viii., fig. 7.

« F. A. Roemer (3), p. 330, pl. xli., figs. 20 21.

The Invertebrate Fauna of the Uitenhage Series. 159

straight upper margin and little-curved lower margin. P. baine

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 Plewromya, to which genus belong also, in all probability, the

majority of the Neocomian forms to which the name Panopea 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 Plewromya that the name proposed by Krauss

must be regarded as a synonym—a view already adopted by Zittel.}

Genus GONIOMYA J. L. R. 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 umbo 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). + Zittel (5), p. 125.

160 Annals of the South African Musewm.

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

22mm. 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. Rogers from the highest beds in

the kloof behind Colchester, Sunday’s River 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 Trigonie of the group of T. vaw

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

Trigome, 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 vaw 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 cost 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 Trigome

mentioned.

Genus THRACIA W. E. Leach.

THRACIA sp.

Plate VIIL,, fig. 5.

A single specimen, which must be ascribed to this genus, is

scarcely so well preserved or so strongly characterised as to allow

The Invertebrate Fauna of the Urtenhage 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. The umbones 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.—

WMenethvaresrsacesestesSec cas aradss seme seoseos 40 mm.

Height, measured from the umbo......... ZO,

Depihrotoneryaliyer-ncccscescetsn. odes tee oseee Sue

Occurrence.—In the railway cutting between milestones 244-242

on the line between Uitenhage and Graaff-Reinet, about three miles

from Uitenhage (331).

Remarks.—Comparison with English specimens of Thracia phil-

linsi Roem.,* 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 loriert. 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 phillipsi F. A.

Roemer, (2), p. 74, Taf. 10, fig. 1 (1841).

t+ Whiteaves (2), p. 140, pl. 17, fig. 7.

t Meek (2), p. 224, pl. 39, fig. 6.

§ Coquand (1), p. 100, pl. 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 (Rolling Downs Formation)

of Queensland, ascribed by Etheridge with some doubt to Thracva

primula Hudleston, and figured under the generic name Corumya.}

Less similarity is shown to the specimen originally named Thracia

primula,{ 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 GASTROCHANA L. Spengler.

GASTROCHEZNA DOMINICALIS Sharpe.

1856. Gastrochena donunicalis D. Sharpe, Trans. Geol. Soc. Lond.,

ser. 2, vol. vii., p. 198, pl. xxiii., fig. 4.

Sharpe described under this name a shell which had bored into

a fragment of wood,§ found on the Sunday’s River ‘near Enon.”

He also mentions the occurrence of Gastrochena boring into

fragments of wood and Trigonia-shell, also from the Sunday’s

River.

Fragments of fossil wood bored by Gastrochena were collected by

Messrs. Rogers and Schwarz from an oyster bed at the base of the

cliff below the old school-house at Dunbrodie, Sunday’s River (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 River below Blue Cliff station.

* VOrbigny (3), p. 380, pl. 372, figs. 1, 2 (1845).

+ Jack and Etheridge (1), p. 481, pl. 28, fig. 11.

+ Hudleston (1), p. 245, pl. 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

Crass GASTEROPODA.

Genus PATELLA Linnzus.

PATELLA CAPERATA Tate.

1867. Patella caperata R. Tate, Quart. Journ. Geol. Soce., vol. xxiii.,

p. 152, pl. vii., fig. 1.

Occurrence.—A single specimen, on Hxogyra 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 Rest” (Sunday’s

River).

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 Hxogyra. In

one place, on the surface of the Hxogyra, in the position which the

serrated margin of the Patella formerly occupied, there are indenta-

tions which correspond with the terminations of the cost on the

Patella.

Genus PLEUROTOMARIA 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 .—

Heights. 2. 2.3tnososteusercneneenaecececeessce 90 mm

Greatest width of base............ eaten 1LOO® .;;

Height of aperture: (inc.nescescoscestaace ay

Approximate breadth of aperture ...... ON

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 Plewrotomaria (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 Plewrotomaria

tardensis Stanton,{+ 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 TURBO Linneus.

TURBO ATHERSTONI Sharpe.

Plate VIII., figs. 6, 6a, 7, 7a, 7b.

1856. Turbo atherstoni D. Sharpe, Trans. Geol. Soc. Lond., ser. 2,

vol. vii., p. 200, pl. xxviii., figs. 15, 16.

1856. Turbo baini D. Sharpe, Ibid., p. 200, pl. 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.

+ Stanton (3), p. 29, pl. vii., figs. 1, 2.

The Invertebrate Fauna of the Uitenhage Series. 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.

Diameter of the fourth whorl.................s00.00. 4...

{ Height of a specimen with five (or six ?) whorls 14 _,,

( Diameter of the body-whorl....................000800 oe

Occurrence.—Found in the cliff below the old school-house at

Dunbrodie on the Sunday’s River (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 Ostree from Sunday River,” as recorded by Sharpe. This form

was obtained by Mr. Rogers in 1905 in a cliff W. 20S. from Comley’s

house, right bank of Sunday’s River (95h).

femarks.—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 atherstoni 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 7’. atherstont, 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 7. bainti 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

reedt 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 TZ. 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 Loriére. +

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 Aucella-beds of Novaya Zemlya, and

was ascribed by Tullberg to Minster’s Turbo capitaneus.t This has

ornaments similar to those of 7. 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.