Petre Le
oes ae)
CCE ey
Pe Oe,
carte
=
+
iT
eens
rer fetitaele®
Seen
2
seers
“+8!
Tats
5 ay
ena
ise atti
2 Herat aye
ast SHE et
ees hee
Petit Telit tates,
tp abet eT titthne
Apart
ave ye
mee
65
* 7 ¢ © \7
a) te ete G soa te
RS Pae iarstetresetts
islet Nate
Prcely +
MN
pe itaets
Ct ie Fae kD
Site Veco Ok Se
*
S
¢ a
Sete
‘ats
See
Rute
Gilde ak
Ae bare
es
SS
\\ |
Neo, fs
SEMIO de ae
JOM BUEN
[s GENER a’
BULLETIN OF ( 2 5 smunes)
THE BRITISH MUSEUM pee
(NATURAL HISTORY)
GEOLOGY
VOL. 23
1973
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1981
DATES OF PUBLICATION OFTHE PARTS
No. 1 . 2 March 1973
No. 2 . 8 March 1973
No. 3 29 June 1973
No. 4 31 May 1973
No. 5 12 July 1973
No. 6 13 July 1973
No. 7 20 July 1973
ISSN 0007-1471
PRINTED IN GREAT BRITAIN
BY HENRY LING LIMITED, AT
THE DORSET PRESS, DORCHESTER DTI IHD
CONTENTS
GEOLOGY VOLUME 23
Marsupialia, Insectivora, Primates, Creodonta and Carnivora from the
Headon Beds (Upper Eocene) of southern England.
P. E. CRAY
Some British Cretaceous gastropods belonging to the families Procerithi-
idae, Cerithiidae and Cerithiopsidae (Cerithiacea).
H. L. ABBASS
A review of some English Palaeogene Nassariidae, formerly referred to
Cominella.
C. P. NUTTALL & J. COOPER
Palaeozoic coral faunas from Venezuela, II. Devonian and Carboniferous
corals from the Sierra de Perija.
C. T. SCRUTTON
Prodeinotherium from Gebel Zelten, Libya.
J. M. HARRIS
Cirripedes from the Upper Cretaceous of Alabama and Mississippi,
eastern Gulf Region, U.S.A.
I. Palaeontology. J. S. H. CoLLINs
II. Geology. F. F. MELLEN
Fenestrate bryozoa from the Viséan of County Fermanagh, Ireland.
R. TAVENER-SMITH
Index to Volume 23
PAGE
103
ET
221
283
349
381
389
495
i pedis. hier? ae
Son paidaiey' | a a
+¢@a>? 2
) a '
5 ty oe > ew Al
‘ : ~~ 2° i
sv ev Le wTiot 4. -..
eee ee |
ftpmect
any? «
y neg?
7
'@
) Sani ok
INDEX TO VOLUME 23
New taxonomic names and the page numbers of the principal references are printed in bold type.
An asterisk (*) denotes a text-figure.
AsBass, H. L. 103-75
Acanthocladiidae 483-90
Actinostoma 409, 411
Acinophyllum stramineum 231
vermetum 230-1
Adapidae 65-6
Adapis 65, 66-73
magnus 16, 66, 67-71, 72-3; pl. 1; pl. 2. fig. 1
minimus 66
parisiensis 27, 66, 71-3; pl. 2, figs 2-3
priscus 66, 71-2
ruetimeyeri 66, 71-2
sciureus 66, 71-2
Adapisorex 38-9
Adelomys depereti 12, 16
siderolithicus 12, 27
Adunator 39
Africa 291
Afrocyon burolleti 288
Ageria 106, 110, 156-60, 169
? angustata 57-8; pl. 8, fig. 13
costata 108, 158, 159; pl. 8, figs 7-12
gaultina 108, 148, 156, 157-9; pl. 8, figs 1-6
Agerina 74
Albian 106-8, 114, 119, 123-4, 127-8, 130, 133-4,
137, 155, 157, 159-60, 167
Alum Bay 18-19, 24, 199
Amphichiromys europaeus 63
Amphilemur 66
Amplexizaphrentis 239-42
cassa 242
sutherlandi 224, 230, 232, 239, 240-2, 241*,
281; pl. 1, figs 9-13
Amplexus intermittens 234
Anchilophus sp. 16
Anchomomys 65-6
Anchura carinella 124
Ancillaria bretzi 208
Andagularia 142
Androconus 48
Annona 351, 358-9, 379, 381-2
Anomia 409
Anoplotherium 71
anthracotheroid incertae sedis 16
Anthracotherium 15
Apatemyidae 61—5
Apatemys 61-2
bellus 63
Aphanolemur 66
Apheliscinae 47
Aphronorus 48
appendicular skeleton 330
Aporrhaidae 107
Aptian 106-9, 111, 113, 118, 120, 122-3, 126-7,
132-3, 138, 162, 165
Aquitanian 288
Archaeofenestella 461
Arcoscalpellum 362-74, 381
bakeri 351-2, 367-8, 381; pl. 3, figs 10-13
campus 351-3, 369-71, 373, 381; pl. 3, figs
14-16; pl. 4, figs 1-4
comptium 371
conradi 352
fossula 353, 366-7, 371, 373
gracile 373
hubrichti 351-3, 357, 363-7, 365*, 370, 373,
381-4; pl. 2, figs 13-20; pl. 3, figs 1-9
maximum 353, 367
michellotianum 363
simplex 362
solidulum 368
sulcatum 367
velutinum 363
withersi 351-3, 366, 370, 371-4, 381; pl. 3, figs
17-18; pl. 4, figs 5-15
Arctica morrisi bed 203, 214
Artiodactyla 16
Arvaldus 44-5, 46-7
stintoni 27, 34, 45-7, 46*
Asia 84, 89-90, 134, 291
Aston Clinton 136
astragalus 295, 340-1
496 INDEX
atlas vertebra 290, 319-21
Atherfield 107, 113, 131-2, 142, 144, 146, 162,
165
Austinian 381
Australia 180
Austria 152, 184
Auversian 183-4, 193, 208
axial skeleton 319
axis vertebra 290, 322-4
Babylonia (Peridipsaccus) archambaulti 205-6;
pl. 5, fig. 8a, b
Barton Beds 182, 186-8, 190-1, 199-201
Bartonia 181
canaliculata 181, 182*, 183, 192
Ballyshannon Limestone 393
Barytherium 344
Barytichisma 239
Bathonian 150
Bathraspira 110, 123, 124-30, 169
beaudouini 126
brightoni 108, 123, 129-33; pl. 3, figs 9, 11, 14,
15
cleevelyi 108, 124, 125*, 127-8; pl. 3, fig. 10
ervynym 129
excavata 127
fouadi 108, 125*, 126, 128-9, 130; pl. 3, figs
7-8, 12-13
multistriata 126
neocomiensis 123-7, 129
ornata 126
shanklinensis 109, 124, 126-7; pl. 3, figs 3-6
subornata 127
tecta 108, 123-6, 125*, 127, 129; pl. 3, figs 1-2
Batillaria concava 22
Belgium 62, 138, 161, 184, 188, 193, 205-6, 212
Benbulben Shales 393
Bessoecetor 48-50
levei 48, 50
Billingsastraea 256, 263
Billingsastraeinae 256-64
bivalves, see lamellibranchs
Blackdown 107, 130, 148, 157, 159-60
Bognor Rock 188, 203
Bordeaux 188, 212
Bowenelasma 231, 242-3, 244-8
breviseptata 224, 230, 245-7, 280; pl. 3
typa 224, 230, 243-5, 253, 280-1; pl. 2
brachiopods 228
Brachylepadidae 380
Brachylepadomorpha 353, 380
Brachylepas 380
angulosa 351-2, 380, 381; pl. 5, fig. 16
niassanti 380
Brachysphingus 180-1, 183, 208-9
gibbosus (208); pl. 8, figs 3-4
Bracklesham Beds 186-8, 193
Bramshaw 187, 193-4
Briantelasma 231, 243, 247-51, 280-1
americanum 248
oliveri 224, 230, 248-51, 249*, 253, 280-1; pl. 4
Brightonella 123
sandlingensis 123
turris 123
British Museum (Natural History) 183, 186, 188,
196
Brockenhurst Bed 187-8, 191, 193, 197, 205,
210-11
Brontops 341
Brook 193-4, 211
Brotia 138
bryozoa 398, 409
Buccinacea 179, 189-215
Buccinanops 188; see Bullia
Buccinidae 179-83
Buccinum acies 201
armata 196
auversiense 208; pl. 7, figs 6, 7
bicorona 212-3, 215
bolli 212
bullatum 187, 193
aspera 189
canaliculatum 180-1, 192
cassidaria 212
cinctum 180
desertum 180, 183, 189-90, 192, 198
desori 201-2
excavatum 190
flexuosum, see Cominella
fusiforme 183, 192-3, 195; pl. 3, figs 7, 8
fusiopsis 193, 195-6
gossardi 180, 212
latum 201
lavatum 192
montense 180, 183-4, 215; pl. 9, fig. 8a, b
ovatum 201
porcatum 180
quoesitum 213-5
semicostata 196
solandri 198; see Cominella solanderi
suturosum 204; pl. 6, fig. la, b
testudineum 180
uniseriale 212
ventricosum, see Cominella
sp. nov. 198
Buccitriton 180, 213
Bullia 181, 188
(Buccinanops) monilifera 181
(Molopophorus) striata 208
Bundoran Shale 393
Bunolistriodon massai 288
Burdigalian 184, 212, 288
Burham 136
Burnupena 180
cincta 180
Buxolestes 48, 50
Caenopithecus 65-6
Calantica saskatchewanensis 352
(Titanolepas) martini 352
Calcaire Grossier 188
Calenscio Serir 286
California 180, 208-9
Callista kickxi horizon 205
Calp Sandstone 393
Campanian 111, 117, 351, 379, 381-2
Campanile 109, 138, 150, 169
breve 152
carezi 152
cenomanica 108, 151-2; pl. 6, figs 1, 2
giganteum 152
morgani 152
Campanilinae 151
Campo Chico Formation 226
Cano Colorado 240, 281
Cano del Noreeste Formation 229
Cano del Norte 227, 252, 281
Cano del Oeste 226, 281
Cano del Sur 280
Cano Grande Formation 226, 280
Cano Indio Formation 229
Cano Los Guineos Formation 228, 280
Cano Pescado 281
carina 354-9, 363-4, 367-75, 378, 380; pls 1-4
Carnivora 16, 89-93
Carrick Lough 392, 394, 397, 399-400, 404
Catodontherium sp. 16, 23
Cebochoerus 12, 16
INDEX
Cenomanian 108, 111, 115, 119, 121, 136, 138,
140-1, 151, 155
Centetodon 44-5
Cerithiella 160, 165, 167-8
atherfieldensis 109, 161*, 162; pl. 6, fig. 10
cloacina 161*, 162
devonica 108, 160-2, 161*; pl. 6, fig. 9
metula 161*
nodoliratum 160, 161*
prealonga 162; pl. 6, fig. 8
semirugatum 161*
Cerithiidae 150-9
Cerithiopsidae 160-7
Cerithiopsis 146
meeki 164
quadrilirata 164
Cerithium 118, 122
aculeatum 109, 112-3
aptiense 133
ascheri 109, 138-40
attenuatum 109, 113-4
belgicum 138, 140
bicostatum 161
binodosum 110, 157-8
chargense 111
cingillatum 152-3
clementinum 109, 141-3, 145
cribriforme 166-7
cuckhamsliense 109, 116
davonstianum 110
delpeyae 109, 137
derignyanum 135
(Fibula?) detectum 122
disparile 133
distinctum 149
dupinianum 145
ervynum 124
excavatum 115
fertile 137
forbesianum 109, 141, 143-5
frickei 109, 114
gallicum 109
gottfriedi 109, 113
gurgitus 125
hector 110, 157
hispidulum 109
inauguratum 109
kaunhoweni 149
leufroyi 165
metula 160
millepunctatum 141
mosense 109, 135-6
navasi 110, 154
neocomiensis 130
ornatissimum 135-7
peregrinorsum 146
Phillipsi 110, 143-5
pseudoclathratum 154
pustulosum 110, 153-4
rhodani 135
ricordeanum 109, 139
rochati 115
sabandianum 137
sanctaecrucis 109, 139
saundersi 110, 153
schrammeri 126
subattenuatum 113-4
497
498 INDEX
subspinosum 118-9 bicoronata 214
terebroides 109, 139 bonnecarrei 201
tectiforme 166 canaliculatum 181
tectum 123 deserta 180-1, 190, 193, 195-6
trimanile 133 solanderi 192, 198
tuberculatum 133 ‘ desori 202
turriculatum 109, 131, 133 flexuosa 208, 210
Cervella cervoidea 435 gossardi 180
cervical vertebra 290, 325-6 montense 185
Chadronia 48 porcatum 180
Chalons-sur-Vesle 188 praecedens 212
Chattian 184 solanderi, solandri 189, 198, 200
Cheilostomata 402, 459 supracostata 201
China 65 suturosum 204
Chinji 305 testudinea 180, 182*; pl. 9, fig. 1la, b
Choeropotamus depereti 16 ventricosa 204, 206; pl. 6, fig. la, b
Cimolithium 109-10, 138-41, 169 (Buccinum) flexuosa 210
ascheri 109, 138-40; pl. 2, figs 6, 7, 9, 10 solanderi 198
belgicum 140-1 ventricosa 204
aff. eleanorae 108, 138, 140-1; pl. 2, fig. 5 Cominellidae 179, 181
gallicum 138 Compressiphyllum 251
hispidulum 138 Condranema 471
insuguratum 138 Coon Creek 352, 383-4
komarense 140 Cooper, J. 177-219
miyakoense 139-41 corals, Middle Devonian 224
Cirripedia 354 Lower Pennsylvanian 224
Cirsocerithium 110, 118-9, 120-2 Corbula bed 203, 214
aptiense 121 Cornulina minax 196-7
gauthieri 119 Cray, P: E. 1-102
harborti 120-1 Creodonta 16, 26, 83-9
kirkaldyi 109, 118, 120-1; pl. 2, fig. 8 Cretiscalpellum 353, 354, 381, 384
nooryi 108, 119, 121-2; pl. 2, figs 3, 4 glabrum 355
peroni 119, 121 harnedi 351-3, 355, 357-8, 381; pl. 1, figs 10-12
reticulatum 118 macrum 351-3, 354-5, 357-8, 381; pl. 1, figs 1-6
subspinosum 108, 118, 119-20, 121-2; pl. 2, striatum 357
figs 1, 2 subcarinatum 357
Clava 122, 153 unguis 354-62
Coffee Formation 351, 358-9, 379, 382 vallum 351-2, 357, 358-62, 361*, 381-4; pl. 1,
CoLuins, J. S. H. 349-80 figs 14-22; pl. 2, figs 1-12
Colombia 231 venustum 351-3, 355, 356-7, 381; pl. 1, figs
Columella 179, 181, 182* 7-9, 13
Colwel] Bay 197-8, 210 Cryptaulax 156
Colwellia 180-1, 208-9, 212 angustatum 158
antiquata 184, 208; pl. 7, fig. 11 Cryptopithecus 48-50
auversiensis 184, 208-9, 211; pl. 7, figs 6, 7 sideroolithicus 49
bretzi 184, 208; pl. 7, figs 8, 9 Cryptostomata 409
cretacea 184, 208 Cuisian 184, 201, 204
flexuosa 184, 208-9; 210-2; pl. 7, figs 1-5 cuneiform 290, 295, 330, 336-7
tejonensis 184, 208 Cyathaxoniicae 233
Cominella 179-81, 182*, 183, 190, 209, 215; see Cyathophyllidae 255-64
Northia Cyathophyllum 257
annandalei 212 goldfussi 269
aturensis 212 limbatum 269
bicorona 214 (Peripaedium) turbinatum 257
venezuelense 252-4, 257
Cylindrophyllum 224, 256, 262-4, 262*
elongatum 230, 232, 262*, 263-4, 281; pl. 7,
figs 3-6
Cyllena 188, 209
Cystiphyllidae 269-72
Cystiphyllina 269-72
americanum 269-70
conifollis 272
Czechoslovakia 157-8
Dacrytherium ovinum 16
Dalmanophyllum dalmani 243
Danian 134
Dartry Limestone 393
Dasyurodon 84
Deinotheriidae 285, 291, 344
Deinotherioidea 291
Deinotherium 285, 292-3, 345
bavaricum 289, 292, 294-5
bozasi 291-3, 296, 312, 345-6
cuvieri 296
giganteum 286, 289, 291-3, 296, 305-6, 308,
312, 337-9, 341-5
gigantissimum 293, 329, 342
hobleyi 292, 296
indicum 291-3, 345
levius 312
pentapotamiae 292
Delaware Basin 399
dentition 290, 295, 313; pls 4—5
Derrygonelly 392
Desorinassa 180-1, 201-2, 208-9, 213
acies 184, 201, 203-4; pl. 5, fig. 5
bonnecarrei 184, 201
desori 184, 201, 202-3, 204; pl. 4, figs 4-7
lata 184, 201; pl. 4, figs 8, 9
ovata 184, 201, 203-4; pl. 5, fig. 4
supracostata 184, 201
williamsi 180, 184, 188, 201, 203-4; pl. 5, figs
1-3
Devonian 229
Lower 231, 251
Middle 224, 231, 233, 248, 251-2
Diatinostoma 150
Dichobune 13
Dichodon 15, 19-20
cervinum 12, 16
cuspidatus 15-16, 21, 27
Didelphidae 27, 28-36
Didelphidectes 28
Didelphinae 28
Didelphis 29
Didymictis 90
INDEX 499
Didymoconus 49
Dinantian 393
Diplocynodon 25
Diploschiza melleni 381
Disphyllidae 255
Ditoecholasma dalmani 243
Ditretus 150
Dor el Talha 289
Dorsanum 185, 188
Dover 135, 155
Durhamina 224, 230, 232, 265-8, 281
cordillerensis 268
sp. nov. 266-8; pl. 8
? sp. 265-6; pl. 7, figs 7-8
Durhaminidae 264-5, 266-8
Dyspterna 48-50, 53, 57-61
helbingi 57
hopwoodi 26-7, 57, 58-61, 59*, 60*
woodi 57-9
Egypt 134
Eifelian 231
Elephas 285, 295, 305, 329, 336, 338, 342, 344-5
maximus 286, 289, 295, 330, 331*, 332, 334-43
Emmonsia 272
carmani 274
radiciformis 274
Emsian 231
Engelhardtia 25
Enniskillen 392
Enniskillenia 239
Eocene 180
Eochiromys 62
Eodevonaria 231
Epapheliscus italicus 57
Eppeslheim 305
skull 306-8, 310
Europe 28, 36, 48, 62, 89-90, 138, 179-80, 189,
291
Exechocirsus 110, 152-6, 169
pustulosus 154; pl. 7, figs 6, 7
saundersi 107-8, 153-5, 169; pl. 7, figs 3—5
aff. saundersi pl. 7, figs 1, 2, 8, 9, 12-14
aff. subpustulosus 108, 155-6; pl. 7, figs 10, 11
Exelissa 169
Favosites 224, 230, 272, 273-5
arbuscula 230-1, 274-5, 280; pl. 10, figs 3-6
digitata 275
gothlandicus 272
venezuelensis 230, 273-4, 280; pl. 10, figs 1-2
Favositidae, Favositina 272 :
femur 290, 295, 330, 339-40
500 INDEX
Fenestella 401-2, 409-10, 424, 440, 459, 462, 465,
472-5, 482, 484
albida 477
cf. albida 406, 451-3; pl. 14, figs 1-6
alternata 477
antiqua 410
cf. arthritica 406, 431-3; pl. 7, figs 8-9; pl. 8,
figs 1-4
benskiensis 470
binodata 468, 470
cerva 448
colymaensis 470
compressa 430-1, 433, 443
cornuta 435
crassa 409-10
crockfordae 450, 462
delicatula 406
cf. delicatula 407, 455-7, 469; pl. 15, figs 5-8;
pl. 16, figs 1-3
demaneti 430-1
eichwaldi 450
ejuncida 428
fanata carrickensis 406-7, 436-8; pl. 9
fenestratum 413, 415
cf. filistriata 406, 444-6; pl. 11, figs 5-10; pl. 12,
fig. 1
frutex 406, 411-3, 416; pl. 1
funicula 443
cf. funicula 406, 441-4; pl. 10, figs 7-10; pl. 11,
figs 1-4
ghzelensis 443, 448
gracilis 462
hemispherica 406-7, 421-3, 425; pl. 4, figs 7-9;
pl. 5, figs 1-4
iowensis 427-8
irregularis 406, 460-2; pl. 17, figs 2-7
ivanovi 406-7, 413-5, 466-7; pl. 2
limbata, limbatus 411, 415
lyelli 411
matheri 413
milleri 410
modesta 406, 418-21; pl. 4, figs 1-6
multiporata 457, 465
multispinosa 406, 416-8; pl. 3
var. karakubensis 418
ninae 466
nododorsalis 433
oblongata 406, 453-5; pl. 14, figs 7-8; pl. 15,
figs 1-4
oculata 470, 472
parallela 406, 423-5; pl. 5, figs 5—9
placida 444
plebeia 406, 410, 428-31, 441; pl. 7, figs 1-7
var. longifenestrata 430
plebeja 428
plummerae 466-7
polyporata 406, 457-60, 462, 465; pl. 16, figs
4-8; pl. 17, fig. 1
praemagna 406, 433-6, 448; pl. 8, figs S—11
praerhomboidea 466-7
prisca 410
pseudovirgosa 406, 448-51; pl. 13
quadradecimalis 462
regalis 450, 477
reticulata 410
rudiformis 427-8
rudis 426-7, 430-1
major 427
multinodosa 406, 425-8; pl. 6
sevillensis 418
cf. spinacristata 406, 439-41; pl. 10, figs 1-6
subantiqua 409-10
subspeciosa 406, 446-8; pl. 12, figs 2-8
tenax 407
tenuifila 421
tenuivirgata 457
triserialis 470
varicosa 473, 475
varifenestrata 436
virgosa 448
Fenestellidae 409, 483
Fenestrella 409
Fenestrellina 409-10
multispinosa 416
Ferruginous Sands 107, 120, 122, 126
Fibula, see Cerithium detectum
fibula 290, 330, 340
Fistulipora 420, 440, 462
Floresta 231
Folkestone 107, 114, 119, 124, 128, 130, 133-5,
BIG IST, WG
Fort Benton 352
France 5, 62, 65-6, 71, 74-5, 80-2, 84, 90-1, 119,
124, 130, 133, 136, 146, 157, 183-4, 192-4,
202, 212, 214-5
Fusus desertus 190, 193
Galba spp. 24-5
Galethylax 48
Gebel Zelten 285-6
Geolabis 44
Germany 48, 63, 74, 80, 91, 119, 124, 126, 133,
136, 138, 184, 187, 205
Gesneropithex 65—6
Glass Mountains 403
Glencar Limestone 393
Goniacodon 49
Gorgonia antiqua 410
INDEX 501
Grewinkia buceros 243
europheum hosholmensis 243
Guildford 132
Gymnocerithium 122
Gymnophyllum wardi 237, 239
Hadrophyllidae 237
Hadrophyllum 230, 232, 237-9, 238*, 281
orbignyi 237
sp. 237-9, 238*; pl. 1, fig. 8
Hamilton Group 230-1
Hampshire Basin 180
Haplaletes 49
Haplobunodon lydekkeri 16, 27
sp. 16, 20
Hapsiphyllidae 239
Harris, J. M. 283-348
Headon Beds 186-8, 191, 193, 205-6, 210, 212
Heliophyllum 256-63
halli 227, 230-1, 253, 256-60, 262*, 280-1; pl.
5, figs 6-8; pl. 6, figs 1-6
wellsi 224, 230, 261-2, 262*, 280; pl. 6, figs
7-8; pl. 7, figs 1-2
Hemicerithium 118
Hemipsalodon 84
Hemitrypa 435, 440, 475-8
burulica 478
hibernica 406, 435, 475-8; pl. 22
oculata 475
reticulata 475
Herne Bay 214
Herpetotherium 28
Heterochiromys 63
fortis 63
gracilis 63
Heterohyus 12, 62-5
armatus 62-5
europaeus 63-5
heufelderi 63-5
nanus 62-5
quercyi 62-5
Heterophrentis 224, 243, 251-5
inflata 253
prolifica 253
simplex 230-1, 253-5, 280; pl. 5, figs 3-5
spissa 253
venezuelensis 227, 230, 252-3; pl. 5, figs 1-2
Highcliffe 199
Hitchin 116, 154
Holland 184, 205
Hordle 187
Howenegg 286, 289, 343
Humbertia 90
angustidens 9
hastingsiae 93
helvetica 90
humerus 290, 295, 330, 332, 334
Huntingbridge 183, 193-4
Hyaenodon 13, 84-9
aimi 15, 85-7
‘hantonensis’ 85
leptorhynchus 84
cf. minor 16, 27, 85-9; pl. 5
Hyaenodontidae, Hyaenodontinae 83-9
Hyainailouros 84
Hyopsodontidae 47
Hypacodon 45
Ictidopappus 90
India 137
Insectivora 16, 36-65
Todictyum 476
Ischnognathus 84
Isle of Wight 107, 120, 122, 127, 131, 144, 146,
165
Japan 118, 127, 133, 138, 209
Jepsenella 61
jugal 301
Kansas 362, 400
Keepingia 180-1, 188, 201, 209, 211-2
annandalei 184, 212
aturensis 184, 212
bolli 184, 212; pl. 8, fig. 8
cassidaria 184, 188, 211-2; pl. 8, fig. 9
cancellata 211
gossardi 184, 188, 211-2; pl. 8, figs 5-7
praecedens 184, 212
tarbellica 184, 212
uniseriale 184, 212
Keriophyllum dahlemense 257
Kiaerophyllum semilunatum 243
Kindle Formation 242
Kotyhaza 294, 343
Labidolemur 61-2
lacrimal 301
lamellibranchs 398, 409
Lantianus 65
Latdorf 193, 205-6
Lattorfian 184, 187, 205, 212
latus, carinal 359-60, 363-4, 366, 369-71, 373,
375-6; pls 2-5
inframedian 358, 362; pl. 2
rostral 358, 362, 371, 373, 375-6; pls 2-3, 5
subcarinal 358-60; pl. 2
upper 358-60, 363-4, 366, 373, 375; pls 1, 3-4
502
Leakitherium 84
Leighton Buzzard 108, 124, 127
Leitrim 393
Lepadomorpha 354
Lepidosteus 14
Leptacodon 36, 38, 44
Jepseni 37
ladae 37-8
minutus 37
tener 37-8, 44
Leptocoelia 231
Levifenestella 463-5, 482
maeve 463
undecimalis 406, 463-5; pl. 17, figs 8-9; pl. 18,
figs 1-5
Lewes 136
Libya 286
Limnaea Marl 8, 10
Lindstroemiidae 233
Litomylus 49
London Clay 188
Lonsdaleia cordillerensis 265
Loricultina texanum 352
Lough Erne 393
Lough Melvin 393
Loxodonta 341
africana 286, 289, 319, 321*, 323*, 324-6, 326*
lunar 337
Lutetian 183-4
Lyndhurst 187, 197
Maastrichtian 111, 122, 149, 161, 166, 351, 353,
357, 367, 371-2, 374-5, 378-82
Maboko Island 342
Madagascar 119, 124, 134, 137
Mainz Basin 188
magnum 295, 337-8
mandible 290, 311; pls 4-5
Manitoba 352
manus 295
Manzati 342
Marada Formation 287
Marisastrum sedgwicki 256
Marmosa 34
Marsupialia 16, 26-36
Mastodon 316, 327, 336, 338, 344
angustidens 289, 295, 319, 333*, 334-5, 337-41,
343-4
Pygmaeus 288
maxilla 290, 300; pl. 5
Megalopterodon 84
Melania scalariformis 141
Melanopis attenuatum 114
MELLEN, F. E. 381-8
INDEX
Melongena (Cornulina) minax 196-7
Merida Andes 229
Merstham 136
Mesalia (Bathraspira) tecta 123
Mesestoma 157
Messelina 38
, metacarpals 295
Metacerithium 109-10, 131-6, 169
abictiforme 134
amudasiensis 134
dentatum 137-8
ornatissimum 108-9, 134-5, 135-7; pl. 4, figs
4-5, 11-12
renngarteni 137
rikuchuense 132-3
stoliczkai 134
trimonile 108, 131-2, 133-5, 137; pl. 4, figs 1-3,
6-8, 10
turriculatum 107, 109, 131-3, 137; pl. 5, figs
8-11
sp. nov. 108, 137-8; pl. 4, fig. 9
Metachriacus 49
Metapterodon 84
Mexico 140
Miacidae 89-93
Microchoerinae 73-83
Microchoerus 13, 21-6, 29, 74-80, 81
edwardsi 75, 80
erinaceus 12, 16, 23, 74, 75-80; pls 3-4
ornatus 75, 80
Mimotricentes 49
Minilya 465-72
binodata 406, 468-70; pl. 19, figs 5-7; pl. 20,
fig. 1
duplaris 465
mimica 468
ninae 467
nodulosa 406-7, 470, 490
oculata 406, 470-2; pl. 20, figs 2-8
plummerae 406, 466-8; pl. 18, figs 6-8; pl. 19,
figs 1-4
praerhomboidea 467
rhomboidea 468
Miocene 287, 291-2, 296
Mississippian 242
Molopophorus 180-1, 183, 209
antiquatus 209; pj. 7, fig. 11
cretacea 209
gabbi 180
striatus 180-1, 208-9; pl. 8, figs 1-2
tejonensis 208
Montian 215
Mooreville, Middle 381-2
Munich skull 308-10
Myolestes dasypelix 37, 44
Myrica 25
Nannopithex 74, 81
Naples, Bay of 182
Nari Stage 184, 212
nasal 301
‘Nassa’ 183; see Nassarius
antiquata 208-9
cretacea 208-9
tarbellica 212
Nassarlidae 177-219
Nassarius 188, 190, 213
arcularius (181); pl. 9, fig. 9
Nassidae 180; see Nassariidae
Nasua 61
Navarroan 381
Necrolemur 74-5, 81
antiquus 75
parvulus 80
raabi 80
Neocomian 109, 143, 159
Neogastropoda 189-215
Neogene 296
Nerinaea 153
Nerineopsis 109, 110-8, 169
aculeatum 109, 112-3, 115; pl. 1, figs 5-8
adeli 108, 111, 116, 117-8; pl. 1, fig. 4
attenuatum 113
claxbiensis 109, 111; pl. 1, fig. 3
coxi 108, 114-5; pl. 1, figs 10-11
cuckhamsliensis 107-8, 111, 116-7; pl. 1, figs
3-4
davonstiana 110-1, 114
gottfriedi 115
longi 115
melburiensis 108, 111, 115—6, 117; pl. 1, figs 1-2
subattenuatum 109, 111, 113-4, 115; pl. 1, figs
9, 12
Nettleton 111, 147, 159
Neurogymnurus major 49-SO, 57
neuropophysis 290
New Forest 186
New Jersey 352
New York 230
New Zealand 179-80, 182-3
Ngorora 296
North America 28, 36, 48, 61-2, 84, 89-90, 184
Northern Ireland 107, 117
Northia (Cominella) angusta 185
bullata f. armata 196
suturosa 205
Nudivagus 110, 122, 123
? cooperensis 122
INDEX 503
morrisi 109, 122-3; pl. 2, figs 11-12
simplicus 122
Nummiulites elegans 199
NUTTALL, C. P. 177-219
Nyctitheriidae, Nyctitheriinae 36-47
Nyctitherium 36-9, 44
daspyelix 37
serotinum 37
velox 44
Oligocene 180
Oliva 209
Onesquethaw 224, 230-1, 236, 252
Onondaga Limestone 230-1, 251
Ontario 231
Opsiclaenodon 48-57, 58
major 12, 16, 49, 50-7, 52*, 53*, 54*, 55*,
57-61
Oregon 180, 208
Orthochetus 165-8; pl. 6, fig. 15
charlesworthi 166*, 167
elongatus 166*, 167
hantoniensis 109, 165-7, 166*; pl. 6, fig. 12
helmyi 108, 167-8; pl. 6, figs 13-14
leufroyi 166*, 167
mapeulensis 166*
tectiformis 166*
Oxyclaenus 49
Pagonomus 48, 50
Pakistan (West) 184, 212
Palaeocene 180, 352
Palaeocoryne 435, 443, 459
Palaeogene 177-219
Palaeosinopa 50
Palaeotherium 13, 15-16, 19, 21, 26-7
medium 15
muehlbergi muehlbergi 27
praecursum 16, 23, 27
palatine 302
Palencia 307
skull 309-10
Palmarito Formation 228-9
Paloplotherium 13
annectens 15
Paludina lenta \7
palynomorphs 229
Pantolestes 48, 50
Pantolestidae, Pantolestinae 47-61
Pantomimus 48, 50
Paracerithium 110, 118-9, 121, 131, 142
parietal 302
Paris Basin 187-8, 194, 204
pelvis 339
504
Penniretepora 440, 489
Pennsylvania 402
Pennsylvanian, see corals
Pentagonia 231
Peratherium 12, 16, 23, 27, 28-36, 30*, 31*, 35*
crassum 28
cuvieri 28
Taurillardi 29
Periconodon 66, 74
Peridipsaccus, see Babylonia
Peripaedium, see Cyathophyllum turbinatum
Perissodactyla 16
Permian 402
Phillipsastraeidae 255
Phos 180, 212-3
Pikermi 343
Pinus 25
Pirinella 147
Plagiolophus annectens 16, 27
minor 16
Planorbina 24-5
Plasmophyllinae 269
Plasmophyllum 224, 230, 269-72, 280
secundum americanum 230-1, 269-71, 280; pl.
9, figs 1-4
conistructum 270-1
sp. 271-2; pl. 9, figs 5-7
Pleistocene 291
Pleurodictyum venezuelense 273
Pliocene 291-2, 296
Pollia 183, 215
lavata 192
montense 185, 215
Polypora 398, 410, 438, 464, 479-82
dendroides 406, 478-80; pl. 23
radialis 413
septata 413
stenostoma 406, 490
verrucosa 406, 480-2; pl. 24
Pontian 292
Potamides 147, 153
(Tympanotonus) radula 113
Potamomya plana 17
Prairie Bluff 351, 375, 378, 381-2
premaxilla 300
Primates 16, 65-83
Priabonian 180, 184, 187, 190, 197, 199, 204-6,
208, 210
Proboscidea 291
Procerithtidae 110-50
Procerithium 110, 141, 146
Procynodictis euzetensis 91
Prodeinotherium 285-6, 293-330
bavaricum 291, 294, 312, 343, 345
INDEX
hobleyi 291, 294, 296, 297*, 298*, 299*, 304*,
SK Sl, Bie BiG, SIG), SAO, Gv Syst
325, 326; 33)l=5332,9395 554,.555—46-mpls
1-5
pentapotamiae 291, 294, 305, 312, 345
Prolibytherium magnieri 288
_ Pronycticebus 65-6
Propalaeosinopa 48-9
Prophet Formation 242
Propterodon 84
Protoadapis 65-6
Protictis 90
Protofusus 110, 142
Pseudamphimeryx hantonensis 16
Pseudocominella 180-1, 184, 188, 189-90, 201-2,
204, 208, 213
armata 184, 188-93, 193-6, 198; pl. 3, figs 1-8
bullata 184, 189, 193, 196, 198; pl. 2, fig. 9a, b
aspera 184, 189
deserta 182*, 184, 188, 190-3, 196-7, 200-1;
pl. 1
semicostata 180, 184, 189-90, 196-8; pl. 3, figs
9-10; pl. 4, figs 1-3
solanderi 184, 188-9, 193, 196, 198-201, 213;
pl. 2, figs 1-8
Pseudoloris 74, 80-3
abderhaldeni 80
parvulus 16, 80, 81-3, 82*
reguanti 80
Pseudophaulactis 243
Pterodon 84
Ptilofenestella 401
carrickensis 406, 467, 490
Ptiloporella 460, 471, 472-5, 489
varicosa 406, 473-5; pl. 21
Ptylopora 398, 471, 473, 475, 483-7
pluma parva 406, 483-7; pl. 25
Quaternary 296
Quercygale 90-3
angustidens 90-1
hastingsiae 16, 91-3; pl. 6
helvetica 91
radius 290, 295, 330, 333*, 334-5
Reculver 202
Retepora polyporata 457
reticulata 410
undulata 452-3
Rhabdocolpus 109-10, 140-6, 169
clementinum 109, 142-3; pl. 5, figs 6, 12-13
forbesianum 109, 143-5, 146; pl. 5, figs 1-5
manselli 142
melvillei 109, 145-6; pl. 5, fig. 7
INDEX 505
quehenensis 142 Siphonalia 190
undulatus 142 bicarinata 190
Rhabdomeson 471 Siphonophrentis 243
Rhinoceros 341 Sirte, Basin 287
rib 290, 327 Gulf of 286
Rio Cachiri Group 226, 230-2 skull 290, 295-6; pls 1-3
Rio Palmar Formation 232, 240, 281 Sligo 393
Ripley Formation 351-2, 357, 367, 369, 371-2, Smilodectes 66
374-5, 378-82 South Africa 180
Rodentia 16 South America 181
rostrum 380; pl. 5 Spain 74-5, 80, 82
Royden 187, 197, 205-6 Spalacodon 13, 37, 41
Rugosa 233-72 Spanoxyodon 49
rugose horn-corals 398 Sparnacian 184
Rupelian 184, 204-5, 212 Sphaeronassa mutabilis 182*
sphenoid 303
Sabaneto Group 229 Spinulicosta 231
Sagenella 180, 213 Squama spissa 352
Santonian 381-2 squamosal 304
Sarmatian Flinzsande 308 Stehlinella 62
Saturninia 36-8, 44—5 Stereolasma 224, 230, 233-4, 280
Scalpellidae 354 sp. 233; pl. 1, figs 6-7
Scalpellum 374 Stewartophyllum 224, 230, 232, 234-5, 281
gabbi 374 intermittens 235
inaequiplicatum 352 sp. 235; pl. 1, figs 4-5
scapula 290, 295, 330-2, 331* Stramentum canadensis 352
Scenopagus 38 haworthi 352
Schloenbachia varians 136 Strepsidura 190
Scraeva 23, 37-43, 44-5 armata 183, 189, 193, 196, 198; pl. 3, figs 9-10;
hatherwoodensis 23, 37, 39-41, 40*, 43 pl. 4, figs 1-3
woodi 12, 16, 37, 40, 41-3, 42*, 45-6 deserta 190, 193
Scrupocellaria 424 armata 196
SCRUTTON, C. T. 221-81 semicostata 193, 198
scutum 354-5, 357-9, 363-4, 369-75, 379; pls 1-5 turgida 211; pl. 9, fig. 10
Seila 160, 162-4, 167-8 (Buccinum) armata 196
dextroversa 164*, 168 semicostata 196
iglali 108, 163-4, 164*, 169; pl. 6, fig. 11 Streptelasma rectum 233
meeki 164* Streptelasmatidae 242
mundula 164 Streptelasmatina 233
quadrilirata 164* Strombodes helianthoides 256
Selma Chalk 351, 381-2, 384 simplex 254
Senonian 108, 134, 157-8, 352, 354, 356, 358-9, Strombus 113, 213
363-4, 368, 379 subcarina 358, 360; pls 1-2
Septopora 392, 487-90 subrostrum 359, 362; pl. 2
cestriensis 487, 489 supraoccipital 302
hibernica 406, 487-90; pl. 26 Switzerland 63, 65-6, 74-5, 91, 124, 133, 136-7
subquandrans 490 Syringaxon 224, 230, 236, 280
Serratocerithium 153 arnoldi 236
Shaftesbury 115 rudis 236
Sierra de Perija 229, 251 sp. 236; pl. 1, figs 1-3
Silurian 231, 251
Simpsonictis 90 Tabulata 272-5
Sinclairella 62 Tabulipora 462
Sinodia suborbicularis 22 Tapocyon 90-1
506 INDEX
tarsal bones 340 Unichinia 62
Tarsiidae 73-83 Unio 10, 17
Tarsius 73 United States 138, 180, 213
TAVENER-SMITH, R. 389-493 Upper Calp 393
Tayloran 381 WESSSaReeIey
teeth 293, 317
Teilhardia 62 Valladolid 289, 343
Teliostoma 153 Venus Bed 188, 197, 206, 210
Tennessee 352 Vincetown Formation 352
Terebellum 209 Virgiscalpellum 374-80, 381-3
Terebrella 139 beisseli 374
tergum 354-60, 363-79; pls 1-5 darwinianum 353, 378
Tertiary 352 gabbi 352-3, 374, 377-80, 383-4
Texas 122, 352 apertus 351-2, 378, 381; pl. 5, fig. 15
Thamnopora 272, 274 gabbi 352-3, 374-8, 377*, 381; pl. 4, figs
venezuelensis 273 16-18; pl. 5, figs 1-11, 14
Thanetian 184, 201, 203, 213-4 hagenowianum 353, 377-8
Thanet Sands 188, 202, 214 radiatum 377
Thanetinassa 180-1, 201, 212-3 spp. 379-80; pl. 5, figs 12-13
bicorona 184, 188, 212, 213-5; pl. 9, figs 1-7 Viverra angustidens 90-1
Thericium 153 Viverravinae 89-93
theridomyid Spp. 12, 16-17, 20, 723}, Dil Viverravus 90
Thoracica 354 angustidens 90-1, 93
Titanolepas, see Calantica euzetensis 91
Togo 184 hastingsiae 93
Tongrian 212 vomer 302
Tortonian 184 Voysa 141
Trionyx 14-18
es horis aie ee Hs 162 Waagenophyllidae 265
a eagie Washington State 180, 208-9
ae idolep ius) 23) Westeregeln 187, 191
Turonian 108, 111, 116, 131, 146-7, 150, 153, 380 Whitecliff Bay 194, 197, 199, 205-6
Turritella angustata 110, 157 Whitecliffia 180-1, 204
CE a ah suturosa 184, 204-6, 208; pl. 5, figs 6-11; pl. 6,
cingulatocostata 157 figs 1-4
Bee recenne nia 1ST tumida 180, 184, 188, 204, 206-8; pl. 6, figs
tusk 290 510
tympanic bulla 305 : :
Tympanotonus 110, 152-3; see Potamides Whitley ese AOE a
(Exechocirsus) subpustulosus 155 Wieht (sec egcu Maem
. Wrotham 127
Uchauxia 108, 110, 141, 145, 146-50, 169 .
badri 108, 147, 148-9, 150; pl. 6, figs 3-4 Yorkshire 112, 138, 155
distinctum 147 Ypresian 180, 184, 203
forbesiana 109
peregrinorsa 110, 146-8 Zaphrenticae 242-52
Phillipsi 149 Zaphrentis curvulena 239
richei 148 prolifica 251-2
solida 147 simplex 254
wiset 109, 147-8, 150; pl. 6, figs 5-6 spatiosa 251-2
sp. nov. 149-50; pl. 6, fig. 7 zoarial features 406
Uintatherium 341 Zonophyllum 271-2
ulna 290, 330, 335-6 zooecial chamber 405, 485
unciforms 338 features 405
os 7. |
a>
a)
{ 2 8 MAR }37
7
MARSUPIALIA, INSECTIVORA, @,,&.~
PRIMATES, CREODONTA AND
_ CARNIVORA FROM THE HEADON
BEDS (UPPER EOCENE) OF
SOUTHERN ENGLAND
Neg ft ;
MARSUPIALIA, INSECTIVORA, PRIMATES,
CREODONTA AND CARNIVORA FROM THE\
HEADON BEDS (UPPER EOCENE) OF
SOUTHERN ENGLAND
lee
PETER EDGAR CRAY
Bath Academy of Art, Contam
Pp. 1-102; 6 Plates, 23 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vols 23) Nos 1
LONDON : 1973
‘
f
2 8)
"Aix iy
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted im 1949, 1s
issued in five series corresponding to the Departments
of the Museum, and an H istorical series.
Parts will appear at irregular intervals as they become
veady. Volumes will contain about three to four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1065 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 23, No. 1 of the Geological
(Palaeontological) series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation
Bull. Br. Mus. nat. Hist. (Geol.).
© Trustees of the British Museum (Natural History), 1973
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 2 March, 1973 Price £4.45
MARSUPIALIA, INSECTIVORA, PRIMATES,
CREODONTA AND CARNIVORA FROM THE
HEADON BEDS (UPPER EOCENE) OF
SOUTHERN ENGLAND
By P. E. CRAY
CONTENTS
I. INTRODUCTION AND ACKNOWLEDGEMENTS . 5
II. FossiL HORIZONS AND FAUNAL LISTS . : : j : 6
(a) Introduction : : : F : 6
(b) Horizons at Hordle Cliff, Heepshire 8
(i) Introduction 8
(ii) Lower Headon Beds . : : : » 3 10
Rodent Bed . . : . : : 5 10
Crocodile Bed : ¢ 0 : : ; 13
Mammal Bed. : : 14
Mammalian fauna of the MererenallGraeadile Bed 16
Minor horizons . : : : 2 17
(c) Horizons at Headon Hill, Isle of Wight c 0 : 18
(i) Introduction . : : : : ; ; 18
(ii) Lower Headon Beds . 5 2 : : : Bik
Minor horizons : 5 : : é : 21
(iii) Upper Headon Beds . 3 0 , : 0 21
Microchoerus Bed . F : ‘ é é 21
Lignite Bed . ; ‘ : 2 : < 24
Il]. SysTEMATIC DESCRIPTIONS c . : ; c : : 27
Order MARSUPIALIA 2 : F ‘ j : j 27
Family DIDELPHIDAE Gray . : . : 5 : 27
Subfamily DIDELPHINAE Simpson : 2 ‘ : 28
Genus Pevatherium Aymard : . : : : 28
‘Peratherium species A’. : ; : : 29
‘Pevatherium species B’ é : : : 34
Order INSECTIVORA : : : : : 36
Family NYCTITHERITDAE Spann ; : : ; 36
Subfamily NYCTITHERIINAE Se : 6 ¢ 36
Genus Scyaeva gen. nov. . : : : : 37
Scvaeva hatherwoodensis sp. nov. - : ; 39
Scraeva woodi sp. nov. : : A é 2 41
Genus Avvaldus gen. nov. . : 2 ° : 6 44
Arvaldus stintoni sp. nov. . c : : 6 45
4 UPPER EOCENE MAMMALIA
Page
Family PANTOLESTIDAE Cope : : : . c 47
Subfamily PANTOLESTINAE Simpson . F : : 47
Genus Opsiclaenodon Butler 5 : : C 48
Opsiclaenodon major (iiss) : : ¢ 50
Genus Dyspterna Hopwood : ; : 2 5 57
Dyspterna hopwoodi sp. nov. : c c 7 58
Family APATEMYIDAE Matthew . : : : 61
Genus Heterohyus Gervais . : : : : ¢ 62
Heterohyus sp. . 2 : : 5 c 63
Order PRIMATES . : : : : 6 65
Family ADAPIDAE piroueesare! : : : : 0 65
Genus Adapis Cuvier . 2 ¢ : : 4 66
Adapis magnus Filhol : ; c A 67
Adapis parisiensis (Blainville) . 4 0 : 71
Family TARSIIDAE Gray : : : : 73
Subfamily MICROCHOERINAE Tedei lees : : : 73
Genus Microchoerus Wood . : c ; ° 5 74
Microchoerus evinaceus Wood. c 2 : 75
Genus Pseudoloris Stehlin . z ¢ : : < 80
Pseudoloris pavvulus Saye : ‘ 5 : 81
Order CREODONTA . : ; : : c 83
Family HYAENODONTIDAE Let : . ¢ 5 83
Subfamily HYAENODONTINAE Trouessart . : : 83
Genus Hyaenodon Laizer & Parieu c : : 84
Hyaenodon c.f. minoy Gervais . - ‘ : 85
Order CARNIVORA . : : : : : é c 89
Family MIACIDAE Cope . : : : o : 89
Subfamily VIVERRAVINAE Matthew : : . 5 89
Genus Quercygale Kretzoi_ . ; : : b : 90
Quercygale hastingsiae (Davies) . : : - QI
IV. CORRELATION AND AGE OF THE HEADON BEDS . : - 5 93
V. REFERENCES . : : 5 ; 5 P é = : 96
SYNOPSIS
The present work incorporates a revision of part of the mammalian fauna of the Headon Beds
(Upper Eocene) of Hampshire and the Isle of Wight.
Stratigraphical and historical aspects of the main mammal-yielding horizons in the Headon
Beds are presented, together with provisional lists of the complete mammalian fauna.
Marsupialia, Insectivora, Primates, Creodonta and Carnivora are considered in the systematic
descriptions. Fourteen species of mammals are described, of which four are new. Two new
insectivore genera, Scraeva and Avvaldus are referred to the Nyctitheriidae. Avvaldus is known
from a single species, A. stintoni; two species of Scrvaeva are recognised, S. hathevwoodensis and
S.woodi. A partial upper dentition has been referred to the pantolestid insectivore Opsiclaenodon
major, previously known only from the lower dentition. Well preserved material of a new species
of Dyspterna, D. hopwoodi, supports reference of the genus, formerly almost a nomen dubium, to
the Pantolestidae, and confirms the close relationship of Dyspteyna and Opsiclaenodon. Hetero-
hyus sp., Adapts parisiensis and Pseudoloris parvulus all constitute new records from the Headon
Beds.
The evidence from those mammals given detailed study, indicates that at present the Upper
Headon Beds are best grouped provisionally with the Lower Headon Beds in the Upper Eocene
(Lower Ludian). The Eocene/Oligocene boundary should be sited at some position above the
Upper Headon Beds.
HEADON BEDS OF ENGLAND 5
I. INTRODUCTION AND ACKNOWLEDGEMENTS
THE first mammals recorded from the Headon Beds were listed by Wood (1844);
these specimens, listed again by Charlesworth (1845), came from the Lower Headon
Beds of Hordle Cliff. In a further paper, Wood (1846-47) listed and excellently
illustrated this and further material from Hordle Cliff. The first record of mammals
from the Upper Headon Beds of Headon Hill was in a stratigraphical account by
Prestwich (1846). The earliest detailed description of mammals from the Headon
Beds was by Owen (1848).
In a preliminary paper in 1848, Barbara, Marchioness of Hastings recorded
mammals from Hordle Cliff. She published, in 1852 and 1853, the results of six
years’ collecting from this section. In addition to mammals, the Marchioness
recorded fish, reptile and bird material, together with valuable information about
occurrence and preservation. Also in 1852, Wright recorded an isolated mandible
from the Lower Headon Beds of Headon Hill; this was described by Owen (18572).
Some of the Marchioness of Hastings’ specimens were described by Owen (1857b)
and Davies (1884).
The only detailed systematic work was by Lydekker in his ‘Catalogue of the Fossil
Mammalia in the British Museum’, published in five parts between 1885-87. The
Headon Beds material was incompletely and sometimes inaccurately listed, but
Lydekker made a valuable contribution, consolidating Owen’s work and describing
more of the Marchioness of Hastings’ collection. Some of Lydekker’s results were
published as a few short papers (1884, 1885b, 1885d).
Stehlin’s monograph ‘Die Saugetiere des schweizerischen Eocaens’ (1903-16)
contained critical discussions of some of the Headon Beds material. In a faunal
paper, Stehlin (910) listed sixteen species from Hordle Cliff. Depéret (1917), in
a revision of the mammalian fauna of Euzet-les-Bains, listed four species from Headon
Hill and sixteen from Hordle Cliff. Contributions on aspects of the fauna were made
by Cooper (1910, 1925, 1926, 1928). The most recent work, on isolated genera, has
been by Butler (1946), Simons (1961) and Franzen (1968).
The present work comprises a revision of part of the mammalian fauna of the
Headon Beds. In the faunal lists, identification of Rodentia, Perissodactyla and
Artiodactyla is tentative; these elements of the Headon fauna have not yet been
studied in detail.
Throughout the systematics, the age of the Headon Beds is considered provisionally
as Upper Eocene (Lower Ludian). This problem is discussed later in the light of the
evidence afforded by study of the mammals.
Traditional stratigraphical nomenclature has been retained for the subdivision of
the succession; as the purpose of the present work is primarily palaeontological, no
attempt has been made to bring this nomenclature into concordance with modern
international stratigraphical practice.
The registration numbers of museum specimens are prefixed as follows:
B.M. British Museum (Natural History)
S.M. Sedgwick Museum, Cambridge
G.S.M. Geological Survey Museum
6 UPPER EOCENE MAMMALIA
D.M.S.W. D.M.S. Watson collection; now in University Museum of
Zoology, Cambridge
In addition, some specimens are from the private collection of Mr F. C. Stinton;
these are prefixed F.C.S. Where differentiation of several specimens grouped under
a single museum registration number has been necessary, numbers in parentheses
have been added after the museum registration number.
In the tables of measurements, all dimensions are in millimetres. Antero-posterior
length (a-p) and transverse width (trs) are maximum values. Measurements in
parentheses represent approximations, based on damaged teeth or taken from
roots.
I am very grateful to Dr R. J. G. Savage, not only for supervision of the initial
research, but also for his continued encouragement, advice and stimulating discussion
since that time. I also wish to thank him for so readily making available his personal
library and for critically reading the draft manuscript. I am indebted also to Dr
D. E. Russell for much encouragement and advice, for his generosity in permitting
examination of type specimens and manuscript from unpublished work and for
critically reading the draft manuscript.
My thanks are due to the following for facilities to study collections or for the loan
of specimens: Dr A. J. Sutcliffe and Dr R. Hamilton of the British Museum (Natural
History); Dr C. L. Forbes and Dr R. B. Rickards and their predecessor Mr A. G.
Brighton of the Sedgwick Museum, Cambridge; Professor J.-P. Lehman of the
Muséum National d’Histoire Naturelle, Paris; Mr J. M. Edmonds of the University
Museum, Oxford, and the late Mr A. T. Grapes of the Museum of Isle of Wight
Geology, Sandown, Isle of Wight. Thanks are due also to Mr F. C. Stinton for the
loan of specimens from his private collection.
I am indebted also to Dr J. W. Murray for helpful discussion of certain aspects of
the stratigraphy; to Dr J. M. Pallot for the processing and identification of pollen
samples and to Dr N. Edwards for much helpful discussion and for the use of
information from unpublished manuscript.
My thanks are due also to Mr J. Martin for the drawing of figs 9, 10 and 12, and to
Mr R. J. Godwin for the photography.
The initial research, of which this paper forms part, was carried out during the
tenure of a studentship from the (then) Department of Scientific and Industrial
Research. This grant and the facilities offered by the late Professor W. F. Whittard
in the Geology Department of the University of Bristol are gratefully acknowledged.
Il. FOSSIL HORIZONS AND FAUNAL LISTS
(a) Introduction
A general account of the geological structure and Tertiary stratigraphy of the
Hampshire Basin can be found in Chatwin (1960). ~— ~— 20 13 —
S.M. C30707 ih On O77 meant ae Oo/75 el: OE T lei 2 he LT nO
S.M. C53476 —- —- — — ~— — 16 135 18 13° —
S.M. C30708 — — 185 105 — —
B.M. 36812b —- —- —> — — 18 o9
B.M. 30348 —- — ~—~— ~—~— ~~ ~ ~ ~ 19 12 #%1:8 0
M2 M3
a-p trs a-p trs
S.M. C54171 — — 18 2:0
S.M. C54170 195 20 — —
34 UPPER EOCENE MAMMALIA
which may indicate a greater individual age. Hough observed (1961), that in recent
didelphids, spaces tend to appear between the premolars in the mature adult. This
feature of the premolars in S.M. C30707, may be similarly a variation due to individual
age.
‘Peratherium species B’
(Text-fig. 12)
MATERIAL. All specimens, except where otherwise stated, are from the Lower
Headon Beds (Mammal/Crocodile Bed), Hordle Cliff.
B.M. 13250 (2). Fragment of left mandibular ramus with Mzy—My and the basal
part of the coronoid. Teeth badly worn.
B.M. M13251. Fragment of left mandibular ramus with M2—M, and alveoli of
C—P3 and M,. Teeth badly worn.
B.M. 13250 (3). Fragment of right mandibular ramus with M3—My. Teeth badly
worn.
S.M. C54155. Fragment of left mandibular ramus with M,—Mg and alveoli of M3.
Teeth badly worn. Lower Headon Beds, Hordle Cliff. The Sedgwick Museum
label gives ‘Rodent Bed’ as the locality but the preservation and adhering white sand
suggests probable derivation from the Mammal/Crocodile Bed.
S.M. C53474. Anterior fragment of left mandibular ramus with Mg and alveoli
of the canine (base), P}-M; and Ms. Teeth moderately worn.
B.M. 44142 (1). Fragment of right mandibular ramus lacking teeth, with alveoli
of Py—M,.
(2). Small fragment of right mandibular ramus with alveoli of uncer-
tain position in the tooth row.
S.M. C53480. Fragment of right mandibular ramus with posterior root of Mg and
the roots of M3—Mg.
S.M. C54169. Isolated trigonid fragment of left lower molar. Badly worn.
Lower Headon Beds (Rodent Bed), Hordle Cliff.
DESCRIPTION. The posterior mental foramen lies below M; (B.M. M13251) or
Mg (S.M. C53474); the anterior mental foramen is situated below P; (B.M. M13251)
or Pz (S.M. C53474). The subhorizontal symphysial surface extends to below Py.
S.M. C53474 possesses a nutrient foramen (as in S.M. C30707, referred to ‘Peratherium
species A’) opening through the symphysial surface.
No description is given of the dentition which is structurally almost identical to
previously described specimens referred to ‘“Peratherium species A’.
REMARKS. Because of the much larger dimensions of the teeth (especially the
length of the tooth row) compared with ‘Peratheriwm species A’, these specimens are
considered to represent a distinct species. Hough stated (1961), that this criterion
is less subject to variation than depth of mandible and spacing of premolars, and has
been used in the determination of Marmosa species. There is some variation in
mandible depth and premolar spacing; as in ‘Pevatheriwm species A’ this probably
reflects individual age differences.
HEADON BEDS OF ENGLAND 35
Two mandibular fragments without teeth (B.M. 44142 (2) and S.M. C53480) are
tentatively referred to this species. B.M. 44142 (1) is considerably longer and more
slender than the other referred specimens with widely spaced anterior premolars; it is
considered tentatively as an old adult of ‘Peratherium species B’.
el
Ml B ily
Al
7 i,
4 i
tae a
Si ,
Dw) WW Nc iN -
\ IAW Pe MZ il RW
Fic. 12. ‘Pevatherium species B’. Fragmentary left mandibular ramus with M2—M4.
B.M. M13251 x6. Lower Headon Beds, Hordle Cliff. (a) Occlusal view. (8) Internal
view. (c) External view.
36 UPPER EOCENE MAMMALIA
A pathological condition is seen in S.M. C53480. The mandible evidently had been
damaged during life internally below Mz and Ms, and shows some formation of callus
in this area.
TABLE 6
Measurements (in mm) of ‘Pevatherium species B’
My M2 Ms M4
a-p trs a-p trs a-p trs a-p trs
B.M. 13250 (2) — — 2°5 75 eee 1:6 2°5 I°4
B.M. M13251 a —- 2°5 1:6 2°55 1:6 2°25 I'4
B.M. 13250 (3) 2:5 ito 2°5 1°35
S.M. C53474 — _ 2:6 I*4
S.M. C54155 2, I°4 2°7 a —
Order INSECTIVORA
Family NYCTITHERIIDAE Simpson, 1928
REMARKS. Simpson’s family Nyctitheriidae was abandoned by McKenna
(r1960a) as an unnatural unit. Van Valen (1967) placed Nyctitheriwm together with
Clinopternodus Clark, 1937, as a subfamily (Nyctitheriinae) within the Adapisoricidae.
On the basis of Robinson’s (then unpublished) work on the group, McKenna (1968)
recognised ‘a restricted family-group taxon based on WNyctitherium and several
referred genera’, into which he placed Leptacodon. In his review of the family,
Robinson (1968) upheld the status of Simpson’s family Nyctitheriidae, subdividing
it into three subfamilies: Nyctitheriinae Simpson, 1928; Geolabidinae McKenna,
1960; and Micropternodontinae (Stirton & Rensberger, 1964). The Geolabidinae,
placed originally by McKenna (1960b) in the Erinaceidae, had been grouped by
Van Valen (1967) within the Adapisoricidae. Robinson’s view is adopted here.
Subfamily NYCTITHERIINAE Simpson, 1928
DIAGNOSIS. Pi semi-molariform, Ps, P3 with two roots each, P; with one root,
jaw long and slender, with no apparent shortening. Probable dental formula
i, ie i, 5. (Robinson, 1968 : 129).
ReMARKS. Three genera were included by Robinson (1968) in the subfamily:
Nyctitheriwm Marsh, 1872. Middle—Late Eocene; North America.
Leptacodon Matthew & Granger, 1921. Middle (?), Late Palaeocene—Early
Eocene; North America.
Saturnimia Stehlin, 1940. Late Eocene; Europe.
To these are added two new genera described below.
Leptacodon has undergone a varied history. McKenna (1960a : 52and 1960b : 156)
removed Leptacodon (sensu stricto) from the Leptictidae; this genus, composed of L.
HEADON BEDS OF ENGLAND 37
teney Matthew & Granger, 1921 (the type species) ; L. Jadae Simpson, 1935 (apparently,
but not mentioned specifically by McKenna); L. minutus Jepsen, 1930, and L.
jepsent McKenna, 1960a, was considered to be near the base of erinaceid phylogeny
and possibly closely related to the ancestry of the Geolabidinae. This tentative
reference of Leptacodon (sensu stricto) to the Geolabidinae was accepted by Russell
(1964 : 46). Van Valen (1967) placed Leptacodon (sensu stricto) in the Adapisoricidae
(subfamily Adapisoricinae) and more recently, McKenna (1968) has stated that the
genus is perhaps best considered as a nyctithere. Robinson (1968) in his review of
the Nyctitheriidae, clarified the systematics of the genus Nyctitherium ; he indicated
that the type specimen of Myolestes dasypelix Matthew, 1909, is a nyctitherine,
although he has agreed with McKenna’s allocation of a referred specimen ‘c.f.
Myolestes dasypelix’ to the Geolabidinae (McKenna, 1960b : 147). In addition to
Nyctitherium dasypelix, Robinson (1968) recognised two further species: the type
species, N. velox Marsh, 1872, and N. serotinum (Marsh, 1872). Inthe same paper, he
rejected Stehlin’s reference of Saturninia to the Soricidae, maintaining for the genus,
a close relationship with Nyctitherium.
Genus SCRAEVA gen. nov.
EtyMoLtocy. Screawa—(Anglo-Saxon), a shrew.
Dracnosis. In Py—-Ms, talonid narrower transversely than trigonid; talonid and
trigonid subequal in length; trigonid with markedly angular outline, with straight
or slightly convex internal margin and moderately to sharply pointed anterior apex;
protoconid larger than metaconid; talonid angular with internal and external
margins subparallel, except on Ms. M,—M3 with a mesoconid; hypoconid and
entoconid subequal in size; hypoconulid median or slightly internal of midline, in Mg
posteriorly projecting as a prominent rounded lobe. Pq hypoconulid minute,
slightly external to midline. Internal face of coronoid process with antero-posterior
ridge at molar alveoli level.
TYPE SPECIES. Scraeva hatherwoodensis sp. nov. Upper Headon Beds (Mzcro-
choerus Bed), Headon Hill.
REMARKS. One other species has been distinguished: Scraeva woodi sp. nov.
Lower Headon Beds (Mammal/Crocodile Bed), Hordle Cliff.
In 1844, Charlesworth (in Wood, 1844 : 350, footnote) recorded a fragmentary
mandibular ramus, obtained by Flower from Hordle Cliff, which he named Spalacodon ;
this was again recorded by Charlesworth (1845 : 50). A year later, the specimen
was figured by Wood (1846, pl. 2, figs 5, 5a-c) and listed as ‘an imperfect lower jaw’.
Charlesworth and Wood did not designate a species and the specimen was referred
to as ‘Spalacodow or ‘Spalacodon sp.’ Because of the inadequate original descriptions
and the loss of the specimen, Spalacodon is considered here to be a nomen nudum.
Comparison of the type material of Scraeva woodi with Wood’s illustrations indicate
the probable specific identity of the two forms. This contention is strengthened by
the probability that Spalacodon and the material of Scraeva woodi were obtained from
the same level, the Crocodile Bed of Hordle Cliff. The specimen described by
38 UPPER EOCENE MAMMALIA
Charlesworth and Wood (from Wood’s figures, a right mandibular ramus with
Pz and P4s—Ms) has not been located during the present study and the material is
considered to be lost.
Scraeva is referred to the Nyctitheriidae mainly on the basis of strong resemblances
to Leptacodon and Nyctitherium. The soricid-like antero-posterior internal ridge on
the coronoid, characteristic of Nyctitherium (Robinson, 1968), is well developed in
Scraeva. A talpid characteristic, occasionally developed in Nyctitherium and
present also in Saturninia, is a continuation of the crista obliqua up the posterior wall
of the metaconid; this is prominently and invariably developed in Scvaeva. The
Headon genus is distinguished from other nyctitheres as follows: In Nyctitherium
(see Robinson, 1968), Pg has a smaller talonid basin and lacks an entoconid; in the
molars, the hypoconulid is more internal in position and smaller and the paraconid
is more internally situated. In this study the type specimen of Saturninia has not
been examined, but from Stehlin’s illustrations (1940, fig. 1c, fig. 2c) the Pg is
relatively smaller with a less basined talonid, and the molar paraconids are more
internal in position. In addition, in both Nyctitherium and Saturninia, the outline of
P4—Mz is more rounded and the P,—Ms trigonid and talonid are of similar transverse
width.
The Py, of Scraeva is very similar to that of the type species of Leptacodon, L. tener,
except for a rounded posterior talonid margin and a sharper, more separated para-
conid in Leptacodon tener. The molars are also similar although in Leptacodon tener,
the metaconid is as high as the protoconid; the paraconid is somewhat sharper and
more separated; the Mg hypoconulid is less projecting posteriorly and the trigonid
cusps are not so elevated above the talonid. Less resemblance is apparent in the P,
of Leptacodon ladae which has a more isolated median paraconid, a more anteriorly
placed protoconid and a much shorter talonid.
Scraeva also shows marked resemblances to Scenopagus McKenna & Simpson,
1959, Messelina Tobien, 1962, and Adapisorex Lemoine, 1883; these genera have been
generally regarded as ‘primitive probable erinaceids’, although extensive regrouping
and familial separation from the Erinaceidae has been proposed by Van Valen
(1967 : 261). Inseveralrespects, particularly the elevated trigonids and the structure
of Mz, Scraeva is similar to Scenopagus but there are significant differences: in
Scenopagus, the talonid cusps of the molars are slightly higher; the trigonid cusps
are compressed antero-posteriorly and the paraconid crest is larger and more oblique,
resulting in a blunter anterior tooth apex; the hypoconid crest runs to the internal
base of the protoconid, not the metaconid as in Scraeva, and the hypoconulid is only
slightly developed in Mj-Mz. Py has not been adequately described or figured. In
Messelina, the lower molars are much narrower transversely ; the talonid and trigonid
are subequal in width and the hypoconulid is internal; the trigonid of the molars is
not elevated, and in M,—Mg the trigonid and talonid cusps are subequal in height.
Also, the Py of Messelina has a rounded outline and lacks a hypoconulid. Greater
resemblance to Scraeva is seen in Adapisorex: in the molars of both genera, the
protoconid is larger than the metaconid (but not so extremely developed in Scraeva)
and the basined talonid has a median hypoconulid. The P4 is similar in Scraeva and
Adapisorex, with a small metaconid, a large protoconid anterior to the metaconid
HEADON BEDS OF ENGLAND 39
which dominates the trigonid (less extremely so in Scrvaeva) and a small median
hypoconulid. However, Adapisorex is distinguished from Scraeva by having much
lower-placed cusps, the trigonid being little elevated above the talonid; the entoconid
and hypoconid are much larger and the trigonid and talonid are subequal in width.
In basic structure, particularly of Ps-Mz, Adunator Russell, 1964 (originally
considered by Russell as a leptictid but placed by Van Valen, 1967, in the Adapisori-
cinae) shows similarities to Scraeva, although the Py paraconid is much more separated
anteriorly as a distinct cusp. Also, in Adwnator the Mz hypoconulid is more internal
and not so projecting as an isolated lobe, and in the molars the metaconid and
entoconid are higher than the protoconid and hypoconid respectively.
Scraeva hatherwoodensis sp. nov.
(Text-fig. 13)
EtymMoLocy. hatherwoodensis—from Hatherwood Point, off Headon Hill,
Isle of Wight, the sea cliff locality.
Dracnosis. The smaller species. P4-Me trigonid with a slight posterior tilt.
P4-M2 with straight internal trigonid margin and protoconid and metaconid more
internally placed than in S. woodi. P4 metaconid and protoconid slightly lower than
corresponding cusps of M;; paraconid median, at same height as molar paraconids.
P4 metaconid more posteriorly placed in relation to protoconid than in S. woodz, in
M, metaconid only slightly posterior to protoconid.
Hototyre. B.M. M26051. Fragment of left mandibular ramus with P,s-M2
and alveoli of Iz-P3 and M3. Lower external surface embedded in matrix. Teeth
slightly worn. Upper Headon Beds (Microchoerus Bed), Headon Hill. The only
specimen of the species.
Description. The long slender mandible has a large oval posterior mental
foramen just below the alveolar level of Ps. There is a small angular process (tip
damaged on the specimen) at molar alveoli level. The coronoid and condyle are not
preserved. The mandibular foramen is visible just below alveolar level. On the
internal face of the coronoid, a prominent antero-posterior ridge runs from the level
of the M3 alveoli towards the mandibular foramen. A similar ridge was observed
by Simpson (7m McGrew, 1959) on a toothless mandibular fragment from the Middle
Eocene of Wyoming, tentatively referred to the Soricidae. This structure also
occurs in Nyctitherium (Robinson, 1968 : 130).
M3 is double-rooted and known only by alveoli. In Mj—Mg the talonid and trigonid
are subequal in length, but the talonid is narrower transversely. The internal and
external talonid margins are straight and subparallel. The trigonid is angular with
a sharp anterior apex and a straight internal margin; sharply elevated, it falls almost
vertically to a low basined talonid. There is a prominent antero-external cingulum.
The protoconid is larger than the metaconid and slightly more anteriorly placed
(on both My and Mg the metaconid is damaged). The paraconid is moderately well
40 UPPER EOCENE MAMMALIA
separated and cuspate, forming the sharp anterior extremity of the trigonid; this
cusp is median in M; but slightly internal to this position in Mz. The hypoconid and
entoconid are subequal; the hypoconulid is a little smaller but comparison is difficult
as the entoconid is almost unworn, whereas the hypoconid and hypoconulid are
badly worn. The median hypoconulid (in Mg slightly internal to the midline) is
posterior to the hypoconid and entoconid, projecting very slightly posteriorly and
joined to the hypoconid and entoconid by smalllow crests. From the entoconid a low
feeble crest runs anteriorly to the metaconid base. A stronger crista obliqua runs
from the hypoconid up the posterior wall of the metaconid; this crest swells into a
prominent mesoconid (stronger on M; than Mg) midway along its length.
P, is very similar to M, and Mg but differs in several respects: the tooth is more
laterally compressed and smaller in all dimensions; the hypoconulid is very small,
situated external to the midline near the hypoconid; the metaconid is more posterior
in relation to the protoconid; the hypoconid is larger than the entoconid and the
protoconid/paraconid crest is smaller, placed slightly external to the midline and less
V-shaped than in the molars. P3 and Pz are represented only by alveoli; these teeth
were apparently double-rooted and closely spaced. Py is single-rooted, represented
only by the alveolus. Known only from the alveolus, the canine was evidently
slightly anteriorly inclined, with a root diameter as that of P}. Is is represented by
the possible base of the alveolus.
REMARKS. This species is clearly related to Scraeva woodi in that both forms
exhibit the characteristic squared angular outlines of the teeth. There are a number
of small but distinct differences between S. hatherwoodensis and S. woodi (see diagnoses)
which indicate at least a specific separation. Also, S. woodi is markedly larger.
Fic. 13. Scvaeva hatherwoodensis gen. et sp. nov. Fragmentary left mandibular ramus
with Py-Mz. Holotype (B.M. M26051) x10. Upper Headon Beds, Headon Hill. (a)
Occlusal view. (8) Internal view.
HEADON BEDS OF ENGLAND 41
TABLE 7
Measurements (in mm) of Scyvaeva hatherwoodensis
Py My Me
a-p trs a-p trs a-p trs
Holotype 1°35 o-9 I°5 Ir I'5 I'2
(B.M. M2605r)
Scraeva woodi sp. nov.
(Text-fig. 14)
ETYMOLOGY. woodi—after Searles Wood who first listed and figured insectivore
material from the Headon Beds as Spalacodon, a name suggested by Charlesworth,
but here considered as a nomen nudem.
Dracnosis. Larger than S. hatherwoodensis. Py, with slightly concave internal
trigonid margin, in molars slightly convex. P4metaconid and protoconid same height
as corresponding cusps of M;; paraconid markedly internal, lower than molar para-
conids. P, metaconid slightly posteriorly placed in relation to protoconid, in My,
metaconid opposite and in Mg—M3 metaconid slightly anterior to protoconid.
HototypPe. S.M. C53475. Fragment of left mandibular ramus with Py-M3 and
alveoli of Ps. Teeth unworn. Lower Headon Beds (Mammal/Crocodile Bed),
Hordle Cliff.
PARATYPE. B.M. M25112. Left mandibular ramus with P2, Py—-M3 and alveoli
of I2—-P;, P3. Badly worn dentition. Lower Headon Beds (Mammal/Crocodile Bed),
Hordle Cliff.
MaTERIAL. S.M. C30709. Fragment of left mandibular ramus with Mj—Mp,
posterior alveolus of P4, and alveoli of Ms. Teeth unworn but metaconids damaged.
Lower Headon Beds (Mammal/Crocodile Bed), Hordle Cliff.
B.M. 36801c. Fragment of left mandibular ramus with Mg and alveoli of Mg
Internal margin of tooth damaged and tips of cusps broken off. Patches of coarse
white sand matrix adhering. Lower Headon Beds (Mammal/Crocodile Bed), Hordle
Cliff.
S.M. C54168. Isolated right Mz. Slightly worn. Lower Headon Beds (Rodent
Bed), Hordle Cliff.
DESCRIPTION. The mandible is a little stouter than in S. hatherwoodensis, with a
large oval posterior mental foramen below the posterior toot of P3. The antero-
posterior ridge on the internal face of the coronoid is present as in S. hatherwoodensis.
Paratype B.M. Mz25112 shows the wedge-shaped symphysial surface with ridged
margins; this occupies nearly the whole depth of the jaw under Is, sloping baso-
posteriorly to below the anterior root of Pe.
The incisors, canine and P; are known only from alveoli in paratype B.M. M25112.
In this specimen, two distinct, well-separated alveoli both slightly smaller than the
P; alveolus, indicate at least two incisors, here considered as Ip-I3. Another more
42 UPPER EOCENE MAMMALIA
anterior tooth may have been present but evidence is lacking. The canine is re-
presented by an alveolus indicating a larger anteriorly-inclined tooth. Pj, single-
rooted and well-separated, is represented by the alveolus which is smaller than that of
the canine. Pg: is double-rooted and ovalin outline. A single large anterior median
cusp slopes antero-basally to a small low antero-internal cuspule; a small heel bears
a minute internal cuspule. Pg is known only from the alveoli; the tooth is double-
rooted and apparently about the same size as Pp».
Fic. 14. Scyvaeva woodi gen. et sp.nov. Fragmentary left mandibular ramus with P4—M3.
Holotype (S.M. C53475) x10. Lower Headon Beds, Hordle Cliff. (a) Occlusal view.
(B) Internal view. (c) External view.
HEADON BEDS OF ENGLAND 43
The trigonid and talonid in P, are subequal in length. In M,—Ms the talonid is
slightly more elongated antero-posteriorly and narrower transversely; the internal
and external margins are fairly straight and subparallel, except in M3 where the
talonid narrows posteriorly (not as extreme as in S. hatherwoodensis). The Mg in
paratype B.M. Mz5112 has a somewhat indented external margin. P, and Mg are
narrower transversely than M; and Mz. In P4—Ms the trigonid is angular with only a
slightly curved internal margin; sharply elevated, it drops steeply to a low basined
talonid. There is a prominent antero-external cingulum as in S. hatherwoodensis.
The metaconid is smaller than the protoconid, situated posterior to the protoconid
in Py, opposite in Mj and slightly anterior in Mz and M3. The paraconid, moderately
well separated and cuspate, forms the fairly sharp antero-internal extremity of the
tooth—this cusp is moderately internal to the midline in all teeth (P4-M3). The
hypoconulid in My; and Mgis a little smaller than the entoconid and median in position ;
in Py, it is very small and slightly external to the midline (but in Ms, hypoconulid,
entoconid and hypoconid are subequal). The P, hypoconulid in paratype B.M.
M25112is indistinct, indicated only bya slight externally situated projection posterior
to the hypoconid and entoconid. Ms has a posteriorly projecting hypoconulid lobe,
slightly internal to the midline; this lobe is very prominent and apically projecting
in paratype B.M. M2511z. In Pg—Mg the hypoconulid is joined to the hypoconid
and entoconid by small low crests; a low feeble crest runs antero-internally from the
entoconid to the metaconid base. A much stronger crista obliqua runs from the
hypoconid up the posterior wall of the metaconid ; this crest bears a distinct mesoconid
in M\—Msz, although the cusp is only feebly developed in Mg.
REMARKS. Comparison with S. hatherwoodensis has been effected in the diagnosis
and description. Only five specimens are known and little idea can be gained of
variation within the species. S.M.C30709 and S.M. C54168 differ from the holotype
only by having a less distinct mesoconid in the molars. Paratype B.M. M25112
represents a much older individual than the holotype and comparison of the two
specimens is difficult because of a high degree of wear on the former specimen.
Reference of B.M. Mz25112 to the species is fairly certain, however; when structural
differences which can be assigned to excessive wear ate discounted, the two specimens
have a high degree of similarity. Also, both specimens are apparently from the same
locality and approximate horizon. B.M. 36801c appears closely similar to the
holotype, although the former specimen is damaged.
TABLE 8
Measurements (in mm) of Scvaeva woodi
Pe, P3 12h My, Me Ms
a-D tis =ph tts! a—p (Ets) d-p) tts) a-p) irs! fa-p)) irs
B.M. M25112 Ti O;0 M1 nT 7 OF NT 75 AS eat Om seICA5 a T8— 1-2
B.M. 36801c SS eS SS SS i a ee YSZ ee
S.M. C54168 — — —«-—-S—s T8255 —— —
S.M. C53475 Ti), TRO) FUO|G} aS} ESI} ga, acts) ago
S.M. C30709 a aS TNS) 1-7) 125) =
44 UPPER EOCENE MAMMALIA
Genus ARVALDUS gen. nov.
EtymoLocy. Arvaldus—Jutian king of the Isle of Wight, A.D. 686.
DraGnosis. P, protoconid markedly anterior to the metaconid; hypoconulid
(damaged) appears more prominent than in Scvaeva. P4-Ms posterior margin some-
what oblique to length of tooth row; paraconid internal; protoconid larger than
metaconid; P, hypoconid opposite, M;—Mg slightly posterior, and in M3 markedly
posterior to entoconid. M3 hypoconulid prominent, internal to midline, slightly
projecting; talonid angular in outline with straight posterior and postero-internal
margins.
TyPE SPECIES. Avrvaldus stintoni sp. nov. Upper Headon Beds (Lignite Bed),
Headon Hill. The only known species.
REMARKS. Avrvaldus shows considerable similarity with Leptacodon, Nyctitherium
and Saturninia. The Py of Leptacodon tener (the type species) seems quite close to
that of Avvaldus with a well differentiated talonid and the protoconid strongly
anterior to the metaconid, although in Py, of L. tener, a hypoconulid is absent and the
posterior talonid margin has a rounded outline. Similarity with the Py of Leptacodon
ladae is shown by the anterior position of the protoconid and the presence of a small
hypoconulid, although in several other respects—the median paraconid, rounded
outline and small talonid, the tooth is rather different. The type species of
Nyctitherium, N. velox, is distinguished from Arvaldus by the Py talonid being much
shorter antero-posteriorly with the protoconid opposite the metaconid ; in the molars,
the paraconid is more internal and the hypoconulid is weaker and placed more inter-
nally, although in this latter respect, Avvaldus is closer to N. serotinum where the
molar hypoconulid is slightly more median in position. In Saturninia the Py,
protoconid is opposite the metaconid and the paraconid of the molars is more
internally situated.
One of the most obvious features which distinguishes Lepiacodon, Nyctitheriwm,
Saturninia and Scraeva from Arvaldus is the structure of M3. In Leptacodon there is
a strong entoconid opposite the hypoconid and a well separated posteriorly projecting
hypoconulid. This tooth in Nyctitheriwm velox is different in outline; the talonid is
shorter with a median hypoconulid and an entoconid which is opposite the hypoconid.
The Mg talonid in Saturninia is shorter and different in outline with a median
posteriorly projecting hypoconulid. In M3 of Scraeva the entoconid and hypoconid
are opposite and there is a median posteriorly projecting hypoconulid.
In this M3 talonid structure, Avvaldus shows similarities with the American
geolabidines, Geolabis, ‘c.f. Myolestes dasypelix’ and Centetodon, particularly the
latter. In Centetodon, the Mg talonid bears only a hypoconid and an internal
crestiiform hypoconulid (fused entoconid and hypoconulid?) ; the internal wall of the
postero-internally projecting hypoconulid curves abruptly baso-posteriorly and there
is no sign of an entoconid. A somewhat similar distinctive Ms; talonid structure is seen
in Arvaldus, although here it is not so pronounced; the hypoconulid is less posteriorly
situated and a small anteriorly placed entoconid is still present. However, in
Centetodon and other geolabidines, the structure of Py is different: in the American
forms, the Py protoconid and metaconid are opposite and the talonid is much reduced,
HEADON BEDS OF ENGLAND 45
with a single large internal cusp. Greater similarity to Avvaldus is seen in the P4
of ‘c.f. Hypacodon sp.’ (= Centetodon sp.) described by McKenna. (1960b : 150).
Here the elongation of the tooth is similar to that in Avvaldus and the protoconid is
anterior to the metaconid; the talonid is more developed and there is a central
hypoconid and a small though distinct entoconid. Robinson (1968 : 134) has
considered this specimen as an adapisoricid.
Arvaldus differs from Scraeva in a number of important aspects (see diagnoses),
the more obvious of these being the non-angular tooth outlines, the structure of P4
and the distinctive Mg talonid; the two genera do not appear to be closely related.
To some extent, the structure of Avvaldus is intermediate between nyctitherines and
geolabidines and on the basis of the Mg structure, the genus could be tegarded as a
primitive relatively unspecialised geolabidine. However, the molarised P4 and the
continuation of the crista obliqua up the posterior wall of the metaconid support
reference to the Nyctitheriinae. The anterior lower dentition and the upper teeth
of Avvaldus are unknown. At present the European record of the family comprises
Saturninia, Scraeva and Arvaldus, all from the Upper Eocene. It is likely that the
phyletic position of Avvaldus will be clarified with the discovery of nyctitheriids
from the Lower Eocene of Europe.
Arvaldus stintoni gen. et sp. nov.
(Text-fig. 15)
EtyMoLocy. stintoni—after Mr F. C. Stinton, the finder of the specimen.
Dracnosis. No other species are known and the diagnosis is the same as that for
the genus.
Hototyre. F.C.S.1. Fragment of right mandibular ramus with Pe, P4sa-Ms3 and
the alveoli of Ps. Upper Headon Beds (Lignite Bed), Headon Hill. The only
known specimen of the species.
DEscrRIPpTION. The mandible has a large posterior mental foramen below P3.
The posterior region is not preserved. Py, is very similar to that of the paratype of
Scraeva woodt (B.M. M25112). The tooth is double-rooted and oval in outline with a
large median anterior cusp, which slopes baso-anteriorly to a small low antero-
external cuspule. A small low heel bears a single minute internal cuspule. Repre-
sented by only alveoli, P3 is double-rooted and apparently a little smaller than Py.
In P4—Ms, the talonid is a little longer antero-posteriorly than the trigonid especially
in Mg and about the same width, with an indented internal and external margin.
The trigonid has a curved internal margin. The posterior wall drops less steeply
than in Scvaeva. There is a prominent antero-external cingulum. The protoconid
is larger than the metaconid and situated opposite to it in M;—M3; in P, the protoconid
is markedly anterior to the metaconid. The cuspate paraconid is moderately well
separated and situated internally in all four teeth as in Scraeva woodi. The Py
paraconid is sharper and more anteriorly projecting than in this latter species. In
P4—M3 the hypoconid is stouter than the entoconid; the hypoconulid is a little smaller
than the entoconid or subequal, except on Mg where it is a little larger; this cusp is
40 UPPER EOCENE MAMMALIA
slightly external to the midline in Py, median in M, and slightly internal in Mz and
Msg; it is posterior to the entoconid and hypoconid and projecting slightly in all four
teeth. The hypoconulid is well separated from the entoconid but is joined to the
hypoconid by a small low external crest. In P4, the hypoconid is opposite to the
entoconid, in My and Mg slightly posterior and in Mg markedly posterior. From the
entoconid, a feeble crest runs antero-internally to the base of the metaconid. A
stronger crista oblique runs antero-internally from the hypoconid up the posterior
wall of the metaconid; this swells up midway along its length into a prominent
mesoconid. The enclosed talonid is deeply basined. The posterior margin of the
talonid is slightly obliqua to the length of the tooth in P4—Mg (a tendency also seen in
paratype B.M. M25112 of Scraeva woodt). In Msg the straight posterior margin runs
from the sharp postero-external extremity of the talonid to the slightly more
posteriorly situated hypoconulid. From this latter cusp, the straight talonid margin
runs obliquely forwards to the relatively anteriorly-placed entoconid.
Fic. 15. Avvaldus stintoni gen. etsp.noy. Fragmentary right mandibular ramus with P2,
Pa-M3. Holotype (F.C.S. 1) x10. Upper Headon Beds, Headon Hill. (a) Occlusal
view. (B) External view. (c) Internal view.
HEADON BEDS OF ENGLAND 47
TABLE 9
Measurements (in mm) of Avvaldus stintoni
Pe Ps P4 M, Me M3
a DECLSE a PRRELS sae biS aD cLSee cd —-Dm cise d—Pees LES
Holotype 1200756 ee O-Guunl-O) Li T-O5met-2 0) To5) | 10:0
(F.C.S. 1)
Family PANTOLESTIDAE Cope, 1884
Diacnosis. Facial part of skull short, basicranial region broad, middle portion
of skull elongate. Angle of mandible a short stout process expanded at the tip.
Mental foramen usually beneath My, sometimes below Py. Dental formula 3, f, 3, +
Molars and premolars with low massive cusps. Canines large, massive, lower
canines often semiprocumbent. Upper incisors spaced and _ peg-like. P ,—-P3
elongate and fairly small. P4 relatively enlarged (after Matthew, 1909).
REMARKS. On the basis mainly of differences in the structure of Pi, two sub-
families have been traditionally distinguished :
Pantolestinae Simpson, 1937
Pentacodontinae Simpson, 1937
Gazin (1959) erected a new subfamily, Apheliscinae; this was referred by McKenna
(1960a) to the condylarths (Hyopsodontidae). Van Valen (1966) supported Gazin’s
reference of the Aphelescinae to the Pantolestidae, but more recently (1967) has
agreed with McKenna’s removal of the group to the Hyopsodontidae.
Van Valen (1967) in his revision of insectivore classification raised the pentacodonts
to family level. As the justification for such a procedure (Van Valen, 1967 : 321)
seems no greater than that for retaining them in the Pantolestidae, the old-established
two fold subdivision of the Pantolestidae is retained here.
Subfamily PANTOLESTINAE Simpson, 1937
Dracnosis. Pj relatively little enlarged. P, compressed, no metaconid, semi-
shearing, heel essentially unicuspid and little or not basined. P4 with compressed
amphicone, no metacone, low, sharp styles, small protocone without cingula (Simpson,
1937).
Remarks. The Pentacodontinae are distinguishable from this subfamily by
Pi being much enlarged; Py has a well developed metaconid and basined talonid; P4
has a distinct metacone, large protocone, with expanded cingula and styles are small
or absent.
48 UPPER EOCENE MAMMALIA
The following genera have been referred, to date, to the Pantolestinae:
Propalaeosinopa Simpson, 1927. Middle-Late Palaeocene; North America.
Bessoecetor Simpson, 1936. Middle-Late Palaeocene; North America. ?Late
Palaeocene; Europe.
Pagonomus Russell, 1964. Middlé-Late Palaeocene, ?Early Eocene; Europe.
Pantomimus Van Valen, 1967. Middle Palaeocene; North America.
Palaeosinopa Matthew, 1901. Late Palaeocene-Early Eocene; North America.
Pantolestes Cope, 1872. Middle Eocene; North America.
Cryptopithecus Schlosser, 1890. Late Eocene; Germany.
Opsiclaenodon Butler, 1946. Late Eocene; Britain.
Dyspterna Hopwood, 1927. Early Oligocene; Britain.
Androconus Quinet, 1965. Early Oligocene; Europe.
Chadronia Cook, 1954. Early Oligocene; North America.
Galethylax Gervais, 1848-52. Late Eocene; Europe.
Buxolestes Jaeger, 1970. Middle Eocene; Europe.
The above generic list follows that of Van Valen (1967) but with the tentative
retention of Opsiclaenodon and Bessoecetory as distinct genera, and the addition of
Buxolestes.
Simpson (1945) listed Propalaeosinopa, Bessoecetor, Palaeosinopa, and Pantolestes.
Van Valen (1967) added a new genus Pantomimus, Pagonomus, Cryptopithecus,
Dyspterna, Androconus, Chadronia and Galethylax. ‘This latter problematical genus
(discussion in Van Valen, 1966 : 73; 1967 : 227) was only provisionally referred to the
family. Van Valen originally listed a new genus Pantinomia from the Middle
Palaeocene of North America, as a pantolestid (1967 : 259) but in a later view
(1967 : 225, footnote) referred it to the arctocyonids. In the same paper, Van Valen
(1967) considered Bessoecetor as a synonym (tentative) of Propalaeosinopa and
Opsiclaenodon as a synonym of Cryptopithecus, although Russell (in Russell, Louis &
Poirier, 1966) retained Bessoecetor as a valid genus, into which he placed tentatively
a new species, B. lever. More recently, Dr D. E. Russell (personal communication,
1971) has reaffirmed his belief in the validity of Bessoecetor. Russell (1964) in his
type description of Pagonomus did not assign the genus to a subfamily; he noted
resemblances of most of the teeth to the pentacodontine, Aphronorus. Additional
confirmation of this view was given by a molarised P4 tentatively referred to the
genus by Russell (im Russell, Louis & Poirier, 1966). Van Valen (1967) considered
the type of Pagonomus (an isolated M2) as a pantolestine.
Genus OPSICLAENODON Butler, 1946
Diacnosis. Lower molars with moderately elevated cusps, molars uncompressed
antero-posteriorly. My, and Mg very similar in size and structure, Mg much smaller
but quite similar structurally. Entoconid moderate-sized and distinct on all molars,
hypoconid slightly larger. Hypoconulid of molars small but distinct, not extended
posteriorly as a lobe on Mg. Upper molars moderately elongated transversely with
HEADON BEDS OF ENGLAND 49
rounded outlines; M! slightly smaller than M?, M? markedly smaller; prominent
hypocone in all molars; conules developed in all molars, situated close to metacone
and paracone, the metaconule being larger than the paraconule. In M2, conules
well developed, but more rudimentary in M! and M’. M1! and M? parastylar region
extended antero-externally, and metastylar region extended postero-externally, as
prominent rounded lobes. No continuous cingulum round base of protocone.
Type SpEcIES. Opsiclaenodon major (Lydekker, 1887). Lower Headon Beds
(Mammal/Crocodile Bed), Hordle Cliff. The only known species.
RemarKs. Butler redescribed (1946) Newrogymnurus major Lydekker known only
from fragmentary lower dentitions, as a new genus Opsiclaenodon. In his original
description, Butler referred (1946 : 691) Opsiclaenodon to the Arctocyonidae,
suggesting a relationship with Dyspterna and Didymoconus. McKenna (im Russell
and McKenna, 1961 : 281, footnote) suggested that Opsiclaenodon was an oxyclaenine
arctocyonid, with affinities to Metachriacus, Mimotricentes and Spanoxyodon.
Russell (1964 : 193) observed that reference of Opsiclaenodon to the Oxyclaeninae
was unlikely. Van Valen (1966) referred Cryptopithecus and Dyspterna to the
Pantolestinae. He later added (1967) Opsiclaenodon as a synonym of Cryptopithecus.
Restudy of Butler’s material and examination of additional specimens, has confirmed
Van Valen’s view that Opsiclaenodon is a pantolestine. The type of Cryptopithecus
sideroolithicus is a fragmentary left mandibular ramus with Mz2—M3 (Bayerische
Staatssammlung fiir Palaontologie, Munich, No. A.S. (128) XI 1). Comparison
with a cast of this specimen (kindly made available by Dr D. E. Russell) reveals
reasonable structural differences (see below) and in the present work, the validity ot
Butler’s genus is accepted provisionally, pending discovery of better more complete
material of Cryptopithecus.
Isolated M1 and M2 (described below) referred to Opsiclaenodon show resemblances
to the oxyclaenine arctocyonid, Oxyclaenus. However, in this genus, the internal
margin of the upper molars is more angular and the hypocone is rudimentary; also
the lower molars show important differences. The same specimens also show
similarity within the arctocyonids to the triisodontine, Goniacodon, except for the
poor hypocone development in the latter; M; and Mg in these two genera also show
similarities. However, these resemblances do not seem significant as Opsiclaenodon
shows none of the peculiar adaptive features characteristic of the Triisodontinae.
Structurally, the referred upper molars of Opsiclaenodon are strikingly similar to
some hyopsodontine condylarths such as Haplaletes and Litomylus. In the latter
genus, however, the conules are stronger; tooth outlines are more angular; M3 is
triangular-shaped and lacks a hypocone. Also, P4 has a metaconid and a small low
paraconid. These characteristics (and those of the lower dentition such as P4 witha
metaconid and a small low paraconid) serve to exclude Opsiclaenodon ftom any close
relationship with the Hyopsodontidae.
Amongst the described members of the Pantolestinae, Propalaeosinopa and
Bessoecetor are easily distinguished from Opsiclaenodon by having slender elevated
trigonid cusps; Mg is much larger and bulbous and Py is very elongated with a strong
anterior basal cusp and incipient basining of the talonid ; M!—M? are very slender and
D
50 UPPER EOCENE MAMMALIA
transversely elongated with a smaller hypocone and the outline of Mg is quite
different. In the European species provisionally referred to Bessoecetor, B. levei, the
M2 and M8 are more slender and transversely elongated, with angular parastylar and
metastylat areas; the lower molars are more angular with more elevated trigonids
and the Mg is longer antero-posteriorly than the other molars, with a posteriorly
projecting hypoconulid. The upper molars of Pantomimus are more elongated
transversely with prominent externally projecting metastylar and parastylar areas; a
hypocone is absent. The M? (type) of Pagomomus is more elongated transversely
with more elevated slender cusps and more angular metastylar and parastylar areas;
the hypoconal area is more angular and posteriorly extended. In Pantolestes the
upper molars are similar in shape to Opsiclaenodon, but the hypocone is very small and
the metastylar and parastylar areas are rounded and less prominent; the lower
canine is massive but lacks striations and a groove; the molar paraconid is vestigal
and the hypoconulid is strong and posteriorly projecting, in M3 forming an enlarged
posterior lobe. In Cryptopithecus, the anterior margin of the molars is more rounded
and blunt, with the M,; paraconid smaller, less elevated and less separated from the
protoconid; the trigonid is compressed antero-posteriorly with crowded elevated
cusps; the M; metaconid is more massive and the talonid cusps are better developed
than in Opsiclaenodon with a very strong hypoconid. The Ms in Palaeosinopa is as
large as M2; the molar trigonids are slender and more elevated, especially Mg and the
antero-external cingulum is smaller. The upper molars are more elongated trans-
versely with a small hypocone and the posterior mental foramen lies under Mb.
Buxolestes has a relatively simple canine section; P4 has a more elongated talonid ; Mg
is more rectangular and antero-posteriorly compressed and M3 is enlarged. In the
upper molars of Buxolestes, M? is more elongated transversely with a less rounded
outline and M3 is triangular-shaped and only slightly reduced with a less developed
parastylar area.
Apart from the closely related Dyspterna (see later discussion), Opsiclaenodon
appears nearest structurally to the contemporaneous Cryptopithecus. The earlier
European history of the family is poorly known and none of the described European
species from the Palaeocene and Early-Middle Eocene appear to be closely related to
Opsiclaenodon and Cryptopithecus. The Late Palaeocene and Early Eocene
Palaeosinopa, however, seems to exhibit the basic dental structure from which
Opsiclaenodon and Cryptopithecus could be derived.
Opsiclaenodon major (Lydekker, 1887)
(Text-figs 16-19)
1887 Neurogymnurus major Lydekker: 302
1967 Cryptopithecus major (Lydekker); Van Valen: 87
Dracnosis. As for the genus.
Lectotype. B.M. 29718a. Fragment of right mandibular ramus with Py-M3
and alveoli of the canine and P2-Ps. Talonids badly worn. Lower Headon Beds
(Mammal/Crocodile Bed), Hordle Cliff. Listed Lydekker, 1887 : 302. Figured
Butler, 1946, fig. 1; fig. 2, (2), (9).
HEADON BEDS OF ENGLAND 51
MateriaL. All material is from the Lower Headon Beds (Mammal/Crocodile
Bed), Hordle Cliff, except where otherwise stated.
B.M. 29718. Posterior fragment of left mandibular ramus with M,-M3. The
coronoid, condyloid and angular processes preserved, though damaged. External
surface embedded in matrix. Wear greater than lectotype. Listed Lydekker
1887 : 302. Figured Butler, 1946, fig. I.
B.M. 36801. Fragment ofright mandibular ramus with Mz—-M3. Listed Lydekker,
1887 : 303. Listed Butler, 1946 : 601.
B.M. 36802a. Fragment of right mandibular ramus with alveoli of M,—Ms3.
Listed Butler, 1946 : 691.
B.M. 36803. Left mandibular ramus with Ps, Ms the base of the canine and the
alveoli of Py; and P3-Mz. Teeth unworn. Listed Lydekker, 1887 : 303. Figured
Butler, 1946, fig. 1; fig. 2, (3), (4), (6), (7), (8), (Zo).
B.M. 36807. A crushed fragment of left mandibular ramus with Mz—-M3 and
alveoli of Pg-My. External surface embedded in brown clayey sand matrix. Teeth
unworn. Listed Lydekker, 1887 : 303. Figured Butler, 1946, fig. 2, (1), (5).
S.M. C53478. Isolated right My. Unworn. Damaged metaconid.
B.M. M12565b. Isolated right Mj. Unworn.
S.M. C30611. Fragment of left mandibular ramus with P3—P, and alveoli of Mj,
bases of alveoli of Pp and the canine. Unworn.
S.M. C54162. Isolated teeth. All from Lower Headon Beds (Rodent Bed),
Hordle Cliff.
a. Right Pz. Unworn.
b. Right Mg. Part worn.
c. Right Mg. Part worn.
d. Right Mg. Part worn.
e. Left M;. Part worn.
B.M. 29863. Anterior fragment of left mandibular ramus with the canine,
Py-P,, and alveolus of P;. Tip of canine broken off. Symphysis preserved.
Matrix of brown clayey sand.
B.M. M25094. Anterior fragment of right mandibular ramus with P3 and alveoli
of the canine, Pj, Pe, Paand M;. Posterior part of symphysis preserved.
B.M. 25084. Isolated right lower canine. Tip damaged.
B.M. M25oq1. Isolated right Mg. Moderately worn.
S.M. C30713. Isolated right upper molar (M2?)._ Unworn.
S.M. C30722. Isolated left upper molar (M??). Badly worn.
S.M. C9677. Isolated left upper molar (M3?). Unworn.
S.M. C54163. Isolated right upper molar (M1?). Moderately worn.
Lower Headon Beds (Rodent Bed), Hordle Cliff.
Description. The mandible is moderately elongated and stout with a slight
anterior deepening. The symphysial surface (B.M. Mz5094, B.M. 36803 and
B.M. 29863) is elongated and oval in outline, ending posteriorly below Pa. The
posterior region of the mandible is preserved only in B.M. 29718: the condyle,
situated above the level of the tooth row is damaged, but appears to have been
52 UPPER EOCENE MAMMALIA
transversely elongated. The angular and coronoid processes are separated from the
condyle by fairly prominent notches. The masseteric fossa extends only slightly
below the level of the tooth row; the mandibular foramen is situated similarly. The
anterior mental foramen occurs below P,/canine (B.M. 36803 and B.M. 29718a) or
below P; (B.M. M25094 and B.M. 29863). The posterior mental foramen is also
variably sited below Py (B.M. 36803 and B.M. 29718a) or M, (B.M. 36802a, B.M.
25094 and S.M. C30611). An additional small foramen occurs below P3 (B.M.
29863 and B.M. 36803) or below Pe (B.M. Mz25094).
Fic. 16. Opsiclaenodon major (Lydekker, 1887). Fragmentary right mandibular ramus with
P4-M3. Lectotype (B.M. 29718a) approx. X35. Lower Headon Beds, Hordle Cliff.
(a) Occlusal view. (B) External view. (c) Internal view.
HEADON BEDS OF ENGLAND 53
The lower incisors ate not known. The lower canine is known from B.M. 36803,
B.M. 29863 and B.M. M25084. The upper half of the tooth is enamel covered; this
enamel is wrinkled externally, forming regular striations. The upper half of the
tooth is triangular in section with sharp postero-external, internal and anteto-
internal crests; the latter two crests bound a prominent groove. In B.M. 36803
and B.M. 29863 the groove, crests and enamel covering occur below alveolar level,
but in these specimens, the tooth is incompletely erupted. These features are
considered to occupy the upper half of the tooth, by analogy with material of
Dyspterna (described later) where an almost identical tooth is fully developed and
completely preserved. Butler, in his type description (1946 : 694) observed that
the canine of B.M. 36803 was grooved and incompletely erupted, but did not record
the curious external striations of the enamel and the enamel distribution. In
B.M. 29718a the canine alveolus extends to the posterior end of the symphysis; in
S.M. C30611 and B.M. M25094, it extends posterior to this.
Fic. 17. Opsiclaenodon major (Lydekker, 1887). Fragmentary left mandibular ramus with
C, Pe-Ps. B.M. 29863 3:5. Lower Headon Beds, Hordle Cliff. (a) Occlusal view.
(B) Internal view. (c) External view.
54 UPPER EOCENE MAMMALIA
P, from its alveolus (B.M. 36803, B.M. 29863 and B.M. Mz25094) was evidently
small, single-rooted and placed immediately posterior to the canine. Pz is preserved
in B.M. 36803, S.M. C54162a and B.M. 29863; the tooth is double-rooted and has a
slight forward inclination. From a single large anterior cusp, steep posterior and
postero-internal crests fall to the antero-external and antero-internal extremities,
respectively of a small slightly basined heel. An anterior crest from the main cusp
curves internally at the anterior tooth margin forming a faint cingulum. On the
antero-internal wall of the tooth, just posterior to the anterior crest, there is a vertical
groove, similar to though less pronounced than that of the canine. Horizontal
striations similar to those of the canine, are well developed externally on the main
cusp in B.M. 29863 and S.M. C54162a, though less prominent on B.M. 36803. P3 is
known from S.M. C3061r and B.M. 29863; these specimens were not listed by
Butler (1946) who described only alveoli of this tooth. The tooth is generally
similar to Pg but larger with a large crested main cusp. The anterior crest curves
internally at the base of the antero-internal groove forming a very faint cingulum-
like cuspule (S.M. C30611) or a strong low cusp (B.M. 29863). The posterior crest
continues antero-posteriorly across the middle of the well developed heel. Externally,
there is some development of irregular, subhorizontal enamel striations. Py, (B.M.
29718a, S.M. C30611 and B.M. 29863) is very similar in structure to P3 but much
larger. The antero-internal groove is marked clearly on $.M. C30611 and B.M.
29863, but less prominent on B.M. 29718a. According to Butler (1946 : 695), the
groove is absent in B.M. 29718a, the only specimen containing Py, listed by him.
The antero-posterior talonid crest in S.M. C30611 swells posteriorly to form a cusp;
in B.M. 29863, the median region of the crest is raised as a distinct cusp. Two small
internal cuspules occur on the talonid of B.M. 29718a; these are absent in the other
specimens.
M,-Ms are structurally very similar with rounded trigonid and talonid margins;
M, and Mp are subequal; M3 is markedly smaller. Identification of isolated teeth
Fic. 18. Opsiclaenodon major (Lydekker, 1887). Composite right M;—Mg all approx. x5.
M, (B.M. M12565b) and Mz (S.M.C 54162d). Lower Headon Beds, Hordle Cliff.
(A) Occlusal view. (B) External view. (c) Internal view.
HEADON BEDS OF ENGLAND 55
as M, or Mgis uncertain ; M, has been distinguished here by a slight anterior narrowing
of the trigonid. The metaconid and protoconid are subequal (when unworn),
opposite and transversely crested. The small though distinct low paraconid is
median or slightly internal to the midline, forming the fairly sharp anterior apex of the
tooth; a crest from this cusp meets the anterior protoconid crest. A fairly strong
shelf-like antero-external cingulum is present. The hypoconid and entoconid are
prominent and opposite, the hypoconid being slightly larger; each cusp is united
to the posterior trigonid wall by sharp crests. The hypoconulid is small, though
distinct; it is strongly posterior to the hypoconid and entoconid, and posteriorly
projecting; the cusp is usually slightly external to the midline and is joined to the
hypoconid and entoconid by low crests. This condition is seen in the M3 of B.M.
36807, but this cusp in the Ms of B.M. 29718 and B.M. 36803 is distinct and median.
The talonid is strongly basined. Small accessory crests and cuspules are variably
developed in the molars: in B.M. 36803, B.M. 36807 and B.M. M25og1, a discon-
tinuous crest swings obliquely down the posterior trigonid wall from the metaconid
towards the crista obliqua; this cannot be distinguished in other specimens owing
to excessive wear. On Mz of B.M. 36807 and S.M. C54162d, there is a small cuspule
in the middle of the talonid at the base of the posterior trigonid wall. This is absent
in S.M. C54162b and S.M. C54162c. In other specimens wear is too great for this
feature to be distinguished.
The upper teeth are known only in the isolated state. S.M. C30713 and S.M.
C30722, almost identical structurally, are the largest teeth; these are considered as
M2. S.M. C54163 is somewhat smaller antero-posteriorly, showing several minor
structural differences; this is tentatively considered as M1. S.M. C9677, markedly
smaller in length and width than the other teeth, probably represents M3. M2? is
elongated transversely and has a rounded outline. The antero-internal crescentic
protocone, the largest cusp, dominates the internal half of the tooth. The conical
paracone is subequal in height with the protocone; the metacone is smaller. There
is a strong shelf-like external and postero-external cingulum round the metacone base,
ending externally as a small mesostyle, and a similar anterior cingulum round the base
of the paracone (not present externally). The parastylar region is extended antero-
externally, the metastylar region extended postero-externally, as rounded lobes.
The sharp postero-externally trending protocone crest is interrupted just internal to
Fic. 19. Opsiclaenodon major (Lydekker, 1887). Occlusal view of composite M!-M3 x5.
Right M1 (S.M. C54163), right M2 (S.M. C30713) and left M® (S.M. C9677), drawing
reversed. Lower Headon Beds, Hordle Cliff.
56 UPPER EOCENE MAMMALIA
the metacone by a prominent metaconule. A smaller paraconule is similarly placed
on the protocone crest just internal to the paracone. There is a small low anterior
cingulum and a very strong shelf-like postero-internal cingulum which supports
externally a very strong hypocone which is subequal in height with the metacone.
S.M. C30722, a well worn M2, has‘a more developed anterior cingulum. M1! is
basically similar to M? and although the tooth is more worn than M2 (S.M C307713)
and damaged externally, significant differences can still be recorded: the tooth is
shorter antero-posteriorly than M2. The hypocone and hypocone-cingulum are less
developed; the metaconule is very small and the paraconule, just distinguishable on
the slightly worn crest must have been very rudimentary on the unworn tooth. The
postero-external cingulum round the base of the metacone is absent and there is only
a trace of the antero-internal cingulum. M3 is basically similar to the other molars
but much smaller and compressed antero-posteriorly. The metaconule is very small,
merging with the antero-external cingulum and an internal paracone crest. The
antero-external trending protocone crest joins the paracone; a small swelling on this
crest just internal to the paracone indicates a rudimentary paraconule. A small
postero-external cingulum is present, but not round the metacone base as in M! and
M2; this latter cusp, slightly smaller than the hypocone, is more antero-internal than
in M! and M2.
RemArRKS. There is considerable variation in the depth of the mandible: B.M.
29718a is greatest, B.M. 29718 intermediate, and B.M. 36803 smallest in size. B.M.
36803 has unworn teeth and an incompletely erupted canine, indicating derivation
from a very young individual. This suggests that the variation in mandible depth
is, at least in part, related to individual age differences. There are slight variations
in the positions of the mental foramina; in the degree of prominence of the antero-
TABLE I0
Measurements (in mm) of Opsiclaenodon major
Pe P3 Pa Mi Me M3
a-p tis a-p ts a-p tts) a-p ts a-p ts a-p iis
B.M. 29718 ee
B.M. 29718a —- — — — 45 2:3 40 34 41 3:3 34 26
B.M. 36807 SE OE
BM 36802 —_ — -— — — 4°3 29 Bz 233
B.M. M12565b — es
B.M. 36803 sys rechs} — = =| SF a6
B.M. 29863 FOR oan s 2 45 24 => =| = => — ==
B.M. M25094 — — 36 2:0 = SO
B.M. Mz25091 Se ia ea iS iS SPEECH
S.M. C53478 SS ee), Se OE ee Se
S.M. C54162a 30 17 — — — — — — ~~ ~—- —- —
» on b eee | ee — 44 32 == =
” » C i SS SS SS iS ST OS
oe Sa a ee Se SS a
oF SS SS ee ee Oe a ee Oe ee oe ae
S.M. C30611 — — 37 19 42 24 — — — — — —
HEADON BEDS OF ENGLAND 57
internal groove and antero-external enamel striations in the premolars, and in the
development of accessory cuspules. These have been considered in the preceding
description.
Genus DYSPTERNA Hopwood, 1927
Dracnosis. Lower molars with low blunt cusps, Mg trigonid rather compressed
antero-posteriorly. M, and M3 probably subequal in size, Mz larger. Entoconid
of molars vestigial or absent with postero-internal talonid margin raised as a crest,
hypoconid large. Hypoconulid of molars median and small to vestigial.
TYPE SPECIES. Dyspterna woodi Hopwood, 1927. Early Oligocene (‘Sannoisian’),
Yarmouth, Isle of Wight.
Remarks. The genus was originally based by Hopwood on a single specimen,
from the Lower Hamstead Beds, a very fragmentary worn and damaged mandibular
ramus with M2—M3 (B.M. M13125), provisionally placed in the Oxyclaeninae. Three
years later, Dal Piaz described (1930) a maxillary fragment with P4-M3 which he
tentatively referred to Hopwood’s species. This specimen was removed by Van
Valen (1966 : 88) from Opsiclaenodon as the type of a new apheliscine, Epapheliscus
ttalicus Van Valen, 1966. Dal Piaz also described a new species (1930 : 13) which he
tentatively referred to the genus as ?Dyspterna helbingi; the holotype (and only
known specimen) is a fragmentary left maxilla with P? and the roots of P4-M?.
Recently, Van Valen has observed (1966 : 88) that from the published figure and
description, the affinities of this specimen cannot be determined.
In his redescription of Neurogymnurus major as Opsiclaenodon major, Butler
suggested (1946 : 698) a possible relationship with Dyspterna woodi. The genus was
not listed by Simpson (1945) being based on inadequate material, and McKenna
stated (7m Russell & McKenna, 1961 : 281, footnote 3) that Dyspterna constitutes
practically a nomen dubium. Van Valen (1966 and 1967) referred Dyspterna to the
Pantolestidae.
In the present work, material collected fairly recently from the Upper Headon
Beds has been referred to Dyspterna (on the basis of molar structure) as a new species.
The anterior lower dentition of this species of Dyspterna is strikingly similar to that of
Opsiclaenodon and the two genera appear to be closely related, thus supporting
Butler’s original view.
Opsiclaenodon is distinguished generically from Dyspterna by the uncompressed
Mg trigonid; the molar entoconid is strongly developed; the hypoconulid is well
defined and the cusps are more trenchant and separated. The prominent hypoconid
and vestigial or absent entoconid sharply distinguishes Dyspterna from all other
pantolestines. These and other specialised features could have arisen by modification
of a dental structure similar to that of Opsiclaenodon.
Van Valen has stated (1967 : 227) that Opsiclaenodon could be directly ancestral
to Dyspterna. This possibility is precluded by the contemporaneous occurrence of
Dyspterna hopwoodi sp. nov. and Opsiclaenodon major in the Headon Beds; both
genera were probably derived from an earlier common ancestry, perhaps in the Middle
Eocene.
58 UPPER EOCENE MAMMALIA
Dyspterna hopwoodi sp. nov.
(Text-figs 20, 21)
ErymoLocy. After A. T. Hopwood.
Dracnosis. My, and M3 hypoconid large, entoconid and hypoconulid vestigial,
appearing as small swellings on postero-internal marginal crest. Crista obliqua in
M, and Ms directed anteriorly to base of prococonid. Mg trigonid more compressed
antero-posteriorly than in D. woodt, with the small paraconid strongly antero-internal
in position, close to metaconid; M3 hypoconulid vestigial. Sharp condyloid and
coronoid crests bordering deep masseteric fossa.
HototyrPe. B.M. M26052. Right mandibular ramus with the canine, P2—P4 and
Ms, and alveoli of I;-Is, P1 and Mj-Mz. Symphysis and posterior region of mandible
are preserved. Slightly worn dentition. Upper Headon Beds (Lignite Bed),
Headon Hill.
PARATYPES. B.M. M26053. Isolated right My. Slightly worn. Upper Headon
Beds (Lignite Bed), Headon Hill.
B.M. M26054. Isolated right P}. Unworn. Upper Headon Beds (Lignite Bed),
Headon Hill.
DESCRIPTION. The mandible is moderately stout with a slight anterior deepening
as in Opsiclaenodon. The symphysis is elongated and rectangular in outline, ending
posteriorly under Pp: and inclined at about 45° to the level of the tooth row. The
posterior region of the mandible is complete in the holotype, except for the condyle
which lay just above the level of the tooth row. The coronoid process is well developed
and massive, the anterior edge rising steeply from the tooth row level. The notch
between the condyle and the coronoid is hardly perceptible; this is in contrast to
Opsiclaenodon, where it appears to have been more developed. The angular is
fairly prominent and well separated from the condyle; the lower edge is deflected
internally as a small flange. The mandibular foramen is situated just below alveolar
level; the ridge above this running from the condyle to a position below Mg is more
prominent than in Opsiclaenodon. The anterior mental foramen is sited below the
canine/P;, the posterior mental foramen below Mj.
The bases of the alveoli of I;-Is indicate that these teeth were closely crowded
and situated immediately anterior to the canine. The canine is almost identical to
that of Opsiclaenodon major with the upper half of the tooth enamel covered;
externally the enamel is wrinkled, forming regular striations. The upper half of the
tooth is triangular-shaped with sharp antero-internal, internal and postero-external
crests, and a prominent deep antero-internal groove. The groove and crests are
confined mainly to the enamel-covered upper half of the tooth. Below this level,
the crests become rounded but the groove, in contrast to Opsiclaenodon major is
continued to alveolar level as a distinct but rounded constriction. From the
alveolus in the holotype, it appears that P, was a little separated from the canine;
the tooth (B.M. M26054) is single-rooted with a main central cusp and a small postero-
internal heel. The cusp is crested anteriorly, postero-internally and posteriorly.
There are small but distinct antero-internal and postero-external cingula and a
cingulum trace antero-externally. A rudimentary antero-internal groove is present.
HEADON BEDS OF ENGLAND 59
Py is slightly larger than P; and double-rooted. There are no antero-external and
postero-external cingula; otherwise the tooth is structurally nearly identical to Pj.
There is the suggestion of a slight antero-internal groove. Pg is identical to Pe,
except for a larger size and a slightly stronger cusp-like antero-internal cingulum.
There is a trace of an antero-internal groove. Py, is structurally quite similar to Ps
but much larger. The sharp anteriorly crested main cusp falls to a prominent low
anterior cusp; this latter cusp is larger than that of Opsiclaenodon and the tooth is
generally more pointed. There are small antero-external and antero-internal
cingula. The antero-posterior talonid crest ends posteriorly inasmall cusp. There
is a slight antero-internal groove.
It is difficult to estimate reliably the relative sizes of My and Mg from the alveoli in
the holotype of D. hopwoodi because of mandible damage. By comparison with the
holotype of D. wood: (about the same size as D. hopwoodt) where M3 is smaller than
Fie. 20. Dyspterna hopwoodi sp. noy. Right mandibular ramus with C, Ps—P4 and Ms.
Holotype (B.M. M26052). Upper Headon Beds, Headon Hill. (a) Occlusal view x2°5.
(B) Internal view x2.
60 UPPER EOCENE MAMMALIA
Mg, it has been concluded that an isolated molar (paratype) of D. hopwoodi is best
considered as My. This latter tooth is elongated with a rounded margin. The
protoconid and metaconid are subequal, opposite and transversely crested. The
small but distinct low median paraconid forms the sharp anterior apex of the tooth.
There is a moderately strong antero-external cingulum and a smaller antero-internal
cingulum. The entoconid and hypoconid are vestigial, appearing as small swellings
on a low curved crest which forms the postero-internal talonid margin. The
median hypoconulid projects slightly posteriorly; the entoconid and hypoconid are
opposite each other. The talonid is slightly basined. Msg is almost the same length
as Mj, with a subequal and opposite protoconid and metaconid. The smaller
paraconid is strongly internal and anterior to the metaconid; the trigonid appears
antero-posteriorly compressed. The almost straight antero-external trigonid margin
is formed by a paraconid and protoconid crest. There is a small antero-external
cingulum but the antero-internal cingulum is absent. The vestigial hypoconulid is
represented by a minute rounded median lobe which is posteriorly projecting and
separated from the larger hypoconid. The entoconid is vestigial; a low rounded
crest runs anteriorly from the hypoconulid-lobe, forming the postero-internal tooth
margin. The talonid is only very slightly basined.
Remarks. In the canine of the holotype (B.M. M26052) distinct wear facets are
visible: part of the postero-external face has been gauged away by the upper canine
in occlusion, to form above alveolar level, a prominent oblique baso-exteriorly
directed shelf. Wear has taken place along the whole vertical length of the tooth
from the tip to this shelf, but becomes progressively greater towards this latter
position; presumably slight wear took place near the canine tip when the tooth
was partially erupted (compare Opsiclaenodon major, B.M. 29863). Anteriorly and
basally to the prominent groove, there is a shallow vertical oval facet, produced
probably by friction against one of the upper incisors.
Fic.21. Dyspternahopwoodisp.nov. Right P; (B.M.M26054) and right Mz (B.M. Mz6053).
Both x4. Upper Headon Beds, Headon Hill. (a) Occlusal view. (8) Internal view.
(c) External view.
HEADON BEDS OF ENGLAND 61
The antero-internal groove does not show any signs of wear; the bordering crests
are very sharp and quite unworn. The same situation exists in the closely similar
canine of Opsiclaenodon major. As Butler noted (1946 : 695), a similar unworn
groove occurs on the lower canine of Nasua, a recent procyonid. No information has
been found about the significance of this groove in Nasua. On the available evidence,
the structure in all three genera would appear to be non-functional, at least in
occlusion.
In molar structure, D. hopwoodi is closely related to the type species, D. woodi
and may be ancestral to the latter. D. hopwoodi sp. nov. appears specifically distinct
on the basis of the following differences: in D. woodi, the hypoconid crest is directed
antero-internally to meet the base of the metaconid, in Mg running up the posterior
wall of the metaconid; the cingula are relatively larger and more shelf-like; the M3
trigonid is less compressed antero-posteriorly with the paraconid slightly more median ;
the hypoconulid appears to have been relatively larger, and the coronoid and condyloid
processes are lower and more rounded, enclosing a shallower masseteric fossa.
More detailed comparison is not possible because of the poor preservation of the
holotype of D. woodi.
TABLE II
Measurements (in mm) of Dyspterna hopwoodi
Py Pe P3 Pa My Mz Ms
a-p ts ap ts a-p tr a-p trs a-p trs a-p trs a-p trs
B.M. M26052 — — 3:6 2:3 4:7 298 66 3:3 — — — — 5:0 3:6
B.M. M26053 — 3 — ~~ —~ —~—~ ~~ ~~ ~~ 54 37 —- ~—- —~— —
B.M. M26054 3°44 29 — — — - -—- —_—- —
Family APATEMYIDAE Matthew, 1909
DiacGnosis. Genera with a reduced dental formula—probably Z 4 5, ;. Jaw
deep and stout in proportion to molars, with posterior mental foramen below M, or
Mz. Marked soricid-like development of lower incisors, the roots of which terminate
below Mg. Pg blade-like and single-rooted. Molars in early forms trenchant; more
typically, bunodont and primate-like in appearance. P%—P4 double-rooted and
unicuspid. Upper molars relatively elongated antero-posteriorly and broadly
expanded postero-internally with a well developed hypocone, but lacking a mesostyle.
Lower molars, except for Mz, have a simple talonid but the trigonid, particularly in
M2-Msz shows a distinctive parallelogram arrangement of cusps and crests. (Mainly
after Gazin, 1958).
Remarks. Of the seven genera assigned to the family, five are known from North
America and two from Europe:
Jepsenella Simpson, 1940. Middle Palaeocene; North America.
Labidolemur Matthew & Granger, 1921. Late Palaeocene; North America.
Apatemys Marsh, 1872. Early-Late Eocene; North America.
62 UPPER EOCENE MAMMALIA
Stehlinella Matthew, 1929. Late Eocene; North America.
Sinclairella Jepsen, 1934. Early-Middle Oligocene; North America.
Eochiromys Teilhard de Chardin, 1927. Early Eocene; Belgium.
Heterohyus Gervais, 1848-52. Middle-Early Eocene; Europe.
Van Valen (1967) considered Teilhardella Jepsen, 1930 was invalid and included it
within Apatemys. In addition to the above generic list, McKenna (1963 : 30)
suggested that Unuchinia Simpson, 1937, previously placed incertae sedis in the
Insectivora, might be an apatemyid. More recently, Van Valen (1967) formerly
erected a new subfamily Unuchiniinae Van Valen & McKenna, for the reception of
Unuchinia.
Stehlinella is known from the anterior portion of the skull with part of the upper
dentition and an almost complete mandibular ramus lacking only Py. Sznclairella
is known from a complete but crushed skull with part of the upper dentition and a
mandibular ramus lacking P3-P, and Mj. The remaining six genera are based on
fragmentary mandibles, although Heterohyus is known also from a palatal fragment.
Owing to incomplete material and the comparative rarity of specimens, classifica-
tion is very tentative. Some genera are probably composite but better material is
needed before significant advances can be made.
The family has oscillated between Primates and Insectivora for many years.
The most recent detailed work on the affinities of this problematical group is McKenna
(1963). From a consideration of molar structure in the earliest genera, McKenna
referred the family to the Insectivora; this view, followed by Van Valen (1967) is
adopted here.
Genus HETEROHYUS Gervais, 1848-52
Diacnosis. Usually single mental foramen below Mg; fossa below Py, generally
reduced. Enamel covering lower incisors extends into alveolus but does not
completely coat the crown; P,4 generally reduced ; Mg talonid usually strongly elongated
and Ms posterior trigonid wall usually oblique to longitudinal axis of tooth.
TYPE SPECIES. Heterohyus armatus Gervais, 1848-52. Middle Eocene (Early
Lutetian) ; Bouxwiller, France.
ReMARKS. Distinguished from the other genera as follows: Jepsenella shows only
incipient development of apatemyid characteristics; in Eochiromys and Labidolemur
the incisor crown is completely covered by enamel. These genera and all other
apatemyids possess either a little-elongated Mg talonid or lower molars with a
transverse posterior trigonid wall.
Five species have been distinguished:
H. armatus Getvais, 1848-52, pl. 35, fig. 14. Middle Eocene (Early Lutetian) ;
France.
H. quercyi (Filhol, 1880), p. 174, figs 1-3. Late Eocene (Early Ludian, Quercy
Phosphorites) ; France.
H. nanus Teilhard de Chardin, 1922, p. 93, fig. 41; pl. 4, fig. 21. Late Eocene
(Early Ludian, Quercy Phosphorites) ; France.
HEADON BEDS OF ENGLAND 63
H. europaeus (Riitimeyer, 1890), p. 346, figs 1-2. Middle Eocene (Early-Middle
Lutetian) ; Switzerland.
H. heufeldert Heller, 1930, p. 32, pl. 5, fig. 4. Middle Eocene (Lutetian) ; Germany.
The most complete species, H. guercy1, is based on a crushed skull and almost com-
plete upper dentition and a fragmentary mandibular ramus; H. manus and H.
aymatus are based on mandibular fragments, H. heufelderi on a complete lower
dentition. The type of H. aymatus (the type species) was not located by Teilhard de
Chardin (1922 : 89) and here, is considered to be lost.
Stehlin (1916 : 1462) erected the genus Heterochiromys for two species from
Egerkingen: H. gracilis known from mandibular rami lacking teeth and isolated
incisors, and H. fortis based on isolated incisors. It is highly probable, as suggested
by Teilhard de Chardin (1922 : 92), that Heterochiromys gracilis Stehlin is synonymous
with Heterohyus quercyi (Filhol) and Heterochivomys fortis Stehlin, is synonymous
with Heterohyus armatus Gervais. Stehlin (1916 : 1434) also placed several mandi-
bular rami lacking teeth, and isolated incisors from Egerkingen in a new genus and
species, Amphichiromys europaeus. This was based on minor differences in the
incisors. Isolated lower molars tentatively assigned by Stehlin to this genus are of
characteristic heterohyid type and apparently do not warrant generic separation.
The genus is considered synonymous with Heterohyus.
The generic diagnosis is unsatisfactory and probably, Heterohyus is a composite
genus. H. nanus does not fit well into this diagnosis and appears to be indistinguish-
able from Apatemys bellus. American material has not been examined, so this view
is only tentative.
The material described below, constitutes the first record of an apatemyid from the
Tertiary of Britain.
Heterohyus sp.
(Text-fig. 22)
Material. S.M.C54172. Isolated left M3. Moderately worn. Lower Headon
Beds (Rodent Bed), Hordle Cliff.
DEscrIPTION. This strongly elongated tooth is composed mainly of a gently
basined talonid, the latter being 2} x the length of the trigonid. The trigonid is
wider than the talonid and externally projecting. The low blunt cusps and crests
of the trigonid exhibit a distinctive parallelogram arrangement.
The metaconid and paraconid are almost equal in size, the antero-internal paraconid
being directly anterior to the metaconid ; these cusps are united by a massive antero-
posterior crest. The smaller protoconid is moderately separated from the metaconid
and paraconid, and more anteriorly placed in relation to the metaconid; the posterior
trigonid wall is thus markedly oblique to the longitudinal axis of the tooth. From
the paraconid a small low crest runs parallel to the posterior trigonid wall, forming
the oblique anterior tooth margin. The antero-external extremity of the tooth,
directly anterior to the protoconid, is sharp and acute-angled ; here, the crest is very
faint and low.
64 UPPER EOCENE MAMMALIA
The talonid basin is bounded externally and internally by rounded fairly strong
crests which are almost parallel. The higher crista obliqua meets the posterior
trigonid wall midway between the metaconid and protoconid; the internal crest is
continued faintly up the posterior wall of the metaconid. The internal talonid
margin is damaged; it may have curved out internally, with a more rounded outline
than the external margin. Around the posterior edge of this damaged area is the
possible trace of a small entoconid. From this position, the straight talonid margin
runs to a relatively high rounded externally shifted hypoconulid-lobe. From the
entoconid, a minute crest dilates as it curves round to meet a prominent hypoconulid,
in the anterior part of the hypoconulid-lobe. No hypoconid is distinguishable,
although the tooth is damaged anterior to the hypoconulid; a small cusp may have
been present.
The total antero-posterior length of Mg is 5-8 mm; the trigonid length is 1-6 mm,
that of the talonid 3-7 mm. The trigonid transverse width is 2-9 mm, the talonid
width is 2-2 mm.
REMARKS. This specimen is distinguished from the Mg of known species as follows:
H. nanus is much smaller with a short talonid and a slightly oblique trigonid; H.
quercyt is much smaller with a slightly oblique trigonid; H. europaeus is larger with a
slightly oblique trigonid; H. heufelderi is smaller with a relatively shorter talonid
and in H. armatus the talonid is less elongated and wider transversely than the tri-
gonid. Also, the tooth outline in all the above is more prominently rounded.
The Headon Beds specimen appears closest to H. armatus. However, little is
known about variation within the genus; the known species are restricted to the
Fic. 22. Hetervohyus sp. Left M3. S.M. C54172 approx. x 8. Lower Headon Beds, Hordle Cliff.
(A) Occlusal view. (8B) Internal view. (c) External view.
HEADON BEDS OF ENGLAND 65
Lutetian and Early Ludian and the genus already may be oversplit. S.M. C54172
could represent a distinct new species but better material is needed. At present the
specimen seems best recorded as Heterohyus sp.
TABLE 12
Measurements (in mm) of M3 of Hetevohyus species
Angle between
Ratio of posterior
trigonid wall of
length to trigonid and
Total a-p talonid longitudinal
length length axis
H. heufelderi (ex Heller, 1930) 4:0 — =
H. quercyi (ex Teilhard de Chardin, 1922) 3°9 1:24 100°
H. nanus (ex Teilhard de Chardin, 1922) 2°5 r:14 105°
H. armatus (ex Gervais, 1848-52) 7:0 1653 7 130°
H. europaeus (ex Stehlin, 1916) 9:0 ig BOs T00°
H. sp. (S.M. C54172) 5:8 T 224 120°
Order PRIMATES
Family ADAPIDAE Trouessart, 1879
Diacnosis. Dental series usually continuous—no marked diastema. Incisors
typically wide-crowned and trenchant; upper canines straight, lower canines short,
subpremolariform. Upper molars tritubercular or quadritubercular with a large
V-shaped protocone; small but true hypocone derived from posterior cingulum;
mesostyle usually absent. Paraconid absent or much reduced on lower molars;
entoconid retarded; hypoconulid absent or rudimentary except on Mg. Mandible
generally short and stout with a greatly expanded angular and a high recurved
coronoid process. Condyle elongated transversely and relatively flattened. Skull
usually with high sagittal and lambdoidal crests. Small ovoid braincase. Prominent
heavy facial region. (after Hill, 1953).
REMARKS. Simons (1962) presented a revised classification of European Eocene
primates. This revision, together with Gazin’s revival (Gazin, 1958 : 31) of the
family Notharctidae Trouessart, 1879, has significantly altered Simpson’s classifica-
tion of 1945. The generic list follows Simons (1962) with the addition of Lantianus
Chow, 1964.
Adapis Cuvier, 1822. Middle-Late Eocene; France, Britain, Switzerland.
Pronycticebus Grandidier, 1904. Late Eocene; France.
Anchomomys Stehlin, 1916. Middle-Late Eocene; France, Switzerland.
Caenopithecus Ritimeyer, 1862. Middle Eocene; Switzerland.
Protoadapis Lemoine, 1878. Early-Late Eocene; France, Germany.
Gesneropithex Hiirzeler, 1946. Late Eocene; Switzerland.
Lantianus Chow, 1964. ?Late Eocene; China.
66 UPPER EOCENE MAMMALIA
In addition to Caenopithecus, Simpson, (1945) considered two further genera as
Adapidae incertae sedis: Aphanolemur Granger & Gregory, 1917 and Amphilemur
Heller, 1935. Aphanolemur is now believed to be a synonym of Smilodectes Wortman,
1903, anotharctid (Gazin, 1958 : 30). . Amphilemur has been considered by McKenna
(1960 : 58) as an erinaceoid insectivore. More recently, Simon’s generic list has been
accepted by McKenna (1967) and Russell, Louis & Savage (1967) although the latter
authors regarded reference of Gesneropithex as provisional and tentatively added
Periconodon Stehlin, placed by Simons (1962) and McKenna (1967) in the Omomyidae.
Genus ADAPIS Cuvier, 1822
Synonomy. See Hill, 1953 : 464.
Diacnosis. Mandible has a greatly expanded angular process; coronoid process
separated from condyle by a fairly shallow notch. P ; present, small in size; P4
submolariform but cusps variably developed ; Mj and Mg similar to Py; Mg considerably
elongated by hypoconulid which is not isolated distinctly as a third lobe; metastylid
present in later species. P! present, small in size; P4 molariform with a bicusped
ectoloph. Upper molars lack mesostyle; hypocone not strongly developed, absent
in M%; metaconule rudimentary or absent; strong external cingulum; internal
cingulum continuous.
TYPE SPECIES. Adapis parisiensis (Blainville, 1841). Late Eocene (Late
Ludian); France.
REMARKS. Anchomomys, Pronycticebus, and Caenopithecus differ from Adapis by
having the Mg hypoconulid more distinctly separated as a lobe and by lacking the
bicusped ectoloph of P, and the continuous internal upper molar cingulum.
Caenopithecus is further distinguished by lacking P! and having a mesostyle. In
Protoadapis Pz is much smaller than Pg and the molar paraconids extend more
internally. Gesneropithex has a median paraconid; the metaconid—protoconid
crest is perpendicular to the longitudinal axis of the tooth and the Mg trigonid is
compressed antero-posteriorly.
The following species are referred to Adapis:
A. parisiensis (Blainville, 1841), p. 112, pl. 9. Late Eocene (Bartonian-Late
Ludian) ; France.
A. magnus Filhol, 1874, p. 11, pl. 8, figs 9-12. Late Eocene (Bartonian-Early
Ludian) ; France, Britain.
A. ruetumeyert Stehlin, 1912, p. 1261, pl. 21, figs 5, 8, 12, 14. pl. 22, figs 17-19.
Middle Eocene (Late Lutetian) ; Switzerland.
A. sciureus Stehlin, 1916, p. 1514, text-figs 368-370, pl. 22, fig. 5. Middle Eocene
(Early Lutetian) ; Switzerland.
A. priscus Stehlin, 1916, p. 1511, text-figs 366-367. Middle Eocene (Early
Lutetian) ; Switzerland.
Simons (1962) observed that the holotype of Adapis minimus Heller, 1930, is
indeterminate owing to removal of enamel by post-mortem chemical absorption.
HEADON BEDS OF ENGLAND 67
Adapis magnus Filhol, 1874
(Ble i e lee tie 1)
Diacnosis. Large size. Molars moderately elongated; metastylid developed.
Upper canines large with grooved crown; P? transversely elongated with prominent
internal cingulum and talonid.
Hototypre. Fairly complete skull lacking the incisors and canines of both sides,
left P!-P2 and M3. ?Late Eocene/Early Oligocene (Quercy Phosphorites) ; Raynal,
France. Described and figured, Filhol, 1874, p. 11, pl. 8, figs 9-12. Specimen not
located.
MaTeERIAL. All specimens from the Lower Headon Beds (Mammal/Crocodile
Bed), Hordle Cliff.
B.M. 30323. Fragment of right mandibular ramus with M\—-M3. Very slightly
worn. Brown shelly sand matrix. Listed, Lydekker, 1885a : 263.
B.M. 30856. Fragment of right mandibular ramus with Mz-M3. Moderately
worn. Brown sand matrix. Listed, Lydekker, 1885a : 263.
B.M. 29864. Fragment of right mandibular ramus with P,—P), and posterior
root of P3. Moderately worn. Listed, Lydekker, 1885a : 263.
B.M. 29741. Fragment of left mandibular ramus with Py, M3, roots of Pz and
part of the canine. Teeth and alveoli of P;-Py are badly damaged. External
surface embedded in brown clayey sand matrix. Listed, Lydekker, 1885a : 263.
B.M. Mio260. Fragment of left mandibular ramus with P3—Mg and alveoli of M3.
Fairly well worn.
B.M. Mi1o269. Fragment of right mandibular ramus with Pa-Mz. Teeth unworn.
B.M. M10267 (3). Fragment of left mandibular ramus with Mj-Mz. Moderately
worn.
D.M.S.W. M556 (41). Fragment of left mandibular ramus with Py—Mg.
B.M. M26057 (1). Right Mp.
(2). Right Mj. Both teeth slightly worn. Wear and preservation
indicates probable derivation from same individual.
B.M. M20560. Right Mg. Badly worn.
B.M. M25093. Left M3. Badly damaged.
S.M. C30680. Fragment of right mandibular ramus with M,;—-Mz. Slightly worn.
S.M. C53483. Left M3. Unworn
S.M. C53477. Right Ms. Moderately worn.
S.M. C54221. Right Me. Unworn.
S.M. C30659. Right My. Moderately worn.
S.M. C30660. Right My. Well worn.
S.M. C30656. Left My. Well worn.
S.M. C30658. Right Mz. Well worn.
S.M. C30657. Right My. Well worn.
S.M. C31345. Left Me (?). Well worn and badly damaged.
B.M. M3572. Fragment of left maxilla with left half of palate and P2-P?, M1-M3
andalveoliofthecanineand P1, P2isdamaged. Teethunworn. Listed, Lydekker,
1887 : 299.
68 UPPER EOCENE MAMMALIA
B.M. Mroz261 (1). Left upper molar (M2?). Moderately worn.
2). Right upper molar (M2?). Moderately worn.
). Upper molar. Moderately worn but badly damaged.
). Left upper molar (M?2?). Slightly worn.
). Left P4. Moderately worn.
S.M. C53484. Left upper molar (M??). Unworn.
S.M. C30661. Right upper molar (M??). Slightly worn.
S.M. C30662. Right upper molar (M!?). Slightly worn. Probably from same
individual as S.M. C30661.
B.M. M25098. Right upper molar (M??). Unworn.
B.M. 25092. Right upper molar (M!?). Moderately worn.
S.M. C53479. Left upper molar (M??). Moderately worn.
B.M. 12564 (1). Left upper molar (M??). Slightly worn.
(2). Right P4. Moderately worn.
M. C30681. Right upper molar (M3?). Well worn.
M. C30679. Left upper molar (M!?). Badly worn.
M. C30673. Left upper molar (M2?). Unworn.
M. C30674. Left upper molar (M!?). Slightly worn. Probably from same
vidual as S$.M. C30673.
M. C30678. Left P4. Moderately worn.
M. C30675. Left upper molar (M2?). Badly worn.
M. C30676. Right upper molar (M?2?). Slightly worn.
.M. C30677. Fragment of upper molar.
B.M. 30346a. Listed, Lydekker, 1885a : 263.
(1) Right upper molar (M?2?). Slightly worn.
(2) Left upper molar (M??). Slightly worn. Probably from same
individual as B.M. 30346a (I).
(3) Left P4. Unworn.
(4) Left upper molar (M??). Unworn.
(5) Right upper molar (M1?). Moderately worn but badly
(
;
B.M. M20560 (1
(2
K
NNNNRNANMNW
damaged.
(6) Left upper molar (M1?). Unworn but slightly damaged.
(7) Left upper molar (M??). Unworn but slightly damaged.
Probably from same individual as B.M. 30346a (6).
(8) Left upper molar (M3?). Unworn.
(9) Left P8. Shghtly worn.
B.M. M25096. Right P2. Slightly worn.
B.M. M10267. (1) Left P?. Unworn.
(2) Right upper canine.
DEscRIPTION. The base of the strong coronoid crest and the shallow masseteric
fossa are preserved in B.M. M1o260. A small mandibular foramen occurs just below
alveolar level. The posterior mental foramen lies between P3 and Pa.
The lower incisors and P;—Ps are unknown from the Headon Beds. The lower
canine is known (B.M. 29741) only by the root. Ps, preserved only in B.M. M1o260,
HEADON BEDS OF ENGLAND 69
is oval and double-rooted, with an almost continuous cingulum; it has a main crested
cusp which falls to a small anterior cusp and a single-cusped postero-internal talonid.
Py (unworn in B.M. M10269) is moderately molarised with the protoconid larger
than the metaconid. The metaconid is median and posterior to the protoconid.
A slight swelling on the postero-internal wall of the metaconid (distinct in B.M.
M10269 but indistinct in B.M. Mro260) indicates a rudimentary metastylid. These
cusps, which are crested transversely, form the oblique posterior trigonid wall.
There are small external and antero-external cingula. A straight crest from the
protoconid falls antero-basally to a small median paraconid which merges into a
prominent antero-internal cingulum. From the prominent hypoconid, a crest runs
antero-internally (crista obliqua) to meet the posterior protoconid wall, and internally
to join a small but distinct entoconid (posterior to the hypoconid). A low crest
from the entoconid runs anteriorly to the base of the metaconid. In B.M. Mro260,
the P, entoconid is vestigial.
In M,, protoconid and metaconid are subequal when unworn. A metastylid is
well differentiated, although it is indistinct in B.M. M1o0260; this cusp, with the
crested paraconid and metaconid forms the strongly oblique posterior trigonid wall.
From the protoconid a crest curves antero-internally to meet the paraconid external
to the midline. This latter cusp, as in Py, merges with a strong antero-internal
cingulum. There are faint antero-external, external and postero-external cingula.
The hypoconid is slightly larger than the more posteriorly situated entoconid; these
cusps are joined by crests to a small but distinct median hypoconulid which is
slightly posterior to the entoconid. There is a prominent crista obliqua. Mg js
closely similar to Mj, except for its larger size. The trigonid of Mg is almost identical
with that of M; and Mg, except for a generally poorly developed metastylid (in B.M.
30323 and B.M. 30856, for example). The hypoconid is strongly developed (as in
M,—Mg), with a prominent crista obliqua. The entoconid is vestigial (B.M. 30856)
or represented by a low swelling (B.M. 30323) posterior to the hypoconid. The
median hypoconulid (external to the midline in S.M. C53483) is as large as the
hypoconid and surmounts a large rounded posteriorly projecting hypoconulid lobe
which greatly extends the length of the talonid. From the hypoconulid, a low
crest curves antero-internally, forming the internal talonid margin. The hypo-
conulid lobe is well separated from the hypoconid.
The upper canine alveolus (B.M. M3572) indicates a fairly large curved tooth.
An isolated canine (B.M. Mro267 (2)) has an almost continuous cingulum,
more prominent internally. The tooth is grooved vertically on the anterior and
posterior faces. P! is known from the Headon Beds only by an alveolus (B.M.
M3572) indicating a very small single-rooted externally placed tooth. P?is oval and
obliquely set in the tooth row (B.M. 25096) with an almost continuous cingulum
which is strong generally except antero-externally. There is a large external
anteriorly-crested cusp, and a small internal rounded lobe which is variably basined.
P3 is larger than P2, with a large antero-external cusp and a smaller antero-internal
cusp. The tooth is set obliquely in the jaw as in P2. The antero-external cusp is
crested anteriorly and posteriorly. There is a distinct parastyle which is connected
to the protocone by a sharp crest. A cingulum is absent round the protocone but
70 UPPER EOCENE MAMMALIA
elsewhere it is prominent (although very reduced in B.M. 30346a (9)). P4is smaller
than M1 but larger and more quadrate than P%, and set more squarely in the tooth
row. The paracone is slightly larger than the metacone; these cusps are sharply
crested anteriorly and posteriorly. The protocone and metacone are subequal. As
in P3, a sharp crest runs from the protocone to a distinct parastyle. The cingulum
is generally well developed but absent anteriorly.
M?! is roughly quadrate in outline and larger than P4. Externally and anteriorly,
the tooth is almost identical with P4, except for a small paraconule midway along the
protocone-parastyle crest and a stronger cingulum round the protocone base. There
is a hypocone sited slightly internal to the protocone and opposite the metacone;
this cusp, the smallest of the major cusps, is joined to the posterior cingulum by a low
curved crest. M2 is larger than M1, but otherwise structurally nearly identical.
M® is similar but slightly narrower antero-posteriorly than M2. The M? internal
TABLE 13
Selected measurements (in mm) of Adapis magnus
P3 Pa M, Me M3
a-p trs a-p trs a-p trs a-p trs a-p trs
B.M. 30323 Se EG BE OH OFS
B.M. 30856 TO— 5:4 Cay
B.M. 29864 — = 6:4 4°8 Felt 5:1 — — ae iad
B.M. M10260 6:3 4:0 7p 4°4 7:0 4°8 7:6 533 — —
B.M. M10269 —— — 6:8 4:8 6:9 5°7 7:2 6:1 — =
B.M. M10267 (3) = = = — 67 5:0 73 54 oe =
D.M.S.W. M556 (41) — — aes 5°6 72 5-0) 8:0) 5-6 -- —
B.M. M26057 (1) — — 76 533 ae —_
B.M. M26057 (2) 7I 8646 == = sess =
B.M. M25093 — = os —_ — — = = 8-7 4:2
S.M. C54221 = a = = = = Q-7 5-7 = =
S.M. C53483 —- -—- |S Fe Oe aD 452
S.M. C53477 ee aS
ips p4 M1 M2 M3
a-p trs a-p trs a-p trs a-p trs a-p trs
B.M. 3572 4°7 5°8 —— — 6:7 FE, Ff) tI G20) 18°8
B.M. M1o261 (1) — — = = == ae 69 ©6883 Se el
B.M. Mroz261 (2) — = = — = =) 0m 53 nas 2s
B.M. M20560 (1) — = = = oe, a= 75 83 eX =
B.M. M20560 (2) —— — 62 7°5 == = — —_ ee =
B.M. M12564 (1) — == —_ == = aa 63 SI = oly
B.M. M12564 (2) — — 6-1 7:2 = —_ = = =—_ =
S.M. C30678 — Gs ae is ae —
$.M. C30674 — — — = 75 9:7 — = —_ =
S.M. C30679 = = — = 6:9 8-3 el _ = il
S.M. C30681 — = = —_ == —_ 770-88
S.M. C53479 = = = = = = = — =O) ny
HEADON BEDS OF ENGLAND 71
cingulum in B.M. 30346a (2), is very rudimentary. In some specimens, the antero-
internal molar cingulum is continuous with a crest from the hypocone; in other
specimens thisisnotso. This feature is also variable in M1—M of the same individual,
as in B.M. 3572.
Remarks. The species has long been recognised from the Lower Headon Beds
of Hordle Cliff. Adapis magnus (sensu lato) is subject to considerable variation and
several distinct varieties have been recognised (Stehlin, 1912).
The species is distinguished from other species as follows: A. sciwreus has shorter
rounded molars; the protoconid-metaconid crest is only slightly oblique and there is
no metastylid; P3 is more simple and Py less molarised with a smaller talonid; the
upper molars have a strong metaconule and protoconule and a stronger hypocone.
A. priscus possesses rounded less elongated molars; the protoconid-metaconid crest is
only slightly oblique and there is no metastylid. In A. ruetimeyert the rounded
molars lack a metastylid; the protoconid-metaconid crest is only slightly oblique
and there is a strong hypocone. A. farisiensis is much smaller with the lower
molars relatively more elongated antero-posteriorly; the upper molars are less
extended transversely; P? is extended antero-posteriorly with a smaller internal
cingulum and talonid, and the upper canine is smaller and lacks grooves.
Adapis parisiensis (Blainville, 1841)
(Pl. 2, figs 2, 3)
Dracnosis. One third smaller than A. magnus, with molars relatively more
elongated ; metastylid variably developed. Upper canine small, generally lacking a
grooved crown; P? elongated antero-posteriorly with small talonid and internal
cingulum.
Hototyre. A skull, according to Hill (1953), now in the Paris Museum. Late
Eocene (Late Ludian); France. Described and figured, Blainville, 1841, p. 112,
pl. 9 (Genus Anoplotherium).
MATERIAL. F.C.S. 7.
(1) Fragment of left mandibular ramus with M;-M3. Msg is damaged.
(2) Fragment of right mandibular ramus with P2—Mj, Ms, and alveoli of the canine,
P; and Mg. Part of the symphysis is preserved.
(3) Right M2-talonid only.
(4) Left P2 (?).
(5) Left P4 with roots embedded in a maxillary fragment.
(6) Left M1 (?)—internal fragment with protocone only. All specimens from the
same individual. Teeth moderately worn. Upper Headon Beds (Lignite Bed),
Headon Hill.
Description. The mandible is fairly long. The posterior mental foramen lies
below the posterior root of P4; the larger anterior foramen is sited between the canine
and P,. A small intermediate foramen lies below the anterior root of Pg. The
symphysis is long and oblique.
72 UPPER EOCENE MAMMALIA
The canine alveolus indicates a semiprocumbent tooth. P, (from the alveolus)
was small and single-rooted. Ps: is an oval double-rooted tooth with a large anteriorly
crested cusp and a small postero-internally projecting talonid. The talonid bears a
strong antero-posterior crest which posteriorly is raised into a small cusp. There isa
fairly strong internal cingulum and a weak postero-external cingulum. Pg3 is
double-rooted and similar to P:, except for a longer talonid and a larger talonid
cusp. In P4, the protoconid and metaconid appear to be subequal but this is
obscured by wear on the one specimen. The metaconid is markedly posterior to the
protoconid, with a distinct postero-internal metastylid. A straight anterior pro-
toconid crest joins a distinct median paraconid. There is a strong antero-internal
shelf-like cingulum and a smaller continuous antero-external, and postero-external
cingulum. The hypoconid is joined to the smaller more posteriorly sited entoconid
by a curved posterior crest, and joined to the posterior trigonid wall by a strong
crista obliqua.
M, is almost identical with Py, except that the entoconid and hypoconid are
subequal, and there is a small median hypoconulid slightly posterior to the entoconid.
M2 differs from Mj only by its larger size. Mg is very similar but broader and more
elongated antero-posteriorly than the other molars; the metastylid is very well
developed, being higher when unworn, than the protoconid. The talonid is almost
the same as in A. magnus: the moderately developed hypoconid is joined to the
posterior trigonid wall by an oblique crest ; the smaller entoconid is slightly posterior
to the hypoconid; the hypoconulid, the largest talonid cusp is extended posteriorly
as a prominent rounded median separated lobe.
P! is ovoid with a large external cusp which is crested anteriorly and posteriorly ;
internally the tooth is basined. There is a strong cingulum except externally where
it is vestigial. P4 is slightly elongated transversely. Of the major cusps, the pro-
tocone appears to be the largest and the metacone smallest, with the paracone
intermediate in size. The paracone and metacone are crested anteriorly and
posteriorly. Astrong broad crest runs from the protocone to a distinct parastyle. A
strong external and posterior cingulum is present. Antero-internally there is a
short cingulum; this does not extend round the protocone base. Postero-internally
there is little sign of a hypocone differentiated from the cingulum. The M? internal
fragment shows a strong cingulum which is continuous round the protocone base and
a small distinct hypocone.
Remarks. Distinguished from other species as follows: A. sciwreus has rounded
shorter lower molars; the protoconid-metaconid crest is only slightly oblique and
there is no metastylid; P3 is more simple; P, is less molarised with a smaller talonid,
and the upper molars have a strong metaconule and protoconule, with a stronger
hypocone. A. friscus possesses rounded less elongated lower molars ; the protoconid-
metaconid crest is slightly oblique and there is no metastylid. In A. ruetimeyert, the
rounded lower molars lack a metastylid; the protoconid-metaconid crest is slightly
oblique and the upper molar hypocone is strong. The presence of Pj, the strongly
oblique protoconid-metaconid crest and a prominent metastylid, support reference
of this specimen to A. magnus or A. parisiensis, However, A. magnus is much larger
HEADON BEDS OF ENGLAND 73
and has relatively less elongated lower molars and a slightly more transverse
protoconid-metaconid crest.
There is considerable variation in the skull, posterior region of the mandible
and molars of A. parisiensis. Filhol (1883) recognised six varieties which were
reduced to three by Stehlin (1912 : 1235). F.C.S. 7 does not agree closely with any of
Stehlin’s figured specimens, differing in the greater development of the metastylid
and in mandible shape. The material is perhaps best considered as a variety of A.
parisiensis ; it is undoubtedly specifically distinct from A. magnus from the Lower
Headon Beds.
TABLE 14
Measurements (in mm) of Adapis parisiensis
Pe P3 P4 Mi Me M3
Gijo) gs} Elf) mS) GIO) aS} IO) tas) Giejo) ams Elejo) tags
BCS. 7 (2) Geir gic OO 7eR oy ACS)
iBeC:S: 7) (2) AiO Mn; OMNES 2 een yc ON 4: ON -ONNA- 2 ey —— ——— OcON 4S
Family TARSIIDAE Gray, 1825
DraGnosis. Zygomatic arches slender. Brain case much expanded transversely.
Bullae much inflated; ectotympanic tubular, extends out of bulla. Foramen
magnum directed largely downwards. Very short narrow muzzle; palate narrow
anteriorly. Tooth rows converge (V-shaped). Upper canine smaller than anterior
incisor. Lower canine larger than upper. Hypocone often small. (after Simons,
1967).
REMARKS. Following Simons (1961), the group is divided into two subfamilies:
Tarsiinae Gray, 1825. Recent; East Indies—Philippines.
Microchoerinae Lydekker, 1887. Middle-Late Eocene; Europe.
On the basis of relationship with Tarsius, Simons (1961 : 53) placed the Microchoerinae
(previously grouped with the Anaptomorphidae) in the Tarsiidae. Following
McKenna (1967) and Russell, Louis & Savage (1967), Microchoerinae Lydekker, 1887
is adopted in preference to the previously more widely used Necrolemurinae Simpson,
1940.
Subfamily MICROCHOERINAE Lydekker, 1887
Diacnosis. Skull with comparatively large anteriorly-directed orbits. Snout
reduced in size with anterior convergence of the tooth rows. Typically the dental
formula is ¢, i i, 3, with total loss of lower incisors except in one species. Lower
canine enlarged and P; small and vestigial. Upper canine small and premolariform,
the upper enlarged tooth being the most anterior incisor, here considered as I?._ P4
not elevated, having a short heel and tending towards a molariform condition.
74 UPPER EOCENE MAMMALIA
P.—P, low and oval in shape. In M2-Ms, paraconid fusing with metaconid, the
talonid basined and subovate. Upper molars quadrate with large hypocone
usually developed on border ofa postero-internal basin. Metaconule and protoconule
developed and in more advanced forms, numerous accessory cuspules. (compiled
from Simpson, 1940, and Simons, 1961).
REMARKS. Distinguished from the other subfamily as follows: The Tarsiinae
has upper molars with a primitive tritubercular crown, a rudimentary hypocone and
a reduced metaconule.
Five genera are currently referred to the Microchoerinae:
Microchoerus Wood, 1884. Late Eocene; Britain, France, Switzerland, Spain.
Necrolemur Filhol, 1873. Middle-Upper Eocene; France, Germany.
Nannopithex Stehlin, 1916. Middle Eocene; Switzerland.
Pseudoloris Stehlin, 1916. Late Eocene; France, Spain.
Agerina Crusafont, 1967. Middle Eocene; Spain.
Microchoerus and Necrolemur have long been placed in this subfamily. Recent
work by Hiirzeler (1948) and Simons (1961 : 61) has added Nannopithex, previously
only tentatively assigned to the group. Simons has also shown (1961 : 54) that
Pseudoloris, formerly placed by Simpson (1940 : 198) in the Anaptomorphidae as the
single member of a separate subfamily, is closely related to the three former genera
and should be placed, therefore, in the Microchoerinae. The genus Periconodon
from the Middle Eocene of Switzerland, was tentatively assigned to the group by Hill
(1955 : 290), but Simons (1962 : 30) referred it to the Omomyidae; more recently,
this genus has been placed tentatively in the Adapidae by Russell, Louis & Savage
(1967). Crusafont (1967) departed from the generally accepted classification by
referring Pseuwdoloris to the Omomyidae.
Genus MICROCHOERUS Wood, 1844
Diacnosis. Upper molars with hypocone, mesostyle and numerous accessory
cuspules ; Mg witha prominent wide rounded posterior lobe, separated from hypoconid
and entoconid by small constrictions; small though distinct paraconid on Mj, not
distinct on Mp and M3. Extensive enamel crenulations on lower molars.
TYPE SPECIES. Mucrochoerus erinaceus Wood, 1844. Lower Headon Beds
(Mammal/Crocodile Bed), Hordle Cliff.
REMARKS. Distinguished from other genera as follows: Pseudoloris is much
smaller with sharp cusps; the molars lack enamel crenulation and accessory cuspules;
upper molars lack a mesostyle; the lower molars have a distinct paraconid and the
M3 hypoconulid lobe is narrow, sharp and well separated. Nannopithex is smaller
with only slight enamel crenulation; no mesostyle is developed and the M3 hypo-
conulid lobe is narrower and more distinctly separated. Necrolemur has only slight
enamel crenulation in the molars and lacks a mesostyle, and the M3 hypoconulid
lobe is less developed.
HEADON BEDS OF ENGLAND 75
Three species have been described:
M. erinaceus Wood, 1844, p. 350. Late Eocene (Early Ludian); Britain, France,
Spain.
M. edwardsi (Filhol, 1880), p. 124. Late Eocene (Late Ludian); Britain, France.
M. ornatus Stehlin, 1916, p. 1377, pl. 22, fig. 12. Late Eocene (Late Ludian);
Switzerland, Spain.
M. erinaceus and M. edwardsi are known from parts of the maxillae and fragmentary
mandibles; M. ovnatus is based on a partial upper dentition.
Crusafont (1967) considered that Microchoerus edwardsi was synonymous with M.
evinaceus. In the same work, Crusafont (1967) referred Microchoerus erinaceus to
Necrolemur and considered Necrolemur antiquus Filhol as a synonym of ‘Necrolemuyr’
evinaceus ; Microchoerus (sensu stricto) was retained for the species M. ornatus.
Microchoerus erinaceus Wood, 1844
(Pls3eePl A)
Dracnosis. Dental formula é i ;, ;. The largest species. Apparently lacks
lower incisors. Upper molars only moderately complex, lacking duplicated
paraconule. P3 with rounded outline and a single large external cusp.
HototyPe. B.M. 25229 (1). Fragment of left mandibular ramus with Py—Ms3.
(2) Palatal fragments with an almost complete dentition, lacking only the left
canine and right I? (latter tooth reconstructed in plaster). Fairly badly worn.
Lower Headon Beds (Mammal/Crocodile Bed), Hordle Cliff. Listed and described,
Wood, 1844 : 350; Figured, Wood, 1846 : 12, figs 1 and 3; Described and figured,
Lydekker, 1885d : 529, fig. (unnumbered); Figured (upper), Lydekker, 1887 : 304,
fig. 45; Described and figured, Cooper, 1910 : 40, pl. 3, fig. 1; Described and figured,
Simons, 1961 : 59, fig. 2.
MaTERIAL. All specimens from the Lower Headon Beds (Mammal/Crocodile
Bed), Hordle Cliff, except where stated otherwise.
B.M. 10768. Fragment of right mandibular ramus with Mj—-M3. Fairly badly
worn; crowns damaged.
B.M. M12563 (x). Fragment of left mandibular ramus with Ps-Mg.
(2). Fragment of right mandibular ramus with M\—Ms3.
(3). Isolated Mg, probably from (1) above.
B.M. 30346. Fragment of left mandibular ramus with Mj—Mg.
B.M. 30346b (x). Fragment of left mandibular ramus with Mz and broken Mj.
(2). Fragment of right mandibular ramus with M3.
(3). Fragment of left mandibular ramus with Py—-M3.
(4). Fragment of right maxilla with M1-M3,
(5). Right M2.
(6). Right P4.
Mentioned by Lydekker, 1887, p. 304. (4) above, listed Cooper, Igto : 40.
70 UPPER EOCENE MAMMALIA
B.M. M2605b. Isolated My, Mz and M3, from same individual.
S.M. C9672. Left mandibular ramus with M;—M3. Posterior region almost
complete. Listed and figured, Cooper, Igro : 39; pl. 3, fig. 6.
S.M. Cg681. Left mandibular ramus with P3—M3 and the alveoli of the canine,
P,; and Pg. The symphysis is preserved.
S.M. Cg670. Left mandibular ramus with P2-M3. Listed and figured, Cooper,
IgI0 : 40; pl. 3, fig. 4.
S.M. C9671. Right mandibular ramus with P3—M3 and roots of the canine, Py and
Py. Symphysis preserved. Listed, Cooper, Ig10 : 39. According to Cooper,
associated with S.M. C9670.
S.M. C9673. Fragmentary right mandibular ramus with Mj—M2, damaged M3
and roots of Py. Listed, Cooper, 1910 : 40.
S.M. C9675. Fragmentary right mandibular ramus with Py—Mg.
S.M. C9678. Right M3.
S.M. C9679. Right Mz. Probably from same individual as S.M. C9678.
S.M. C9680. Fragment of right mandibular ramus with Py and M;. Probably
from same individual as S.M. C9678.
S.M. C54174 (1). Right P3. Unworn, slightly damaged.
(2). Left M,;. Talonid only. Almost unworn. Lower Headon
Beds (Rodent Bed), Hordle Cliff.
S.M. C53482. Fragment of left mandibular ramus with roots of Pa, Mj, and alveoli
of the canine and P;—P3.
B.M. M26055. Fragment of left mandibular ramus with Ps-Ms. P3 damaged.
Teeth badly worn. Upper Headon Beds (Microchoerus Bed), Headon Hill.
F.C.S. 3. Left Ms. Upper Headon Beds (Lignite Bed), Headon Hill.
B.M. M12563 (1). Right M2.
(2). Left M1.
S.M. Cg669. Palatal fragment with right and left M?-M%. Listed, Cooper,
I9IO, p. 39.
S.M. C30710-27. Teeth mounted on a board:
S.M. C30712. Left M2.
S.M. C30714. Left M?.
S.M. C30715. Left M!.
S.M. C30716. Right Mj.
S.M. C30717. Right Mp.
S.M C30718. Left M}.
S.M. C30719. Right P4.
SAM. €30720., cette.
S.M. C30723. Left Mj.
S.M. C30724. Right Mg.
S.M. C30727. Right upper canine.
B.M M12565a =(x). Right Mj.
(2). Right My; (or Mg).
(3). Right M1.
HEADON BEDS OF ENGLAND 77
(4). Right I? (?).
(5). Right upper canine (?).
B.M. M12565c (1). Left upper canine (?).
(2). Right P§. Partially embedded in matrix.
B.M. M25090 (i) ettilien(@):
(2). Left I3 (?). Both moderately worn.
S.M. C54175 (x). Left P§. Moderately worn.
(2). Left upper canine (?). Lower Headon Beds (Rodent
Bed), Hordle Cliff.
DEscrIPTION. The mandible is fairly short and stout; the oval-shaped symphysis
(S.M. Cg681, S.M. Cg671, and S.M. Cg670) is steeply inclined, following the line of the
canine root. Below P3, the lower edge of the mandible projects internally as a small
shelf. The smaller posterior mental foramen lies below Py, (S.M. C9671) and S.M.
Cg68r), or slightly anterior to this (S.M. C9670 and S.M. C53482). The position of
the larger anterior mental foramen varies from below P3 (S.M. Cg671 and S.M.
C9681) to below Pz (S.M. C9670). The posterior region is preserved only in S.M.
C9672 (wax restoration of the type excepted). The moderately developed rounded
angular process falls considerably below the lower anterior margin of the mandible.
The mandibular foramen lies just below alveolar level. The coronoid is small, the
posterior margin rather damaged on the specimen. The slender condyle has an
articulating surface which is slightly elongated transversely; it lies considerably
above the tooth row level. The coronoid crest is more prominent than the condyloid
crest, although both are not well developed.
The tooth row is compact and continuous. In Ms the protoconid and metaconid
are subequal and larger than the hypoconid and entoconid. A small crest which
unites the protoconid and metaconid follows the rounded anterior tooth margin;
there is no distinct cuspate paraconid. A crest from the hypoconid runs antero-
internally towards the metaconid. The posterior region of the talonid is separated
into a prominent rounded hypoconulid lobe which is slightly displaced internally;
this lobe greatly increases the length of the talonid. In B.M. 10768, the hypoconulid
lobe is considerably enlarged relative to the other specimens. There is no distinct
cuspate hypoconulid but the lobe bears an internal and an external accessory cusp;
these accessory cusps are rather indistinct in some specimens especially after wear.
There is a fairly strong external cingulum (only weakly developed in S.M. C9672).
Several specimens show slight traces of an internal cingulum.
M2 is more quadrate than Mg but otherwise very similar without the hypoconulid
lobe. No paraconid is distinguishable as in M3; the protoconid, metaconid and
hypoconid are subequal in size, the entoconid smaller. There is a small bilobed
median hypoconulid on the hypoconid-entoconid crest (indistinct on S.M. Cg68r).
In most specimens, there is an accessory cuspule (mesoconid) on the prominent
crista obliqua. A strong external cingulum is present (weak in S.M. Cg672) but there
is no trace of an internal cingulum.
M;, is similar structurally to Me, with a small duplicated hypoconulid on most
specimens (not distinguishable on S.M. C9671) anda mesoconid. There is no internal
cingulum. The tooth differs from Mz and Mg by having a more squared anterior
78 UPPER EOCENE MAMMALIA
margin and a small distinct paraconid situated directly anterior to the metaconid;
the paraconid is separated from the metaconid but united to the protoconid by a low
crest.
P,is smaller and subtriangular with a smalllow heel. The tooth is dominated by a
large median protoconid. A curved antero-basally directed crest from the protoconid
forms the anterior tooth margin; the posterior face of the protoconid bears two crests.
The smaller separated metaconid in S.M. Cg680, S.M. C9670, S.M. Cg671 and S.M.
C9675 is more posteriorly placed than the protoconid and falls to a strong antero-
internal cingulum. In the holotype (B.M. 25229) and S.M. C9681, the protoconid
and metaconid are opposite and the internal cingulum is only weakly developed.
S.M. C9681 and S.M. Cg670 have a well developed external cingulum; this is poorly
developed in S.M. C9680, S.M. Cg674 and S.M. Co68r.
P3 (well preserved in S.M. Cg681, S.M. C9671 and S.M. C9670) bears a large median
anteriorly crested cusp and a small cingulum-like heel; there is a strong internal
cingulum. In S.M. Cq67zr and S.M. Co9670 the anterior crest runs parallel to the
length of the tooth; the main cusp slopes moderately basally towards the heel,
bounded by two sharp crests; an external cingulum is moderately developed. In
S.M. Cg681, the anterior crest curves round towards the internal cingulum; the
main cusp is more posteriorly situated, falling more steeply to the heel and the two
posterior crests are less prominent.
Pz (S.M. C9670 and B.M. M26055) is made up of an anteriorly-crested cusp and a
small cingulum-like heel. The cusp is concave internally and convex externally.
There is a prominent internal cingulum and a smaller external cingulum.
Pj, represented only by an alveolus (S.M. Cg681) and a broken root (S.M. C9671)
was evidently a small anteriorly inclined tooth, situated on the external edge of the
tooth row. The broken root (S.M. Cg671) and a large oval deep alveolus (S.M.
Cg681) of the lower canine indicate a large anteriorly-inclined tooth, lying at a much
lower level in the jaw than the other teeth.
I2-P4 are associated only in the holotype. The most anterior tooth I?, is enlarged
with almost vertical sides. I? is considerably smaller with an antero-internally
sloping crown and a small external cingulum. The semi-procumbent canine is much
enlarged (but not as high as I?) and has an external cingulum. P? is single-rooted
and semi-procumbent with a small heel; the tooth is smaller than the canine.
P3 and P4 are quadrate and larger than P2; both have a single large external cusp
and a strong external cingulum. P4 has a small antero-internal and a small postero-
internal cusp which are sharp and unworn in S.M. C30719 and S.M. C30720; these
are worn down to an antero-posterior ridge in the holotype and B.M. 30346b (6).
M! and M2 are quadrate and subequal in size; the metacone lies on a straight
antero-posterior line with the paracone. In the smaller M°, with a more rounded
posterior margin, the metacone is more internal to the paracone. The hypocone,
which in M1! is almost equal to the protocone, becomes progressively smaller in M2
and M3. A paraconule and metaconule are well developed in all molars, the former
sited anterior of a line between protocone and paracone, the latter anterior of a line
between metacone and hypocone. The metaconule, equal to the paraconule in M},
becomes progressively smaller in M2 and M8. In all molars, a small accessory cusp
HEADON BEDS OF ENGLAND 79
is situated midway between the metaconule and the protocone; this can be regarded
as a duplication of the metaconule. In M! and M®?, this accessory cusp is smaller
than the main metaconule, but in M it is larger. There is a prominent mesostyle
in M! and M? external to a line between the paracone and metacone. This cusp is
represented in M3 by a slight cingular swelling. A small but definite hypostyle occurs
on M! and M? of B.M. 30346b (4). A parastyle and metastyle are encountered more
frequently, appearing as upgrowths from the prominent external cingulum: a
parastyle occurs on M? of S.M. C30712 and S.M. C30714 and M! of B.M. 30346b (4),
B.M. M12563 (2), S.M. C30715 and S.M. C30718; the metastyle is present on M2? of
the holotype (B.M. 25229), B.M. 30346b (4), S.M. C30712 and S.M. C30714 and the
M! of B.M. 25229 and B.M. 30346b (4). In all other specimens, these three accessory
cusps are indistinct.
Remarks. The number of specimens is really inadequate for any satisfactory
statistical treatment of variation within the species to be carried out. However,
some tentative conclusions may be drawn from the available material. The following
variates were considered: length and width of Ms, Me, M; and Py; length of tooth
rows M;—Ms and P,—M3 and the internal depth of the mandible below the posterior
TABLE 15
Selected measurements (in mm) of Micvochoerus erinaceus
Mi-
Pe P3 Py, Mi Me M3 M3
aap ciSua-p LES! a-p) tts) ap) trs) a-p) tks! a=p trs ‘a=p
B.M. 25229 (1) 3 Onee2 Cra 3 + ON——0(3'-4) 4-3 Ol LTO)
B.M. 10768 —- — — — — — (2:5) (2:9) 3°83 3:2 4:5 27 —
B.M. M12563 (1) —- — — — 3:0 26 41 3:9 42 39 — — —
B.M. M12563 (2) 1-23 AC SSO) AO 3-2
B.M. 30346 ee tee oo ety ALO 3240 AO, Brg
B.M. M26055 312 rn 27S 2 IASON 13-0) -4cAy 2-8) ——
S.M. C9670 Ones 2g ee 397 340 8-7 34) (45 93°C TTS
S.M. C9671 ae ao SS 9 85 87 3540 4-0 3 :P Io
S.M. C9681 a — eS 2A 3-4 2-0) 42 3°50 3°Q)) 3-4) 4-2 2:5) 12-0
S.M. C9672 At -OmmrA-ONNnS "5 4- On 3 - Onl 225
S.M. C9673 —- — — ~— —~— — 4:0 3:2 40 3:2 — — —
S.M. C9675 BoP eS Oma SMES OM AGNES) a Se
p2 P3 p4 M1 M2 M3
B.M. 25229 (2) aan 220 Ooh Soh 4rON 74.0) 93:00 5:0) 3°01) 3°90
B.M. 25229 (2) ZOE ZAR 27S 35) SiO) AZ) 40) VAT 3:9» <4°9. 3°2. -Ar0
B.M. 30346b (4) ai Gaia aa 5On 40 4:9) 2:8) 470
B.M. 30346b (5) 1 a.
B.M. 30346b (6) Sg I re ee
B.M. M12563 (1) —- — — 39 51 — —
B.M. Mr2563 (2) 4:2) 4-0 = — =
S.M. C9669 SS SS 39 (48) 31 42
S.M. C9669 —- —- —- —- —- — — 3-5 SOS 4:2
S.M. C30715 = ee SS a 7
S.M. C30719 —- —- — — — 38 — —- —- ~—- —- —
S.M. C30720 ee
80 UPPER EOCENE MAMMALIA
margins of Ms, M; and P3. The mandible depth in M. erinaceus appears to be
highly variable and cannot be correlated with individual age as reflected by dental
wear. Frequency distributions utilising dimensions of the posterior lower teeth
indicate a separation of the larger specimen B.M. 303,46; this is removed by secondary
grouping of the data and is, in all probability, due only to incomplete sampling.
P3_P4 of the type specimen (B.M. 25229 (1)) and S.M. C9681 are different structurally
from the rest of the material. These differences might be sexual or could possibly
indicate an anterior milk dentition; they are unlikely to be of specific importance.
Simons (1961) observed an alveolus anterior to the lower canine in some French
specimens of M. edwards, indicating at least one lower incisor. There is no evidence
for similar teeth in the specimens of M. evinaceus from the Headon Beds. But
preservation of this region of the mandible is rare and their absence in M. erinaceus
should be treated with caution. Apart from the probable structural difference in
dentition, M. evinaceus differs from M. edwardsi by its greater size.
Several features of the upper dentition distinguish the species from M. ornatus
(known only from upper dentition): P? in M. evinaceus has a rounded outline and a
single major external cusp whilst this tooth in M. oyvnatus has a squared outline and
twoexternalcusps. Inaddition to the duplicated metaconule present in both species,
the M2? of M. ornatus is further complicated by a duplicated paraconule and a
prominent hypostyle. MM. erinaceus is significantly larger than M. ornatus: in the
holotype of MW. evinaceus, the length of the tooth row, M2-M$ is 11-0 mm, whereas
in the holotype of M. ornatus, this measurement is 9:0 mm.
Genus PSEUDOLORIS Stehlin, 1916
Diacnosis. Small size. No accessory cuspules and enamel crenulations.
Upper molars relatively elongated transversely, lacking mesostyles; cingulum-
hypocone distinct on M! and M2, indistinct on M3; lower molar cusps relatively sharp
and separated. Mg entoconid and hypoconulid high and separated with distinct
narrow hypoconulid-lobe. Distinct small paraconid on My and Mg, reduced on M3.
Type species. Pseudoloris parvulus (Filhol, 1890). ? Late Eocene (Quercy
Phosphorites) ; France.
REMARKS. Thegenus was proposed by Stehlin (1916 : 1397) for a species originally
described by Filhol, (1890a) as Necrolemur parvulus. The validity of this genus has
been subsequently accepted generally.
Simpson (1940: 198) agreeing with Teilhard de Chardin (1916—1921a) that
Pseudoloris was significantly Tarsius-like, erected a new subfamily, Pseudolorisinae,
for this monospecific genus. Recently Simons (1961 : 54) has shown that the
similarities between Psewdoloris and other microchoerines are so great, that separation
at subfamily level is not possible.
Weigelt (1933) described P. abderhaldeni from the Middle Eocene brown coal
deposits of Geiseltal, Germany. Simons (1961 : 61) stated that the type of this
species is conspecific with ‘“Necrolemur’ vaabi Heller, the latter species being referable
to Nannopithex. Crusafont (1967) described a new species, Pseudoloris reguanti
from the Upper Ludian of Spain.
HEADON BEDS OF ENGLAND 81
Pseudoloris is distinguished from other genera as follows: Microchoerus has a
mesostyle and is markedly larger with lower rounded cusps; the M3 hypoconulid-
lobe is wide and rounded and the molars possess numerous accessory cuspules and
enamel crenulations. In Necrolemur the M3 hypoconulid-lobe is wide and rounded;
the cusps are lower, rounded and less distinct and the genus is much larger. Nanno-
pithex has slight enamel crenulations in all teeth; P4 is much larger than the lower
molars; the molar cusps are more rounded and lower, and Mg entoconid and hypo-
conulid are not as high and separated.
Pseudoloris parvulus (Filhol, 1890)
(Text-fig. 23)
1890 Necrolemur pavvulus Filhol.
Diacnosis. As for genus.
HoLotyrPe. Fragment of left mandibular ramus with P3—Ms3 and alveoli of the
canine, P;-Py. ?Late Eocene (?Ludian, Quercy Phosphorites); Caylux, France.
Described and figured, Filhol (1890a : 39, text-fig., unnumbered). Specimen not
located by Teilhard de Chardin (1916—-192I1a: 5). Abundant referred material
described and figured by Stehlin (1916 : 1397) and Teilhard de Chardin (1916-
1g21a : 4).
MATERIAL. B.M. 36812f. Fragment of right mandibular ramus with M2—Ms3.
Teeth slightly worn. Lower Headon Beds (Mammal/Crocodile Bed), Hordle Cliff.
DESCRIPTION. Mg and Mg are elongated antero-posteriorly. The trigonid and
basined talonid are well differentiated, but the cusps are low, the trigonid cusps
being only slightly higher than those of the talonid.
In Mg, the trigonid and talonid are subequal in width but the trigonid 1s compressed
antero-posteriorly and is shorter. The protoconid is slightly higher than the
metaconid and markedly more anteriorly placed; both cusps are crested transversely.
The metaconid is internal in position; the protoconid tip lies just external to the
longitudinal midline. There is a well developed antero-external cingulum. A
small median cuspate paraconid forms the sharp anterior trigonid apex; this cusp is
joined by a small low crest from the metaconid and from the protoconid. The
talonid is strongly basined. The entoconid is slightly smaller than the metaconid;
a sharp crest runs anteriorly from the entoconid to meet, at the base of the posterior
trigonid wall, a sharp metaconid crest. Although the tooth is damaged externally,
it appears that the hypoconid was opposite, or just anterior to the entoconid, a
strong crista obliqua running from this cusp to the base of the posterior protoconid
wall. A rudimentary hypoconulid is represented as a very small projection on the
posterior talonid margin, just external to the midline; this is joined to the hypoconid
and entoconid by small crests. Owing to damage, the extent posteriorly of the
cingulum cannot be observed; it may have been continuous round the base of the
hypoconid as in M3.
F
82 UPPER EOCENE MAMMALIA
The anterior part of Mgis very similar to that of Mg, differing only in minor features:
the protoconid is slightly more externally placed, and is subequal with the metaconid;
the posterior trigonid margin is less oblique than in Mg and the paraconid is slightly
less developed and more crestiform in appearance, forming a more rounded anterior
trigonid apex. The anterior part of the talonid is also very similar. Mg differs
markedly from Mz by having a hypoconulid-lobe. This long prominent separated
lobe is narrow and mainly median although the posterior extremity is slightly
internally directed. The lobe, consisting of a single large crestiform cusp, is con-
nected with the hypoconid and entoconid by crests. A prominent continuous antero-
external, external and postero-external cingulum is present.
REMARKS. Recently Pseudoloris parvulus has been recorded by Crusafont from
the Early Ludian of Sosis, Spain (1965, 1967) and Euzet-les-Bains, France (1967).
The single specimen from the Headon Beds appears to agree precisely with additional
material of Pseudoloris parvulus figured by Stehlin (1916), Teilhard de Chardin
(1916—1g921a) and Piveteau (1957); it constitutes the first record of the genus from
the Early Ludian of Britain.
Fic. 23. Pseudoloris parvulus (Filhol, 1890). Fragmentary right mandibular ramus with
Me-M3. B.M. 36812f x12. Lower Headon Beds, Hordle Cliff. (a) Occlusal view.
(sp) External view. (c) Internal view.
HEADON BEDS OF ENGLAND 83
TABLE 16
Measurements (in mm) of Pseudolovis parvulus
M2 M3
a-p trs a-p trs
B.M. 36812f 16 1:25 19 Te.
Order CREODONTA
Family HYAENODONTIDAE Leidy, 1869
Diacnosis. Py double-rooted except in some specialized genera; P3 lacking
distinct protocone; Mz or Mz functioning as main carnassials; M3 present or absent;
cheek teeth row moderate to long; Mg when present transversely extended. Jaw
long and shallow to moderately short and deep, with symphysis weak to fairly robust,
and angle long, slender, projecting and upcurved at tip. Face long to moderately
short; basicranial region narrow to moderately wide; no preglenoid crest (mainly
from Denison, 1938 : 117).
Remarks. The order Creodonta is retained here in a restricted sense, following
Romer, (1966 : 229). The family is known from the Early Eocene-Middle Oligocene
of North America; Early Eocene-Late Oligocene of Europe; Late Eocene-Oligocene,
Early Pliocene of Asia and the Early Oligocene, Early Miocene of Africa.
Denison (1938) distinguished four subfamilies: Proviverrinae, Hyaenodontinae,
Limnocyoninae and Machaeroidinae; this was followed by Simpson, (1945). Gazin
(1946) proposed a new family, Limnocyonidae, to accommodate the latter two
subfamilies. Van Valen (1966) placed all four subfamilies in the Hyaenodontidae
but reduced them to the rank of tribes; Gazin’s two fold subdivision was retained as
two subfamilies, the Hyaenodontinae and the Limnocyoninae. More recently,
Van Valen (1967) has abandoned the use of tribes within the Hyaenodontinae,
grouping together the Hyaenodontini and the Proviverrini. In the present work,
the earlier scheme of Denison is utilized as it provides a simple working arrangement
(see also Savage, 1965).
Subfamily HYAENODONTINAE Trouessart, 1885
Diacnosis. Long faced, narrow skulled genera; main carnassials M3 with
carnassial specialization very advanced; M3 present, M8 variably present or absent;
molars sectorial, length greater than width; M3, where present, small and transverse ;
M! and M? with paracone and metacone completely or nearly connate and protocone
reduced or absent ; lower molars lack metaconid, talonid vestigial or absent. Body
compact, robust; limbs and feet cursorial (after Matthew, 1909 and Savage, 1965).
84 UPPER EOCENE MAMMALIA
Remarks. Distinguished from the other subfamilies as follows: in the Proviver-
rinae, the metaconid is retained; the carnassial specialization is less advanced and M3
are invariably present. The Limnocyoninae and Machaeroidinae both have M3
functioning as the main carnassials.- Savage (1965) listed five good genera in the
Hyaenodontinae:
Hyaenodon Laizer & Parieu, 1838. Late Eocene-Late Oligocene; Europe. Late
Eocene-Middle Oligocene; North America, Asia. Early Oligocene, Early
Miocene; Africa.
Pterodon Blainville, 1839. Late Eocene-Early Oligocene; Europe. Late Eocene-
Early Oligocene; North America. Early Oligocene, Early Miocene; Africa.
Late Eocene; Asia.
Apterodon Fischer, 1881. Early-Middle Oligocene; Europe. Early Oligocene;
Africa.
Metapterodon Stromer, 1926. Early Miocene; Africa. Early Oligocene; Europe.
Leakitherium Savage, 1965. Early Miocene; Africa.
Simpson (1945) listed Hemipsalodon Cope, 1885; Dasyurodon Andreae, 1887 and
Propterodon Martin, 1906. Savage (1965) has considered that Hemipsalodon is a
synonym of Pterodon and Dasyurodon a synonym of Apterodon. More recently, Van
Valen (1966 and 1967) has asserted the generic validity of Hemipsalodon and has
stated (1967) that Metapterodon is a synonym of Pterodon ; he also adds Ischnognathus
Stovall, 1948 (Van Valen, 1966 and 1967) and (tentatively) Hyainatlouros Bieder-
mann, 1863 (Van Valen, 1967). Pvopterodon is not well known (see Van Valen,
1966 : 75) and Savage has stated (personal communication) that this latter genus
will probably prove to be invalid. According to Van Valen (1967), Megalopterodon
Dashzeveg, 1964, is synonymous (provisionally) with Pterodon. Apterodon was
referred by Van Valen (1966) to the Mesonychidae; the genus was returned to the
Hyaenodontidae by Van Valen (1967) following the work of Szalay (1967).
Genus HYAENODON Laizer & Parieu, 1838
Diacnosis. Dental formula 3, s i ; ; M? with shallow groove on completely
connate paracone and metacone; molars without protocone; metastyle strongly
elongated, especially on M2. Main carnassial pair Mj, with M3 as accessory carnas-
sials. Ms without talonid, M; +2 with or without vestigial talonid (Savage, 1965).
TYPE SPECIES. Hyaenodonleptorhynchus Laizer & Parieu, 1838. Middle Oligocene
(Chattian) ; France.
REMARKS. Distinguished from other genera as follows: in Pterodon, M? is present
and a small protocone and large parastyle occur on M1! and M?. In Apterodon, a
protocone is present on M1 and M2, and Mj, possesses a prominent three-cusped
talonid. M2? is present in Metapterodon and M1 and M? possess a strong protocone.
Leakitherium has a strong protocone on P4—M2?.
HEADON BEDS OF ENGLAND 85
A complete list of species attributed to Hyaenodon has been given by Savage (1965).
Some 36 species have been described: 9 are from North America, 3 from Asia, 20
from Europe and 4 from Africa, the latter being grouped by Savage as a distinct
subgenus. Specific distinctions are based considerably on size differences.
Numerous structural features have also been utilized (listed in Savage, 1965), but
have nearly all proved highly variable. Many of the early European types are lost
and the existing figures and descriptions are frequently quite inadequate. Of the
described species, many are undoubted synonyms; several true genera probably are
involved. In the absence of a greatly needed thorough revision of the genus, some
attempt is made below to clarify the systematics of the Headon Beds specimens.
Hyaenodon c.f. minor Gervais, 1848-52
(Pl. 5)
1926 Hyaenodon aimi Cooper: 370, fig. B.
Diacnosis. M3 with talonid trace usually present but variably developed.
Mj—-Mz2 usually with distinct cuspule at base of antero-internal paraconid face.
P; with a single root.
HototyrPe. Right mandibular ramus with P2—-M3 and alveoli of the canine and
P;. Molars badly worn. Late Eocene (Early Ludian); France. Described and
figured, Gervais, 1848-52 : 129; pl. 25, fig. 9. Specimen not located.
MateriaL. All specimens from Lower Headon Beds (Mammal/Crocodile Bed),
Hordle Cliff, except where stated otherwise.
B.M. 29752. Complete mandible attached at symphysis. With right canine,
P.—M3, broken root of P; and alveolus of I3; and left Ps—M3, broken root of canine
and alveolus of Pj. Teeth very badly worn—often to root level. The posterior
region of both ramiis complete. Labelled in British Museum (N.H.) as “Hyaenodon
hantonensis’. Listed by Lydekker, 1884 : 444, 1885a : 25.
S.M. C31347 a-c, d, k. Probably all from same individual. Fine greenish-white
sand matrix. Teeth well worn.
a. Right mandibular ramus with canine, Py—-Ms3, and alveoli of Pj-P3. M1 is
badly damaged. Posterior region preserved. Figured, Cooper, 1926 : 372, fig. A.
b. Left mandibular ramus with canine, Pz, Py-M3. Ps3 represented by broken
roots.
c. Fragment of right maxilla with P4, damaged fragment of M!, alveoli of P%
and M2. Anterior part of the zygomatic arch preserved.
d. Fragment of left maxilla with P4-M2 and roots of P3.
k. Isolated left upper canine. Tip damaged.
S.M. C31344. Fragment of left mandibular ramus with Py-Mz. Moderately
worn.
B.M. 29814. Left M3. Badly worn. Listed by Lydekker, 1885a : 26.
(erroneously as right Ms).
86 UPPER EOCENE MAMMALIA
S.M. C31346. Left Ps. Moderately worn.
S.M. C30775. Fragment of left maxilla with I8, P3—-P4, alveoli of the canine,
M1—M2 and roots of P! and P?. Teeth damaged. Internal surface embedded in
fine greenish sand matrix. 7
B.M. M25073. Fragment of left maxilla with P1-P3, posterior alveoli of P4 and
part of the canine alveolus.
B.M. M25074. Fragment of right mandibular ramus with Ps—M3. Moderately
worn.
B.M. M13075. Fragmentary right and left mandibular rami, not attached at
symphysis. Same individual. Adhering matrix of shelly orange sand. Holotype
of ‘Hyaenodon aims’. Described and figured Cooper, 1926 : 370, fig. B.
Right ramus with Iz and the canine, Po-M3 and alveoli of Iz and P,. Tip of canine
missing.
Left ramus with the canine, P2—M3 and alveolus of P}. Tip of canine missing.
B.M. M25075. Fragment of left maxilla with P2-M?2. Unworn. Upper Headon
Beds (Lignite Bed), Headon Hill.
DESCRIPTION. The mandible is moderately elongated. The posterior mental
foramen lies under the posterior root of P3, the anterior mental foramen at a higher
level, under the root of Py. The symphysial surface is inclined at 45° to the tooth
tow and ends posteriorly under Pz. The posterior mandibular region is well pre-
served in B.M. 29752. The prominent transversely elongated condyle, situated
below the level of the tooth row, is separated from a small narrow angular process by
astrong notch. The angular projects downwards just below the level of the anterior
mandibular margin. The coronoid is moderately well-developed with a slightly
notched posterior margin. The mandibular foramen is sited markedly posteriorly,
just below condyle level. The coronoid and condyloid crests are prominent.
Iz is represented in the holotype of ‘Hyaenodon aimi’ (B.M. M13075) by a small
alveolus. Is is a small semiprocumbent tooth immediately external to the I,
alveolus and pressed up close against the canine. 1; may have been present but no
evidence remains. Cooper (1926 : 371) stated that the incisors were not preserved.
The lower canine is large and sharp with an ovoid section. In the right ramus of
S.M. C31347, this tooth is probably somewhat displaced from its alveolus. P, is
represented by the alveolus; this single-rooted tooth was slightly separated from
the canine. In S.M. C31347b, the left ramus anomalously lacks a P, alveolus and
there is a larger gap between P2 and thecanine. Contrary to the statement by Cooper
(1926 : 371, footnote) P; was originally present in the right ramus of B.M. 29752, asa
root-filled alveolus is visible. Pz, is double-rooted, posteriorly-directed and
immediately posterior to P;; it bears a large central cusp which is postero-externally
crested. There is usually a continuous internal and postero-internal cingulum, but
in B.M. M13075, it is small and antero-internal in position. Pg is similar to Pp:
except that the tooth is longer antero-posteriorly and has a small posterior longi-
tudinally crested cusp; there is no distinct posterior cingulum in B.M. M13075. Pa
is similar to Ps but larger transversely and strongly inclined posteriorly. The
cingulum is usually prominent antero-internally and postero-internally, although
HEADON BEDS OF ENGLAND 87
indistinct in B.M. M13075. Where the anterior crest of the main cusp merges with
the antero-internal cingulum there is an irregular cuspule (S.M. C31347 and B.M.
M25074) or a distinct cusp (S.M. C31344, B.M. M13075 and B.M. 29752).
Mj is very small. The protoconid is slightly larger than the paraconid when un-
worn; these crestiform cusps are separated by a cleft. A small low talonid bears an
elongated cusp. The antero-internal paraconid face (S.M. C31344) bears a small
projecting cuspule which is closely pressed against the postero-internal margin of
P,; this cuspule is indistinct in B.M. M13075. On the antero-external paraconid
face there is a vertical buttress pressed up against the postero-external margin of P4.
The cingulum is vestigial. In B.M. 29752, only worn rounded stumps of this tooth
remain.
Mg is similar to Mj, but of larger size. The crested paraconid and protoconid, well
separated by a cleft, form good shearing facets. The antero-internal cuspule and
antero-external buttress are usually well developed; the cuspule is higher than in My
and is not in contact with the adjacent anterior tooth. The cuspule is indistinct in
B.M. M13075. The posterior talonid cusp is prominent. In B.M. 25074, this tooth
is structurally similar but very small, being about the same length as Mj.
M3 is markedly elongated in length; the crested paraconid is separated from the
larger protoconid by a deep cleft; these cusps together form an effective slightly
oblique shearing blade. Postero-internally, there is a small talonid rudiment in
B.M. M13075 and S.M. C31347 (more strongly developed in B.M. 29752). Antero-
externally, a buttress is developed as in the other molars. A trace of the antero-
internal cuspule situated high on the antero-internal paraconid face is present in the
least worn specimens.
Anterior to the orbits, the maxillaries have a great height, as is seen in S.M.
C30775. The infra-orbital foramen is narrow and slit-like, opening above P?
(B.M. M25073 and S.M. C30775). S.M. C31347c and d indicate a fairly narrow
palatal region.
I3 is a very small pointed tooth, slightly separated from the canine alveolus
(S.M. C30775). The upper canine (S.M. C31347k, S.M. C30775 and B.M. M25073)
is a large curved tooth with an ovoid section and a long vertically-striated root. Pl
(B.M. M25073) is double-rooted, elongated antero-posteriorly and situated immediate-
ly posterior to the canine; the tooth has a main cusp over the anterior root, and a
strong continuous internal cingulum. P2? is larger than P!, but otherwise similar;
it is separated from P! by a gap.
P3 is separated from P2 by a gap. This tooth is slightly larger than P? (B.M.
Mz25073) and has two roots. The main cusp is crested antero-internally and
posteriorly. There is a distinct crested posterior cusp, separated from the main
cusp by a cleft, and a prominent internal cingulum.
P4 (S.M. C31347c and d, and S.M. C30775) has three roots and lies immediately
posterior to P3. There is a strong protocone lobe but no distinct cusp; the transverse
width is greater than the antero-posterior length. The large main cusp is separated
from a small though distinct posterior cusp by a cleft; there is a small anterior
cusp (more developed in B.M. M13075 and S.M. C30775 than in S.M. C31347c and d).
M1 is much smaller than P4 and situated immediately posterior to it; the tooth is
88 UPPER EOCENE MAMMALIA
triangular-shaped, the antero-posterior length being about the same as the transverse
width. The crested anterior cusp (fused metacone and paracone) and a crested
metastyle form an effective shearing blade; the cusps are separated internally by a
small cleft and externally by a shallow valley. There is a small indistinct parastyle
in S.M. C31347d; this is better developed in B.M. M13075.
M? is very similar structurally but much larger and more trenchant. The crested
metastyle is tapered and elongated posteriorly ; with the crested metacone/paracone,
this forms internally an elongated almost vertical shear plane.
REMARKS. With the exception of B.M. M13075, most of the material listed above,
for many years has been referred to the Late Eocene species, Hyaenodon minor.
The length of the tooth row and the single-rooted P; appear to exclude reference to
any of the other European species of Late Eocene age (listed in Savage, 1965 : 280);
with the present chaotic state in the systematics of the genus, comparison of the
Headon specimens with Hyaenodon minor seems the most reasonable course.
Cooper (1926) described a new species, Hyaenodon aimi, based on a well-preserved
mandible from the Lower Headon Beds of Hordle Cliff. In the present work, the
holotype (and only specimen) of ‘Hyaenodon aimi’ is not considered specifically
distinct from the other Headon Beds material. The length of the lower tooth row
in the holotype of ‘Hyaenodon aimi’ and in the specimens of Hyaenodon c.f. minor is
almost identical (see table of measurements) ; dimensions of individual teeth in the
two species overlap considerably. The only difference in dentition worthy of mention
is the presence of a distinct cuspule antero-internal to the paraconid on My; and Mg in
Hyaenodon c.f. minor, this cuspule is indistinct in the holotype of “Hyaenodon aim’,
but its presence or absence is unlikely to be of any significance.
Cooper (1926) noted the closely similar length of the lower tooth row in ‘Hyaenodon
aima’ and the other Headon Beds specimens, but cited as differences the smaller
more slender mandible, the closed tooth row and the curved lower margin of the
mandible in ‘Hyaenodon aimi.’ In particular, Cooper emphasised the differences in
mandible depth below Mg and considered the possibility of this variation being caused
by differences in the sex of individuals; he concluded, by comparison with living
carnivores, that the discrepancy in dimensions was too great to be accounted for in
this way.
It is highly probable that these differences in mandible shape and size are due to
variations in individual age. The holotype of ‘Hyaenodon aimi’ is of a young
individual with only a slightly worn dentition. Nearly all of the remaining Headon
Beds specimens of lower dentitions exhibit greater degrees of wear, indicating more
mature individuals; in these animals, increasing age has been accompanied by many
changes such as a deepening of the mandible, flattening of the lower margin of the
mandible and spacing of the premolars.
B.M. M25074 is rather smaller than most specimens, with a curiously small Mp.
The specimen may be specifically distinct, but as little is known about intraspecific
size variation in Hyaenodon species, it is best grouped at present with the other
material. In general aspect, the teeth in B.M. M13075 and B.M. M25075 are slightly
more slender and delicate; these specimens may represent females of the species.
HEADON BEDS OF ENGLAND 89
TABLE 17
Measurements (in mm) of Hyaenodon c.f. minor
P2 P3 Pa My Me M3 Ps3-Ms3
Gisj) wes} Glejo) wes) A) Ww) ely) eas lee) ws} EI)
B.M. 29814 — — I6I 70 —
B.M. 29752 94 46 12:2 5:0 (Ci) ESS (TET IS
B.M. 29752 Ae eee ee OA let 4) a (248)! 1(672)115575
S.M. C31347a —- — — — 12:2 63 (7:99) — I04 5:0 15:0 66 —
S.M. C31347b 96 47 — — 128 61 7:6 4:2 10:5 4:8 14:5 6:3 (57-0)
S.M. C31344 I22 61 7:6 4:0 10:0 5:2 — — —
S.M. C31346 — — 163 5:0 — —~— —~ ~ — — —_—- — —
B.M. M25074 Sa ae ce neg 2. ante 7-3 F4rS) LE QM" 597. 45%5
B.M. M13075 OP FO uOG Fee wu OF why Foe) aso GA avies) Fite) Foro)
B.M. M13075 WOO BO wine Bet wits xo kyl fle! aetsy ES} auitey (Os GSH)
P2 p3 p4 M1 M2 P4-M2
Gi=jo) GS) lef) aes) I=) ass BI=jo) was} jo) was} IR99)
B.M. M25073 76 36 ITO 5:2 124 63 — — — —~— —
S.M. C31347¢ —- — — ~~ ~~ — 113 107 — ~~ —
S.M. C31347d —- — — — Ito t0:2 9:8 7:3 (14:0) 9°3 (32°5)
S.M. C30775 pie aetna arn aS) 0.94105, (0:7) = =
B.M. M25075 1 OMe ACO Gu hs >2 eI li-O) LO 7-O)) L745 O-O) Aly
Order CARNIVORA
Family MIACIDAE Cope, 1880
Dracnosis. P4, M; carnassials as in modern carnivores. Post-carnassial teeth
more or less tubercular. Metacone smaller than protocone (Matthew, 19009).
REMARKS. The family is recorded from the Middle Palaeocene-Late Eocene of
North America; Late Eocene of Europe and Late Eocene—?Early Oligocene of Asia.
The group has been traditionally divided into two subfamilies (Simpson, 1945 and
Piveteau, 1961): Viverravinae Matthew, 1909 and Miacinae Trouessert, 1885.
Subfamily VIVERRAVINAE Matthew, 1909
Dracnosis. Molars 5. Antero-external cusp of P4 and posterior accessory cusp
of Pyprominent. Elongated oval outline of M2. Mabsent. Skull much elongated.
(Partly from Matthew, 1900).
Remarks. Distinguished from the other subfamily as follows: in the Miacinae
the skull is short or moderately elongated; molars 5 and the antero-external cusp of
P4 and the posterior accessory cusp of P4 rudimentary or absent.
90 UPPER EOCENE MAMMALIA
The Viverravinae contains five genera:
Didymictis Cope, 1875. Middle Palaeocene-Early Eocene; North America.
Ictidopappus Simpson, 1935. Early-Middle Palaeocene; North America.
Protictis Matthew, 1937. (incl. Simpsonictis MacIntyre, 1962.). Middle (and
Late?) Palaeocene; North America.
Viverravus Marsh, 1872. Early-Middle Eocene; North America. ?Early Oligo-
cene; Asia.
Quercygale Kretzoi, 1945. Late Eocene; Europe.
In 1962, MacIntyre described a new genus Simpsonictis. More recently, MacIntyre
(1965 : 3544) has separated Protictis Matthew, 1937, from Didymictis as a distinct
genus: Protictis was considered to include Simpsonictis MacIntyre, 1962.
Genus QUERCYGALE Kretzoi, 1945
1965 Humbertia Beaumont: 142.
DraGnosis. Viverravinae of medium to large size. P4 with or without parastyle,
deuterocone strong and projected. M1 with strong parastyle, paracone robust and
flattened externally, metacone smaller and more conical, no true metaconule but a
paraconule present. Protocone near the anterior border bearing two crests. No
hypocone. M? much smaller than M! and very elongated transversely. Lower
premolars have a posterior cuspule low and trenchant. My, high with a metaconid
higher than the paraconid. Talonid short with hypoconid trenchant and dominant.
Mz small with trigonid complete and talonid reduced and trenchant. Symphysis high
and short, coronoid process more or less reduced (modified from Beaumont, 1965).
TYPE SPECIES. Quercygale angustidens (Filhol, 1872). ?Early Ludian (Quercy
Phosphorites) ; France.
REMARKS. Teilhard de Chardin (1914-15) first recognised the viverravine affinities
of Viverra angustidens Filhol, placing it in the American genus Viverravus. This
European material was separated by Kretzoi (1945) as a new genus Quercygale.
Beaumont (1965) placed ‘Viverravus’ angustidens into a new genus Humbertia and
described a new species Humbertia helvetica (Ritimeyer, 1862). Humbertia Beau-
mont, 1965, is synonymous with Quercygale Kretzoi, 1945 (Beaumont, 1966 : 282).
Quercygale differs from the other genera as follows: in Viverravus, the Mg trigonid
is high and the talonid is long with a distinct hypoconulid; M1 has a hypocone
(usually), a paraconule is absent and the P‘4 parastyle is invariably present.
Ictidopappus has upper teeth with a pronounced transverse elongation; the talonid
of Py is short with a very small cusp and the lower molar talonids are basined. In
Protictis, the My; and Mg trigonid is higher and the talonid is larger and wider trans-
versely; Pz and P, have a well separated anterior cusp. As observed by Beaumont
(1965), Quercygale appears closest to Didymictis but in this latter genus, the My and
Mz talonid is basined and the P, parastyle is invariably present. Beaumont (1965)
observed that apart from the absence of Mg, Quercygale is very similar to Tapocyon
Stock, 1934, a miacine from the American Late Eocene. Van Valen (1967) has
considered that this presence or absence of Ms is inadequate for generic distinction,
HEADON BEDS OF ENGLAND gl
and that Quercygale is best considered as a subgenus of Tapocyon. Following this,
Van Valen has concluded (1967 : 255) that . . . ‘the Miacinae can no longer be
unambiguously distinguished from the Viverravinae on the basis of the presence of
three molars rather than two’.
Three species are recognised here:
Q. angustidens (Filhol, 1872), pl. 17, figs 33-35. Late Eocene (Bartonian-Early
Ludian) ; France.
Q. helvetica (Riitimeyer, 1862), p. 86, pl. 5, fig. 86. Middle Eocene (Lutetian) ;
Switzerland.
Q. hastingsiae (Davies, 1884), p. 433, pl. 15, figs 1-4. Late Eocene (Early Ludian) ;
Britain.
Beaumont (1966) noted that Miacis? macintyri Van Valen, 1965, from the Lutetian
of Germany, is very similar to Quercygale helvetica. Reference of Miacis? macintyri
to Quercygale was supported by Van Valen (1967). The species is not well known
(based on an isolated P4 and M!) and may well be synonymous with Quercygale
helvetica.
Depéret (1917) described a new species Procynodictis euzetensis (later in the same
work described as Viverravus euzetensis) from the Early Ludian of France. This
species was considered by Gauthier & Favre (1948) to fall within the range of
variation of ‘Viverravus angustidens’. This latter view was followed by Beaumont
(1965 : 143) who recorded Procynodictis euzetensis as a synonym of Humbertia
angustidens. Although the holotype of Procynodictis euzetensis has not been examined
and Depéret’s illustrations (1917, text fig. 2; pl. 24, figs 3-6) make precise comparison
difficult, Procynodictis euzetensis is synonymised with Humbertia angustidens in the
present work, on the authority of Gauthier & Favre, and Beaumont.
Quercygale hastingsiae (Davies, 1884)
(Pl. 6)
1884 Viverva hastingsiae Davies: 433, pl. 15, figs. 1-4.
1885 Viverva hastingsiae Davies; Lydekker: too, figs 11-12.
1915 Vivervavus angustidens (Filhol); Teilhard de Chardin (partim): 117.
1917 Vivervavus hastingsiae (Davies); Depéret: 248.
1965 Humbertia angustidens (Filhol); Beaumont (partim): 143.
Diacnosis. About the same size as Q. angustidens. My trigonid somewhat wider
and larger than talonid, with fairly blunt anterior apex; talonid larger than that of
_ Q. angustidens with squared outline. Outline of upper teeth rounded; internal lobe
of P8 large; posterior margin of M! with strong embayment.
Hototyre. B.M. 30203. Skull and mandible in occlusion. Badly crushed
and damaged. The outline of the skull is preserved in fine greenish sand matrix,
but only the lower part is distinct and relatively undamaged. The right and left
halves of the skull have been displaced by crushing along the palatal sutures; the
occluded mandibular rami are also similarly displaced relative to each other. To
92 UPPER EOCENE MAMMALIA
facilitate examination, the posterior part of the right ramus has been removed (in
the past) from its original occluded position. The removed right ramus shows
P3—Mg, the exposed right maxilla and premaxilla, I-C-M2. The upper and lower
teeth anterior to P3 are damaged, displaced and partly obscured by matrix.
Described and figured, Davies, 1884 : 433, pl. 15, figs 1-4. Lydekker, 1885a : 100,
figs 11-12. From Lower Headon Beds (Mammal/Crocodile Bed), Hordle Cliff.
The only known specimen from the Headon Beds.
DESCRIPTION. Pg is rectangular in outline and elongated antero-posteriorly,
with a slight anterior narrowing. The anterior margin is squared, the posterior
margin more rounded. There is a large crested central cusp and a prominent talonid
which bears a small cusp. A slight postero-external and antero-internal cingulum
is present. Py, is generally similar to P3 but much larger, and the anterior narrowing
is more prominent. The talonid bears an antero-posteriorly aligned crestiform cusp
and is basined internally.
The largest tooth is Mj; this has a stout bulbous trigonid which is much wider and
longer than the talonid. The protoconid, the largest cusp, is opposite the metaconid ;
both cusps are anteriorly and transversely crested. The paraconid, the smallest
trigonid cusp, is crestiform with its internal edge strongly internal in position.
There is a short but wide antero-external cingulum. The small talonid is structurally
similar to P4; the internal region is slightly basined and there is an antero-posteriorly
aligned crestiform cusp. Mg is very short antero-posteriorly. The trigonid is
almost as wide transversely as the talonid of M;, with very low cusps; the protoconid
is largest and slightly anterior to the metaconid. A low curved crest running from
the internal paraconid to the protoconid, forms the squared anterior tooth margin.
The antero-external part of this crest is badly worn; it may represent the site of a
small cusp. The talonid, small and narrowing posteriorly, is well worn and shows
no evidence of any cusps.
The upper teeth anterior to and including P2 have been much displaced from their
original positions. One upper incisor is preserved; it is small and simple. The
upper canine is large and curved witha strong root. P1is embedded in matrix; it is
smaller than the incisor with a single central cusp. The anterior region of P? is
missing. The tooth has a central major cusp but is larger and more elongated
antero-posteriorly than P!. P% is strongly elongated antero-posteriorly and very
much larger than P2. The tooth narrows anteriorly and is dominated by a large
central crested cusp. Posteriorly there is a low rounded separated cusp and a
postero-external shelf-like cingulum. P# is triangular-shaped in outline and much
larger than P38. The tooth is dominated by a large central crested paracone;
antero-externally this falls as a crest towards a low rounded parastyle. There is a
low rounded protocone which is larger than the parastyle. The anterior margin
between parastyle and protocone is strongly embayed. The crestiform metacone is
separated from the paracone by a deep cleft which continues externally as a narrow
valley; these cusps together form a sharp oblique postero-internal shearing edge.
There is a small distinct antero-external cingulum and a faint postero-external
cingulum.
HEADON BEDS OF ENGLAND 93
Ml is the same width transversely as P4. The metacone is directly posterior to the
larger paracone; a posterior metacone crest curves antero-externally along the
external tooth margin to the parastyle. The crested protocone is crescentic in
shape, an anterior crest running antero-externally towards the paracone; a low
paraconule is developed on this crest just antero-internal to the paracone. The
parastylar area extends externally as a wing which is separated from the postero-
external area by a smallembayment. The parastyle is crestiform; it is joined to an
external paracone crest. There is a distinct internal and antero-internal cingulum.
M2 is much smaller than M! and a little smaller than P?. The ovoid tooth lies
obliquely in the tooth row. The antero-external margin is in contact with M! just
posterior to the M1 metacone. The crown of the tooth is badly damaged and no
details can be discerned.
RemMARKS. Teilhard de Chardin (1915 : 117, footnote 1) considered that ‘Viver-
yavus’ hastingsiae was synonymous with ‘Viverravus’ angustidens. Depéret (1917 :
248) recorded several differences, on the basis of which he believed the two species
to be distinct. Gauthier & Favre (1948 : 116) emphasised the larger Mj talonid of
‘Viverravus’ hastingsiae but were uncertain whether this merited specific separation.
More recently Beaumont (1965 : 143) on the evidence of the more elongated Mj
talonid, considered ‘Humbertia’ hastingsiae as a distinct subspecies of ‘Humbertia’
angustidens. Depéret’s view is followed here. In addition to the considerably
more elongated talonid of M;, the P? of Quercygale hastingsiae (from comparison
with illustrations of Q. angustidens) has a smaller internal heel as Depéret observed.
Although this single specimen from the Headon Beds may be aberrant, it seems
reasonable at present to regard it as specifically distinct.
TABLE 18
Measurements (in mm) of Quercygale hastingsiae
P3 P4 Mi M2
a-p trs a-p trs a-p trs a-p trs
B.M. 30203 7:0 2:8 Q:1 317) 9°7 6:2 51 3:6
Pps Pp M1 M2
a-p trs a-p trs a-p trs a-p trs
B.M. 30203 7:8 — I13 — 6:7 — — —-
IV. CORRELATION AND AGE OF THE HEADON BEDS
Although there is general agreement concerning the broad correlation of Upper
Eocene and Lower Oligocene deposits in Europe, no unanimity has been reached yet
regarding the detailed correlation of marine and non-marine sediments in the different
depositional basins. One particular problem is the exact siting of the Eocene/
Oligocene boundary. The situation is hindered by a confused stage and substage
94 UPPER EOCENE MAMMALIA
terminology, which despite many recent attempts at clarification by authors, has
still to be resolved.
It is not intended here to present a critical review of this problem, detailed aspects
of which have been considered recently by Rey (1964), Krutzsch & Lotsch (1964),
Blondeau, Cavelier, Feugueur & Pomerol (1965), Thaler (1966), Denizot (1968),
Cavelier (1968), Curry, Gulinck & Pomerol (1969) and others. The complete
mammalian fauna of the Headon Beds has not yet been studied in detail, and
authoritative use of the mammals in correlation and dating is clearly premature.
Nevertheless, in view of the importance of the Headon Beds in current stratigraphical
research, some interim conclusions are presented here.
The exact age of the Headon Beds and indeed of most of Forbes’ ‘Fluvio-marine
series’ has long been controversial. The uppermost beds of the British succession,
the Hamstead Beds, are generally accepted to be Oligocene in age. No unanimity
has been reached regarding the age of the underlying Bembridge, Osborne and
Headon Beds. Three schemes have been proposed in the past for the age of the
Headon Beds; two schemes are supported currently by European stratigraphers and
palaeontologists, but the third, that proposed by Reid & Strahan (1889) is now
obsolete.
Reid & Strahan (1889) and Arkell (1947) considered that the Lower, Middle and
Upper Headon Beds were Oligocene (Lattorfian of Arkell) and the underlying Barton
Beds were uppermost Eocene. The Eocene/Oligocene boundary lay at the base of
the Lower Headon Beds. This view, in which is emphasised the general change from
marine to brackish and freshwater conditions at the beginning of Lower Headon
times, has not found further support.
In a currently held view, supported traditionally by most British workers, the
Lower Headon Beds are considered as uppermost Eocene (Bartonian) and the Middle
and Upper Headon Beds are Oligocene (Lattorfian, Tongrian). The Eocene/
Oligocene boundary is placed, therefore, at the base of the Middle Headon Beds.
This view, based on correlation by mollusca and foraminiferida of the Middle
Headon Brockenhurst Bed with the type Lattorfian (Lower Oligocene) of Germany,
has been adopted by Wrigley & Davies (1937), Bhatia (1955), Denizot (1957, 1968),
Curry (1958a, 1966), Chandler (1g61a, 1963), Rey (1964), Franzen (1968) and others.
In the second currently supported arrangement, the Lower, Middle and Upper
Headon Beds are placed together with the Barton Beds in the Upper Eocene (Lower
Ludian). This was adopted by Stehlin (1910) and followed more recently by
Krutzsch & Lotsch (1964), Blondeau, Cavelier, Feugueur & Pomerol (1965), Thaler
(1966) and Cavelier (1968). In this scheme, the Eocene/Oligocene junction is placed
at some position above the Upper Headon Beds. Stehlin (1910) on the basis of
what he believed to be a fundamental break in the mammal faunas (‘grande coupure’),
considered that the Eocene/Oligocene junction in Britain should be placed at the
base of the Hamstead Beds; this has been followed more recently by Krutzsch &
Lotsch (1964), Blondeau et al (1965) and Cavelier (1968). Thaler (1966) preferred
to place the boundary at the base of the Bembridge Beds.
Stehlin (1910) compared the mammalian fauna of St.-Hippolyte-de-Caton (=Euzet-
les-Bains) with the Lower Headon Beds, considering them Lower Ludian in age.
HEADON BEDS OF ENGLAND 95
The same faunas were compared by Depéret (1917) who also assigned them a Lower
Ludian age. Stehlin (1910) also compared the mammalian fauna of Montmartre
with a similar fauna from the Bembridge Limestone; he considered these faunas to
be Upper Ludianinage. According to Stehlin (1910) the fauna from the Bembridge
Marls was similar to that from the Bembridge Limestone. He observed that the
Upper Ludian (Bembridge Beds) fauna differed from that of the Lower Ludian
(Lower Headon Beds) fauna by the extinction of several species and the arrival of
several forms unknown in the Lower Ludian. Stehlin’s correlations of the Lower
Ludian faunas of Euzet-les-Bains with the Lower Headon Beds, and the Upper Ludian
faunas of Montmartre with the Bembridge Limestone, have now become well
established.
Recently, Thaler (1965, 1966) proposed to replace the old classification by a series
of seven zones defined by theridomyids. Although some of Thaler’s zones transgress
the traditional stratigraphical subdivisions, the Euzet zone, Montmartre zone and
Ronzon zone correspond exactly with the Lower Ludian, Upper Ludian and
‘Sannoisian’ respectively.
Stehlin (1910) noted that as the faunas of the beds above the Lower Headon Beds
and below the Bembridge Limestone were poorly known, it was not possible in
Britain to determine the precise junction of the Lower and Upper Ludian faunas.
More recently, Krutzsch & Lotsch (1964) grouped the Upper Headon Beds and
Osborne Beds in the Montmartre zone with the Bembridge Beds; for them, the
junction of Lower and Upper Ludian faunas lay at the base of the Middle Headon
Beds.
The mammalian fauna obtained recently from the Upper Headon Beds appears to
be similar to that from the Lower Headon Beds and apparently distinct from the
Bembridge fauna. While the sample from the Upper Headon Beds is clearly still
incomplete, the Upper Headon Beds, on present evidence, should be grouped, at
least provisionally with the Lower Headon Beds in the Upper Eocene (Lower
Ludian). The Upper Headon fauna should be placed in the Euzet zone, rather than
the Montmartre zone as suggested by Krutzsch & Lotsch (1964). It follows that,
on the basis of the mammalian faunas, the traditional British practice of siting the
Eocene/Oligocene boundary at the base of the Middle Headon Beds, cannot be
sustained. This present view is in agreement with Martini (1970) whose study of
calcareous nannoplanckton from the Brockenhurst Bed, has indicated that the
Brockenhurst Bed is uppermost Eocene in age, and not equivalent to the type
Lattorfian (Lower Oligocene).
The Eocene/Oligocene boundary therefore, must be situated at some position
above the Upper Headon Beds. Owing to the poorly known mammalian fauna,
the exact stratigraphical position of the overlying Osborne Beds is uncertain at
present. However, study of this fauna, together with the fauna of the Bembridge
Beds, currently in progress by Mr A. Insole at the University of Bristol, may clarify
this situation.
96 UPPER EOCENE MAMMALIA
V. REFERENCES
ARKELL, W. J. 1947. The geology of the country around Weymouth, Swanage, Corfe and
Lulworth. Mem. geol. Suvv. U.K., London, 386 pp.
BraumontT, G. DE. 1965. Les Viverravinae (Carnivora, Miacidae) de l’Eocéne de la Suisse.
Bull. Soc. vaud. Sci. nat., Lausanne, 69 : 133-146, pls. 1-2.
1966. Les Miacinae (Carnivora, Miacidae) de l’Eocéne de la Suisse. Bull. Soc. vaud. Sci.
nat., Lausanne, 69 : 273-285, pl. r.
BuatTiA, S. B. 1955. The foraminiferal fauna of the late Palaeogene sediments of the Isle of
Wight, England. J. Paleont., Chicago, 29 (4) : 665-693, pls. 66-67.
BraInviL_te, H. M. DE. 1839-64. Ostéogyaphie ou description iconographique comparée du
squeletie et du systéme dentaive des mammiferes vécents et fossiles pour servir de base a la
Zoologie et a la Géologie. Paris.
BLonDEau, A., CAVELIER, C., FEUGUEUR, L. & PoMEROL,C. 1965. Stratigraphie du Paléogéne
du Bassin de Paris en relation avec les bassins avoisinants. Bull. Soc. géol. Fy., Paris, (7)
7 : 200-221.
Butrer, P. M. 1946. An arctocyonid from the English Ludian. Ann. Mag. nat. Hist.,
London, (11) 13 : 691-701, 2 figs.
Cavetier, C. 1968. L’Eocéne supérieur et la base de l’Oligocéne en Europe occidentale.
im Colloque sur l’Eocéne, Paris, 1968. Mém. Bur. Rech. géol. minier., Paris, 58 : 507-527.
CHANDLER, M. E. J. 1922. A recent exposure of the ‘Marine Bed’, Hordle, Hants. Geol.
Mag., London, 59 : 224-229, 1 fig.
1925. The Upper Eocene flora of Hordle, Hants, Part 1. Palaeontogy. Soc. (Monogy.),
London, 1-32, pls. 1-4.
1926. The Upper Eocene flora of Hordle, Hants, Part 2. Palaeontogy. Soc. (Monogy.),
London, 33-52, pls. 5-8.
—— 1961a. The Lower Tertiary floras of Southern England. 1. Palaeocene floras. London
Clay flora (supplement). 354 pp., 34 pls. Brit. Mus. (Nat. Hist.), London.
1961b. Flora of the Lower Headon Beds of Hampshire and the Isle of Wight. Bull
Br. Mus. nat. Hist., London, 5 (5) : 91-158, pls. 24-30.
1963. Revision of the Oligocene floras of the Isle of Wight. Bull. Br. Mus. nat. Hist.,
London, 6 (3) : 321-384, pls. 27-35.
CHARLESWORTH, E. 1845. On the discovery, by Mr Searles Wood, of an alligator in the
freshwater cliff at Hordwell, associated with extinct mammalia. Rep. Br. Ass. Adumt Sci.,
London, 14 : 50.
Cuatwin, C. P. 1960. The Hampshire Basin and adjoining areas. By. veg. Geol., 3rd ed,
99 pp., 42 figs. London.
CuHow, M. 1964. Rhabdocolpus
at one horizon in the Lower Greensand [Crackers], requires further consideration and
palaeoecological interpretation.
Haas, after investigating South American Triassic material, concluded that the
relationship of the genera Protofusus, Paracerithiwm and Rhabdocolpus is much closer
than that implied by Cossmann’s classification, although as he pointed out (pp. 212
& 293), their exact position can only be discovered from a thorough investigation of
all Triassic Procerithiidae. He showed that the morphological changes occurring in
these genera, particularly in their ornament, could be interpreted as a series, with
Rhabdocolpus the most highly specialized of the three. However, the evidence and
determination of the earliest representatives of these genera is confusing and such a
lineage remains doubtful. A further complication is the close resemblance, especially
of ornament, of Rhabdocolpus to the Triassic genus Andagularia Haas, which is
considered to belong to the Purpurinidae.
?Rhabdocolpus ?clementinum (d’Orbigny)
(Pl. 5, figs 6, 12 & 13)
?1843 Cerithium clementinum d’Orbigny: 357, pl. 228, figs 1-3.
1845 Cerithium clementinum d’Orbigny; Forbes: 352, pl. 4, fig. 9.
MATERIAL, LOCALITY AND HORIZON. BM(NH) GG 5692-5694, GG 20841-20843,
GG 20844 (5), Crackers, Lower Greensand; GG 5708 (5 specimens), Lower Lobster
Bed, Lower Greensand; Atherfield, Isle of Wight.
DescripTion. A very small, turriculate shell with a narrow spire, flat-sided
whorls and a grooved linear suture. The whorls are ornamented by 16-20 axial ribs,
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 143
that are orthocline on the early whorls but soon become strongly opisthocline.
Generally, five primary spiral cords are visible on the whorl side, with occasionally a
secondary cord between the second and third. Small rounded tubercules occur at the
intersections of the axial ribs towards the base of the whorl; these tubercules become
somewhat rectangular on the abapical cords.
The base is convex and has a varying number of spiral cords ornamenting its
surface, although there are always a pair of primary cords forming small carinae
some distance from the periphery and another pair of strong cords close to the
columella. The entire semi-rounded aperture is flat at its base and has a straight
inner lip.
In complete specimens, the growth lines are visible on the body whorl and these
are initially strongly prosocline, but at the second spiral cord swing back to become
strongly opisthocline, producing a rather asymmetric opisthocyrt shape.
Discussion. Unfortunately, it has not been possible to discover Forbes’ original
specimen in the Geological Survey Collection. The various specimens from the
same locality listed above, bear a strong resemblance to his figure of a turriculate,
gastropod with cancellate ornament. Judging from the specimens in the British
Museum (Natural History) collection, it would appear that in his figure, the ribs are
too numerous, and that the dimensions of the whorls are incorrectly drawn.
The only other figures of C. clementinum are those given by d’Orbigny (1843 : 357,
pl. 228, figs 1-3) and these do not show the typically cancellate and tuberculate
ornament of the British specimens. D’Orbigny’s figures show smooth cords and
ribs that are orthocline, or only opisthocyrt rather than the strongly opisthocline
ribs of the British Aptian fossils. In fact, d’Orbigny’s figure could equally as well be
interpreted as an attempt to represent the ornament of C. forbesianum d’Orbigny
[= C. phillipst Leymerie of Forbes].
Wollemann (1900 : 169) suspected that Forbes’ Aptian fossil was not identical to
those described by d’Orbigny from the Neocomian of Marolles (Dept. d’Aube),
although he did not explain his doubts. In his description of C. clementinum, he
mentioned that the axial ribs of subsequent whorls are irregularly alternate and that
the whorls are convex, while he described the aperture as being quadrangular and
depressed. Such a description could not be applied to the specimens referred here to
Forbes’ 1845 figure.
It is not advisable to introduce a new specific name for the British material, until
d’Orbigny’s material has been re-examined and his types recognized. Peron
(I900 : 192) commented that the ornament of C. clementinum is extremely variable
and that the specimen figured in the ‘Paléontologie francaise’ is a very rare form.
? Rhabdocolpus forbesianum d’Orbigny, 1850
(Pl. 5, figs 1-5)
1845 Cevithium phillipsi Leymerie; Forbes: 352, pl. 4, fig. 12.
1850 Cerithium forbesianum d’Orbigny: 116
1854 Cervithium forbesanum d’Orbigny; Pictet & Renevier: 52, pl. 5, fig. 6.
1864 Cerithium forbesianum d’Orb.; Pictet and & Campiche; 286.
144 BRITISH CRETACEOUS GASTROPODS
1868 Cevithium forbesianum d’Orb.; Verneuil & Loriére: 15, pl. 2, figs 7a-e.
1906 Uchauxia forbesiana (d’Orb.); Cossmann: 57 & 252.
1961 Uchauxia forbesiana (d’Orb.); Casey: 607.
MATERIAL AND OCCURRENCE. Four specimens in the collections of the Geological
Survey Museum have been accepted as the original material in the Geological
Society Collection that was described by Forbes (1845) as C. phillipst. Two of the
specimens bear the Geological Society number 2267 and the other two are associated
with a label numbered. 2275 and all are from the Lower Greensand of Atherfield,
Isle of Wight. Blake (1902) lists R 2267 in his catalogue of type material at the
Geological Society and, it is true, that a specimen with this number in a small rock
fragment, bears ornament similar to that shown in the original figure on Forbes’
plate 4. However, not one of these specimens is exactly like the figure, which is
probably a composite representation of the features shown by these incomplete and
poorly preserved specimens. This material must, therefore, be accepted as a type
series and one of them, GSM Geol. Soc. 2275 (a)—an incomplete specimen but clearly
showing the ornament of later whorls,—is selected here as a lectotype.
I have used better preserved, topotype material from the Crackers, Lower Green-
sand, in the British Museum (Natural History) collections to describe this species;
GG 20845-20849, GG 20850 (6), GG 5609, GG 5695-5697, G 20898 and G 20908.
Another specimen G 20983 comes from East Shalford in Surrey, an Atherfield Clay
locality mentioned by Casey (1960 : 547).
DeEscRIpTION. A very small shell (on average 2-4 mm in height), composed of
convex whorls that are separated by a grooved linear suture; in the early whorls the
suture may appear to undulate owing to the effect of the axial ribs. The ornament
of the early whorls is quite different from that of the later whorls. Initially,
ornament consists of several spiral cords and 10-14 opisthocline axial ribs, with
slight tubercules occurring where the cords cross the ribs. After 3 or 4 whorls the
axial ribs disappear but continue to influence the cords, first as the tubercules
briefly persist and then by producing slight crenulations and, or, thickenings in the
smooth ribbon-like spiral cords. On later whorls, six raised-primary cords are
generally visible but some variation in this number occurs. Frequently, the third
and fourth cords from the adapical suture are the strongest and these occur at the
centre of the whorl. In other examples, the strongest cords occur towards the
abapical suture with the weakest on the adapical shelf. The interspaces between the
spirals are considerably wider than the width of the cords. Secondary cords develop
in these interspaces, but not according to any definite pattern.
The rounded base is ornamented by a varying number of spiral cords with a primary
cord close to the periphery and two other primary cords close to the columella.
Growth lines are only visible on the whorl sides as striae between the cords, but they
are easily seen on the base and swing back towards the columella.
The semi-rounded aperture is entire, has a flat basal lip, a thin outer lip, straight
inner lip and a smooth columella.
Discussion. Several authors have followed d’Orbigny and used the name
forbesianum to describe this British Aptian fossil, but few have endeavoured to
PROCERITHIIDAE, CERITHIIDAE & CERIDHIOPSIDAE 145
distinguish it from Cerithium phillips: Leymerie 1842. D’Orbigny first used the
name forbesianum in his Prodrome (1850 : 116) when recording the British material,
which he considered Forbes had misidentified, but did not provide any detailed
explanation. Pictet and Campiche (1864 : 287) suggested that the slender elongate
form of both species may have led to the confusion, but commented that the rare
varices that occurred in C. phillips: provided an easy method of distinguishing the
two species. These varices are easily seen in d’Orbigny’s figure of the French species
(1843 : pl. 227, fig. 10) but are not shown in Leymerie’s own figure (1842 : pl. 17,
figs Ira & b) of this more elongate shell. Verneuil and Loriére (1868 : 15) recognized
that C. forbestanum was quite different from C. phillipsi and provided the only
adequate figure (pl. 2, figs 7a-e) of the species to show its variable ornament, when
describing its occurrence at several localities in the Aptian of the Spanish province of
Teruel.
Cossmann (1906 : 57) obviously accepted, or also observed, the varices in
C. phillipsi Leymerie, for he included the species in his new genus Uchauxza in which
the possession of varices is a diagnostic feature. Yet, this evidence still needs to be
verified by examining type material, and, in order to ensure that phillips: does
belong to Uchauxzia, it is also necessary to ascertain that an anterior canal is present
for one has not been figured. At the same time, Cossmann (1906) referred
C. forbestanum to Uchauxia commenting that it wasclose to phillipsiand quoted Pictet
as his source. The only figure produced by Pictet is that in Pictet and Renevier
(1854 : pl. 5, fig. 6) which depicts a shell with ornament between that of forbesianum
and pluillips: [as understood from the figures quoted above]. However, the presence
of an entire aperture in several of the British specimens of forbesianum listed above,
shows that this species cannot belong to Uchauxia (as interpreted by Cossmann,
1906).
The turreted and flat-sided whorls of C. clementinum d’Orbigny; Forbes, together
with their persistent cancellate ornament, easily distinguish this species from the
differently ornamented forbesianum with its strong spiral cords and convex whorls.
Yet both have several features in common, such as a grooved linear suture, and a
semi-rounded entire aperture, while their early whorls can be confused if the
ornament is not adequately preserved.
The greater convexity of the whorls and their persistent orthocline axial ribs
distinguish C. dupinianum d’Orbigny (1843 : 357, pl. 227, figs 4 & 5) a similar species
to forbesianum, described from the Neocomian of Marolles (Dept. d’Aube).
? Rhabdocolpus melvillei sp. nov.
(Ele Spatie7)
Diacnosis. A small procerithiid with reticulate ornament on its early whorls but
which loses axial ornament on later whorls, and has a wider spiral angle than usual
in the genus.
Hototyre. In the Sedgwick Museum, B. 27340.
OTHER MATERIAL. Four paratypes in the Sedgwick Museum, Cambridge,
B. 27341-27344, all belonging to the Wiltshire Collection.
146 BRITISH CRETACEOUS GASTROPODS
LOCALITY AND HORIZON. Crackers, Atherfield Clay Series, Lower Aptian, Forbesi
Zone, near Atherfield Point, Isle of Wight.
DescripTIon. The shell is very small and has a wide spiral angle. The whorl
outline is strongly convex and the suture impressed. The earlier whorls show about
twelve opisthocyrt axial ribs. On laterwhorls, threespirals appearand small rounded
tubercles are developed where these intersect the axials. Later, a gap develops
between the first and second spiral cords and two further cords are produced on
either side of the first. The new spirals eventually become as strong as the earlier
formed ones. On the body whorl, the spirals are separated by interspaces greater
than their width and the axials are separated by interspaces greater than those
between the spirals. The rounded base is ornamented by several less-tuberculated
spiral cords.
Discussion. This species can be distinguished from ? R. forbestanum by its more
strongly-rounded whorl outline, by its greater number of axial ribs, slightly stronger
development of tubercles and wider spiral angle. The few specimens of ? R. melvilles
that are available do not show any indication of secondary spirals between the
primaries, nor the marked differentiation in ornament that occurs in forbesianum.
However, the full range of variation within forbestanum is not known and it could be
possible that ? R. melvillei is, merely, a variety of that species.
Family ? PROCERITHIIDAE
Genus ? UCHAUXIA Cossmann, 1906
TYPE SPECIES. By original designation Cossmann (1906: 56), Cersthiwm
peregrinorsum d’Orbigny 1843, from the Turonian of Uchaux, Vaucluse, France.
Discussion. The material described below, undoubtedly belongs to a common
genus and because of considerable similarity in form and ornament to species of
Uchauxia, they are regarded as belonging to that genus. However, owing to their
incomplete preservation this determination must remain provisional.
The genus Uchauxia is also subject to uncertainty, for a crucial factor in its
recognition is the apertural shape of its type species, U. peregrinorsa. The various
figures of that species provided by d’Orbigny (1843: pl. 231, fig. 3), Cossmann
(1906 : pl. 6, figs 37-40) and Roman and Mazeran (1920 : pl. 5, figs 42-44) do not
provide sufficient reliable evidence of this feature, although each author mentions a
rounded aperture with a short anterior canal in his description. In order to verify
these features it 1s necessary to refer to the type material. Unfortunately, this has
not been possible during this investigation, nor have other specimens of the species
been available for comparison.
For the moment, following Cossmann (1906 : 57), it is assumed that these small
Cretaceous shells are fully grown and that they resemble Procerithium and
Rhabdocolpus, but have a distinct, short anterior canal.
Uchauxia, as here understood, is generally larger than Cerithiopsis Forbes & Hanley
but does have a similar whorl shape and tends to have the same range of ornament.
Cerithiopsis, however, usually has a more advanced type of siphonal canal, which
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 147
does not project above the basal plane and resembles a keyhole in basal view.
Cossmann (1906 : 57) pointed out that Uchauxia only occurs in strata that are
believed to be entirely marine and for this reason, differs from both Potamides
Brongniart 1810 and Pirinella Gray, 1847.
Uchausia wisei sp. nov.
(Pl. 6, figs 5 & 6)
Diacnosis. Uchauxia with four prominently beaded spirals, the abapical
penultimate spiral being weaker than the others, and weakly developed axial
ornament.
Hototyre. BM(NH) GG 6161, C. W. Wright collection.
OTHER MATERIAL. Eight Paratypes BM(NH) GG 5611, GG 6158-6160 & GG 6162—
6165. C. W. Wright collection. All incompletely preserved.
LOCALITY AND HORIZON. Claxby Ironstone, Valanginian-Hauterivian, Nettleton,
Lincs.
Description. The shell is small, narrow and turriculate, slightly cyrtoconoid,
with a flat-sided whorl outline and linear suture. On the earlier whorls, three
primary spiral cords are developed with interspaces of equal width. The anterior
and posterior spirals become more prominent on later whorls, with the central spiral
moving posteriorly as a secondary spiral cord makes its appearance abapically to it.
A secondary spiral is also developed behind the posterior primary spiral. On still
later whorls, a secondary spiral cord may appear between the two posterior primary
spirals. The spirals cross 20 to 24 axial ribs, producing rounded tubercles at each
intersection. The axials are separated by interspaces almost equal to their own
width and are orthocline on the early whorls but become opisthocyrt.
The base is convex and bordered by two spiral cords; the axials do not extend on
to it and the surface is almost smooth. The small, oval aperture has a curved
anterior canal. The growth lines follow the axial ridges.
Dimensions. Holotype BM(NH) GG 6161.
Height (approx. 5 whorls). : . . ‘ 10-0 mm
Spiral angle ‘ : : Q : : ‘ a
Penultimate whorl height. ' ; : : 1-8 mm
Penultimate whorl diameter . ; ; ; : 3°8 mm
Discussion. The most characteristic feature of this species, when compared
with other species of Uchauxia that have four spiral cords, is the weak influence of
any axial component. In the Turonian type species, U. peregrinorsa, the axials are
quite strong, and similarly, in the Gosau species U. solida (Zekeli, 1852 : pl. 20, fig. 3)
and U. distinctum (Zekeli, 1852 : pl. 19, fig. 6), the exact age of which needs to be
investigated (see Kuhn, 1947 : 188).
U. wiser is distinguished from the two species described below, in that Uchauxia
sp. nov. has only three spiral cords of tubercles ornamenting the whorl and that its
tubercules have some axial alignment; while the prosocline axial ornament of
U. badri is quite diagnostic.
148 BRITISH CRETACEOUS GASTROPODS
The specimen described by Roman & Mazeran (1920: pl. 5, fig. 45) as Uchauxia
vichet, although having predominantly spiral ormament, has many more cords,
which have a lower degree of tuberculation, than U. wisei. Originally, their speci-
men was figured by Cossmann in his examples of U. peregrinorsa and it may still be
found to come within the range of variation of that species, when this aspect is fully
investigated.
Although the figures of the type species in both Cossmann (1906) and Roman &
Mazeran (1920) are inadequate for exact comparison, it is possible to decide that
peregrinorsa (d’Orb). has several more tuberculate spirals than this specimen of
U. wisei. The Turonian species described by Roman & Mazeran, U. richei, has three
granular cords but possesses many more primary cords and has a slightly different
whorl outline.
The British Museum (Natural History) collection included a number of Ageria
gaultina sp. nov. from the Gault of Folkestone, which at first sight, have some
resemblance to this fossil but the existence of a greater number of axial ribs and a
differing growth line pattern provide adequate distinguishing characters.
Uchauxsia badri sp. nov.
(Pl. 6, figs 3 & 4)
Diacnosis. A small, slender, cyrtoconoid, turriculate shell that is ornamented
by four tuberculate spiral cords, the tubercules of which are aligned along prosocline
axial ribs, giving a distinctive twisted appearance to the shell.
HototypPe. Sedgwick Museum, Cambridge, B 44632.
HORIZON AND LocALiTy. Upper Greensand, Albian, Blackdown, Devon.
Description. A small cyrtoconoid shell with a slender turriculate spire. The
whorl outline is slightly convex and the shell has a grooved linear suture. The
ornament of the earlier whorls is dominated by strong axial ribs which cross three
unequally-spaced spiral cords. On subsequent whorls, the number of cords increases
to four and eventually five, producing rounded tubercles at their intersections with
the axial ribs. The ribs consequently become less dominant and contribute to the
cancellate effect of the ornament. The interspaces between the spiral cords are then
equal and also equal to the width of the cords but considerably narrower than the
spaces between the axial ribs. The prosocline axial ribs are partly offset in relation
to one another on subsequent whorls and this gives a twisted look to the shell. On
later whorls, these ribs become reduced in strength, although this may, in part, be
due to wear and poor preservation. On the penultimate and body whorl, it is also
possible to see the fifth spiral cord, which occurs close to the abapical suture. The
axial ribs influence this, but although they may cross it, they then weaken towards
the suture.
The convex base is covered by faint collabral threads and ornamented by five
spiral cords, two of which produce small carinae. The small rounded aperture has a
partly twisted anterior canal.
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 149
Dimensions. Holotype, Sedgwick Museum, B 44632.
Height 10-6 mm
Mean spiral angle ; : : : ; : 20°
Height of last whorl . : : : : , 3°0mm
Maximum diameter. : : ; 4 2-9 mm
Discussion. The distinctive prosocline axial ribs, that give the ornament of this
species a slightly twisted look, have been mentioned in the discussion of the pre-
ceding species. It is this feature that immediately distinguishes U. badvi from the
type species U. peregrinorsa despite the figures of the latter being unsuitable for any
further detailed comparison. However, it is also possible to observe that the axial
tibs of pevegrinorsa are opisthocyrt and that its spiral cords are much thinner (i.e.
weaker) than those of badri.
One of the Lower Cretaceous species, forbesianum (d’Orbigny), that Cossmann
included in this genus, is now thought to belong to Rhabdocolpus Cossmann, 1906.
There is still some confusion concerning the characters of U. phillipsi (Leymerie) and
as suggested elsewhere in this paper, type material needs to be re-examined in order
to verify the validity of Cossmann’s determination. The ornament shown in the
few figures of this slender species is quite different from U. bad, but Leymerie’s
figure (1842, pl. 17, figs 11a & b) although not showing varices, does show orthocline
axial ribbing.
Cerithium kaunhowent Cossmann (1902 : 61), from the Maastrichtian of Maastricht,
differs in that, although having cancellate ornament, this is much finer and composed
of at least 6 untuberculated spiral cords and numerous opisthocyrt axial threads,
while the shell also has a definite anterior canal. Cerithium distinctum Zekeli
(1852: pl. 19, fig. 6) from Gosau, although having four predominate tuberculate
cords between the sutures, also has a thin cord in the centre of each interspace and
has strongly orthocline axial ribs.
Uchausia sp. nov.
(EL 6. fig 7)
MATERIAL AND LOCALITY. BM(NH) G 73788 J. S. Gardner Collection, Gault,
Folkestone, Kent. *
DEscriPTION. Unfortunately, this unique specimen is rather crushed and
measurements are not reliable, but it has been possible to discover details of the
ornament by developing out its reverse side. This gastropod can be described as a
small, more or less subulate, slender shell with convex whorls. Its moderately deep,
linear suture lies at the bottom of a concave furrow formed by adjacent whorls. The
initial whorls show bicarinate ornament, but a third spiral cord soon appears just
behind the posterior of the two initial cords, and others develop later. These spiral
cords are separated by wider interspaces and each interspace is occupied by at least
one almost smooth spiral thread. The primary cords are tuberculate and the
rounded tubercles are separated by intertubercular spaces that are nearly equal to
their width. Usually these are of the same number and strength on each cord, but
sometimes those on the posterior cord are slightly weaker. On the early whorls, the
150 BRITISH CRETACEOUS GASTROPODS
axial ridges are as dominant as the spiral cords, yet, later, with the increase in size of
the whorl they appear to become slightly disrupted and if there is any connection at
all, only a thin curved ridge joins the tubercules. These ridges tend to follow the
opisthocyrt growth lines. Towards the base the whorl flattens repeatedly; first to a
serrated cord making a slight carina and then again to another smooth cord. Only
the former is visible on previous whorls and would appear to mark the periphery of
the whorl. Traces of a short columella and an aperture with a short, shallow
anterior canal are present.
Discussion. Cerithiaceans with this general form are common amongst both
European and American gastropods and include many marine and non-marine
species. However, the Cretaceous age of the specimen and its occurrence in an
undoubtedly marine deposit, prompt its inclusion in the genus Uchauxia.
The ornament of the specimen serves to distinguish it from both U. wisei and U.
badri. It differs from wisez in the number of spiral cords and their interspace width,
while the possession of fewer tubercles on each cord and the stronger axial ridges
emphasize the differences between the two species. The more reticulate pattern
produced by cords, tubercles and ribs, assists in separating it from badri, although it
is the markedly prosocline ribs of the latter which separate the two species and
which give U. badri a slightly twisted look.
Family CERITHIIDAE Fleming, 1828
REMARKS. Cossmann (1906) discussed the relationship of the Procerithiidae and
the Cerithiidae (p. 20-21) and the derivation and evolution (p. 60-61) of the very
prolific Cerithiidae in detail. He concluded that the anterior canal was the most
important diagnostic character in separating the two families, even though the form
of the canal was extremely variable within the Cerithiidae and that the demarcation
between the subcanaliculate beak of the Metacerithiinae and the truncated canal of
the Potamidiinae would appear to be very slight but quite distinct. In his view, the
two families can easily be separated by the character of the canal, for even in the
most truncated cerithiid the anterior canal always forms a slight ridge or projection
beyond the surface of the base, while conversely, the beak of the Procerithiidae
never protrudes below the base of the shell. In the Cerithiidae the canal is also
generally twisted.
Although Cossmann was satisfied that the Cerithiidae were derived directly from
the Procerithiidae, it is possible that they may have arisen from the Jurassic family
Eustomidae (see Cossmann, 1906 & 1913). Members of this family, consisting of the
two genera Diatinostoma and Duitretus, possess a well-marked, but straight,
cerithiid-like canal and generally occur in the coral-rudist assemblage at horizons
from the Bathonian to the Turonian, (see Cossmann 1906 : 12-14). If this alterna-
tive should prove to be correct, the relationship of the Cerithiidae with holostomatous
forms might be further apart than Cossmann suspected. On the other hand, the
apparent relationship may be explained by homeomorphy.
It is particularly noticeable that from Upper Cretaceous to Recent times, the
members of the Cerithiidae have frequently been associated with corals. In the
PROGR RaGh ED AB, (Ch Rit ri DIA & CHRTTHTORSTDAE 151
Cretaceous, the richly coralline areas occurred further to the South and consequently,
the British record of the Cerithiidae at this time is very sparse. The available
evidence, which undoubtedly suggests that the Cretaceous was a period of experi-
mentation within the family, represents differing degrees of specialisation of the
entrance to the mantle cavity and of the anterior siphon. It is interesting to note
that typical holostomatous procerithiid forms also occurred at these times but only
persisted until the end of the Cretaceous.
Sub-family > CAMPANILINAE Wenz, 1940
Genus ? CAMPANILE Bayle (in Fischer), 1884
? Campanile cenomanica sp. nov.
(Pl. 6, figs 1 & 2)
Diacnosis. High, conical shell with whorls ornamented throughout by an adapi-
cal carina and three prominent spiral cords; the twisted columella lacks any folds.
MATERIAL AND OCCURRENCE. BM(NH) GG 18686; its history not certain, for
although the original label states “Chalk Marl, Craie Chloritique, Dorset’ and gives
the registered number 73597, the information in the register given with that number
could not appertain to this particular specimen. Its preservation does suggest that
it may come from the Cenomanian of Dorset, yet efforts to find similar material
both in collections and in the field have not been successful.
DEscriPTION. Unique turreted, conical, slender shell of medium size that is not
completely preserved. Flat whorls, having straight whorl outline with a slight
adapical shoulder. Adpressed linear suture. Whorl ornament characterized by a
high adapical astragal with a rounded edge and bearing beads that are separated by
intervals equal to their own width. Three prominent spiral cords are equally
distributed below this and in each interspace there is a faint spiral thread. The
central cord is somewhat weaker. Beaded ornament gradually develops on the
astragal and on the cords. A fourth, but smooth spiral cord can be seen at the
basal edge of the penultimate whorl. This cord marks the limit of the shallow
convex base, ornamented by numerous equally spaced spiral cords.
The spiral ornament is crossed by frequent opisthocyrt growth lines, which
straighten a little towards the abapical suture, but after crossing on to the base,
curve forward and then finally swing back towards the columella. At intervals
corresponding to the occurrence of the beaded tubercles, and following the growth
lines, there is a slight alteration in the growth direction of the shell.
The poorly preserved aperture is not large in proportion to the remainder of the
whorl, as in some species of Campanile. It shows a slightly twisted columella with a
distinct anterior canal and has a callused parietal lip. There are indications of a
small posterior ‘gutter’ or neck, and the form of the outer lip cannot be interpreted
with any certainty.
Discussion. The apertural features shown by this specimen, undoubtedly,
indicate a strong affinity with the Cerithiidae. Its ornament and general shell
morphology are comparable to those of the genus Campanile. However, the
D
152 BRITISH CRETACEOUS GASTROPODS
apparent atypical growth lines, the shorter anterior canal and lack of columella folds
could be interpreted as an indication that it would be better placed in the Cerithiinae.
Yet, on referring to the literature, one recognizes that the features generally accepted
as those of Campanile, because of its Eocene type species C. gigantewm (Lamarck)
1804, are only one end of the range occurring within that genus. In fact, another
Eocene species C. cornucopiae (J. Sow.) 1818, is basically very similar to the British
Cretaceous specimen, having a sub-rhomboidal aperture, a small twisted anterior
canal below a callused parietal lip, initially a flat whorl outline, a linear suture and
the same slight straightening of the opisthocyrt growth lines. The presence of two
columellar folds is the only difference between the two species.
Two Maastrichtian species from Iran described by H. Douville (1904 : 312, pl. 43),
C. morgami and C. breve show the rather simpler ornament of C. cenomanica. Another
Maastrichtian species C. cavezi Vidal, (1917; 7, pl. 3, fig. 6) from Sensui, Spain, has
the typical coronet ornament of later species of Campanile. All of these, however,
possess the distinctive columellar folds of the genus.
Whether the British fossil is a juvenile, the early whorls of an incomplete specimen,
or an early form of the genus Campanile, remains to be discovered. It certainly
does not show the characteristic two fold ornament that develops in Tertiary species
of the genus. More material is also necessary before one can attempt to investigate
its relationships with other Cretaceous species, or to make any conclusions as to its
possible position in the evolution of the campanilids. Cossmann recognized that the
genus first appeared towards the end of the Cretaceous, but C. cenomanica, if
correctly identified, would probably place their origin farther back than he suspected.
Genus EXECHOCIRSUS Cossmann, 1906
TYPE SPECIES. By original designation, Cossmann (1906: 121), Cerithium
cingillatum Zekeli, 1852 (pl. 18, fig. 6), from the Upper Cretaceous of Austria.
DESCRIPTION. Moderate-sized shells with weakly convex to flat-sided whorls.
Distinct adpressed suture, often undulates and is occasionally grooved. Ornament
extremely variable in size, strength and pattern, both within the genus and within
species; consists basically of tuberculate spiral cords and axial ribs. Adapical
ornament frequently prominent and separated from the remainder. Opisthocyrt to
opisthocline growth lines. Genus is characterized by the presence of a prominent
varix diametrically opposite the aperture on the body whorl. In some species,
varices occur on earlier whorls but these are never aligned as in other Cerithiidae. A
varix thins towards the adapical and abapical sutures. Small, low, rounded to
quadrangular aperture with a short anterior canal and a weakly developed adapical
‘gutter’, or channel. Smooth, curved columella with callused inner lip; basal lip
nearly flat, outer lip not expanded outwards.
DIscussION. Cossmann (1906: 121) considered Exechociyvsus a subgenus of
Tympanotonus Schumacher, 1817 on account of some similarity in ornament, but
more particularly, because of their apertural characters. However, despite these
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 153
resemblances it would seem reasonable to reserve the genus Tympanotonus for non-
marine gastropods and to place the marine Exechocirsus in a separate genus.
Exechocirsus also has certain similarities with other Upper Cretaceous and Lower
Tertiary cerithiid genera, e.g. Thericium, Clava (Clavocerithium), Clava (Pseudo-
vertagus), Serratocerithium and Teliostoma. There is a common element in shell
shape, size and in ornament patterns (in many cases varices occur at intervals),
while apertural features are also alike. Further studies may show that Exechocirsus
is a precursor of these genera and probably of other groups of the Cerithiacea such as
Potamides and even Cerithiwm itself. Certainly, from amongst the few Cretaceous
cerithiids, Exechocirsus appears to be the most likely ancestor for many of the
Tertiary forms and this warrants its consideration as a separate genus. The
possession of essentially cerithiid characters suggests inclusion in the sub-family
Cerithiinae (Cerithiidae) rather than to accompany Tympanotonus in the
Potamididae.
Péhelincev (1953 : 192) has attempted to explain the strong and varied ornament
occurring within the genus, by suggesting that it probably increased the mechanical
stability of the shells and he cites their prolific occurrence at Gosau, in company with
thick shelled Opisthobranchs and Nerinaea, as evidence.
With regard to the type species of the genus, Cerithium cingillatum Zekeli, it is
possible that this may be the early whorls of Cevithiwm pustulosum J. de C. Sowerby,
1835, a species which is also recorded from the Upper Cretaceous at the same locality
(Gosau, Austria).
Exechocirsus saundersi (Woods)
(Pl. 7, figs 3-5)
1896 Cevithium saunderst Woods: 92, pl. 4. figs 12 & I2a.
1921 Cervithiwm saundersi Woods, Ravn: 47.
HoLotyPe. Sedgwick Museum, B 4459, Cuckhamsley, North Farnborough,
Berks., Turonian, Holaster planus Zone.
OTHER MATERIAL. Chalk Rock, G 66724, Latimer, Bucks.; G 69915, Henley
Quarry, ? Medmenham, Bucks; GG 20821, GG 20825-20826, Hitch Wood Pit (Hill
End Farm), nr. Hitchin, Herts.; G 67704 H. planus Zone Dover; G 11523 Missen-
den, Herts.; GG 20829, Kensworth Quarry, Beds. Forbes (1960: p. 237) records a
large specimen from H. planus Zone, Western Colville, Cambs., SM. B 81559.
Description. Medium-sized, slightly elongate shell with almost flat-sided
whorls. Distinct impressed linear suture, which owing to the shell ornament appears
to undulate a little. There are four equal spiral cords, with interspaces from one
half to equal width between them. These cords are ornamented by rounded
tubercles, which are longitudinally aligned, occur at the junctions of the cords and
ribs and conform to the orthocline (or very slightly opisthocline) axial ribs. In
general, the axial ribs number about sixteen per whorl, but this varies according to
growth, and their interspaces are equal to their width. There are a varying number
of fine spiral threads between the tuberculated cords. In some specimens, a smooth
154 BRITISH CRETACEOUS GASTROPRODS
partly undulating cord occurs below the main spiral and just adapical to the suture.
Woods had described and figured this, but in the holotype this cord appears to be
composed of much smaller and more numerous tubercles (see pl. 7, fig. 5).
The base is flat to very slightly convex, has a sixth granular, sub-carinate cord
near the border and numerous spiral threads crossing its surface. The aperture is
only partially exposed but appears to be obovate to quadrangular. It shows a
short, well-formed anterior canal that is slightly curved, a callused inner lip and a
concave columella. A strong varix occurs opposite the aperture on the last whorl
and in a few specimens, there are indications of others on earlier whorls.
Discussion. The occurrence of a strong varix on the last whorl diametrically
opposite the aperture is characteristic of the genus Exechocirsus. In his original
description, Woods distinguished the new species from C. pustulosum, J. de C.
Sowerby (1835: pl. 39, fig. 19) by its smaller spiral angle and fewer tubercles.
Another feature is that the axial ribs are more definite ; they are also opisthocyrt and
only incorporate the tubercles of the lower three cords, those on the adapical cord
remaining unconnected. The tubercles on the abapical cord become somewhat
elliptical and on successive whorls they gradually diminish in size (see Zekeli, 1852,
pl. 19, fig. 5). Finally, there are undoubtedly more longitudinal rows of tubercles,
i.e. ribs, per whorl, in E. pustulosum (J. de C. Sowerby).
Cerithium pustulosum d’Orbigny (non Sowerby), 1843 resembles E. sawndersi in
ornament, but its longitudinal rows of tubercles are strongly opisthocline and are
also said to be farther apart (Zekeli, 1852 : 100). In addition, the suture is grooved
and the aperture more ovate.
Woods also separated E. sawndersi from C. pseudoclathratum d’Orbigny, 1850 (as
figured in Geinitz, 1874, pl. 31, fig. 5), yet, although the latter has similar ornament,
its true generic position is uncertain. Sohl (1960 : 83) included C. pseudoclathratum
with two American, Upper Cretaceous species that he has since (1964 : 364),
mentioned as being closely related to Ceritiella Verrill, 1882.
The Upper Cretaceous, Cerithium navasi Vidal, 1917 from Sensui, Spain, is easily
distinguished from E. saunders, in only having three tuberculate spiral cords and a
less prominent, finely-beaded, cord at the abapical suture.
It is difficult to decide the relationship of several British specimens which show
slight differences from typical sawndersi ornament, owing to the lack of knowledge of
the range of ornament variation occurring in that species. In a number of these
examples, the tubercles appear to be more numerous and this obscures any regular
axial alignment that they may have.
Two undescribed, unlocalized specimens from the Sowerby Collection G 60542—
60543, and others in material collected by C. W. Wright from Hitch Wood Pit,
Hitchin, Herts. (GG 20822-20824) show distinctive features (see pl. 7, figs 8 & 9g;
I,2&12). Basically there is a stronger development of both primary and secondary
spiral cords, while the axial ribs are fewer (12-14 per whorl) and consequently wider
apart. Among other features, the adapical cord is separated from the others by a
much wider interspace, the suture occurs in a groove formed between adjoining
whorls and varices frequently occur on earlier whorls.
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 155
In BM(NH) G 67704, (pl. 7, fig. 4) from the Chalk Rock of Dover, some of the very
earliest whorls are preserved. Initially, the whorls are ornamented by two tubercu-
late spiral cords, this eventually increases to three and within a further two whorls,
to four. The spire in this specimen is considerably more slender than those of other
species included in Exechocirsus. In another specimen, BM(NH) GG 20821, pl. 7,
fig. x) the tubercles are weakly developed but the axial ribs produce a rather longi-
tudinal step-like appearance in the shell, which also has a more distinctive suture.
Other material BM(NH) GG 5932-5934, GG 5982-5983, GG 5987 and GG 5771
(see pl. 7, figs 13 & 14) collected by C. W. Wright, from the Holaster planus Zone
at Kiplingcotes in East Yorkshire, is obviously related to E. saundersit. The slender
shape of the shells, their markedly opisthocyrt growth lines and opisthocyrt (rather
than opisthocline) ribs suggest they are a different form. Additional support to
such a view, is provided by their intermediate spiral threads, which are stronger than
in typical saundersi and by the fact, that in general, they are much smaller specimens.
It is possible that these features are the result of a phenotypic response to differences
in environment and for the moment, the specimens are regarded as Exechocirsus aff.
saunderst (Woods).
? Exechocirsus aft. subpustulosus Pcéhelincev, 1953
(Pl. 7, figs 10 & 11)
1953 Tympanotonus (Exechocivsus) subpustulosus Péhelincev: 193-194, pl. 33, figs 13-16.
1960 Tympanotonos (Exechocirsus) subpustulosus Pchelincev, in Orlov, pl. 19, fig. 13.
MATERIAL. BM(NH) G 737908 and G 16134, Upper Greensand, Upper Albian,
Haldon, Devon.
DEscrIPTION. Poorly preserved medium-sized turriculate shells with feebly
convex whorls that have a very small adapical sutural ramp, a slightly grooved
undulating suture and opisthocyrt growth lines.
The whorls are ornamented by about 18 narrow axial ribs, that are orthocline on
the early whorls but rapidly become opisthocyrt. These ribs are separated by inter-
spaces that are at least equal to their width. On the earlier whorls a narrow spiral
groove cuts the ribs to produce an adapical row of tubercles. Other similar spiral
furrows occur abapically to this, on later whorls, and these dissect the ribs even more,
with the result that the original predominantly axial ornament becomes changed to
a spiral pattern. A single spiral thread occurs in each groove. Numerous fine
spiral threads can be seen in the interspaces between the ribs. Further spiral
grooves occur towards the base on the last whorl. Neither the base of the shell or
the aperture can be seen.
Discussion. Lacking knowledge of apertural details and without any indication
that these specimens possess the characteristic varix of Exechocirsus, it is not
possible to refer them to that genus with certainty. They are provisionally referred
to the Transcaucasian Cenomanian Exechocirsus subpustulosus Pthelincev (1953) on
the basis of ornament resemblance. The axial ribs of swbpustulosus are also dissected -
by spirals to produce four rows of tubercles and both species have a rounded base.
156 BRITISH CRETACEOUS GASTROPODS
Although the adapical rows of tubercles is the most prominent and is also separated
by a wide groove from the lower rows, it has fewer tubercles than that of the British
specimens. Pchelincev remarked that there is a tendency for the ornament of
subpustulosus to become weaker on later whorls and this effect is also apparent in the
Haldon material. It is only the lack of varices on the earlier whorls, together with
the narrow apical angle, that enabled Pcthelincev to distinguish his species from
E. pustulosus (J. de C. Sowerby).
Family ? CERITHIIDAE
Genus AGERIA nov.
Named in honour of Dr D. V. Ager, Professor of Geology at the University of
Swansea, in recognition of his help and advice.
TYPE SPECIES. Ageria gaultina sp. nov.
DiaGnosis. Turriculate shells, ornamented by spiral cords and axial ribs; the
cords are frequently beaded and the abapical cord is characteristically strongly
developed and ornamented.
GENERIC CHARACTERS. Small-medium sized, slender, turriculate shells. Convex
whorls with a slight concavity before an abapical astragal. Flush, slightly grooved,
undulating suture. Regular ornament formed by axial ribs and spiral cords. In
some species the axial ribs weaken on later whorls, but in all species they have a
characteristic opisthocyrt shape. In most species, the primary spiral cords are
beaded, but in some the cords are smooth. A strong cord forms a distinctive
abapical feature which delimits the base. Opisthocline growth lines influence the
spiral ornament and show a median sinus, they swing back towards the columella on
the uniform basal slope. Axially elongate, quadrangular aperture and indication of
a short anterior canal with a smooth, slightly curved columella.
Discussion. The general shell shape and ornament, together with the slight
indication of a short anterior canal, suggest that this distinctive group should be
included in the Cerithiidae. Yet, the lack of adequately preserved material showing
the aperture prevents a definite conclusion being formed; particularly as the indica-
tions of an anterior canal in the type species A. gaultina sp. nov., could be the result
of crushing. Consequently, it is not possible to assign Agevia to a particular sub-
family of the Cerithiidae. Furthermore, its similarity in ornament to the procer-
ithiid genus Cryptaulax Tate, 1869 is noticeable and the possibility that Ageria
could be derived from this cannot be dismissed.
In many respects, the generic characters resemble those found in members of the
Turritellidae, i.e. the turreted shell, the whorl outline and the possession of a thin
parietal lip with a smooth columella. However, the holostomous, sub-circular
aperture of the Turritellidae makes any further comparison rather pointless, while
the characteristic lack of axial ornament in that family together with the marked
sinuosity of its growth lines and outer lip and the deeply excavate columella provide
further distinguishing characters.
Two species that can now be referred to Ageria are Turnitella costata J. de C.
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 157
Sowerby, 1827, Upper Greensand, Albian, Blackdown, Devon; and Cerithiwm
binodosum Roemer in Frié (1894, text-fig. 87), Lower Senonian, Biezno Beds
(= Priesener Schichten) Brezno, Czechoslovakia.
Questionably referred species are Turritella angustata d’Orbigny, 1850 from the
Neocomian of France (see p. 158) and Cerithiwm hector d’Orb. (1850, p. 156) in
Guéranger, 1867 (pl. 14, fig. 2), Cenomanian from Le Mans, France.
The original figure of Turritella cingulatolineata J. Miller (1851, p. 33, figured as
cingulatocostata, pl. 6, fig. 15) from Aachen appears to be that of an Ageria, having a
characteristic prominent abapical cord. Yet, the ornament shown in the figure
provided by Holzapfel (1888, pl. 14, fig. 1) when he interpreted Miuller’s species as
belonging to the genus Mesostoma, is quite unlike that of Ageria. This figure shows
a median row of prominent tubercles, while its quadrangular aperture hints at the
presence of a strong abapical cord.
Ageria gaultina sp. nov.
(Pl. 8, figs 1-5)
DiaGnosis. Narrow, tutriculate shell, ornamented by orthocline axial ribs which
do not persist on later whorls; its strong abapical cord is carinate.
HoLotyPeE. BM(NH) GG 6473 Albian, Gault, Folkestone, Kent. J. S. Gardner
Collection.
OTHER MATERIAL. BM(NH) G 4362 (26 specimens), G 11318, G 11544, G 20741-
20742, GG 20857-20862, 38085 (2), 48136 (3), Albian, Gault, Folkestone. G 71026,
Gault, East Wear Bay, nr. Folkestone, Kent and G 49804, Albian, Osmington,
Dorset.
Dr H. G. Owen of the Dept. of Palaeontology, BM(NH) considers that the matrix
of GG 20742 is consistent with that of Bed V of the Lower Gault at Folkestone,
in the Lautus-Nitidus Subzone.
DEscrIPTION. Small to medium-sized, turriculate shell, with an acute spire
composed of moderately high whorls (height equal to ? of width). Flush, linear
suture. Whorl outline slightly convex but briefly becoming concave before an
abapical peripheral carina.
Whorls ornamented by nine orthocline axial ribs, that are crossed by numerous
beaded spiral cords (22 from adapical suture to the peripheral carina in the holotype),
with the interspaces separating the ribs equal to twice the width of a rib. These
axial ribs are successively displaced from whorl to whorl and this produces a counter
spiral pattern in relation to the direction of coiling (see pl. 8).
Such ornament occurs in the earliest whorls, but initially the orthocline ribs are
much stronger than the simple spiral cords and therefore, in cross section, the whorls
are angular. The basal beaded spiral is the first to become differentiated with other
cords soon developing and varying in strength and composition. A pattern of five
strongly beaded primary cords witha varying number of more finely beaded secondary
cords occurring in their interspaces is formed. In some cords the beads are rounded,
in others they become elongated in the direction of the growth lines, while the ‘beads’
158 BRITISH CRETACEOUS GASTROPODS
of some secondary cords are the result of the growth lines simply dissecting the cord.
As the ornament of the spirals strengthens, the ribs tend to diminish and are almost
obsolete on later whorls, where they are only indicated by the presence of strong
tubercles on the primary cords. The opisthocline growth lines influence the spiral
cords, but do not affect the direction of the axial ribs.
The base is delimited by the prominent basal spiral cord and this is followed
inwards by a second, less prominent, primary cord, the base then slopes uniformly to
the columella. The remainder of the surface is ornamented by alternating granulose
spiral cords and smooth spiral threads. After crossing the basal carina, the growth
lines swing sharply back towards the columella.
Unfortunately, a completely preserved aperture is not available, but it is probably
somewhat quadrangular and axially elongate. There are some indications of a
short anterior canal but this may have resulted from crushing of the shell. The outer
lip is virtually straight in cross section, but the growth lines show a slight median
sinus. The inner lip is smooth, partly callused and rather narrow. The columella
is very slightly curved, but this could also be a result of crushing.
Discussion. There is a little variation within the ornament of this species. A
specimen of 5 whorls, found at Osmington, Dorset, BM(NH) G 49804 (see pl. 8, fig. 5)
has only thirteen spiral cords between the adapical suture and the basal carina and
not one of these cords is beaded, yet it does show a typical sequence of primary cords.
Among ornament variations shown by material collected from the Gault of Folke-
stone, Kent are more equidistant primary cords; a less prominent peripheral beaded
carina; broader (i.e. thicker) spiral cords ornamenting the base and a considerable
range of differences in the strength and dominance of both spiral and axial elements
on the early whorls (see pl. 8, figs 2, 3, 4 & 6, BM(NH) GG 20741-20742, GG 20857,
GG 20862).
Agena gaultina is easily distinguished from A. costata (J. de C. Sowerby), described
below, by its orthocline axial ribs and the fact that these ribs do not persist on to
later whorls. Its more carinate abapical cord also serves to identify the species.
The specimen figured as Cerithiwm binodosum Roemer, by Fri€ (1894, text-fig. 87),
from the Lower Senonian, Biezno Beds (Priesener Schichten) of Selten, nr. Louny,
N. W. Bohemia, Czechoslovakia, is not like any other specimen figured as C.
binodosum Roemer, 1841 and undoubtedly belongs to the genus Agevia. It possesses
the distinctive, strongly ornamented abapical spiral cord of that genus and has very
similar whorl and apertural shapes. The cancellate pattern formed by its axial ribs
and spiral cords separates it from other species of Agerza.
Cryptaulax angustatum (d’Orbigny), Peron (1900 : 93, pl. 1, fig. 14) from the
Neocomian of France, is probably another species of Agevia. The ornament, whorl
shape, axially elongated quadrangular aperture, and smooth, slightly curved
columella, are features in common with other members of that genus. Yet, its
aligned (initially orthocline) prosocline axial ribs are also strongly reminiscent of the
genus Cryptaulax Tate, 1869. The apparent lack of a strong abapical cord adds to
the uncertainty, while its smooth spiral cords would readily distinguish it from the
recognized species of Ageria. All of these features are discernible in the figures
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 159
provided by both d’Orbigny (1842, pl. 151, figs 4 & 6) and Peron, and in a British
Neocomian specimen BM(NH) GG 20740, from the Claxby Beds, Nettleton, Lincs.
(see pl. 8, fig. 13).
Ageria costata (J. de C. Sowerby)
(Pl. 8, figs 7-12)
1827 Turritella costata J. de C. Sowerby: 126, pl. 565, fig. 4.
1900 Turritella costaia Sow., Jukes-Browne: 464.
Diacnosis. Acicular shell with persistent prosocline axial ribs and a shelf-like
abapical cord.
Hototyre. BM(NH) 43674, Albian, Upper Greensand, Blackdown, Devon,
Sowerby Collection.
OTHER MATERIAL. BM(NH) 34844 (2 specimens), G 20830-20832, G 71382,
G 16142 (2 specimens) from Upper Greensand, Blackdown, Devon; G 71090-71095,
from Bed 12 Peak Hill, nr. Sidmouth, Devon, H. F. Metcalfe Collection.
DESCRIPTION. A small, slender, somewhat subulate shell with convex whorls that
appear to be twice their height in width. Flush, undulating suture occurs in a very
slight groove beneath the abapical astragal. The whorl outline becomes concave
just before this astragal and causes a small constriction in its shape. The growth
lines are opisthocline and show a definite median sinus.
The shell is characteristically ornamented by a consistent pattern of axial ribs and
spiral cords. Initially, the eight, or ten ribs are orthocline and dominate the
ornament. However, the ribs soon become prosocline and terminate above the
basal astragal as described by Sowerby. The spiral ornament consists of four to
five beaded to tuberculate primary cords with both weaker beaded cords and smooth
threads between these. The peripheral carina, or astragal, is shelf-like and dissected
by numerous growth lines. The flattish base is ornamented by rather broader spiral
cords than those of the whorl side. A strong cord, just in from the carina, forms the
main feature.
The aperture (see pl. 8, fig. 9) is quadrangular and elongated axially, with a notch
in the outer lip just above the astragal. The parietal lip is quite thin and the
columella is not callused. The base of the aperture is flat and an anterior canal is
not apparent.
Discussion. Although this species is very similar to Agevia gaultina sp. nov. in
_ ornament, there are a number of differences between them. The orientation of its
axial ribs is markedly prosocline as opposed to the orthocline ribs of gaultina. A
further contrast is that in costata the ribs persist and occur on later whorls. The
shell also appears to be more needle-like in shape, even though its whorl diameter:
whorl height ratio seems lower than that of A. gaultina. The whorls of costata are
more convex, while the abapical astragal that delimits the base of the two species,
is shelf-like in costata and carinate in gaultina.
160 BRITISH CRETACEOUS GASTROPODS
Family CERITHIOPSIDAE
Genus CERITHIELLA Verrill, 1882
[Lovenella Sars, 1878, non Hincks, 1869; Newtonia Cossmann, 1892, non Schlegel, 1866;
Newtoniella Cossmann, 1893; Cevithiolinum Locard, 1903].
Type by original designation, Cerithium metula, Loven, 1846, Recent.
REMARKS. Wrigley (1940 : 13) considered that Cerithiella has a moderately deep
sunken suture and that its subordinate axial riblets cross the spiral carinae frequently
producing nodes at the intersections. In his view, the related genus Sela has an
inconspicuous suture, is composed of essentially flat-sided whorls and its fine axial
threads are only visible in the interspaces between the spirals. However, in
instances where species have ornamental features similar to those of Seda, the
truncate concave columella and short twisted anterior canal of Cerithiella is easily
distinguishable from the virtually straight columella occurring in Seda. The
aperture of Cervithiella has been described as sub-quadrangular as opposed to the
sub-ovate shape found in Sez/a.
The Cretaceous species mentioned in the discussion below, were originally assigned
to the genus by Sohl (1964 : 364).
Cerithiella (Cerithiella) devonica sp. nov.
(Pl. 6, fig. 9)
MATERIAL. Unique specimen in the Sedgwick Museum, Cambridge, B 44633 from
the Upper Albian, Blackdown, Devon.
Diacnosis. Acuminate cerithiopsid with cancellate ornament consisting of three
equally-spaced tuberculate spiral ribbons and numerous fine axial threads and
possessing a short twisted anterior canal.
Dimensions. Height = 11 mm; Maximum diameter 3 mm.
DESCRIPTION. Small slender shell with turriculate spire. Whorls almost flat-
sided and with a very slight adapical ramp. Linear suture in moderately-deep
asymmetric channel formed by adjacent whorls. Ornament consists of three
equally-spaced spiral ribbons, the adapical ribbon being very slightly weaker. Very
faint spiral threads occur on the short ramp below the adapical suture. The concave
interspaces are equal to the width of the spirals. The fine axial threads are slightly
opisthocline and produce angular tubercles where they cross the spiral ribbons.
Ornament of the thirteen preserved whorls is consistent. The basal ornament,
growth lines and protoconch are not adequately preserved for description. The
aperture is incomplete but there are indications of a short twisted anterior canal.
Discussion. The characters of ornament, whorl shape, suture and aperture
appear to be sufficient to indicate that this specimen belongs to the genus Cerithiella.
Its cancellate ornament of three spirals and numerous axial threads is very close to
that of the North American Upper Cretaceous species C. nodolivatum (Wade), 1926.
On comparison with the figures given by Sohl (1960; pl. 9, figs 15-16, 24) only a
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 161
slightly longer anterior canal in nodolivatwm and the presence of the fine spirals on
the short adapical ramp of devonica could be used as distinguishing features. Yet,
these small differences when considered together with the widely separate geo-
graphical and stratigraphical occurrence of the two forms would seem to support the
belief that they are distinct species. Unfortunately, the basal ornament of the
British specimen is not preserved and consequently, it is not possible to verify the
distinctive growth line pattern on the base of the shell, described by Sohl (1964 : 365)
for the American cerithiopsids.
C. semirugatum (Wade) 1926 and Cerithiella sp. nov. Sohl, 1964 are easily dis-
tinguished from C. devonica by their ornament of four spiral ribbons. Similarly, the
ornament of four variable cords in Cerithium bicostatum Kaunhowen (1808 : pl. 6,
fig. 17) from the Maastrichtian of Belgium, separates that species from devonica.
Kaunhowen also mentions a short, but straight canal, in his description.
Fic. 2. 1, Cerithiella devonica sp. nov., Cretaceous, Upper Greensand, Albian, Blackdown,
Devon. x4. Cerithiella metula (Lovén). Type species. Recent, North Sea. 2:6.
3, Cerithiella nodolivatum Wade, Cretaceous, Maastrichtian, Ripley Formation.
Tennessee, U.S.A. x5. 4, Cerithiella semivugatum Wade, Cretaceous, Maastr., Ripley
Formation, Tennessee. U.S.A. 4:6. 5, Cerithiella cloacina Wrigley, Eocene, London
Clay, Whitton, Middx., England. x5. 6, Cerithiella atherfieldensis sp. nov., Cretaceous,
Lower Greensand, Aptian, Atherfield, Isle of Wight. x2.
162 BRITISH CRETACEOUS GASTROPODS
The more turreted character of the Lower Aptian, Cerithiella atherfieldensis sp. nov.
described below, together with the variation in strength of its spiral cords, its sub-
carinate whorls and limited axial ornament provide distinguishing characters. In
some Tertiary examples of the genus, C. cloacina Wrigley and C. praelonga (Deshayes)
the spiral cords are smooth. [Text-fig. 2]
Cerithiella (Cerithiella) atherfieldensis sp. nov.
(Pl. 6, fig. ro)
HoLotyPe. Sedgwick Museum, Cambridge B 27334 from the Lower Aptian,
Forbesi Zone, Crackers, Lower Greensand, Atherfield, Isle of Wight.
DiaGnosis. Narrow, turreted shell, ornamented by five spiral cords of varying
strength (the fourth producing a slight carination before the grooved suture) and
limited axial threads.
DESCRIPTION. Small, acutely spiral turreted shell of slightly convex, sub-
carinate whorls. Distinct suture in narrow groove. Protoconch not preserved.
Aperture sub-ovate with a strongly twisted anterior canal; short truncated columella.
Shell ornamented by five, more or less equidistant primary spiral cords and fine
axial threads. A cord occurs on either side of the suture with three stronger ones in
the centre of the whorl. The more abapical of these is the strongest and almost
forms a carina. The interspaces are much wider than the width of the cords and a
fine secondary spiral cord occurs in each of the two central interspaces. On the
base, a prominent interspace separates numerous fine spiral cords from the most
abapical primary cord. The fine axial threads are slightly opisthocyrt on the whorl
side and are not visible on the base. These axial threads are more prominent on the
earlier whorls but always weaker than the spiral cords.
Discussion. In this species, the small spiral angle, the limited axial ornament,
the somewhat turreted appearance, the variation in strength of its spiral cords, and
the abapical carination of the whorls, all serve to distinguish C. atherfieldensis from
the other Cretaceous species mentioned in the discussion of C. devonica. It is also
the earliest recorded species of this genus.
Genus SEILA Adams, 1861
[Cinctella Monterosato, 1884]
Type by subsequent designation, Dall (1889 : 250), Tviphoris dextroversus Adams
& Reeve, 1848, Recent.
Diacnosis. (After Sohl, 1960 : 84 and 1964 : 364). Moderately small to very
small, slender, turriculate, thin shell. Smooth, swollen, paucispiral protoconch.
Whorls flat-sided and ornamented by prominent, smooth spiral, ribbon-like cords
with numerous fine growth-line threads in their interspaces. Indistinct suture.
Last whorl moderately high and with a carina at basal periphery. Aperture
subovate, short anterior canal with a swelling on base of almost straight columellar
lip. Outer lip thin.
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 163
Seila iglali sp. nov.
(Bl 6) fig 12)
MATERIAL. Unique specimen, BM(NH) G 16084, from the Upper Greensand,
Albian, Combe Raleigh Hill, nr. Honiton, Devon, W. Vicary Collection.
DiaGnosis. Conical shell, large for genus, ornamented by four raised, flat-topped
spiral ribbons on whorl side and numerous smooth spiral cords on base, with trans-
verse threads occurring in the interspaces.
DESCRIPTION. Small, conical shell (Height: 13:5mm; Maximum diameter:
4:8 mm) with a flat-sided whorl outline. Whorls ornamented by raised, flat-topped
spiral cords (i.e. ‘ribbons’) with fine transverse threads in the concave interspaces
between these. Distinct linear suture occurs in a symmetrical groove between
adjacent whorls.
Early whorls appear to be smooth and rather convex (although this may be the
result of poor preservation). Traces of axial ornament can be seen adapically on the
fourth whorl, while the fifth shows faintly the typical ornament of subsequent
whorls. This consists of four prominent flat spiral ribbons that are approximately
equal in width but vary in their distance apart (see Text-fig. 5). The fourth ribbon
is slightly narrower and forms a basal carina. The concave interspaces are crossed
by regularly-spaced, easily visible, axial threads. Interspaces roughly three times
the width of a thread. Axial threads essentially orthocline, but towards the base,
i.e. between the third and fourth spiral ribbons, and particularly on later whorls,
they tend to become opisthocline. On close examination, it is possible to see that
the axial threads cross over the spiral ribbons.
Within the basal carina, and after a further concave interspace, the flat surface of
the base is ornamented by numerous (15-16) identical smooth spiral cords that
continue on to the columella and are separated by intervals equal to their width.
Growth lines are not visible on the base.
Aperture small and sub-ovate ; columella broad and equal to one third of the whorl
diameter, and is twisted anteriorly. Anterior canal short and narrow.
Discussion. The general flat whorl outline, together with its ornament of raised,
ribbon-like spirals, leads one to consider this species as a member of the genus Sezla.
However, its size, which is greater than that of most species assigned to this genus,
and its distinct linear suture, cause some uncertainty. In addition to these doubtful
features, its conical shell shape is also rather unusual and quite unlike the slender,
sub-cylindrical form of the type species Sezla dextroversa (Adams & Reeve). Ina
number of Recent species of the genus, the base of the anterior canal forms a slit,
which extends for one third of the whorl’s circumference. This feature is certainly
lacking in S. iglali and while this could be due to poor preservation, it also appears to
be lacking in North American Upper Cretaceous species which suggests that it is not
present in these earlier forms. Obviously, in order to confirm that this Albian fossil
is correctly assigned to the genus Seila, it is necessary to obtain a well-preserved
protoconch.
Sohl (1960 : 84) refers the North American, Upper Cretaceous species Cerithiopsis
164 BRITISH CRETACEOUS GASTROPODS
meeki (Wade, 1926) and Cerithiopsis quadrilirata (Wade, 1926) found in the Ripley
Formation (Maestrichtian), to the genus Sez/a, on the basis of their ornament and
protoconch features. Since the columella fold found in both of these species is much
stronger than in typical Sezla, and, in fact, more reminiscent of other Cerithiopsidae,
he qualifies this decision. Yet, as none of these genera possess ornament similar to
that of meekt and quadrilivata, he refers the two species to Sezla.
Seila iglali is very close to S. meeki (Wade) and apart from their considerable
difference in size, the only feature that will separate the two species is the presence of a
fourth spiral ribbon in zglali. In both Sela quadrilirata (Wade) and Sela sp. Sohl,
(1960 : 85, pl. 9, fig. 23), some of the spiral ribbons are very close together. Among
other distinguishing features, the axial threads of S. quadvilivata are finer and more
numerous and also described as being prosocline; while in Sez/a sp. no. axial threads
are present. Sohl (1964 : 364) has also recorded S. meeki from a slightly older
horizon in the Upper Cretaceous [Coffee Sands of Mississippi (Campanian) | and noted
that its spiral ribbons were much thinner and consequently farther apart. The
limited amount of material available, prevented him from considering the possible
significance in this.
Among Eocene species, Seila mundula (Deshayes, 1865 : 222) is the closest to
iglali, but can be separated by its convex base and very much finer axial threads.
In contrast, to the Cretaceous specimens referred to ?Cerithiella, there is no indica-
tion of secondary spirals developing in the interspaces of the primary ribbons in
S. iglali.
If the determination of this specimen is correct, it will be the earliest record of the
genus.
Fic. 3. 1, Seta dextroversa (Adams & Reeve), type species of genus. Recent. China
Seas. xg. 2, Seila iglali sp. nov., Cretaceous, Albian, Devon, England. x2-6. 3,
Seila mundula (Deshayes), Eocene, London Clay. Highgate. 3-3. 4, Seila meeki
(Wade), Cretaceous, Maastrichtian. U.S.A. x5 approx. 5, Sela quadrilivata (Wade)
Cretaceous, Maastrichtian, U.S.A. 3-3 approx.
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 165
Genus ORTHOCHETUS Cossmann, 1889
TYPE SPECIES. By original designation, Cerithium leufroyi Michelin in Deshayes,
(1833 : 380), Lutetian, France.
DiaGnosis. (After Wenz, 1940 : 779). Medium-sized, slender, turriculate regu-
larly conical shell, composed of numerous low whorls that have an abapical keel.
Whorls ornamented by spiral cords and axial riblets, frequently producing a reticu-
late network. Last whorl approximately one third of shell height. Aperture sub-
quadrangular, with a strong, moderately long, truncated canal; thin outer wall,
slightly sinuous adapically. Columella straight, with base a little twisted; may be
smooth, or with several oblique folds. Growth lines opisthocline to opisthocyrt, but
swing back towards the columella on the base.
Orthochetus hantoniensis sp. nov.
(EIS Osdigs 2)
MATERIAL. BM(NH) G 70130, Aptian, Atherfield, Isle of Wight.
Diacnosis. Slender, turreted cerithiopsid with an abapical carina and strongly
developed axial ribs (fewer in number than normal), which influence five of the six
spiral cords to produce a frilled appearance. Columella straight, but uncharacter-
istically short and apparently without plications.
DESCRIPTION. Small, slender, turreted shell with fairly low whorls. The suture
is subcarinate and linear, but axial ribs may give it an undulate appearance. The
protoconch is not completely preserved, but early ornament seems to be identical to
that of later whorls. This ornament consists of strong, straight axial ribs (15-16 per
whorl) that are separated by interspaces equal to their width; and six spiral cords of
varying strength. A slight adapical ramp occurs immediately beneath the suture
and this is bounded by a strong spiral cord. At equal distances below this follows a
very weak cord, a moderately strong cord, then at the abapical carina another strong
cord, and, finally, another weak cord.
All of these five cords are influenced by the axial ribs, the apparent thickening at
their intersections producing a frilled, or tuberculate appearance to the shell. A
sixth spiral cord close to the abapical suture, limits the base and is unaffected by the
axial ribs. The axial ribs diminish soon after the abapical carina.
Numerous faint opisthocline growth lines occur on the whorl side and on the base
these swing back towards the columella. Columella moderately short and straight
with faint oblique ornament along its length. Small aperture subquadrangular,
showing a well-developed, twisted and truncated anterior canal. Thin sinuous outer
lip.
Discussion. The distinctive ornament and slender turreted spire of this species,
together with its general apertural features, are sufficient to refer it to the genus
Orthochetus. However, the columella is somewhat shorter than is usual, while its
twisted anterior canal is less inclined; in fact, such features are reminiscent of the
genus Cervithiella. The strongly developed axial ornament and its consequent
166 BRITISH CRETACEOUS GASTROPODS
influence on the shell’s appearance are adequate diagnostic features to separate this
species from others that have been described. The lack of plications on the
columella might be construed as uncharacteristic of Orthochetus, yet, Wrigley
(1940 : 1r & 12) mentioned a number of instances in Eocene species where this
feature is obscured by a reflected columella border.
Its diagnostic features distinguish O. hantoniensis from the two Maastrichtian
species mentioned by Cossmann (1906 : 97) Cerithium tectiforme Binkhorst (1861:
24 pl. 1 figs 3a-c) from Limbourg and O. mapeulensis Douvillé (1904 : 303 pl. 41
fig. 12) from Kouh Mapeul, Persia. The ornament of both these Maastrichtian
species is more reticulate, due to the presence of smaller and more numerous axial
ribs. Although both O. hantoniensis and O. tectiformis have three primary spiral
cords, the presence of secondary spirals in the former and of tubercles in tne latter
assist in separating the two species.
Douvillé (1904 : 303) referred to Ovthochetus as an ancient group and quoted the
occurrence in the Chalk of Cerithium cribriforme Zekeli (1852 : pl. 20, figs 2 & 2’) as
evidence. In many respects, this turreted species is very like O. hantoniensis,
Fic. 4. 1, Ovthochetus tectifoymis ( Binkhorst), Cretaceous, (Maastrichtian), Limbourg
Netherlands. x1-3. 2, Ovthochetus hantoniensis sp. nov., Cretaceous (Aptian).
Atherfield, Isle of Wight. 3:3. 3, Orthochetus leufroyi; (Michelin in Deshayes 1833).
Eocene. Chaumont, France. Type species. x0-6. 4, Orthochetus charlesworthi
(Prestwich), Eocene, London Clay. Sheppey, Kent. 0-6. 5, Orthochetus mapeulensis
Douville. Cretaceous, (Maastrichtian). Louristan, S. Persia. 1-3. 6, Ovthochetus
elongatus Wrigley. Eocene, London Clay, Sheppey. xo-6.
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 167
having strong axial ribs and three smooth spiral cords undulating over them. Yet
its aperture is not shown, and there are no indications of secondary spirals in the
figure which does show that C. cribriforme is rather awl-shaped, i.e. subulate. The
existence of O. hantoniensis would appear to substantiate Douville’s claim that the
genus Orthochetus arose in the Cretaceous, even if C. cribriforme Zekeli should prove
to be mis-identified.
Several of the larger Eocene species show that the anterior canal and the columella
are not necessarily identical to, or as long as that of the type species O. lewfroyi.
Comparison of O. charleswortli (Prestwich) and O. elongatus Wrigley illustrates that
there is considerable variation in spiral angle and whorl diameter. Wrigley
(1940 : 12) also showed that within a species there is a wide variation in the strength
and position of its ornament components, e.g. see his figures of O. elongatus (p. 17,
figs 13-15).
Incertae sedis
? ‘Orthochetus’ helmyi sp. nov.
(Pl. 6, figs 13 & 14)
MatTerrIAL. BM(NH) G 10500 and G 71988 from the Gault, Albian of Folkestone,
Kent.
Diacnosis. Turreted, conical cerithiopsid, ornamented by four prominent flat-
topped ribbon-like spiral cords and numerous fine opisthocyrt axial threads.
Description. Medium-sized, slightly turreted, conical shell. Distinct linear
suture lies in a furrow between the abapical and adapical primary spiral cords of
adjacent whorls. Protoconch not preserved.
Whorl side ornamented by four strong, prominent, flat, ribbon-like spiral cords,
with a fifth weaker primary cord occurring at the abapical suture and delimiting the
base. The two lower ribbons, i.e. the third and fourth abapically, are stronger and
wider. Secondary spiral threads occur in the unequal interspaces between the
primary ribbons, but can only be seen easily on the lower halves of later whorls.
Fine axial growth threads occur in the interspaces and also continue over the spiral
ribbons. These opisthocyrt threads show a broad shallow sinus, which has its apex
at mid-whorl; each thread is separated from its neighbour by a space equal to double
the width of a thread.
The base is flat to feebly convex and within the delimiting fifth primary cord, it is
ornamented by numerous thin secondary spirals. The opisthocyrt growth lines,
after crossing on to the base, tend to straighten and then swing back towards the
columella. The aperture is sub-quadrangular and has a flat basal lip. The short
columella is smooth.
Discussion. It is difficult to assign this species to one particular genus of the
Cerithiopsidae without having a specimen that shows the important details of its
aperture and protoconch, for its ornament and whorl shape have characteristics
similar to those of Cerithiella, Seila and Orthochetus. Reference to the literature
and to collections of Tertiary representatives of these genera does not provide any
E
168 BRITISH CRETACEOUS GASTROPODS
useful guide, since it is apparent that there is some confusion in distinguishing the
genera and in assigning species to them. The criteria that Wrigley (1940 : 13) used,
have been referred to under Cerithiella (page 160), but these are often inadequate, e.g.
the inconspicuous suture that is considered a diagnostic feature of the genus Sezla.
Undoubtedly, a reappraisal of the criteria used to distinguish the genera of the
Cerithiopsidae is necessary and the review of species assignations suggested by Sohl
could be undertaken at the same time.
Some of the features present in ‘helmyz’, i.e. the distinct linear suture and the
crossing of the spirals by the axial threads, indicate possible affinities with Cerithiella.
The moderate size of the specimen and the fact that its whorls could hardly be
described as flat-sided, indicate that it should not be assigned to Sezla, but could
easily be recognized as a possible member of Ovthochetus. The presence of promi-
nent, ribbon-like spiral cords and the fine axial opisthocyrt threads and the basal
features of the whorl, together with the indications of a short straight columella are
reminiscent of several species of Seda, including the type species S. dextroversa.
However, these features could equally be those of Orvthochetus and therefore after
recognizing that the linear suture is very like that of Eocene species of this genus,
helmyi is provisionally placed in Orthochetus. In the majority of described species of
this genus, and particularly in the earliest member O. hantoniensis, the axial orna-
ment is quite prominent and contributes to the typical reticulate ornament pattern.
The size, ribbon-like spirals, super-imposed axial ribs and turreted whorl shape of
helmyi are considered to be the most important indicators of its probable relationship
with Orthochetus.
Similar difficulties prevent the assignation of a unique specimen, preserved as an
external mould in an ironstone concretion from the Sandgate Beds, found at Parham
Park, Sussex (BM(NH) 9145, G. A. Mantell Collection, (see pl. 6, fig. 15).
The ornament of this medium-sized, turreted gastropod consists of three primary
and three secondary spiral cords. It has a distinct linear, impressed suture and
opisthocyrt growth lines between the sutures. The whorls are not flat-sided, (in
fact they are concavo-convex) and the spiral cords are carinate, or rounded, not
‘ribbon-like’. Axial ornament, apart from the collabral threads, is lacking. A
slight distortion in the smooth opisthocyrt curve wherever the growth line crosses a
spiral cord produces a wavy effect. Unfortunately, neither the abapical portion of
the body whorl, nor the aperture are preserved.
VI. CONCLUSIONS
The need for exhaustive re-examination and re-evaluation of existing collections
and type material has been repeatedly mentioned in the discussion above and can
only be emphasized again here. Before the taxonomy of the Cerithiacea can be
revised, it is necessary to improve upon the incomplete information currently
available by collecting additional material. Admittedly the task of revision is
complicated by the limitations of the characters that are used to distinguish fossil
gastropods, while their poor preservation adds to this difficulty. Yet, the careful
collecting advocated by present-day palaeontologists should enable these problems
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 169
to be surmounted. The important morphological and ornamental variation of
Cretaceous gastropods can be satisfactorily assessed, once accurate information on
their stratigraphical occurrence is produced. These conclusions would then con-
tribute to and facilitate our understanding of their phylogenetic and palaeoecological
relationships.
The family Cerithiidae became conspicuous during the Cretaceous and during that
period experimented with the form of the aperture and anterior canal. Fossil
evidence is sparse and consequently, our knowledge of the forms which are thought
to be transitional between either the Procerithiidae, or the more cerithiid-like
Eustomidae, and true cerithiids is very limited. A similar situation occurs with
their descendants. In order to achieve a re-assessment of these families, a full
understanding of the derivation of the anterior canal is vital. Critical genera such
as Ageria, Cimolithium and Uchauxia require further investigation, and determina-
tion of the phylogenetic position of Metacerithium, Exechocirsus and Campanile is
equally important. Cox (1965 : 158) briefly mentioned the possibility that difficult
groups might prove to be polyphyletic (he referred to this while discussing the
origin of Exelissa, a genus of the Procerithiidae), but did not proceed to discuss the
taxonomic implications of such a suggestion. It would provide an easy explanation
of the morphological diversity apparent in ‘Rhabdocolpus’ but careful consideration
is necessary before such a theory can be followed.
The variation in ornament shown by several of the Cretaceous gastropods dis-
cussed here, is not thoroughly understood. In some genera, e.g. Metacerithium,
ornament is said to be basically stable, while in others, e.g. Bathraspira it would
appear to alter very quickly. Further, the significance of sculptural variation
within a species, such as that shown by Exechocirsus saundersi (Woods), is also not
fully appreciated. It has been suggested that the smaller Yorkshire specimens of
this species might be explained as an example of ecotypic variation. The results of
the detailed study recommended above should assist in providing an answer to
these questions.
Among more detailed points for consideration is the verification and explanation
of the occurrence of several species of the same genus at a particular horizon, e.g.
three species of ?Rhabdocolpus in the Crackers Bed of the Lower Aptian in the Isle
of Wight. There are also certain elements of doubt concerning the generic identi-
fication of several species that are considerably larger than the normal size of the
genus to which they are assigned. In classifying Cretaceous gastropods ‘size’ is
frequently used as a guide, in a subconscious and perhaps, quite illogical manner.
If some disparity occurs, the correctness of the determination begins to be doubted.
This is certainly true for the Upper Cretaceous specimens of Nerineopsis described in
the paper and also for the Upper Albian species Sezla iglalz.
A better understanding of the relationship of Tertiary and Recent gastropods is to
a large extent dependent upon earlier forms. The origin and early evolution of many
Recent prosobranch families occurred during the Cretaceous period and the past
neglect and inadequate descriptions of gastropods of this age needs to be remedied.
It is hoped that this paper will provide some of the preliminary data necessary for
170 BRITISH CRETACEOUS GASTROPODS
a more detailed and sophisticated appraisal of Cretaceous gastropods, and also that it
will serve to pose a few of the questions needing investigation before such a study can
be undertaken.
VI. REFERENCES
Aspass, H. L. 1961. The evolutionary characters of certain turreted gastropod families and
their significance in ontogeny and phylogeny. A review. J. Geol. u. Avab Repub, Cairo,
5, 1, 1961 : 39-54.
1962. The English Cretaceous Turritellidae and Mathildidae (Gastropoda). Bull. Brit.
Mus. (Nat. Hist.), Geol., London, 7, 6, 1962 : 173-196, 5 pls.
1963. A monograph on the Egyptian Cretaceous gastropods. Monogr. geol. Surv. un.
Arab Repub., Cairo, Pal .Ser., 2, 1963 : xii, 146, 12 pls.
Apams, A. 1848. The zoology of the voyage of H.M.S. Samavang, under the command of Captain
Siv E. Belcher . . . 1843-1846. London, 1848-1849.
1861. On some new species of Eulima, Leiostvaca and Cevithiopsis from Japan. Amn.
Mag. nat. Hist., London, 3, 7, 1861 : 125-131.
ApaAMs, A. & REEVE, L. A. 1848. ‘Mollusca’ in Adams, A. 1848-1849.
Aiison, E. C. 1955. Middle Cretaceous gastropoda from Punta China, Baja California,
Mexico. J. Paleont., Tulsa, 29, 3, 1955 : 400-432, 5 pls.
ARKELL, W. J. 1941. The gastropods of the Purbeck Beds. Q. Jl. geol. Soc., London, 97, 1,
1941 : 79-128.
ASCHER, E. 1906. Die Gastropoden, Bivalven und Brachiopoden der Grodischter Schichten.
Beitr. Paléont. Geol. Ost-Ungans, Wien. 19, 2-3, 1906 : 135-172, 3 pls.
BAYLE, E. 1884. in FISCHER, P. 1880-1887.
Bennison, G. N. & Wricut, A. E. 1968. The geological history of the British Isles. Heffer,
Cambridge, 1968 : x, 406.
Binkuorst, J. T. van den B. 1861. Monographie des gastéropodes et des cephalopodes de la
craie supérieure du Limbourg. 4° Miller Bros. Maastricht, 1861 : vi, 44, 18 pls.
Brake, J. F. 1902. List of the types and figured specimens vecognized by C. D. Sherborn, F.G.S.,
in the collection of the Geological Society of London. Geol. Soc. Lond., 1902 : 100, XxXxii.
BRONGNIART, A. 1810. Sur les terrains qui paroissent avoir formés sous l’eau douce. Ann.
Mus. Hist. nat., Paris, 15, 1810 : 357-405. see under 1822, CUVIER, G.
Buvienigr, A. 1852. Statistique géologique, minévalogique, minévallurgique et paléontologique
du département de la Meuse. Atlas : 52 pp., 32 pls., Paris.
CasEy, R. 1961. The stratigraphical paleontology of the Lower Greensand. Paleontology,
London, 3, 4, 1961 : 487-621, 8 pls.
Corticnon, M. 1949. Recherches sur les faunes Albiennes de Madagascar. 1. L’Albien
d’Ambarimanga. Ayn. géol. Serv. Mines, Madagascar, Paris, 16, 1949 : 128, 22 pls.
CossmMann, M. 1889. Catalogue illustré des coquilles fossiles de l’Eocene des environs de Paris.
V & Suppl. Annis. Soc. v. malacol. Belg., Bruxelles, 24, 1889 : 3-385, 12 pls.
1892. Revue de paléontologie pour l’année 1891. Amn. géol. univ. Rev. géol. paléont.,
Paris, 8, 1892 : 718-721.
1893. Revision sommaire de la faune du terrain Oligocene marin aux environs d’Etampes
(suite). J. conchyliol., Paris, 41, (Ser. 3, 33), 1893 : 203-363, pl. Io.
1894. Revue de paléontologie pour l’année 1892. Amn. géol. univ. Rev. géol. paléont.,
Paris, 9, 1894 : 763-768.
1895-1925 Essais de Paléoconchologie comparée. Paris. 13 vols. [Volume 7, 1906.
Includes Cerithiacea].
1902. Rectifications de nomenclature. Revue crit. Paléozool., Paris, 6, 3, 1902 : 160-162.
1906. See Essais de paléoconchologie comparée. Vol. 7 : 261, 15 pls.
—— 1913. Contribution a la paléontologie frangaise des terrains Jurassique. III.
Cerithiacea et Loxenematacea. Mem. Soc. géol. Fr., Paléont., Paris, 46, (19), 3-4 : 88,
4 pls; 46, (20), 3-4 : 89-264, 7 pls.
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 171
CotTEAu, G. 1854. Paléontologie de l)Yonne. Prodrome des mollusques. Gasteropodes.
Bull. Soc. Sci. hist. nat. Yonne, Auxerre, 8, 1854 : 201-231.
Cox, L. R. 1930. The fossil fauna of the Samana Range and some neighbouring areas.
Part IV. Lower Albian Gastropoda and Lamellibranchia. Mem. geol. Surv. India,
Calcutta, NS 15, 1930 : 39-49, 1 pl.
—— 1960. The British Cretaceous Pleurotomariidae. Bull. Brit. Mus. nat. Hist., (Geol.),
London, 4, 8, 1960 : 385-423, 17 pls.
1965. Jurassic bivalvia and gastropoda from Tanganyika and Kenya. Bull. Brit. (Nat.
Hist.), London, Suppl. 1, 1965 : 208.
Cuvier, G. L.C. F. D. & Bronentart, A. 1822. Description géologique des environs de Paris.
4° Paris, 1822 : 428, 16 pls.
Datt W. H. 1889. Report on the Mollusca II. Gastropoda (Report on dredgings....
by U.S. Coast Survey Steamer Blake....). Bull. Mus. comp. Zool. Harvard, Cambridge,
Mass., 18 : 1-492.
Davis, A. G. 1923. Contributions to the geology and palaeontology of the Croydon regional
survey area. No. 1. The Lower Chalk, zone of Ammonites (Schloenbachia) varians.
Proc. Tvans. Croydon Nat. Hist. Soc., 9, 1923 : 125-136.
DELPEY, G. 1948. Gastéropodes mésozoiques de l’ouest de Madagascar. Ann. géol. Surv.
Mines, Madagascar, Paris, 15, 1948 : 25, 7 pls.
DESHAYES, G. P. 1824-1837. Description des coquilles fossiles des environs de Paris, Paris,
Vol. 2 : 1-178 (1824), 179-306 (1833), 307-434 (1834), 435-562 (1835), 563-690 (1836),
691-814 (1837).
1830-1832. Encyclopédie méthodique ou par ordre de matiéves histoive naturelles des Vers et
Mollusques. 3 vols., Paris, 1830-1832 : 1152.
1842. In Leymerie, A.
1864-1865. Description des animaux sans vertebves du Bassin de Paris. Paris, Vol. 3 : I~
200 (1864), 201-656 (1865).
Dovuvitte, H. 1904. Mission scientifique en Perse. J.de Morgan. Vol. 3, Etudes géologiques,
Partie IV, Paléontologie, Mollusques fossiles. 4° Paris, 1904 : 191-380. 25 pls.
EuprEs-DrEsLonccHamps, E. 1842a, Mémoire sur les cérites fossiles des terrains secondaire du
Calvados. Mém. Soc. linn. Normandie, Caen, 7, 1842 : 189-214.
1842b, Mémoire sur les melanies fsssiles des terrains secondaires du Calvados. Mém. Soc.
linn. Normandie, Caen, 7, 1842 : 215-232, pls 11-12.
FIscHER, P. 1880-1887. Manuel de conchyliologie et de paleontologie conchyliologique. 8°
F.Savy, Paris, 1880-1887 : xxv, 1369, 23 pls.
Fitton, W. H. 1836. Observations on some of the strata between the Chalk and the Oxford
Oolite in the southeast of England. Tvans. Geol Soc., London, 4, 2, 1836 : 103-389.
FLEMING, J. 1828. A history of British animals exhibiting the descriptive characters and
systematical avvangement of the geneva and species of . . . of the United Kingdom. 8°
Edinburgh and London, 1828 : xxiii, 565.
Forbes, C. 1960. Field meeting in the Cambridge district. Pvoc. geol. Ass., London, 71,
2 : 234-241.
ForBes, E. 1845. Catalogue of Lower Greensand fossils in the Museum of the Geological
Society with notices of species new to Britain. Q. J1 geol. Soc., London, 1, 1845 : 237-250,
345-355, 4 pls.
Fric, A. 1894. Studien im Gebiete der Béhmischen Kreideformation Palaeontologische
Untersuchungen der einzelnen Schichten v. Priesener Schichten. Aychiv. Naturw. Land.
Durchforsch., Bohm., Prag, 9, 1, 1894 : 1-134.
GeInitz, H. B. 1874. Das Elbthalgebirge in Sachsen. Pt. 2. Der mittlere und obere
Quader, V. Gasteropoden und Cephalopoden. Palaeontographica, Stuttgart, 20, abt. 2,
1874 : 161-198, pls 29-36.
GMELIN, J. F. 1788-1793. C.alLinné.. . ‘Systema Naturae .. .’,3 vols. 8° Lipsiae, 13th
edition, 1788-1793 (in 7 volumes).
172 BRITISH CRETACEOUS GASTROPODS
GoipFuss, G. A. 1844. Petvefacta Germaniae, Vol. 3, Folio, Dusseldorf, 1844 : 128, pls 166—
200.
Grey, J. E. 1847. A list of the genera of recent mollusca, their synonyms and types. Pyoc.
zool. Soc., London, 15, 1847 : 129-219.
GUERANGER, M. E. 1867. Album paléontologique du Département de la Sarthe veprésentant au
moyen de la photographie les fossiles veceuilles dans cette civconscription. Folio, Le Mans,
1867 : 20, 25 pls.
Haas, O. 1953. Mesozoic invertebrate faunas of Peru. Bull. Am. Mus. nat. Hist., New
York, 101, 1953 : xi, 328, 18 pls.
HOoLzapFEL, E. 1887-1888. Die Mollusken der Aachener Kreide. Palaeontographica,
Stuttgart, 34, 1887-1888 : 29-180, pls 4-20.
HUCKRIEDE, R. 1967. Molluskenfaunen mit limnischen und brackishen Elementen aus Jura,
Serpulit und Wealden N.W.-Deutschlands und ihre palaogeographische Bedeutung. Beth.
geol. Jb., Hannover, 67, 1967 : 263, 25 pls.
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. Inteyvnational code of zoological
nomenclature. Int. Trust. Zool. Nomen., London, 1961 : 176.
Ivanova, A. N. 1959. [Bivalves, gastropods and belemnites in the Jurassic and Cretaceous
deposits of Saratov on the Volga]. Tvudy VNIGRI, Leningrad, 137, 1959 : 267-460,
27 pls. [In Russian].
Jupp, J. W. 1868. On the Speeton Clay. Q. Jl geol. Soc., London, 24, 1868 : 218-250.
JuKEs-Browne, A. J. 1900. The Cretaceous vocks of Britain. Vol. 1. The Gault and Upper
Greensand of Britain. Mem. Geol. Surv. Gt. Britain, H.M.S.O., London : xiv, 499.
KAUNHOWEN, F. 1898. Die Gastropoden der Maestrichter Kreide. Paldont. Abh., Berlin, 8,
1898 : 132, 13 pls.
KENNEDY, W. J. 1969. The correlation of the Lower Chalk of South-East England. Proc.
Geol. Ass., London, 80, 4, 1969 : 459-560.
KIRKALDY, J. F. 1939. History of the Lower Cretaceous period in England. Pyvoc. Geol. Ass.,
London, 50, 3, 1939 : 379-417.
1963. The Wealden and marine Lower Cretaceous beds of England. Pyoc. Geol. Ass.,
London, 74, 2, 1963 : 127-146.
Ktun, O. 1947. Zur Stratigraphie und Tektonik der Gosauschichten. Sitzber. Akad. Wiss.,
Wien, 156, Abt. I, 1947 : 181-200.
Lamarck, J.B.P.A. de M. de. 1802-1806. Mémoive sur les fossiles des envivons de Paris,
comprenant la détevmination des espéces qui appartiennent aux animaux marins sans
vertébres, . . . Ann. Mus. Hist. Nat., Paris, 1802-1806 : 284.
LampLuGu, G.W. 1889. On the subdivisions of the Speeton Clay. Q. Jl geol. Soc., London, 45,
4, 1889 : 575-618.
LEYMERIE, A. 1842. Mémoire sur le terrain Crétacé du Département de |’Aube contenant des
considerations générales sur le terrain Néocomien. Mem. Soc. geol. Fr., Paris, 5, 1, 1842:
127, 18 pls.
LinNaEus, C. 1788-1793. Systema Naturae, sive regna tria naturae systematice proposita
per classes, ordines, genera and species. Editio decima tertia, aucta, reformata, cura J. F.
Gmelin, 3 Vols. [in 7] illust. 8° Lipsiae. 1788-1793.
Locarp, A. 1903. Les Cerithium et les Cerithiidae des mers d’Europe. Anmnls. Soc. Agric.
Lyon, 10, 1903 : 95-128.
Loriot, P. de. 1874-1875. Description des fossiles. In Loriot, P de and Perxtrat, E.
Monographie paléontologique et géologique des étages supérieurs de la formation jurassique
des environs de Boulogne-sur-Mer. Mem. Soc. Phys., Genéve, 23, 1874 : 261-407, I0 pls,
24, 1875 : 1-326, 16 pls.
Loven, S. L. 1846. Index Molluscorum litora Scandinaviae occidentalis habitantium.
Faunas prodromum. Ofvers. K. VetenskAkad. Forh., Stockholm, 3 1846 : 135-160, 183-204.
MIcHELIN, H. 1823. In DESHAYES, G. P. 1824-1837.
1838. Sur une argile dépendant du Gault, observée au Gaty Commune de Gérodot,
Departement de l’Aube. Mem. Soc. geol. Fr., Paris, 3, 5, 1838 : 97-103, 1 pl.
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 173
Moore, R. C. and others. 1968. Developments, trends, and outlooks in Palaeontology. J.
Paleont., Tulsa, 42, 6, 1968 : 1327-1377.
Murer, J. 1851. Monographie der Petrefacten der Aachener Kreideformation. Zweite
Abteil. 8° Bonn, 1851 : 88, 4 pls.
Munster, G. 1844. In Gotpruss,G. A. 1844. Vol. 3.
Nacao, T. 1934. Cretaceous mollusca from the Miyako District, Honshu, Japan.
(Lamellibranchiata and Gastropoda). J. Fac. Sci. Hokkaido Univ., Ser. IV. Geol. & Min.,
Sapporo, 2, 3, 1934 : 177-277, 10 pls.
Oprtnton 417, I.C.Z.N. 1956. Rejection for nomenclatorial purposes of volume 3 (Zoologie)
of the work by Lorenz Oken entitled “Okens Lehrbuch dey Naturgeschichte’ published in
1815-1816. Opin. & Decl. 1.C.Z.N., London, 14, 1, 1956 : 1-42.
Oxen, L. V. 1815-1816. Okens Lehrbuch der Naturgeschichte, Vol. 3 (Zoologie). Jena,
Abt. 1 : xxviii, 842, 40 pls; Abt. 2 : xvi, 1270.
Orpicny, A. D. 1842-1843. Paléontologie Francaise, terrains crétacés 2. Gasteropodes.
8° Paris : 456, pls 149-2306.
1850. Pyrodvome de paléontologie stratigvaphique universelle des animaux mollusques et
vayonnés. Vol. 2. 12° Paris : 428.
PéHELINCEV, V. 1927. [The Gault gastropoda from Sokolova Gora near Saratov]. Izv. geol.
Inst., Petvogrvad Bull. Com. Geol. Leningrad 45, 9, (49), 1926 : 991-999, 1 pl. [In Russian]
1927a. [The Jurassic and Lower Cretaceous fauna of the Crimea and the Caucasus.
Gastropoda of the Cretaceous sandstones in the neighbourhood of Balaclava]. Mém. Com.
Géol., Leningrad, 172, 1927 : 148-177, 3 pls. [English summary].
1953. [Gastropod fauna of the Upper Cretaceous strata in Transcaucasia and Central
Asia}. Izd. Akad. Nauk. SSSR, Moscow, Geol. Mus. A.P. Karpinskogo, Ser. Monograph,
1, 1953 : 391, 51 pls. [In Russian].
PCéHELINCEV, V. F. & Korospxov,I. A. 1960. In Or tov, Y. A. [Edit.], ‘Osnovoi Paleontologii,
Part 4—Gastropoda’. Izd Akad. Nauk Moscow, 1960 : 360, 28 pls.
Peron, M. 1899-1900. Etudes paléontologiques sur les terrains du Département de L’ Yonne.
Cephalopodes et gastropodes de l’étage Néocomien. Bull. Soc. Sci. hist. nat. Yonne
Auxerre, 53, 3, 1899-1900 : 67-219, 4 pls.
Pictet, F. J. & CampicHE, G. 1861-1864. Description des fossiles du terrain Crétacé des
environs de Sainte-Croix. Mater. Paléont. Suisse, Geneve, 3, 1861-1864 : 752, 55 pls.
Pictet, F. J. & RENEvIER, E. 1854-1858. Descriptions des fossiles du terrain Aptien de la
Perte-du-Rhéne et des environs de Sainte-Croix. Mater. Paléont. Suisse, Genéve, Ser. 1,
No. 1, 1854-1858 : 184, 23 pls.
Pictet, F. J. & Roux, W. 1847-1853. Description des mollusques fossiles qui se trouvent
dans les Grés Verts des environs de Genéve. Mém. Soc. phys. Hist. nat. Genéve, 11, 2 : 257—
412; 12 : 157-287; 13 : 73-173, 489-558, 51 pls.
Price, F.G.H. 1874. On the Lower Greensand and Gault of Folkestone. Pyoc. Geol. Ass.,
London, 4, 1874 : 135-150.
1875. On the Gault of Folkestone. Q. Jl. geol. Soc., London, 30, 1875 : 342-366.
1879. The Gault. 8°. London : viii, 81.
Quaas, A. 1902. Beitrage zur Kenntniss der Fauna der obersten Kreidebildung in der
libyschen Wiiste. II. Die Fauna der Overwegischichten und der Blatterthone in der
libyschen Wiiste. Palaeontogvaphica, Stuttgart, 30, Part 2, 1902 : 153-334, 33 pls.
Ravn, J. P. J. 1921. Kridtaflejringerne paa Bornholms Sydvestkyst og deres Fauna. III.
Senonet. Mus. Minér. Geol. Univ. Copenhague, Commun. Paléont., No. 17, 1921 : 52, 3 pls.
Rayner, D. 1967. The stvatigvaghy of the British Isles. Camb. Univ. Press, Cambridge,
1967 : X, 453.
Reuss, A. E. 1845. Die Versteinerungen dev Bohmischen Kreideformation. 4°, Stuttgart,
1845 : iv, 148, 51 pls.
Roemer, F. A. 1840-1841. Die Versteinerungen des Norddeutschen Kreidegeberges. 4°,
Hannover, 1840-1841 : 145, 16 pls.
174 BRITISH CRETACEOUS GASTROPODS
Roman, F. & MazErRANn, P. 1920. Monographie paléontologique de la faune du Turonien du
Bassin d’Uchaux et de ses dépendances. Avychiv. Mus. Hist. nat. Lyon, 12, 2, 1920 : 137,
11 pls.
Sars,G.O. 1878. Mollusca vegionis arcticae Norvegiae Oversight over dei Norges arktiske vegion
fovekommende Bloddyy. Christiana, 1878 : 466, 52 pls.
SCHUMACHER, H.C. F. 1817. Essai d’un nouveau systéme des habitations de vers testacés. 4°,
Copenhague. 1817 : iv, 287, 22 pls.
SepGwick, A. & Murcuison, R. I. 1833. A sketch of the structure of the Eastern Alps.
Trans. geol. Soc., London, 2nd Ser., 3, 2, 1833 : 301-420, pls 35-40.
SEELEY, H.G. 1864. Cuttings from a note-book on Chalk gastropods. The Geologist, London,
7, 1864 : 89-93, 1 pl.
SHARMAN, G. & NrEwtTon, E. T. 1896. Note on some Cretaceous fossils from the drift of
Moreseat, Aberdeen. Geol. Mag., London, Dec. IV, 3, 1896 : 247-254.
Sout, N. F. 1960. Archaeogastropoda, Mesogastropoda and stratigraphy of the Ripley, Owl
Creek, and Prairie Bluff Formations. Late Cretaceous gastropods in Tennessee and
Mississippi. Prof. Pap. U.S. Geol. Suvv., Washington 331-A, 1960 : 151, 18 pls.
1964. Gastropods from the Coffee Sand (Upper Cretaceous) of Mississippi. Pyvof. Pap.
U.S. Geol. Surv., Washington, 331-C, 1964 : 341-394, Pls 53-57.
1968. Post-Paleozoic Gastropoda. In R. C. Moore and Oruers 1968. J. Paleont.,
Tulsa, 42, 6, 1968 : 1363-1364.
1969. Cretaceous biotic provinces delineated by gastropods. [Abstract]. J. Paleont.,
Tulsa, 43, 3, 1969 : 899.
SoweERBy, J. & SowERBY, J.deC. 1812-1846. The Minerval Conchology of Great Britain. 8°,
London, 7 Vols, pls 1-383 (1812-1822) by J. Sowerby; pls 384-648 (1823-1846) by J. de C.
Sowerby.
SoweERBy, J.deC. 1833. Plates and Plate Descriptions in SepGwick, A. & Murcuison, R. I.
1833.
SowErRBy, J. de C. 1836. Descriptive notes respecting the shells figured in plates 11-23. In
Fitton, W. H. 1836. [see under Fitton, W. H. 1836].
STEPHENSON, L. W. 1941. The larger invertebrate fossils of the Navarro Group of Texas.
Bull. Univ. Texas. Publ. Bur. Econ. Geol. Austin, 4101, 1941 : 641, 95 pls.
1952. Larger invertebrate fossils of the Woodbine Formation (Cenomanian) of Texas.
Prof. Pap. U.S. Geol. Surv., Washington, 242, 1952 : 226, pls 8-59.
Stoticzka, F. A. 1867-1868. The Cretaceous fauna of Southern India. 2. The Gastropoda
of the Cretaceous rocks of Southern India. Mem. geol. Surv. India, Calcutta, Palaeont.
Indica, Ser. V. 2, 13, 1867-1868 : xiii, 500, 28 pls.
Tate, R. 1869. Contributions to Jurassic palaeontology. 1. Cvryptaulax, a new genus of
Cerithiadae. Ann. Mag. nat. Hist., London, Ser. 4, 4, 1869 : 417-419.
Tryon, G. W. 1887. Manual of conchology, structural and systematic, Vol. 9, Philadelphia,
1887 : 488, 71 pls.
VERNEUIL, P. E. P. de & LoriErRE, G. de. 1868. Description des fossiles du Néocomien
supérieur de Utrillas et ses envivons (Province de Tervel). 4°, Le Mans. 1868 : vii, 30,
3 pls.
VERRILL, A. E. 1882. Catalogue of marine molluscs added to the fauna of New England
during the past ten years. Tvans. Conn. Acad Arts Sci., New Haven, 5, 1882 : 447-587,
5 pls.
Vipat, L. M. 1917. Nota paleontolégica sobre el Cretaceo de Catulufia. Assoc. espan.
Progr. Cienc., Congr. Sevilla, Barcelona, 5, 1917 : 19, 3 pls.
Wane, B. 1917. New and little-known Gastropoda from the Upper Cretaceous of Tennessee.
Proc. Acad. nat. Sci., Philadelphia, 69, 1917 : 280-304, 3 pls.
1926. The fauna of the Ripley formation of Coon Creek, Tennessee. Prof. Pap. U.S. geol.
Surv., Washington, 137, 1926 : 272, 72 pls.
WANNER, J. 1902. Die Fauna der obersten weissen Kreide der libyschen Wiiste. Palaeonto-
graphica, Stuttgart, 30, Pt. 2, 1902 : 91-152, 7 pls.
PROCERITHIIDAE, CERITHIIDAE & CERITHIOPSIDAE 175
Wenz, W. 1938-1944. Handbuch dev Paldozoologie. Band 6 Gastropoda, in 7 vols., 8°, Berlin,
1, 1938; 2, 1938; 3, 1939; 4, 1940; 5, 1941; 6, 1943; 7, 1944 : 1639 pp, 4211 figs.
WoLLEemMaANN, A. tI900. Die Bivalven und Gastropoden des deutschen und hollandischen
Neocoms. Abh. K. Preuss. geol. Landes, Berlin, 31, 1900 : 180, 8 pls.
1903. Die Fauna des mittleren Gaults von Algermissen. Jahrb. K. Preuss. geol. Landes,
Berlin, 24, 1903 : 22—42, 2 pls.
1906. Die Bivalven und Gastropoden des nordeutschen Gaults (Aptiens und Albiens).
Jahrb. K. Preuss. geol. Landes, Berlin, 27, 1906 : 259-300, 5 pls.
1908. Nachtrag zu meinen Abhandlungen tiber die Bivalven und Gastropoden der
Unteren Kreide Norddeutschenlands. Jahrb. K. Preuss. geol. Landes, Berlin, 29, Pt 2,
1908 : 151-193, 5 pls.
Woop, C. J. 1969. Notes on the sedimentology of the Chalk. In Worssam, B. C. & Taytor,
J. H. ‘Geology of the country around Cambridge’. Mem. Geol. Surv. G. B., Sheet 188, New
Series, H.M.S.O., London, 1969 : 43-45.
Woops, H. 1896. The mollusca of the Chalk Rock, Pt 1. Q. Jl. geol. Soc., London, 52, 1,
1896 : 68-08, 3 pls.
WricLey, A. 1940. Some Eocene mollusca, with descriptions of new species. Pyoc. Malac.
Soc., London, 24, 1. 1940 : 6-17.
ZEKELI, F. 1852. Die Gastropoden der Gosaugebilde. Abh. K.K. geol. Reichsanst., Wien, 1,
2, 1852 : 124, 24 pls.
H. L. ApsBass
Dept. of Geology,
FACULTY OF SCIENCE,
Arn SHAMS UNIVERSITY,
CAIRO.
F
PLATE x
Fics. 1 & 2. Nerineopsis melburiensis sp. nov.
Lower Cenomanian, Chloritic Marl, Melbury Park, nr. Shaftesbury, Dorset.
Holotype GSM 93752
Fig. 1. Apertural view. X2°5.
Fig. 2. To show ornament of spiral cords. X2°5.
Fics 3 & 4. Nerineopsis cuckhamsliensis (Woods)
Fig. 3. Holotype, Sedgwick Museum B 4443, Chalk Rock, Turonian, Cuckhamsley, Wilts.
Xe
Fig. 4. BM(NH) G 48978, Chalk Rock, Holaster planus Zone, Hill End Pit, nr. Hitchin,
Herts. 3:5. Silicone rubber mould from natural external shell impression.
Fics 5-8. Nerineopsis aculeatum (Sharman & Newton), 1896
Fig. 5. BM(NH) G 55220, To show later ornament. x6.
Fig. 6. BM(NH) G 55191, To show general ornament. x6.
Fig. 7. BM(NH) G 55222, Lectotype. x6.
All from Speeton Clay, Speeton, Yorks., G.W. Lamplugh Coll’n.
Fig. 8. GSM 93749, General ornament and columella. 4. Speeton Clay, Speeton, Yorks.
Fics9 & 12. Nerineopsis subattenuatum (d’Orbigny)
Fig.9. GSM 93687. x3.
Fig. 12. BM(NH) GG 5610, C.W. Wright Coll’n. Both from Lower Aptian, Forbesi Zone,
Crackers Bed, Lower Greensand, Chale Bay, Isle of Wight.
Fics 10 & 11. Nerineopsis coxi sp. nov.
Gault, Folkestone, Kent, J.S. Gardner Coll’n.
Fig. 10. BM(NH) GG 6474. x4.
Fig. 11. BM(NH) GG 6475, Holotype. x4.
Fic. 13. Nerineopsis claxbiensis sp. nov.
Claxby Beds, Lower Hauterivian, Nettleton, Lincs.
BM(NH) GG 5612, Holotype. x4.
Fic. 14. Nerineopsis adeli sp. nov.
Upper Chalk, Springbank, nr. Coagh, County Derry, Northern Ireland.
GSM 93753, Holotype. x3.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 2 IPIL ANID, i
PATE ee
Fics 1 & 2. Cirsocerithium subspinosum (Deshayes)
Gault, Albian, Folkestone, Kent.
Fig. 1. Sedgwick Museum B 32530. x5.
Fig. 2. BM(NH) GG 20746, (specimen is now slightly damaged). » 6.
Fics 3 & 4. Cirsocerithium nooryi sp. nov.
Lower Chalk, Cenomanian, Ditchling, Sussex.
BM(NH) 98208, Holotype. x3.
Fic. 5. Cimolithium aff. eleanorae Allison.
Cenomanian, Charlton Bay, Devon.
BM(NH) G 74550. x2.
Fic. 6. Cimolithium ascheri Wollemann.
Neocomian, Speeton Clay, Speeton, Yorks.
BM(NH) G 67942. x2.
Fic. 7. Cimolithium cf. ascheri Wollemann.
Barremian, Cementstone Group, Cayton Bay, Yorks.
BM(NH) GG 20820. x7.
Fic. 8. ? Cirsocerithium kirkaldyi sp. nov.
Ferruginous Sands, Lower Greensand, Aptian, Shanklin, Isle of Wight.
BM(NH) GG 5603 Holotype. » 4.
Fics 9 & 10. Cimolithium ascheri Wollemann.
Holotype 414-415, x 2, in Geologisch-Palaeontologisches Institut, Georg-August Universitat,
Gottingen; Neocomian, Bohnenkamp nr. Querum, Sarstedt, North Germany.
Fics 11 & 12. Nudivagus morrisi sp. nov.
Ferruginous Sands, Upper Aptian, ? Group xiv, Lower Greensand, Shanklin,
Isle of Wight.
Holotype, Geological Survey Museum 2252. 2.
Fig. 11. Silicone rubber mould from natural impression of external ornament. Note growth
lines.
Fig. 12. Latex rubber mould from same specimen, which shows spiral threads.
Bull. By. Mus. nat. Hist. (Geol.) 23, 2
PLATE 3
Fic. 1. Bathraspira tecta (d’Orbigny)
BM(NH) GG 6227.
Silicone rubber cast of external mould preserved in nodule from the Regularis Subzone, Tarde-
furcata Zone, Albian, Arnold’s Pit, Billington Crossing, Leighton Buzzard, Beds. C.W. Wright
Collection. x4.
Fic. 2. Bathraspira ? tecta (d’Orb)
BM(NH) GG 20790, Gault, Albian, Folkestone, Kent. J.S. Gardner Collection. 6.
Fic. 3. Bathraspira shanklinensis sp. nov.
BM(NH) G 20956 Lower Greensand, Ferruginous Sands, ? Perna Bed, Aptian,
Redcliff, Sandown, Isle of Wight. Caleb Evans Collection. x2.
Fics 4-6. Bathraspira shanklinensis sp. nov.
BM(NH) GG 5602, Ferruginous Sands, Lower Greensand, Upper Aptian,
Shanklin, Isle of Wight. Holotype. C. W. Wright Collection.
Euger4n | <4e
Biges55 <3:
lene, ©, <3}.
Fic. 7. Bathraspira fouadi sp. nov.
Paratype, Sedgwick Museum B 32524, Gault Albian, Folkestone, Kent, Wiltshire
Collection. x3.
Fic. 8. Bathraspira fouadi sp. nov.
BM(NH) G 71562 Gault, Albian, Folkestone, Kent. x 3.
Fic. 9. Bathraspira brightoni sp. nov.
BM(NH) 34843 Upper Greensand, Blackdown, Devon. 3 approx.
Fic. 10. Bathraspira cleevelyi sp. nov.
BM(NH) GG 5607, Holotype ? Regularis Subzone, Tardefurcata Zone,
Lower Albian, Leighton Buzzard, Beds., or
Bed 5, Ford Place, nr. Wrotham, Kent. C.W. Wright collection.
Silicone rubber cast of external mould in nodule. 2°5.
Fics 11 & 15. Bathraspira cf. brightoni sp. nov.
BM(NH) GG 20791 Gault, Albian, St. Florentin, Yonne, France.
Fig. 11 shows strength of spirals and indications of growth lines. x 4.
Fig. 15 shows spiral ornament. » 4.
Fic. 12. Bathraspira fouadi sp. nov.
BM(NH) G 73795 Hoiotype. Gault, Albian, Folkestone, Kent. J. S. Gardner
Collection. 5.
Fic. 13. Bathraspira fouadi sp. nov. ? var.
BM(NH) G 73797 Gault, Albian, Folkestone, Kent. J.S. Gardner Collection. x2.
Fic. 14. Bathraspira brightoni sp. nov.
BM(NH) G 73796 Holotype. Gault, Albian, Folkestone, Kent. J. 5S. Gardner Collection.
<3 approx.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 2 PLATE 3
PILATE 4
Fics 1 & 6. Metacerithium aff. trimonile (Michelin)
Gault, Albian, Folkestone, Kent.
Fig. 1. BM(NH) G 73793. 3:25.
Fig. 6. Geological Survey Museum. 93725. x3.
Fics 2, 3, 7,8 & 10. Metacerithium trimonile (Michelin)
Gault, Albian, Folkestone, Kent. J. S. Gardner Collection, BM(NH)
Fig. 2. G 73794. X3-
Eilon 3G G647 2x 3
Higa 7p. GIO“ Os:
leis fo, (GG Ogi, <3}.
Fig. 10. GG 6476. x4.
Fic. 9. Metacerithium ? sp. nov
Sedgwick Museum, B 32530, Albian, Gault, Folkestone, Kent. 3.
Fic. 4. Metacerithium aff. ornatissimum (Deshayes)
Sedgwick Museum B 32518, Gault, Albian, Folkestone, Kent. x3.
Fic. 5. Metacerithium ornatissimum (Deshayes)
BM(NH) GG 5601. C. W. Wright Collection, Cenomanian,
Schloenbachia varians Zone. Aston Clinton, Bucks. x3.
Fic. 11. Metacerithium aff. ornatissimum (Desh.)
Geological Survey Museum 1698; Gault, Albian, Folkestone, Kent. 3.
Fic. 12. Metacerithium ornatissimum (Desh.)
BM(NH) G 28908. x3. Cenomanian varians Zone. Bed.1. Merstham, Surrey.
Collected. A. G. Davis & Ref’d. Davis (1923).
Bull. By. Mus. nat. Hist. (Geol.)
ye PLATE 4
PLATE 5
Fics 1-5 ? Rhabdocolpus forbesianum (dOrbigny)
Lower Greensand, Crackers Bed, Lower Aptian, Forbesi Zone, Atherfield, Isle of Wight.
Figs 1-4. Stereoscan.
Fig. 1. BM(NH) GG 20850. x20.
Fig. BM(NH) GG 20848, showing the ornament of the early whorls 23.
Fig Geological Survey Museum 2275, Lectotype from type series. 23.
2.
Be
Fig. 4. BM(NH) GG 20845. 56.
Fig. 5. BM(NH) GG 5609. 5.
Fic. 7. ? Rhabdocolpus melvillei sp. nov.
Lower Greensand, Crackers Bed, nr. Atherfield Point, Isle of Wight.
Holotype, Sedgwick Museum B 27340, Wiltshire Collection. 5.
Fics 6, 12 & 13. 2? Rhabdocolpus ? clementinum (d’Orb)
Lower Greensand, Crackers Bed, Aptian, Atherfield, Isle of Wight. Stereoscan. x 23.
Fig. 6. BM(NH) GG 20841.
Fig. 12. BM(NH) GG 20842.
Fig. 13. BM(NH) GG 5694.
Fics 8-11. Metacerithium turriculatum (Forbes)
Lower Greensand, Crackers Bed, Atherfield, Isle of Wight.
Fig. 8. BM(NH) G 74602. x5.
Figs 9 & 11. BM(NH) GG 20814, J. E. Lee Collection. x5, x7.
Fig. 10. Geological Survey Museum 2253, Specimen on left is the Holotype. 3.
nu
a)
a
a
(e)
o
S
~
az)
~
S
=
=
a
x
3
~~
|
BS
ica)
Ss
=
isa)
Pe ATI 6
Fics 1 & 2. ? Campanile cenomanica sp. nov.
BM(NH) GG 18686. ? Cenomanian, Dorset. x 2.
Fics 3 & 4. Uchausxia badri sp. nov.
Sedgwick Museum B 44632. Upper Greensand, Albian, Blackdown, Devon. x6.
Fics 5 & 6. Uchausxia wisei sp. nov.
C. W. Wright Collection. Claxby Ironstone, Valanginian-Hauterivian, Nettleton, Lincs.
Fig.5. BM(NH) GG 5611. x3°5.
Fig. 6. BM(NH) GG 6161 Holotype. 3.
Fic. 7. Uchausia sp.
BM(NH) G 73788. Gault, Albian, Folkestone, Kent. J.S. Gardner Collection. x6.
Fic. 8. Cerithiella praelonga (Deshayes)
BM(NH) 72024. Middle Eocene, Bracklesham Beds, Bracklesham Bay, Hants. 5.
Fic. 9. Cerithiella devonica sp. nov.
Sedgwick Museum B 44633. Upper Greensand, Albian, Blackdown, Devon. x5.
Fic. 10. Cerithiella atherfieldensis sp. nov.
Sedgwick Museum B 27334. Lower Greensand, Crackers Bed, Lower Aptian,
Forbesi Zone, Atherfield, Isle of Wight. 3.
Fic. 11. Seila iglali sp. nov.
BM(NH) G 16084. Upper Greensand, Albian, Combe Raleigh Hill,
nr. Honiton, Devon. W. Vicary Collection. x 4.
Fic. 12. Orthochetus hantoniensis sp. nov.
BM(NH) G 70130. Aptian, Atherfield, Isle of Wight. P. Cambridge Coilection. 5.
Fics 13 & 14. ‘Orthochetus’ helmyi sp. nov.
Gault, Albian, Folkestone, Kent. x 3.
Fig. 13. BM(NH) G 10500, Holotype.
Fig. 14. BM(NH) G 71988.
Fic. 15. ‘Orthochetus’ sp. nov.
BM(NH) 9145, Lower Greensand, Sandgate Beds, Upper Aptian,
Parham Park, Sussex. G. A. Mantell Collection. x 3.
Bull. By. Mus. nat. Hist. (Geol.) 23, 2 PLATE 6
PLATE 7
Fics 1 & 2. Exechocirsus aff. saundersi (Woods)
Cretaceous, Turonian, Chalk Rock, Holaster planus Zone;
Hitch Wood Pit, near Hitchin, Herts.
Fig. 1. BM(NH) GG 20821; shows variation in ornament i.e. weakly developed tubercles
and the presence of intermediate (or secondary) spirals. Both C. W. Wright Collection, and
silicone rubber moulds from external impressions. 2.
Fics 3-5. Exechocirsus saundersi (Woods)
Fig. 3. BM(NH) GG 66724; Turonian, Chalk Rock, Blackwell Hall Farm, Latimer, Bucks.
Silicone rubber mould. x 2.
Fig. 4. BM(NH) G 67704; Turonian, Chalk Rock, Dover, Kent. A. W. Rowe Collection.
Silicone rubber mould. x 4.
Fig. 5. Sedgwick Museum B 4459, Holotype; Turonian, Holaster planus Zone, Cuckhamsley,
North Farnborough, Berks. Silicone rubber mould. 3.
Fics 6-7. Exechocirsus pustulosus (J. de C. Sowerby) 1833.
BM(NH) G 17916; Holotype; Upper Cretaceous, Gosau, Austria. TI.
Fics 8 & 9. Exechocirsus aff. saundersi (Woods)
BM(NH) G 60542-60543.
Silicone rubber moulds from impressions of each half of the same shell. No other information
available. x3.
Fics 10 & 11. Exechocirsus aff. subpustulosus Pchelinceyv
Fig. 10. BM(NH) G 16134.
Fig. 11. BM(NH) G 73798.
Both from the Albian, Haldon, Devon; W. Vicary Collection. x 2.
Fics 12-14. Exechocirsus aff. saundersi (Woods)
Fig. 12. BM(NH) GG 20823, Hitch Wood Pit, nr. Hitchin, Herts. C. W. Wright Collection.
Silicone rubber mould. x 2.
Fig. 13. BM(NH) GG 5934. Turonian, Holaster planus Zone, Kiplingcotes, E. Yorks.
C. W. Wright Collection. Silicone rubber mould. x 2.
Fig. 14. BM(NH) GG 5983. Same locality as above; natural mould. » 4.
7
JPL AD 18
Bull. By. Mus. nat. Hist. (Geol.) 23, 2
PAC aS
Fics 1-4 & 6. Ageria gaultina sp. nov.
Albian, Gault, Folkestone, Kent. J. S. Gardner Collection. BM(NH) specimens
Fig. 1. GG 6473. Holotype, x4.
ime, 2, (CG a7 33}.
Fig. 3. GG 20742. 3. ? Bedv, Lower Gault (fide Dr. H. G. Owen), shows slight variation
in ornament.
Fig. 4. GG 20862. Shows the ornament of the earlier whorls. » 4.
Fig. 6. GG 20857. x3.
Fic. 5. Ageria gaultina sp. nov.
Albian, Osmington, Dorset, BM (NH) G49804. 3°5.
Fics 7-10. Ageria costata (J. de C. Sowerby)
Upper Greensand, Albian; Blackdown, Devon. BM(NH) specimens
Fig. 7. 71382. Shows aperture. x4.
Fig. 8. 43674. Holotype, Sowerby Collection. 5.
Fig. 9. 34844. Also shows aperture. x6.
JES, Ti@; (EKG; BOS, >< 3}
Fics 12. Ageria costata (J. de C. Sowerby)
BM(NH) G 71095, Albian Upper Greensand; Bed 12, Peak Hill,
nr. Sidmouth, Devon. H. F. Metcalfe Collection. x 3.
Fic. 13. ? Ageria angustata (d Orbigny)
BM(NH) GG 20740. Neocomian, Claxby Beds, Nettleton, nr. Caiston, Lincs
C. W. Wright Collection. 5.
Bull. By. Mus. nat. Hist.
BOE:
= é =
—@ eS
A LIST OF SUPPLEMENTS
TO THE GEOLOGICAL SERIES
OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya.
Pp. 213; 30 Plates; 2 Text-figures. 1965. 6.
. Et-Naccar, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper
Cretaceous—Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley,
Egypt, U.A.R. Pp. 291; 23 Plates; 18 Text-figures. 1966. {10.
. Davey, R. J., Downie, C., SaRGEANT, W. A. S. & Witiiams, G. L. Studies on
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248; 28 Plates; 64 Text-
figures. 1966. {7.
. APPENDIX. Davey, R. J., Downie, C., SARGEANT, W. A. S. & WiLLias, G. L.
Appendix to Studies on Mesozoic and Cainozoic an meckne Cysts. Pp.ige
1969. 8op.
. Ertiott, G. F. Permian to Palaeocene Calcareous Aisi (Dasycladaceae) of the
Middle East. Pp. 111; 24 Plates; 17 Text-figures. 1968. £5.12.
. RHOovDES, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni-_
ferous) Conodont faunas, and their value in local and continental correlation.
Pp. 315; 31 Plates; 92 Text-figures. 1969. 11.
. Cuitps, A. Upper Jurassic Rhynchonellid Brachiopods from Northwestern
Europe. Pp. 119; 12 Plates; 40 Text-figures. 1969. £4.75.
. Goopy, P. C. The relationships of certain Upper Cretaceous Teleosts with
special reference to the Myctophorids. Pp. 255; 102 Text-figures. 1969. £6.50.
. OwEN, H. G. Middle Albian Stratigraphy in the Paris Basin. Pp. 164;
3 Plates; 52 Text-figures. 1971. 6.
. Sippigu1, Q. A. Early Tertiary Ostracoda of the family Trachyleberididae
from West Pakistan. Pp. 98; 42 Plates; 7 Text-figures. 1971. {8.
id
Ser Printed in England by Staples Printers Limited at their Kettering, Northants, establishment
£
q
*
A REVIEW OF SOME ENGLISH
-PALAEOGENE NASSARIIDAE,
_ FORMERLY REFERRED TO
COMINELLA
4 on MUs
BR
4 JULI973
4, &
Y, RAL WSS
©. P; NUTTALL
‘ AND
_ J. COOPER
im rie
“fd iM ¥ uly
nye .
%
eA
\
° a ;
vel ,
¥ :
rs
rr ts, i N, +
ys Zak “ ;
ye, Cea
Ry
Sa e - BULLETIN OF
€ BRITISH MUSEUM amas HISTORY)
Vol. 23 No. 3
Ub a 3
LONDON: 1973
A REVIEW OF SOME ENGLISH PALAEOGENE ASA OS
NASSARITDAE, FORMERLY REFERRED TO a i
COMINELLA ( 4 JULI973 |
&%, e/
NAL WS
BY
CLIVE PATRICK NUTTALL. ...(
AND
JOHN COOPER
Pp. 177-219; 9 Plates; 1 Text-figure
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 23 No. 3
LONDON : 1973
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY) instituted im 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
veady. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper 1s Vol. 23, No. 3 of the Geological
(Palaeontological) series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation
Bull. Br. Mus. nat. Hist. (Geol.)
© Trustees of the British Museum (Natural History), 1973
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 29 June, 1973 Price {2°50
A REVIEW OF SOME ENGLISH PALAEOGENE
NASSARIIDAE, FORMERLY REFERRED TO
COMINELLA
By
CLIVE PATRICK NUTTALL
and
JOHN COOPER
CONTENTS
Page
SYNOPSIS ‘ : : : : : : : : ; ‘ 179
INTRODUCTION : : ‘ : : : : ‘ : . 180
MATERIAL STUDIED . : a ; ‘ : 183
TABLE I. Distribution of some early Nassariidae : 4 : 184
Explanation of abbreviations referring to sources of saosin ; : 186
LOCALITIES AND HORIZONS 5 ¢ . é : ¢ : : 186
MODE OF LIFE . : < 0 . . : : c 5 : 188
ACKNOWLEDGEMENTS : : 2 ; : : : : : 188
SYSTEMATIC DESCRIPTIONS : : : 3 : , : ‘ 189
Pseudocominella gen. nov. : 2 : : 5 3 : ; 189
P. deserta (Solander) . : : 5 : , G c 190
P. avmata (J. de C. Gowerby) : 6 : . : : : 193
P. semicostata sp. nov. : é c : : : : : 196
P. solandevt (Cossmann) 5 : c : c : cl . 198
Desovinassa gen. nov. . : : : 5 : ; : ‘ 201
D. desori (Deshayes) . F ‘ ; F ‘ ; : : 202
D. williamsi sp. nov. : . - : : i 4 : 203
Whiteclifia gen. nov. . : 3 : : c : c : 204
W. suturosa (Nyst) . : 2 Z : : 3 : : 204
W.tumida sp.nov. . : 3 : : ¢ . : : 206
Colwellia gen. nov. : 5 5 , 0 ; : 208
Colwellia flexuosa (Edwards) ° : c ; c . : 210
Keepingia gen. nov. ‘ ‘ : j é : : : 6 212
Thanetinassa gen. nov. . : : : : ° : : : 212
T. bicorona (Melleville) : : - : : : 2 213
REFERENCES . : : . 5 5 : : : . 5 215
SYNOPSIS
English Palaeogene prosobranch gastropod species, previously assigned to the Tertiary and
living New Zealand genus Cominella (Buccinidae or Cominellidae of Buccinacea), are reviewed.
New genera are described to accommodate these English species and their European relatives.
They are assigned to the Nassariidae (Buccinacea) on the characters of their fascioles and
columellae, in particular the presence of a terminal columellar plait. Like living Nassariidae,
180 SOME ENGLISH
they probably lived in warm seas, in shallow water, possibly intertidally. Their known distri-
bution, both in Europe and beyond, is summarized. Colwellia nov. gen. occurs both in the
European and the United States west coast Eocene (California to Washington State) where it is
represented by species usually assigned to Molopophorus. Most other species assigned to the
latter genus also appear to belong to the Nassariidae. Its type species, M. stviatus (Gabb)
seems not to belong to the family but to be a synonym of Brachysphingus which is shown to
lack the characteristic terminal columellar plait. Palaeocene Thanetinassa nov. gen. shows
certain similarities to the later Palaeogene Phos-like (Buccinidae) genera Tritiavia, Buccitriton
and to Sagenella (Nassariidae), all from the U.S. south-eastern province. This publication is a
preliminary to a more intensive review of the Nassariidae and their possible relationship with the
Phos-like group.
The following new genera of Nassariidae are described:—Pseudocominella, Desorinassa,
Whitechffia, Colwellia, Keepingia and Thanetinassa. The following new species are described
from the Hampshire Basin. Eocene (Ypresian):—Desorinassa williamsi; Late Priabonian :—
Pseudocominella semicostata, and Whitecliffia tumida. Several Palaeogene species, including
Buccinum canaliculatum J. de C. Sowerby and B. montense Briart & Cornet are shown to be
unrelated to the Nassariidae.
INTRODUCTION
THE present study started as a result of a request for identification of a specimen
from the English Lower Eocene which proved to belong to an undescribed species.
It is one of a small group of species in need of revision which are normally assigned
to the living New Zealand genus Cominella. ~
The genus Cominella was erected by Gray (1850: 72) who assigned to it nine
living species from New Zealand, Australia and South Africa. Iredale (1918 : 34)
designated as type species the first mentioned, Buccinum testudineum Lamarck from
New Zealand. He also selected the living South African species Buccimum cinctum
Réding (= porcatum Gmelin, preoccupied) as type species of his new genus
Burnupena.
During the latter part of the nineteenth century the name Cominella came to be
almost universally applied to a group of European species, most of which are now
assigned to the new genera described here. The designation by Cossman (Ig0I : 149)
of Buccinum porcatum Gmelin as type species of Comznella is invalid as it was not
among those listed by Gray. Cossmann, unfortunately, also based his definition of
the genus partly on two European Palaeogene species which he regarded as ‘plesio-
types’, Buccinum gossardit Nyst and Buccinum desertum Solander. He regarded
Molopophorus from the Eocene of California as a probable synonym of Cominella,
and (Igor : 201) stated that Molopophorus should be eliminated from the Nassidae
(now known as Nassarlidae), though expressing doubts as to the accuracy of Gabb’s
figure of the lectotype of M. striatus Gabb, the type species of Molopophorus. Later
Cossmann indicated that he had no remaining doubts as to this synonymy after
examining the type figures of another species, Molopophorus gabbi Dall (1909 : 45,
pl. 3, fig. 8), from the Oligocene of Oregon (see Weaver, 1942 : 466). Subsequent
American workers (Stewart, 1926 : 389; Vokes, 1939 : 140-141) have suggested a
connection between Molopophorus and European Palaeogene species assigned to
Cominella. In the present study four United States West Coast species, previously
PALAEOGENE NASSARIIDAE r81
assigned to Molpophorus, are recognized as belonging to Colwellia. A re-examination
of the lectotype of Molopophorus striatus suggests that it is a juvenile Brachysphingus,
and that neither belong to the Nassariidae (see p. 209). All other species normally
regarded as belonging to Molopophorus appear to have little in common either with
its type species or with Brachysphingus and seem to belong to more than one
undescribed genus. Gabb (1869 : 156, 157, pl. 26, fig. 36) in describing Molo-
pophorus as a subgenus of Bullia, compared it with Bullia (Buccinanops) monilifera
(Kiener) now living on the Atlantic Coast of South America. Not only the type
species of Molopophorus, but also species assigned to that genus later, and Brachy-
sphingus have been generally accepted by American authorities as members of the
Nassarilidae.
Some of the gross shell characters of these American Tertiary forms and the
European Tertiary genera described in this paper closely resemble those found in
modern Nassariidae and in Cominella which have fasciole regions of a broadly
similar type (see Text-fig. 1). The growth lines on the neck region are bent aborally
to form a ridge confluent with the adapical margin of the track of the anterior
notch. The deepest point of the asymmetrically U-shaped notch is usually fairly
close to the ridge. Below the ridge there is some variation. The growth lines range
from regular to imbricate and are reverse S-shaped, but the upper and lower halves
of the S may be of varying proportions and curvature. In true Cominella the notch
corresponds with a raised ridge, as opposed to a groove observed in many definite
Nassariidae and the fossil genera under consideration. A more important feature
that the latter share with definite Nassariidae is that the columella is bent strongly
to the left and bears a terminal plait. Comunella lacks this plait and its columella
is not so strongly bent.
On these grounds these fossil genera may be assigned with confidence to the
Nassariidae rather than to the Buccinidae (as constituted by Wenz, 1941 : II51I-—
1200; 1943 : 1201-1210), which includes the Cominellidae of Powell (1929 : 57).
Homoeomorphic, superficial, similarities between these genera and Comuinella
include the frequent development of a subsutural platform and concave ramp and the
rather flexuous but basically orthocline growth lines.
Buccinum canaliculatum (J. de C. Sowerby, 1823 : 14, pl. 415, fig. 2) from the
Barton Beds of the English Upper Eocene was placed in Cominella by Newton
(1891 : 168). Cossmann (1901 : 121-123, pl. 8, figs 13, 14) selected it as the type
species of his monotypic genus Bartonia. He compared it with ‘Cominella’ deserta
from the same formation, pointing out the differences in the features of their aper-
tures and canals. Bartonia canaliculata lacks a columellar plait, its columella
curves to the left and is not truncated, no ridge is formed at the adapical margin of
its fasciole and the growth lines in the fasciole region are less strongly curved than in
Pseudocominella (Text-fig. 1). It is hoped that Bartonia, which remains assigned
to the Buccinidae (sensu Wenz, 1941-1943) will be treated in a future paper.
In this paper, five genera, Pseudocominella, Desorinassa, Colwellia, Whitecliffia
and Thanetinassa are described to accommodate British and European Tertiary
species previously assigned to Cominella. The opportunity is also taken to describe
a sixth genus of the group, Keepingia, though it does not occur in England. The
182 SOME ENGLISH
Ab
Fic. 1. Fasciole and columellar details of 1, Cominella testudinea (Lamarck), type species
of Cominella (fam. Buccinidae) Recent, New Zealand, BMZD 1844.7.29.36, showing
ridge above fasciole and no columellar plait (also see pl. 9, figs 11a, b), x5. 2, Sphaero-
nassa mutabilis (Linné), type species of Sphaeronassa (fam. Nassariidae), Recent, Bay of
Naples, BMZD 19722, showing strong ridge above fasciole, strong terminal columellar
plait with weaker plaits above, x5. 3, Pseudocominella deserta (Solander) type species
of Pseudocominella, Upper Eocene, Lower Barton Beds Bed B, Barton, GG 19675 (Ru),
showing strong ridge above fasciole, imbricate growth lines of fasciole, strong terminal
columellar plait with weaker plaits above (see also pl. 1 figs 7a, b), x5. 4a, b, Bartonia
canaliculata (J. de C. Sowerby), type species of Bavtonia (fam. Buccinidae), Upper Eocene
Lower Barton Beds, Barton. 4a, GG 6897/1 (Wr), apertural view, x 2-5; 4b, GG 6897/2
detail, showing no sharp break between neck and fasciole and no columellar plait, «7-5.
PALAEOGENE NASSARIIDAE 183
gross shell characters of these new genera are fairly similar, the main differences
being in the form of the apical whorls, strength of sculpture and the strength and
character of the ridging in the fasciole region.
The known distribution of these genera, in Europe and beyond, is summarized on
Table 1. We have examined numerous references to Cominella, ‘Nassa’, Molo-
pophorus and Brachysphingus but have found few plausible extra-European
occurrences of the genera newly described here.
The recognition of these additional Palaeogene members of the Nassariidae
affects our understanding of the evolution of the family. It is intended to pursue this
topic in a future paper.
Although it is hoped that the specific and generic diagnoses will be sufficient to
distinguish different taxa, the descriptions given here are detailed. In addition,
the variation in a species from locality to locality has been illustrated and commented
on as fully as possible. This policy is justified because apparently good illustrations
and descriptions have proved inadequate and misleading, so that important points
have only been resolved by the study of actual specimens. To quote only a few
examples, it seems incredible that the European Tertiary species discussed herein
should have been accepted for about one hundred years as members of the Recent
New Zealand genus Cominella which belongs to another family, that Buccinum
montense Briart & Cornet was not recognized as being congeneric with many Eocene
species assigned to Pollia of the Buccinidae, that Bartonia canaliculata (J. de C.
Sowerby) should have been considered congeneric with Buccinum desertum Solander,
and that the French Lutetian and Auversian Buccinum fusiforme Deshayes should
have been considered synonymous with the Bartonian B. desertwm Solander rather
than with the Auversian Stvepsidura armata J. de C. Sowerby.
MATERIAL STUDIED
This study of English specimens is based primarily on the collections in the British
Museum (Natural History). Material in the Sedgwick Museum, Cambridge, the
Geological Survey Museum, London, the Sandown Museum, Isle of Wight, Institut
Royal des Sciences Naturelles de Belgique, Bruxelles, Institut de Paléontologie,
Muséum National d’Histoire Naturelle, Paris, and Ecole des Mines, Paris has also
been examined. In addition, much valuable information has been obtained from the
stratigraphically detailed labels of several private collectors. To indicate the authen-
ticity of various records, the collector is named when known.
The most important single collection is that of F. E. Edwards in the British
Museum (Natural History). Much of his collection was obtained from workers such
as Henry Keeping, who also collected the fossils from Huntingbridge listed by
Fisher (1862 : 79), and it was enriched by exchange with foreign palaeontologists.
It forms the basis of many published faunal lists (von Koenen, 1864; Judd, 1880;
Keeping & Tawney, 1881). The British material in the Edwards collection was
catalogued by Newton (1891), who included many of Edwards’ manuscript names.
Both the Edwards collection and Newton’s catalogue may be presumed to be the
SOME ENGLISH
184
(‘poururexe useq JOU DALY »% PAYIeUI soroeds yo suourtoads :930u)
(viuLsny) vyvssoo0rdns *‘Gixx —_ -- —
(OD0]) tassv9aunng ‘Tixx — —
DAIDIAAI “Dd
pyonbyun “>
susuauola, “Dd
— 12404q *9D DIAISAD * I —
DSOANANS “fy -VSOANANS *
_ — DAagsy DI0]1]NG ‘q wpawssos “yy
DIONE “d DIV1IN4 ‘d
(eseqys Len) DSOANINS * {4 VSOANNS *
(NVISINVG “M) 2a/DpUDUUD “Sixx — DUADPISSYI “SJ PADSSOS “3
BIVUAISWUMN ° ST
4 re DIADPISSDI * ST 1199 “M
4199 “ST
SHONAAANIDOQ (LSvOD Isa) GNVTIOPL
UTHLO VoIuany ANVWUaD walotag
HLUON
TT
1A0sap *‘T
DYUOAOIIG * T.
AA
DIDUAD “I
DIDULAD *
asuarssannn *
OR
UpADSSOs “ST
SUSUGANJD “ST
suaparavad “Mex
91112940} “MY
FONVUT
sepliiesseN ApIee oulos Jo WOIZNqIIysSIG
I alavy
1Aaosap *‘
DUO0AOIIG * I.
smpyyim °C
DIDUAD * J
uvijouey
uereureds
uersoid x
welsing
eon Ty
uvISIOANY
iapunjos ‘q (‘uemoqerig
vyaasap “I
vpyuny * M
DSOANINS *
vsonxayf "9D
pIDsoIMas * qT
wyaasap *q
ANVIOND
Ayres)
ueuoyieg
(uvtuoqetig
oyeT)
ueyI0}}eT[
ueredny
uenyeyD
uerueqyinby
uelesiping
UeIN0}IOT,
HOVLS
HNAOOAV TVd
HNAHOO4
AHNAOVOSITO
AHNHOOIWN
PALAEOGENE NASSARIIDAE 185
EXPLANATION TO TABLE 1
The table is compiled from:
(a) England (this paper).
(b) France: Cossmann, 1901; Deshayes, 1865; Glibert, 1960, 1963; Peyrot, 1927.
(c) Belgium & Holland: Albrecht & van der Valk, 1943; Glibert, 1954, 1957.
) Germany: Beyrich, 1854; Glibert, 1960, 1963; Sandberger, 1863; von Koenen, 1889.
) North America: Vokes, 1939; Weaver, 1942.
f) Austria: Traub, 1938.
g) Togo: Furon & Kouriatchy, 1948.
h) West Pakistan: Vredenburg, 1925.
Notes. An examination of the holotype and paratype (Bayerisches Geol. Landesamt no.
3080, 3081) of Northia (Cominella) angusta H6lzl (1958 : 244, pl. 20, figs 16-16a) suggests that the
species should be referred to Dovsanum. Buccinum montense Briart & Cornet (1871 : 30-31),
referred by subsequent authors to Cominella is a Pollia (Buccinidae). Many North American
species ranging from Paleocene to Miocene in age are omitted from this table. They belong
to undescribed genera (see p. 209) close to those described herein. An examination of the
holotype of Cominella devtonensis Bellardi (1882: 3, pl. 1 figs ra, rb) from the Middle Miocene
(Tortonian) of St Agata-fossili, Italy (Instituto di Geologia, Turin, no. 11436) shows that it is
‘not a member of the Nassariidae and is in no way related to Pseodocominella.
Abbreviations of generic names
C., Colwellia; D., Desovinassa; K., Keepingia; P., Pseudocominella; T., Thanetinassa; W.,
Whitecliffia.
186 SOME ENGLISH
main sources of the records of species in Britain quoted by Continental authorities
such as von Koenen and Cossmann. Keeping’s own collection in the Sedgwick
Museum, Cambridge, is of particular value because of the reliability of his labels.
The publication (1933) of the late E. St. J. Burton’s letter classification of horizons
in the Barton Beds at the type locality Barton-on-Sea has enabled later collectors to
indicate horizons more precisely than hitherto. The faunal list published by Burton
was by no means complete. His own collection, now in the British Museum (Natural
History), is not as valuable as it might have been since he often arranged specimens
of a particular species from several horizons in the same box.
In the systematic descriptions, the localities and horizons of material which we
have examined are given after the synonymy, under the heading ‘Material Studied’.
It has not been considered necessary to list here material in other Museums which
merely complements the British Museum collections unless the record is of particular
interest.
Under the subsequent heading ‘Further Occurrences’, we give records in the
literature of those other localities and horizons from which we have not studied
material. To avoid undue repetition, we have omitted those records that were
probably based on the specimens which we have studied.
Explanation of abbreviations referring to sources of specimens
ANSP, Academy of Natural Sciences of Philadelphia; CAS, California Academy of
Science; Cu, D. Curry Colln.; Ed, F. E. Edwards Colln.; EMP, Ecole des Mines,
Paris—invertebrate collections now being transferred to Laboratoire de Paléontologie,
Institut de Géologie, Université de Paris, Orsay-91, France, under direction of
M. J. Manivit; Ho, M. J. Hoare Colln.; G or GG (followed by number), Department
of Palaeontology, British Museum (Natural History); JRScNB, Institut Royal des
Sciences Naturelles de Belgique, Bruxelles; Je, P. Jennings Colln.; K, H. Keeping
Colln.; KGéT, H. Keeping & E. B. Tawney Colln.; Le, D. N. Lewis Colln.; MNP,
Institut de Paléontologie, Muséum National d’Histoire Naturelle, Paris; Nw,
C. P. Nuttall Colln.; Ru, A. J. Rundle Colln.; St. J. B, E. St. J. Burton Colln. ; Sedg.
Mus., Sedgwick Museum, Cambridge; Sand Mus., Sandown Museum & Public
Library, Sandown, I.0.W.; S, F. C. Stinton Colln.; U.Cal., Museum of Paleontology,
University of California, Berkeley; Wr, A. Wrigley Colln.; ZD, Recent Mollusca
Section, Department of Zoology, British Museum (Natural History).
LOCALITIES AND HORIZONS
Many of the British localities are still in existence, are well-known and need no
further introduction. Some others, notably the New Forest (Hampshire) localities
of the Bracklesham and Middle Headon Beds are either poorly known or are no
longer accessible. Curry (1958) gives details and map references to most.
The following points should be noted about locality names attached to the Edwards
Collection:
PALAEOGENE NASSARIIDAE 187
1. Bramshaw, Upper Bracklesham Beds, is described (Curry, 1958 : 70) under
Shepherd’s Gutter Bed.
2. Roydon, Middle Headon Beds, refers to Royden Manor brick pit (now over-
grown) at Nat. Grid SU 320005. Curry (1958 : 29) discusses this locality under the
heading ‘Brockenhurst Beds’. The section was described by Keeping & Tawney
(r88r : 113 and 114 footnote), the latter refers to the fact that Keeping collected the
material in the Edwards Collection.
3. Brockenhurst, Middle Headon Beds. Specimens in the Edwards Collection
are labelled either Brockenhurst or Whitley Ridge. Keeping & Tawney (1881 :
10g) write ‘The greater part of the fossils from Brockenhurst were collected by the
hands of one of the authors (i.e. Keeping) and . . . dispersed into the various public
and private collections. They were obtained from the (railway) cutting at Whitley
... about (1858, when an extra track was being laid). Ina footnote (pp. 109-110),
they point out that they both visited the cutting in the summer of 1880, and that the
rich (fossiliferous) zone would never be seen here again but that fossils could still
be obtained from the spoil heaps (left from the cutting for the original single-track
railway in about 1838). As it is clear from a further footnote (p. 114) that Keeping
collected the Royden and Lyndhurst fossils in the Edwards Collection, it is more than
likely that the same applied for Whitley Ridge. A possible explanation for some of
Edwards’ material being labelled ‘Brockenhurst’ is that it was obtained from the
spoil-heaps. It seems extremely unlikely that the material came from the Victoria
Tile Works at SU 316034. This locality was not mentioned by Keeping & Tawney
(1881). It was in existence by 1890 and was normally worked to only a shallow
depth as the weathered material near the surface produced the best-coloured products,
so that the Brockenhurst shell bed was seldom exposed (Davis, 1952 : 215-216).
The foregoing, however, suggests that Davis might have been mistaken in stating
that Edwards had worked the local exposures. Whitley Ridge Cutting is the type
locality of the Brockenhurst Beds (Curry, 1958 : 29).
4. Lyndhurst, Middle Headon Beds. Edwards’ specimens were probably
collected by Keeping in 1858 from pits dug on Lyndhurst Hill, to the west of Lynd-
hurst, the Brockenhurst Bed outcropping at about the 200 ft contour (Tawney,
1883).
The Brander Collection locality ‘Hordwell’ (= Hordle) quoted by Solander
(1766) was the nearest village to the Barton coastal section before Barton itself was
built and named.
The Middle Headon Beds have been regarded by most British workers as being of
Lower Oligocene (Lattorfian) age. Recent nannoplankton zonation work by
Martini & Ritzkowski (1968 : 244-247), Martini (1969 : 117-159), and Martini &
Moorkens (1969 : 125-127) show that the Headon Beds are late Priabonian and that
some German localities, previously regarded as Oligocene are distinctly pre-
Lattorfian. Westeregeln, in part, is now regarded as being approximately equivalent
to the Upper Barton Beds.
The ages accorded here to French Paris Basin localities are based on the lists
given by Glibert (1960 : 3-5), and by reference to Fritel (1910).
188 SOME ENGLISH
MODE OF LIFE
Pseudocominella and its relatives appear to have lived in shallow water.
Thanetinassa bicorona, according to Briart & Cornet (1871 : 31), is very common
at some French localities, notably Chalons-sur-Vesle in the Sables Inférieurs—a
shallow-water sandy deposit. The species also occurs in the coarsely-grained Thanet
Sands in England. No species of the group are known from the London Clay.
The only British species occurring in rocks of equivalent age is Desorinassa williamsi
from the shallow-water Bognor Rock. Pseudocominella armata is rare and occurs
in the rich fauna of the Upper and more clayey part of the Bracklesham Beds.
No species is known from the more sandy Lower Bracklesham Beds. In the Barton
Beds, species of Pseudocominella are known from Horizons Az to H. P. deserta
is never common. From the appearance of the specimens it would seem that most
were obtained from the extremely fossiliferous Horizon E. P. solanderi, however,
is extremely common in the lenticular, sandy, shell drifts of Horizon A3. Members
of the group also occur in the Brockenhurst and Venus Beds of the Headon Series.
These two latter beds represent periods of shallow marine deposition in a largely
fresh and brackish water succession.
Turning again to the European Continent, many of the occurrences are consistent
with a shallow-water mode of life. Examples include P. avmata in the Calcaire
Grossier and Keepingia gossardi in the Sables Supérieurs of the Paris Basin, K.
gossardi and Whitecliffia tumida in the Belgian Oligocene and K. cassidavia in the
late Oligocene deposits of the Mainz Basin. Finally, we note the occurrence of
species of Keepingia in the Lower Miocene of the Bordeaux region in conjunction
with Nassarius, Dorsanum and Cyllene, all living genera of the Nassariidae.
Nearly all the present day Nassariidae live in very shallow water and many inter-
tidal species of the three most important genera, Nassarius s.l., Bullia and Buccinanops
are known. Dorsanum and Cyllene are rare but are known only from shallow-water.
Considering only the British species of the Pseudocominella group, it would seem
unlikely that either P. aymata or P. deserta was intertidal. The evidence in the case
of the other species is not clear cut; any of them could well have been intertidal
but there is no proof. Such a mode of life would help to explain the rather sporadic
fossil record of the group; inshore deposits are relatively often eroded after deposition.
The seas in which these forms lived would have been considerably warmer than those
around Northern Europe now. At the present day the family is important in warmer
waters, mostly within 40° North or South of the Equator. The occurrence of a few
species of Nassarius s.l. in Northern European seas may be due to the warming
influence of the Gulf Stream. Modern members of the family are carrion-eaters
(Fretter & Graham, 1962 : 522).
ACKNOWLEDGEMENTS
Many collectors have allowed us to study their well authenticated material and
have willingly presented to the British Museum (Natural History) those specimens
that have been asked for. For such help we therefore wish to thank Messrs D.
PALAEOGENE NASSARIIDAE 189
Curry, M. J. Hoare, D. N. Lewis, A. J. Rundle and F. C. Stinton as well as Mr and
Mrs R. Jennings and Mr and Mrs B. A. Williams. We thank Mr D. Curry also for
useful advice on some nomenclatorial problems. We extend our thanks to Dr
Horace G. Richards, Academy of Natural Sciences of Philadelphia; Dr Leo G.
Hertlein, Academy of Science, San Francisco; Dr Joseph H. Peck, Museum of
Paleontology, University of California at Berkeley; Dr Annie Dhondt and Dr M.
Glibert, Institut Royal des Sciences Naturelles de Belgique, Bruxelles; Drs J. C.
Fischer and P. Brebion, Muséum National d’Histoire Naturelle, Paris; Dr Denise
Petitbois, Ecole des Mines, Paris and M. J. Manivit, Université de Paris, Orsay;
Dr C. L. Forbes and Dr B. R. Rickards, Sedgwick Museum, Cambridge; and Mr
A. E. Baker of Sandown Museum, I.O.W., all of whom either have given access to the
collections in their charge or have loaned specimens.
SYSTEMATIC DESCRIPTIONS
Order NEOGASTROPODA
Superfamily BUCCINACEA
Family NASSARIIDAE
PSEUDOCOMINELLA gen. nov.
(Pls 1-3; Pl. 4, figs 1-3)
TYPE SPECIES. Buccinum desertum Solander, 1766.
DiacGnosis. Bucciniform; moderate sized, normally attaining an adult height
of between 20 and 35 mm; protoconch of three convex, naticoid whorls; form of
transition into teleoconch unknown; teleoconch of up to seven whorls; aperture
broad, only slightly constricted anteriorly, about two-thirds height; posterior
siphonal notch present; fasciole narrow, rugose and bounded adapically by a strong
ridge and abapically by a strong fold and corresponding with a groove on the external
shell surface; spiral ribbing on fasciole sometimes seen on juvenile shells; columella
bent to the left and terminating in a raised diagonal plait abapically; columellar
callus weak, often not quite obscuring ridge and fold of fasciole which may therefore
appear as weak columellar plaits; outer lip usually spirally ribbed within, internal
ribs unrelated to external ribs; growth lines wavy, essentially orthocline but bent
adorally in sutural region and aborally to meet ridge of fasciole; weak sinus (strom-
boid notch) present in abapical part of outer lip; flat subsutural platform present;
ramp concave; spiral ribbing fairly strong; collabral ribbing usually developed, with
a tendency to become spinose at periphery.
OTHER SPECIES ASSIGNED. Strepsidura armata J. de C. Sowerby, 1850; Buccinum
bullatum Philippi, 1847; B. bullatwm aspera von Koenen, 1889; Pseudocominella
semicostata nov. sp.; Cominella solandert Cossmann, 1889.
GEOLOGICAL RANGE. Middle Eocene (Lutetian) to Lower Oligocene (Lattorfian).
190 SOME ENGLISH
REMARKS. Vokes (1939 : 141) suggested that ‘Cominella’ deserta might be
congeneric with a group of species from the United States west coast Eocene,
usually referred to Siphonalia (fam. Buccinidae). S. bicavinata Dickerson, in fact,
strongly resembles both P. avmata and P. semicostata. Its familial position is obscure.
Like Recent Szphonalia, it possesses a very weak terminal columellar plait and
reverse S-shaped growth lines on the fasciole. It is definitely not a Pseudocominella,
which has a strongly grooved fasciole and an immediately obvious columellar plait.
Some species of Nassarius (s.l.) have fairly similar apices and sculpture to Pseudo-
cominella. They differ in often having a constrict:on of the neck above the fasciole
whilst the fasciole itself may correspond with a convex fold which is often strongly
spirally ribbed. The fasciole of Strepsidura (pl. 9, fig. 10) is similar to that of
Pseudocominella but the former genus may be distinguished by its more twisted
columella and by its aperture which is approximately four-fifths shell-height.
In addition, its protoconch is larger and rather flattened apically. Pseudocominella
differs from Comuinella in the form of the fasciole and the lack of a columellar plait;
the two genera are not co-familial.
Pseudocominella deserta (Solander, 1766)
(Pl. 1, figs 1-12)
1766 Buccinum desertum Solander: 13 (pars), pl. 1, fig. 15 (non figs 18, 19)
1823 Buccinum desevtum Brander; J. de C. Sowerby: 14, pl. 415, fig. 1.
1850 Fusus desevtus Morris; d’Orbigny: 363.
1854 Buccinum excavatum Beyrich: 444, pl. to (7), figs ta, 1b, tc.
1864 Stvepsidura deserta (Solander) von Koenen: 100.
1866 Buccinum (Cominella) desertum Lowry et al., pl. 3.
1880 Cominella (Buccinum) deserta (Solander) Judd: 154.
1881 Cominella (Buccinum) deserta (Solander); Keeping & Tawney: 116.
1888 Buccinum desertum Solander; Gardner, Keeping & Monckton: 626.
1889 Buccinum desevtum Brander; Bristow et al.: 289.
1889 Buccinum (Cominella) desertum Sowerby; von Koenen: 241 (pars).
1889 Cominella desevta (Solander); Cossmann: 136 (pars).
1891 Coméinella desevta (Solander); Newton: 168.
1901 Comunella deserta (Solander); Cossmann: 149, 150 (pars).
1901 Coménella deservta (Solander); Cossmann, pl. 6, fig. 4.
1933 Comunella desevta (Solander); St. J. Burton: 156.
1963 Comunella deserta (Solander); Glibert: 67 (pars).
1968 Cominella deserta (Solander); Martini & Ritzkowski: 244-247.
LECTOTYPE (designated herein). Upper Eocene, early Priabonian, Barton-on-
Sea, Hants, figured Solander, 1766, pl. 1, fig. 15, GG 19667 (Brander Colln.).
PARALECTOTYPE (designated herein). GG 19668, same details, but not figured by
Solander.
MATERIAL STUDIED. ENGLAND, Upper Eocene, early Priabonian, Barton Beds,
coastal section between Highcliffe and Barton-on-Sea, Hants. Horizon not
indicated: GG 19669/1 (figd. J. de C. Sowerby 1823, pl. 415, fig. 1, the central
figure); GG 19670/1-25 (Ed). Lower Barton Beds, horizon Az, GG 19671 (Ru).
PALAEOGENE NASSARIIDAE 191
Horizon A3, GG 19672/1-3 (Le); GG 19673 (S); GG 19674/1-13 (Wr). Horizon B,
GG 19675 (Ru). Middle Barton Beds, GG 19676/1-g (St/B); GG 19676/1-6 (Wr).
Horizon C, (Je); (S). Horizon E, (Cu); (S). Upper Barton Beds, Horizon H,
GG 19678/1-3 (Wr); GGu1151 (H. A. Toombs); GG 19725 (Cu). Lower Barton
Beds, Alum Bay, I.0.W., Sand. Mus. 3554 (J. F. Jackson Colln.); same (near top),
GG 19732 (Wr). Upper Eocene, late Priabonian, Middle Headon Beds, Brocken-
hurst, GG 19679 (Ed). Brockenhurst Bed, Whitecliff Bay, I.0.W., Sedg. Mus.
C 29122 (K. & T.). GERMANY, Upper Eocene, early Priabonian, Westeregeln,
GG 19680/1-3 (Ed).
FURTHER OCCURRENCES. GERMANY, Westeregeln, (Beyrich, 1854 : 444; von
Koenen, 1869 : 241; Glibert, 1963 : 67; Martini & Ritzkowski, 1968 : 244-247, with
nannoflora indicating a similar age to that of the Upper Barton Beds (Bed H).
Diaenosis. Pseudocominella up to 36 mm high; up to seven whorls of teleoconch;
mean spire angle 55°—60° ; prominent subsutural platform and concave ramp develop-
ed; spiral sculpture of narrow ridges often alternately of primary and secondary
strength and separated by wide interspaces ;collabral ribbing variable in strength and
often rather irregular; outer lip ribbed internally; columellar callus developed.
Description. Each whorl is enveloped by the succeeding one just below the
periphery. A prominent subsutural platform is developed on the third whorl
(first of teleoconch) and reaches a width of 1-5 mm on specimens over 30 mm high.
It is normally coronate, being crossed by the axial ribs; in axially multirugose
specimens, however, it is often flat on later whorls. It is bordered by a collar
bearing on later whorls up to three wavy spiral ribs (two on both the lectotype and
paralectotype). The platform is undercut, the sutural ramp being noticeably
concave. The whorl shoulder is distinctly adapical to the periphery. On the
first and second whorl of the teleoconch one spiral rib marks the margin of the
subsutural platform, another the shoulder, and three or four more lie on the rather
flat abapical portion of the whorl. These ribs are broad, rather flattened in section,
and are separated by narrow grooves. On the third whorl the interspaces broaden
and spiral ribs of secondary strength may appear, first on the sutural ramp, then
between the primary rib marking the shoulder and the one immediately abapical to
it. On later whorls five or six primary spiral ribs of intermediate strength may
be distinguished on the increasingly convexly-sided portion of the whorl abapical to
the shoulder. These ribs are narrow and are separated by interspaces about double
their width. Secondary and sometimes tertiary ribs are often developed with no
discernible pattern. On the body whorl between fifteen and twenty-two primary
ribs are developed between the shoulder and the adapical margin of the fasciole; up
to twenty-five ribs of secondary and tertiary strength may also be present in the same
area. On the lectotype there are fifteen primary and fifteen secondary ribs in this
region. In the same region there are thirty-nine ribs on the paralectotype but it is
difficult to separate them into different orders of strength. The first whorl of the
teleoconch bears about thirteen collabral ribs which are well-rounded in section.
On the next two whorls the number increases to between fifteen and twenty-five
and thereafter remains fairly constant on all succeeding whorls. The lectotype is a
192 SOME ENGLISH
particularly multicostate specimen (see table). The spacing and strength of the
collabral ribs often varies on a particular shell as well as between different ones.
Their strength is normally inversely proportionate to their number. They are
strongest at the shoulder which is often slightly nodose. The ribs are usually
weaker on the subsutural ramp but stronger on the subsutural platform. They
decrease in strength below the periphery. A variable number of internal spiral
ribs occur on the inner surface of the outer lip. They are rather irregularly arranged,
stretch back into the aperture for varying distances, and some are discontinuous.
DIMENSIONS I 2 3 4 5 7 8 9 10 II
GG 19667 LT 26°8 14:1 15°6 6+ 25 30 9 23 I-90 1-72
GG 19668 PLT 22 13°4 14'1 6 Ir+ 39 13 19 1-64 1°56
GG 19671 8:8 533 5:0 44 12 23 6 15 1-66 1-76
GG 19672/1 12:6 7-4 TPP 5 20 29 II 16 I-70 1°75
GG 19673/1 17-0 1073 9°4 54 ° 36 II 14 1°66 1-82
GG 19677/11 22°9 12-7 13:2 6 15 36 9 21 1°80 1-74
GG 19677/1 30°9 15 16°5 64 14 26 II 16 2:06 1-87
GG 19670/5 341 19:2 19:64 97, I5+ 32 Eh) <= ~ ae 7s
Key (also used in all following descriptions)
HT, Holotype; LT, Lectotype; NT, Neotype; PLT, paralectotype; PT, paratype.
1, total height (mm)
2, height of aperture
3, breadth
4, number of teleoconch whorls
5, number of collabral ribs on last whorl
6, number of collabral ribs on penultimate whorl (not shown for this species)
7, number of spiral ribs on last whorl
8, number of spiral ribs on penultimate whorl
g, number of spiral ribs inside outer lip
10, height: height of aperture
11, height: breadth
REMARKS. Buccinum fusiforme Deshayes, from the Lutetian and Auversian of the
Paris Basin, was placed in the synonymy of this species by Cossmann (1889 : 136).
A re-examination of Deshayes’ specimens in the Ecole des Mines (see p. 195) shows
that they all clearly belong to P. armata. All subsequent continental authors,
however, have accepted Cossmann’s views. In the synonymy given here it is
assumed that their records of French specimens are of P. aymata and not of P.
deserta, and that P. deserta does not occur in France.
From examining the lectotype and paralectotype it would appear that other
authors have correctly interpreted English specimens of the species. The other
specimens figured by Solander (pl. 1, figs 18, 19) as Buccinum desertum are both lost.
J. de C. Sowerby (1823, pl. 415, fig. 2) thought that Solander’s fig. 18, and possibly
fig. 19 belonged to his new species Buccinum canaliculatum (now Bartonia see p. 181);
these determinations have since been disputed. Cossmann (1889 : 137) identified
Solander’s figure 18 as Cominella deserta var. solanderi (Edwards MS). D. Curry
(personal communication) points out that Solander’s fig. 19 is much more likely to
be of Buccinum lavatum J. de C Sowerby (now Pollia).
PALAEOGENE NASSARIIDAE 193
In the adult stage P. deserta most clearly resembles P. bullata (Philippi) (pl. 2,
figs ga, b) from the Lower Oligocene of Latdorf, Germany. The whorl profile of the
latter is more evenly convex, the subsuturalramp being barely grooved. Itssculpture,
particularly the collabral elements, is weaker, and the first four whorls of the teleo-
conch are much smoother. In some specimens the collabral ribs on the last body
whorl are relatively strong in the peripheral region where they form definite nodes.
The spiral ribs on the inner surface of the outer lip are more numerous. Comparisons
with P. armata and P. solanderi are discussed under the latter named.
The records by Morris (1843 : 146) from Bracklesham (as Fusus desertus Brander),
by Forbes (1856 : 86) from the Headon Beds of Colwell Bay and by Nyst (1836 : 36)
from the Tongrian of Klein Spauwen (Belgium) cannot be confirmed and are almost
certainly incorrect. Bristow ef al. (1889 : 289) erroneously state that Keeping &
Tawney (1881) record this species from the Bracklesham Beds of the Isle of Wight.
This record, in fact, is from the Brockenhurst Bed of the Headon Series (see ‘Material
Studied’).
Giebel (1864 : 17, pl 1, fig 1) described and figured Buccinum bullatum Philippi
from Latdorf. Von Koenen (1865 : 466) redetermined this as Strepsidura deserta
(Solander) From the figure it is clear that Giebel’s determination was right and
von Koenen’s was wrong. In his more authoritative later publication (1889 :
241) von Koenen does not record P. deserta from Latdorf.
Pseudocominella armata (J. de C. Sowerby, 1850)
(Pl. 3, figs 1-8)
1835 Buccinum fusiforme Deshayes: 653.
1837 Buccinum fusiforme Deshayes; Deshayes, pl. 87, figs 15, 16, 17.
1850 Buccinum fusiforme Deshayes: d’Orbigny: 420.
1850 Stvepsiduva avmata J. de C. Sowerby in Dixon: 104, 186, pl. 7, fig. 11.
1865 Buccinum fusiopsis Deshayes: 499.
1889 Comunella deservta (Solander) Cossmann: 136 (pars).
1891 Strepsidura avmata J. de C. Sowerby; Newton: 165 (pars).
1901 Comunella deserta (Solander); Cossmann: 149, 150, (pars).
1911 Cominella deseyta (Solander); Cossmann & Pissarro, pl. 37, fig. 178-1.
1948 Cominella deserta (Solander); Morellet, L. & J.: 30, 133, 179, 331, 343-
1963 Cominella desevta (Solander); Glibert: 67 (pars).
NEOTYPE (proposed herein). Middle Eocene, Auversian, exact horizon unknown,
(probably Bed 19 (Brook Bed) of Fisher 1862 : 74) Upper Bracklesham Beds,
Bracklesham Bay, Sussex, GG 19681/1 (Ed).
MATERIAL STUDIED. ENGLAND, All from Upper Bracklesham Beds, Auversian.
Bracklesham Bay, GG 19681/2~-4 (Ed) (probably same horizon as Neotype) ; G 8869/1—
6 (Ogle Colln.) ; Bed 19 (Brook Bed) of Fisher (1862 : 74), Sedg. Mus. C 64860 (Fisher
Colin.) ; G 75897, G 75998 (E. M. Venables Colin.). Brook, Hampshire GG 19683/
1-4 (Ed); Sedg. Mus. C 67793. Huntingbridge, Hampshire, GG 19684 (Ed).
Bramshaw, Hampshire, GG 19685/1-4 (Ed). ‘Fossil bed’ at Bramshaw of Fisher
(1862 : 81, lines 2-4), 1 juvenile (Cu). FRANCE, Two separate samples in Deshayes
B
194 SOME ENGLISH
Colln., Ecole des Mines, Paris (see p. 195). One labelled Le Fayel, Mary and
Caumont, the other labelled Vendrest, Auvers and Acy, with no indication as to
which shell comes from which locality. All these localities were quoted by Deshayes
(1865 : 499), none of them were mentioned by him in his original description (1835 :
653-654).
FURTHER OCCURRENCES. FRANCE, Lutetian:—Grignon (Deshayes, 1835 : 654),
(Cossmann 1889 : 137). Auversian: Valmondois (Deshayes 1835 : 654) (1865 :
499); Le Guépelle (Deshayes, 1865 : 499); Bezu-le-Guery, Le Fayel (Glibert, 1963 :
67); Auvers (p. 133), Nanteuil-le Haudouin (p. 179), Acy (p. 331), Ru d’Alland valley
(p. 343) Morellet (1948) ; Acy (Cossmann & Pissarro, ro1I, pl. 37, fig. 178-1). Exact
horizon unknown. Senlis (Deshayes, 1835 : 654).
Diacnosis. Pseudocominella with teleoconch of up to six whorls; mean spire
angle between 60° and 65°; up to twenty-five collabral ribs per whorl on early part
of teleoconch, reducing to about ten increasingly nodose ribs on each of the last
two to three whorls; whorl profile convex on early whorls, shoulder becoming
increasingly angular on later whorls; spiral sculpture of varying strength; margins
of fasciole strongly ridged; fasciole very rugose; outer lip denticulate within in some
individuals.
DESCRIPTION. All the specimens are damaged and those from the type locality
are partly crushed. The protoconch (pl. 3, fig. 4) is typical for the genus and is
fairly well-preserved on specimens from Brook and Bramshaw. It consists of about
three apparently smooth naticoid whorls. The subsutural platform and close-set
collabral ribbing first appear towards the end of the third whorl. On the first three
or four whorls of the teleoconch the subsutural platform is bordered by a raised
rim which eventually becomes obsolete, finally, on the body whorl the platform
tends to be downward sloping. The early whorls of the teleoconch are broadest
at the adapical suture and are evenly convex in profile except for a groove under the
subsutural platform. In later whorls this groove develops into a broad concave
ramp. The profile is further modified by the development of nodes at the shoulder,
below which the whorl-sides are vertical.
The two strongest spiral ribs on the early teleoconch mark the edge of the sub-
sutural platform and the shoulder. On later whorls there are normally two strong
ribs in each of these positions. They are joined by others on the ramp itself and
increase in number to between seven and fifteen on the ramp of the last whorl. The
spiral sculpture differs considerably on specimens from different localities. The
patterns of ribbing on the ramp of shells from Bracklesham Bay, Whitecliff Bay and
Brook are similar; Bramshaw and Huntingbridge shells differ in having much
stronger ribs which may be differentiated into those of primary and secondary
strength. Below the shoulder on the peripheral region, up to ten weak spiral
ribs may be seen with difficulty on the spire whorls of specimens from Bracklesham
Bay. These usually die out on the body whorl of Bracklesham specimens (including
the neotype) but always persist on specimens from other localities. The ribs are
barely stronger on the Whitecliff Bay and Brook specimens. The Bramshaw and
Huntingbridge shells have over forty considerably stronger and more clear-cut ribs of
PALAEOGENE NASSARIIDAE 195
both primary and secondary strength on the body whorl. The neck region of
specimens from all localities bears between eleven and fifteen spiral ribs.
There are between twenty and twenty-five collabral ribs on each of the first
two whorls of the teleoconch. These become more irregular in strength and spacing
on the next whorls, reducing eventually in number to about nine evenly spaced
nodes on the fifth and sixth whorl of the teleoconch. On the earlier whorls they are
well-rounded in section, separated by rather broader, smoothly concave interspaces.
They are moderately strong and of even strength below the shoulder; above it they
markedly decrease in strength and are almost obsolete at the sub-sutural ramp. On
the body whorl the collabral sculpture is confined to the area just above and below
the shoulder. It consists of strong nodes or sometimes blunt spines. Six of the
twelve specimens from the type locality have, on the last whorl, indications of
a growth halt during which an aperture similar to the terminal one was developed.
Here, the growth lines are more prominent, the outer lip was apparently slightly
flared and parietal callus remains above the adapical suture of the previous whorl.
One specimen has two such halts and on two of the larger shells these halts occur
about 180° before the terminal aperture. Signs of preterminal apertures are common
but much less prominent on specimens from other localities. The track of the
fasciole is rather deeply grooved, and its bordering ridges, particularly the anterior,
are strong. Growth rugae in this region are noticeably imbricate especially in the
larger shells. The aperture is damaged in all specimens; none of those from
the type locality and only one out of the four from Brook shows any indications of the
outer lip being denticulate within. Internal ribs stretch back for about 45° within
the aperture in specimens from other localities. In one specimen from Bramshaw
these break down into numerous intermittent dashes as the aperture is approached’
DIMENSIONS. I 2 3 4 5 6 9 Io II
GG 19681/1 NT 25°5 15'7 16+ 6 9 Io — 1:6 1°57
GG 19681/2 22+ — I5"1 6 be) 10 — — 1°47
GG 19681/3 22 13 I2°9 6 9 12 — 1-69 1-70
GG 19681/4 17+ Ti+ 1073 6 8 ae) — — _
GG 19685/4 12 7 65 5+ 13 18 — I-71 1°84
GG 19685/3 16 9:2 9'7 5+ 12 19 12 1°74 1-65
GG 19685/1 26°7 16:2 16:7 6+ 9 9 15 1°65 1:60
GG 19685/2 28+ 16°5 17'7 6+ 10 9 some I-70 1°58
GG 19684 15°5 933 9°33 6 9 12 13 1-67 1-67
(see p. 192 for key)
ReMARKS. The appearance and matrix of the neotype and associated specimens
GG 19681/1-6 (Ed) matches that of the specimens listed which are known to come
from the Bed 19 (The Brook Bed) at Bracklesham Bay.
All the French records of ‘Cominella’ deserta are considered herein to be of P.
armata (see pp. 192 and 194). Specimens of Buccinum fusiopsis in the Deshayes
Collection (EMP) (no specimens from Deshayes original (1835 : 653-654) localities
survive) and the specimen from Acy figured as Cominella deserta by Cossmann and
Pissarro (1911, pl. 37, fig. 178-1) have strong sculpture very similar in character to
196 SOME ENGLISH
that of specimens from Bramshaw and Huntingbridge in the New Forest (Hampshire).
The specimen which Cossmann figures (1go1, pl. 6, fig. 4) as Cominella deserta is
stated to be from Barton and the specific identification appears to be correct. All
the English specimens of P. avmata have stronger nodes at the periphery of the last
whorl than any of the French specimens examined; the closest comparison is between
them and the specimen figured from Huntingbridge (cf. pl. 3, figs 6 and 8). P.
deservta always has a more rounded whorl profile and a narrower, more concave
ramp. The only previous figure of an English specimen of the species is of the
relatively smooth form originally described by Sowerby from Bracklesham Bay
(1850, pl. 7, fig. 11). Without knowledge of the Bramshaw and Huntingbridge speci-
mens, it is hardly surprising that French authorities considered their specimens
either to be specifically distinct from P. armata or to be synonymous with P. deserta.
The exposures at Bramshaw and Huntingbridge are slightly younger than the
Brook Bed (Bed 19) of Brook and of Bracklesham Bay (Fisher, 1862 : 92-93;
Curry 1958: 70-71). It seems likely that P. armata gave rise to P. deserta.
Changes include an increase in strength of spiral sculpture, an increase in the
number of collabral ribs on the later whorls, gradual disappearance of peripheral
nodes and the narrowing of the ramp.
Deshayes proposed the name Buccinum fusiopsis (1865 : 499) as his earlier
name B. fusiforme (1835 : 653) was preoccupied (Borson, 1820 : 222). Sowerby’s
name (1850 : 104, 186) therefore stands. The surviving parts of both Dixon’s and
the Sowerby family’s collections are housed in the British Museum (Natural History),
but the whereabouts of this figured specimen is unknown. Comparisons with P.
deserta and P. solanderi follow the description of the latter, whilst those with P.
bullata (Philippi) and P. semicostata are discussed under the last named. The name
armata has often been applied erroneously to P. semicostata (see its synonymy). The
record from the Belgian Oligocene by Glibert & de Heinzelin (1954 : 367) of Northia
(Cominella) bullata {. armata is of P. bullata.
Pseudocominella semicostata sp. nov.
(Pl. 3, figs 9-10; Pl. 4, figs 1-3)
1864 Strepsidura deserta var. armata J. de C. Sowerby; von Koenen: roo.
1880 Strepsidura (Buccinum) armata J. de C. Sowerby; Judd: 154.
1880 Strepsidura (Buccinum) semicostata Edwards MS; Judd: 154 (nom. nud.).
1881 Strepsiduva (Buccinum) armata J. de C. Sowerby: Keeping & Tawney: III, 113, 116.
1881 Strepsiduva (Buccinum) semicostata Edwards MS; Keeping & Tawney: 116 (nom. nud.).
1882 Strepsiduva avmata J. de C. Sowerby; Judd: 477.
1891 Strepsiduva avmata J. de C. Sowerby: Newton (pars): 165.
1891 Strepsidura semicostata Edwards MS; Newton: 165 (nom. nud.).
1901 Strepsiduva arvmata J. de C. Sowerby; Cossmann: 133.
tgo1 Strepsidura semicostata Edwards MS; Cossmann: 133 (nom. nud.).
1902 Strepsiduva avmata J. de C. Sowerby; Reid: 4o.
1915 Strepsiduva avmata J. de C. Sowerby; White: 44.
1926 Melongena (Cornulina) minax (Solander) Jackson: 360 (pars).
1926 Strvepsiduva armata J. de C. Sowerby; Jackson: 364.
1960 Stvrepsiduva avmata J. de C. Sowerby; Glibert: 9.
PALAEOGENE NASSARIIDAE 197
Hototype. Upper Eocene, Late Priabonian, Middle Headon Beds, Whitley
Ridge, railway cutting, Brockenhurst, Hampshire GG 19687/1 (Ed ex K).
ParaTyPeEs. All Upper Eocene, Late Priabonian, Middle Headon Beds. Whitley
Ridge Railway cutting, Brockenhurst, Hampshire, GG 19687/2-3 (Ed ex kK);
Sedg. Mus. C 26453-8 (K). Victoria Tilery, Brockenhurst, (S). ‘Brockenhurst’,
exact locality unknown, GG 19688/1-14 (Ed); GG 19689/1-2 (St]B); G7760
(Beckles Colln.). Lyndhurst, Hampshire, GG 19690/1-4 (Ed). Royden, Hampshire
GG 19686/1-9 (Ed ex K); Sedg. Mus. C 25070-90 (KG&T). Brockenhurst Bed,
Whitecliff Bay, 1.0.W., GG 19692/1-5 (Wr); Sedg. Mus. C 29270-3 (KGT); Sand.
Mus. 1930 (J. F. Jackson), recorded as Melongena minax. ?Venus Bed, Colwell Bay,
1.0.W., GG r9691 (Ed).
FURTHER OCCURRENCES. Unknown.
Diacnosis. Stout Pseudocominella with between six and seven whorls of teleo-
conch; mean spiral angle increasing from 60° to 80° with growth; ramp broad;
shoulder angular except for early teleoconch, low, at or below mid-height of
spire whorls; about twenty-one collabral ribs on early part of teleoconch reducing
to nine or ten very nodose ribs on last three whorls; aperture broad; fasciole
bordered by strong ridges; fasciole very rugose; outer lip denticulate within in some
specimens.
DeEscriPTION. The protoconch is decorticated or missing in all specimens;
its gross size and shape is similar to that of P. deseyta (Solander). The first two to
three whorls of the teleoconch are broadest at the adapical suture and are convex
in profile except for a subsutural constriction which rapidly grows into a broad
concave ramp. The profile becomes strongly carinate on the fifth and sixth whorl
of the teleoconch, with the periphery coinciding with the shoulder. The subsutural
platform is horizontal, on later whorls it becomes wavy under the influence of the
axial nodes. At the earliest point in the development of the teleoconch at which
it is possible to determine detail there are three prominent spiral ribs, and the whorl
profile is naticoid. During this whorl, two new ribs develop between the two most
adapical. The opisthocyrt collabral costae are of similar strength and form a
reticulate pattern with slight nodes at the intersections; they do not extend on to
the subsutural platform which first develops on this whorl and is bordered by a
spiral rib. The following whorl (third of the teleoconch) is similar except for the
broadening of the subsutural ramp which lacks spiral ribbing. The shoulder
is marked by the second most adapical spiral rib. The reticulate pattern is made of
increasingly spirally-elongate rectangles. On the fourth teleoconch whorl the
increasing strength of the collabral ribs compared with the spiral ribs becomes
noticeable; within the next whorl or so this leads to the loss of the reticulate appear-
ance. Theramp gradually becomes concave and bears two new spiral ribs of secondary
strength. The edge of the subsutural platform is marked by two strong spiral ribs
and another pair lies on the shoulder. Between the fifth and seventh teleoconch
whorls seven to ten spiral ribs are developed on the ramp. Apart from those on
the shoulder and bordering the subsutural platform these are seldom of greater
strength than those on earlier whorls and are often difficult to follow even on
198 SOME ENGLISH
well-preservedshells. On immature specimens with a teleoconch of four whorls, there
are on the body whorl between eighteen and twenty spiral ribs of varying strength
below the shoulder, seven or eight of these being on the neck and rather stronger.
The full adult number of spiral ribs, about twenty-nine in all, of which fourteen are
on the neck, is reached by the sixth teleoconch whorl. There are between eighteen
and twenty-one collabral ribs on the first four teleoconch whorls; these die away
below the peripheral region of the body whorl even in juvenile shells. On the next
whorl these weaken on the ramp and become increasingly nodose at the shoulder
and grow further apart. From here on nine or ten axial nodes or spines occur on each
whorl. The fasciole is strongly ridged and is very rugose in large specimens.
Specimens showing ribs on the inside of the outer lip are very rare.
DIMENSIONS. I 2 3 4 5 6 7 8 9 Io II
GG 19687/1 HT
estimated 27°0 — — —- —- — — — — 1-86 1°52
actual 26°1 14°5 17°8 6 13 12 — — 16 — —
GG 19686/2 PT 2053 12°8 13'7 53 9 II 36 12 10 1°58 1-48
GG 19686/3 PT 13°8 8-5 8-5 5 12 21 33 10 fo) 1-62 1°62
GG 19686/4 PT 8-7 595 5:2 44 16 20 23 7 ria 1°58 1:67
GG 19686/5 PT 74 4°7 4°3 =n REDS ED 5 O° E57 ear?
4
GG 19688/1 PT 32+ 19°5 21°1 64 8 12 42
(see p. 192 for key)
REMARKS. Edwards gave the manuscript name Strepsidura semicostata to his
single specimen from Colwell Bay (GG 19691). This was so badly worn that he
failed to recognize that it was conspecific with numerous other specimens (listed
here) which he identified as S. armata. P. semicostata differs from P. armata in being
relatively stouter, having a broader, less sloping ramp, more prominent axial nodes,
and a lower shoulder. It has also been confused by von Koenen (1864 : 100) and by
Judd (1880 : 154) with P. bullata (Philippi) which may be easily distinguished by its
much weaker collabral sculpture on the early teleoconch; strong axial ribbing and
nodes are normally confined to its body whorl.
Pseudocominella solanderi (Cossmann, 1889)
(Pl. 2, figs 1-8)
1766 Buccinum desertum Solander: 13 (pars), pl. 1, fig. 18 (non figs 15, 19).
1887 Cominella solandvi Edwards; Keeping: 71 (nom. nud.).
1888 Buccinum solandvi Gardner, Keeping & Monckton: 620 (nom. nud.).
1888 Buccinum sp. nov.; Gardner, Keeping & Monckton: 627.
1889 Cominella solandvi Edwards; Bristow: 120, 290 (nom. nud.).
1889 Cominella deserta (Solander) var. solandevi Edwards; Cossmann: 137.
1901 Cominella (Buccinum) solanderi (Edwards) Cossmann: 150.
1926 Cominella solandvi Edwards MS; Jackson: 356.
1933 Cominella sp.; Burton: 156.
PALAEOGENE NASSARIIDAE 199
NEOTYPE (selected herein). Upper Eocene, early Priabonian, Lower Barton Beds,
Horizon A3, coastal section between Highcliffe and Barton, Hampshire, GG 19693/1
(Ho).
MATERIAL STUDIED. Same locality, Lower Barton Beds, horizon unspecified
GG 19696/1-20 (Ed); 19697/1-10 (Wr). Horizon Az, GG 19703 (Ru). Horizon Az
(near top) GG 19698/1-20 (Nu). Horizon A3, GG 19699/1-50 (S); GG 19700/1-3
(StJB); GG 19693/2-3 (Ho); GG 19694/1-18 (Le); GG 19695/1-3 (S); (Cu); (Je).
Horizon B, 2 specimens (Ru).
Alum Bay, I.0.W., Barton Beds, horizon unknown, Sedg. Mus. C 59712-4; (10
specimens) Sand. Mus. 4720. Lower Barton Beds (near top) GG 19731/1-8 (W7);
(2 specimens) Sand. Mus. 1084 (J. F. Jackson Colln.). Whitecliff Bay, I.0.W.
Barton Beds, lowest 13 inches with Nummulites elegans, Sedg. Mus. C 64008 (K).
FURTHER OCCURRENCES. Lower Barton Beds, Barton-on-Sea, as Cominella sp.,
horizons A3 and B (Burton, 1933 : 156).
Dracnosis. Moderate sized, fairly elongate Pseuwdocominella with a marked
horizontal and undercut subsutural platform; mean spire angle between 50° and
60°; collabral ribs present only on early whorls of neoconch; spiral sculpture of close-
spaced low ribs; outer lip ribbed internally.
DeEscRIPTION. The protoconch is about 1-2 mm high and broad. Its first
half whorl is minute; it seems to be disjunct with a semi-erect tip, but this appearance
is certainly due to weathering. It is followed by two and a half smooth naticoid
whorls. It seems very unlikely that the transition to the teleoconch was abrupt;
orthocline collabral ribs of increasing strength occur, these soon become nodose
and are joined by spiral ribbing within a quarter whorl. Most adult specimens are
badly worn; the sculpture of the body whorl is abraded and the early whorls either
decorticated or even missing, the main exceptions are immature specimens under
8 mm high. Teleoconchs of shells 20 mm high would have been of six whorls.
Each whorl is enveloped by the succeeding one just below the periphery. The first
two-and-a-half to three teleoconch whorls bear twelve, increasing to sixteen rather
angular, orthocline, collabral ribs. These die out by a shell height of between 6 and
8 mm, except on a very few of the specimens seen, where they persist above the
periphery sometimes until full growth is reached. On succeeding whorls the only
collabral sculptural elements are rugae which are distinctly stronger than the
growth lines seen on well preserved shells. Spiral ribbing and the subsutural
platform appear about a quarter whorl after the first collabral ribs. The subsutural
platform, which reaches a width of r mm on the last whorl of large shells is normally
horizontal but may slope either upwards or downwards away from the suture.
Another fairly strong spiral rib marks the shoulder, which is nodose and carinate
for the first two or three whorls of the teleconch. Within the next two whorls
these are joined by a weaker spiral rib on the concavely-sloped sutural ramp and
three or four more between the shoulder and the suture below. By the fifth teleo-
conch whorl the latter have increased in number to eight or nine. Two spiral ribs of
lower strength commonly occur on the sutural ramp. On the last whorl other
weak ribs are sometimes irregularly disposed, mainly above the periphery. The
200 SOME ENGLISH
number of spiral ribs on the last whorl increases with size (see table), most adult
shells having between thirty-eight and forty ribs of varying strength. The nine or
ten on the neck region are the strongest, apart from the primary ribs on the shoulder
and on the edge of the subsutural platform. The spiral ribs are about the same width
as the interspaces except on the neck region where they are more widely spaced.
In section they are usually gently convex and often almost flat-topped. The inner
surface of the outer lip bears ribs which do not correspond with the external ones.
They stretch back inside the aperture for nearly a quarter whorl in large individuals,
less so in small ones and are resorbed as the shell grows. The number of internal
ribs is about half that found on the outer surface and is dependent on size. Two
spiral colour-bands may be seen on the neotype, one, about a millimetre broad, on
the ramp with a narrower one just below the shoulder, other specimens also show this
phenomenon (pl. 2, figs 4, 6, 7, 9).
DIMENSIONS. I 2 3 4 5 6 7 8 9 10 II
GG 19693/1 NT 14°7 76 7°8 53 fo) ° 28 10 18 1°93 1-89
GG 19695/3 52 370 Oi = 953 19 614) «17 5) 13. 178 9
GG 19694/17 6:2 35 35 4 9 16 24 7, Cees
GG 19693/2 8-6 4°7 4°5 44 fe) 14 25 Io Il 1°83 I-QI
GG 19694/14 10°8 6-1 655 5 6 15 32+ II 14 72 1-66
GG 19694/10 12-6 74 6:8 5 fo) fo) 38+ II 17 1-70 I-61
GG 19694/6 14°6 8-2 8-2 5 4 15 41 14 17 1°78 1-78
GG 19694/5 17°8 9°5 92 58 ) © 35 19 10 =:8q ao
GG 19694/1 20:0 12:0 Ten 6 fo) 15 41 16 ze) 1-67 I-80
GG 19700/3 25°5 13°5 13°8 — fo) — -—— — 20 1:89 1°85
(see p. 192 for key)
REMARKS. Cossmann (1889 : 137) in discussing P. deserta wrote ‘les figures qu’en
donne Brander sont tres exactes, sauf la figure 18, qui représente une variété pour
laquelle a été proposé le nom Solanderi, Edw.’. The specimen figured by Solander
is lost. None of the specimens GG 19696/1—20 in the Edwards Collection, which are
labelled as solandn, have been chosen as neotype as they are all badly worn. In the
circumstances it seems justifiable to select a neotype from better preserved material
whose horizon is known more precisely.
This species is unknown except from the Lower Barton Beds. One sample,
GG 19701/1—-4 (Wr) was labelled as being from the Middle Barton Beds. Examina-
tion by several workers suggested that the specimens did in fact, come from the
Lower Barton Beds. Moreover glue was present on the specimens, unlike others in
Wrigley’s Collection, suggesting that he might have acquired the specimens from
an old collection and had assumed that any information with the specimens was
correct. Another specimen, GG 19702 (Wr) was found in the same box as three
specimens of P. deserta, GG 19678/1-3 (Wr) from the Upper Barton Chama Bed
(Bed H). Again, examination by several workers showed that the preservation and
matrix of the specimen of P. solanderi was not the same as that of its companions
and that it was almost certainly from the Lower Barton Beds.
This species is among the twenty most common prosobranchs in horizon A3,
of the Lower Barton Beds where P. deserta is extremely rare. The latter species,
PALAEOGENE NASSARIIDAE 201
however, is more common in the Middle and Upper Barton Beds in which P. solanderi
is unknown. Both species usually attain a height of 20 mm in Bed A3 of the Lower
Barton Beds. In the Middle Barton Beds, P. deserta often exceeds 30 mm in
height. This increase in size corresponds with almost an extra whorl (making a
total of seven for the teleoconch).
P. deserta is relatively broader and its sculpture is stronger. The spiral sculpture
is the only completely consistent difference between the two species. In P. deserta
the interspaces are wider and the ribs are usually alternately of primary and
secondary strength. A small proportion of specimens of one species may show one
or more of the features listed below which normally distinguish it from the other.
In P. deserta the collabral ribs tend to increase in strength with size, whilst they be-
come obsolete by the third whorl in all but a few P. solanderi. The subsutural
platform is broader and more strongly grooved below in P. deseryta and the carina
on the shoulder persists to adulthood in most individuals.
In all four of the British species of Psewdocominella, the early teleoconch has such
similar sculpture, shape and rate of whorl expansion that juveniles are difficult
to tell apart. The nodiferous, carinate, shoulder of this stage is most noticeable
and of shortest duration in P. solanderi which is the least strongly sculptured
when adult.
DESORINASSA gen. nov.
(Pl. 4, figs 4-9, Pl. 5, figs 1-5)
TYPE SPECIES. Buccinum desor Deshayes, 1865.
DracGnosis. Agrees with Pseudocominella except in the following respects:
subsutural platform not always developed; sculpture very weak; columella curved
to the right above columellar plait; growth lines of fasciole not particularly rugose
and not very sinuous; ridge above fasciole not strong; outer lip smooth within.
OTHER SPECIES ASSIGNED. *Buccinum acies Watelet, 1853 (I[RScNB) ; **Cominella
bonnecarret Furon 1948; *Buccinum latum Deshayes, 1865 (EMP); Buccinum ovatum
Deshayes 1835 (EMP, MNP). **Cominella supracostata Traub, 1938; *Desorinassa
williamst sp. nov. (see footnote for explanation).
GEOLOGICAL RANGE. Palaeocene (Thanetian) to Lower Eocene (Cuisian).
REMARKS. Desorinassa may be distinguished from Pseudocominella by its curved
columella and its weaker sculpture, in particular the almost complete lack of collabral
elements. The fasciole is more like that of Keepingia and Thanetinassa.
The specimens of Buccinum latum Deshayes, (Deshayes Colln., EMP) figured
herein as pl. 4, fig. 8 and pl. 4, fig. 9 are selected respectively as LEcToTYPE, and
PARALECTOTYPE of this species. They are labelled as being from Abbecourt and
Bracheux with no indication as to which specimen comes from which locality.
These are the only localities quoted by Deshayes in his original description (1865 :
In these lists * denotes species in which the protoconch has not been studied, ** denotes that no
specimens have been examined.
202 SOME ENGLISH
501). It would appear that his illustration of the apertural view of the species
(1866, pl. 93, fig. 27) is composed of the outline of the lectotype upon which has been
superimposed the better preserved outer lip and stronger columellar and parietal
callus of the paralectotype.
Desorinassa desori (Deshayes, 1865)
(Pl. 4, figs 4-6)
1865 Buccinum desori Deshayes; 501.
1866 Buccinum desort Deshayes; Deshayes, pl. 93, figs 16, 17.
1889 Cominella desovi (Deshayes) Cossmann: 137.
1901 Cominella desovi (Deshayes); Cossmann: 150.
1911 Cominella desovi (Deshayes); Cossmann & Pissaro, pl. 37, fig. 178-3.
1934 Comznella desov1 (Deshayes); Cooper: 8.
1936 Cominella desort (Deshayes); Farchad: 22.
1963 Cominella desovi (Deshayes); Glibert: 67.
MATERIAL STUDIED. ENGLAND, Palaeocene, Thanetian, Thanet Sand, Reculver,
Kent GG 19704 (Wr) (see remarks p. 203). FRANCE, Thanetian, Jonchery-sur-
Vesle, France, GG 19705/1-3 (Wr); G 19438/1, 2 (L. Staadt).
FURTHER OCCURRENCES. ENGLAND, Thanetian, Bishopstone, near Reculver,
(Cooper, 1934 : 8) (see remarks, p. 203). FRANCE, ‘Sables inferieurs’ (= Thanetian),
Abbecourt; Chalons-sur-Vesle; Jonchery; Saint Martin-aux-Bois (Deshayes, 1865 :
501). Chalons; Chenay; Jonchery; Noailles (Glibert 1963 : 67). Bracheux, near
Beauvais (Farchad, 1926 : 22).
DiaGnosis. Moderately large, stout, almost smooth Desovinassa with broad,
sloping subsutural platform; mean spire angle about 60°; sutural slope reversed just
prior to adult aperture; ridge on adapical side of fasciole very weak; outer lip not
striate within.
DESCRIPTION. The protoconch is worn but appears to have been similar to that of
Pseudocominella, consisting of about three smooth, naticoid whorls; initially it
may have been somewhat flattened. The teleoconch has up to six whorls which are
convex-sided with the periphery lying just above the suture. A cord may be seen
below the suture on the first whorl of the teleoconch. The material is not well
enough preserved to show the development of the subsutural platform. It is first
recognizable by the third whorl of the teleoconch where either two (or in some
specimens three) spiral ribs, situated high on the ramp near the suture, form an
angulation in the whorl profile. Subsequently the platform becomes flatter, though
the slope is never less than 45°. It reaches a breadth of nearly 1-5 mm. No groove
is developed but the profile is slightly concave just below the platform. Spiral
sculpture is poorly developed, there are two or three weak grooves just below the
subsutural platform and a mixture of about fifteen weak grooves and step-like ribs
on the neck region of the body whorl. Collabral sculpture is absent except for
growth rugae, these are strongest away from the periphery, becoming quite notice-
able on and just below the subsutural platform and to a lesser extent on the neck
PALAEOGENE NASSARIIDAE 203
region. The ridge marking the adapical margin of the fasciole can hardly be distin-
guished in smaller specimens. No ribbing has been seen inside the outer lip. The
sutural slope becomes reversed immediately prior to the final aperture.
DIMENSIONS. I 2 3 4 7 10 II
GG 19704 22+ 11+ 14+ — a= — —
GG 19705/3 II+3 6:5 63 4+ 12+ 1°74 1°79
GG 19705/1 1373 8-2 7:9 44 17+ 1-62 1:68
G 19438/2 16°5 99 9°7 = = 1-67 1°70
G 19438/1 17°6 10-0 10°6 5+ 18+ 1:76 1°66
(see p. 192 for explanation)
Remarks. D. desori differs from D. acies (Watelet) (pl. 5 fig. 5) and D. ovata
(Deshayes) (pl. 5, fig. 4) in being larger, having a subsutural platform and lacking
regular spiral sculpture. A comparison with D. williamsi is given under the latter.
The single English specimen of D. desort seen is badly damaged but agrees in all
essential respects with the French specimens.
It is thought that Wrigley acquired many of the more interesting specimens of
J. E. Cooper’s collection and GG 19704 was found in a box which was probably made
by Cooper. It is therefore possible that this specimen is that listed by Cooper (1934 :
8) and that only one specimen has so far been discovered in English rocks. However,
A. J. Rundle (personal communication) after examining its matrix reports that it is
a silicified specimen from the Corbula bed in the cliff at Reculver and not from the
morrisi bed, as listed by Cooper.
Desorinassa williamsi sp. nov.
(Pl. 5, figs 1-3)
HoLotyPe. Palaeocene, Ypresian, Bognor Rock, foreshore at Bognor, Sussex,
GG 19706 (B. A. Williams Colln.).
PARATYPES. Some horizon and locality, GG 19707, (Rev. A. Fuller Colln.), and
two internal moulds GG 19708/1-2 (Aistory unrecorded).
FURTHER MATERIAL AND RECORDS. Unknown.
DiacGnosis. Large, stout, almost smooth Desorinassa; mean spire angle of about
60°; subsutural platform lacking; whorl profile convex; interior of outer lip not
denticulate.
DESCRIPTION. The protoconch is unknown. Four whorls of the teleoconch
survive; at least one or two earlier whorls are missing in all specimens. The whorls,
which are broadest at the abapical suture, are convex except where recurved at the
adapical margin to form an adpressed or almost flush suture. Spiral sculptural
elements are lacking except for between six and eight weak ridges on the neck of the
body whorl. Irregularly spaced collabral growth rugae of varying strength are
developed. These, though weak, are distinctly stronger than growth lines. No
ribs are present on the inner surface of the outer lip. Both internal moulds have a
collabral groove about a quarter whorl before the aperture.
204 SOME ENGLISH
DIMENSIONS. I 2 3 7 10 II
GG 19706 HT 27°8+ 17°3 17:0 8+ — —
estimated 29 — — — 1-68 I-71
GG 19707 PT 27°2+ 17°2 179 5+ -- —
estimated 30 —_ — — 1-74 1:67
(see p. 192 for key)
REMARKS. This species may be easily distinguished from D. desort which is
smaller, and has a broad, sloping, subsutural platform with spiral ribbing on and
near it. Two Cuisian species from the Paris Basin, D. ovata (Deshayes) (pl. 5, fig. 5)
and D. acies (Watelet) (pl. 5, fig. 6) are much smaller than D. williamsi and possess
fine spiral ribbing. The former is of a similar shape to D. williamsi but the latter
is distinctly narrower.
WHITECLIFFIA gen. nov.
(Pl. 5, figs 6-12; Pl. 6)
TYPE SPECIES. Buccinum suturosum Nyst, 1836.
Diacnosis. Differs from Pseudocominella only in the following respects: height
less than 20 mm; protoconch of one and half whorls, initially deviated and bulbous;
suture either weakly canaliculate or with narrow, flat, subsutural platform; no
shoulder developed; collabral sculpture either absent or of weak grooves or ribs;
columellar and parietal callus distinctly raised above general level of shell surface;
fasciole similar to that of Pseudocominella but ridges and growth lines less prominent ;
outer lip seldom ribbed within.
OTHER SPECIES ASSIGNED. Whitecliffia twmida sp. nov.
GEOLOGICAL RANGE. Upper Eocene (Late Priabonian) to Middle Oligocene
(Rupelian).
REMARKS. This genus seems likely to be an offshoot of Pseudocominella. It may
readily be distinguished by its protoconch and by the relative smoothness of the
teleoconch.
Whitecliffia suturosa (Nyst, 1836)
(Pl. 5, figs 6-11; Pl. 6, figs 1-4)
1836 Buccinum suturosum Nyst: 36, pl. 3, fig. 96.
1843 Buccinum sutuyosum Nyst; Nyst: pl. 43, fig. 16.
1845 Buccinum suturosum Nyst; Nyst: 579.
1867 Buccinum suturosum Nyst; von Koenen: 83, pl. 6 (1), fig. 3a, b, c.
1880 Cominella (Buccinum) ventricosa Edwards MS; Judd: 154.
1880 Stvepsidura suturosa (Nyst) Bosquet: 467.
1881 Cominella (Buccinum) ventricosa Edwards MS; Keeping & Tawney: 116.
1886 Stvepsiduva suturosa (Bosquet); Vincent: 7.
1889 Buccinum (Cominella) suturosum Nyst; von Koenen: 242, pl. 21, figs 10a, b, c, Ifa, b.
1901 Cominella suturosum (Nyst); Cossmann: 150.
PALAEOGENE NASSARIIDAE 205
1943 Novthia (Cominella) suturosa (Nyst) Albrecht & Valk: 62, pl. 5, figs 140, 141.
1954 Northia (Cominella) suturosa (Nyst); Glibert & de Heinzelin: 367, 382, 392, 393, pl. 7,
figs 4a, b.
1954 Babylonia (Peridipsaccus) avchambaulti Meunier; Glibert & de Heinzelin: 368, 382, pl. 7,
fig. 6.
1963 Cominella suturosa (Nyst); Glibert: 68.
LectotyPE. Middle Oligocene, Lower Rupelian, Callista kickxi horizon; IRScNB
IG 3862, Klein Spauwen, Belgium (selected Glibert, 1963 : 68, figured Glibert,
& de Heinzelin 1954, pl. 7, fig. 4b).
PARALECTOTYPE. Same details, 7RScNB IG 3863, (figured Glibert & de Heinzelin,
1954, pl. 7, fig. 4a).
MATERIAL STUDIED. ENGLAND, Upper Eocene, late Priabonian, Middle Headon
Beds, Royden, GGuig9717 (Ed). Brockenhurst Bed, Whitecliff Bay, I.0.W.,
GG 19718 (Wr); (Cu). GERMANY, Lower Oligocene, Lattorfian, Latdorf, GG 19719/
1-2 (Ed); GG 19720/1-2 (history unknown). Brtcium, Lower Oligocene, Lattorfian
(= Lower Tongrian), Grimmertingen [RScNB sample IG 5002 (de Looz). Middle
Oligocene, Lower Rupelian, Klein Spauwen, GG 19721 (de Koninck). Berg, IRScNB
sample IG 4285 (Bosquet).
FURTHER OCCURRENCES. HOLLAND, Lower and Middle Oligocene, South Limburg,
in mine shafts (Albrecht & Valk, 1943 : 62). GERMANY, Middle Oligocene, Rupelian,
Stettin Sands, Magdeburg, (von Koenen, 1889 : 242).
Diacnosis. Whiteclifia with canaliculate suture; mean spire angle between 43°
and 80°; whorls evenly convex with no shoulder developed; sculpture absent except
for occasional weak spiral grooving; outer lip denticulate within.
Description. All the Belgian specimens and Wrigley’s specimen from Whitecliff
Bay are too badly worn to show sculptural details. The description is based mainly
on the shells from Royden (GG 19717), Latdorf (GG 19719—20) and a specimen from
Whitecliff Bay (Cz).
The mean spire angles of the lectotype, paralectotype and GG 19721 from Klein
Spauwen are all under 50°. The spires of several specimens from Berg are also
more acute than average. The English, Grimmertingen and Latdorf specimens are
normally more tumid with the spire angle between 50° and 55°. The appearance of
the suture is so considerably changed by wear, that this feature cannot be reliably
used as a taxonomic feature. The better-preserved English and German specimens
as well as most of the Belgian specimens have a narrow canaliculate suture and no
subsutural ramp. Many of the Grimmertingen specimens, however, seem to have a
fairly broad ramp and a less incised suture. In most specimens the sutures are
not channelled for the first three or four whorls.
Below the suture the whorl profile is rather gently convex, and its periphery
coincides with the abapical suture in most specimens. In shells with relatively
acute spires, however, whorls are enveloped well below the periphery. About half
the specimens have a relatively strong pad of callus in the parietal and columellar
region which stands out above the rest of the shell-surface. The fasciole is not
206 SOME ENGLISH
very rugose but the ridges on either side are fairly sharp when unworn. Only one
specimen, the largest in sample JRScNB 1G 5002 from Grimmertingen, has ribbing on
the inner surface of its outer lip; this feature is also shown in one of von Koenen’s
figures (1889, pl. 21, fig. rob). All the spire whorls of GG 19717 bear weak spiral
grooves; eight may be counted on the penultimate whorl, but they become obsolete
on the body whorl, which is smooth except for ten ribs on the neck. On another
larger specimen (Cz), spiral grooves which are too poorly preserved to count, are
present on the fourth and fifth whorls whilst the last two are smooth. GG 19719/1
and GG 19720/1, from Latdorf, however, have twelve grooves on the penultimate
whorl, twenty-four on the body whorl and thirteen ribs on the neck below. Another
specimen from Latdorf GG 19719/2 whose surface is also well preserved shows traces
of spiral sculpture only on the neck.
Grimmertingen specimens are remarkable in having two or three relatively
strong grooves just below the subsutural platform; below this weaker grooves may
sometimes be distinguished. All other Belgian specimens seen from Berg and
Klein Spauwen lack this grooving. No specimens show any trace of collabral
sculpture other than growth lines.
DIMENSIONS. I 2 3 10 II
factual 14°5 8-3 8-6 — —
eeony ‘Lestimated 15°5 — = 1:87 1:80
GG 19720/1 16-1 QI gl 1°77 777
GG 19720/2 16-9 10:0 8-9 I-69 I-90
f actual 16:8 9:2 8-6 — =
COs ‘estimated 17°5 — — 1:90 2°03
GG 19719/2 18-8 Io-l 9:0 1-88 2:09
(see p. 192 for key)
REMARKS. GG 19717 from Royden was labelled as Cominella ventricosa Edwards
MS by Edwards; Keeping & Tawney’s (1881 : 116) record is of this specimen.
Judd’s list (1880 : 154) was also compiled from the Edwards Collection, so his record
of C. ventricosa from Colwell Bay, not Royden, is almost certainly a clerical error.
Von Koenen (1867 : 83; 1889 : 242) quotes Royden asa locality. It therefore seems
quite likely that he also examined this specimen, recognizing it to belong to W.
suturosa. Grimmertingen specimens seem to represent a fairly marked variety,
distinguished by the strength of the spiral grooving and by the usually less acute spire.
A damaged specimen, /RScNB figd. spec. no. 4070 I.S.T. from Grimmertingen (pl. 5,
figs 8a, b) identified as Babylonia (Pertdipsaccus) archambaulti (Meunier) by Glibert &
de Heinzelin (1954 : 368, 382, pl. 7, fig. 6) is definitely a Whiteclifia and may be an
exceptionally large, broad example of the present species with a spiral angle of 80°.
Whitecliffia tumida sp. nov.
(Pl. 6, figs 5-10)
HoLotyre. Upper Eocene, Late Priabonian, Middle Headon Beds (Venus Bed),
Whitecliff Bay, 1.O.W. GG 19722/r (Cu).
PALAEOGENE NASSARIIDAE 207
PARATYPES. Same locality and horizon, GG 19722/2-7 (Cu); GG 19724 (Cu);
GG 19723/1-3 (Wr); Sedg. Mus. C 29305 (KGT); (Cu); (S); GG 19756/1-3 (S) from
Bed O of Stinton (1971 : 405).
FURTHER OCCURRENCES. Unknown.
Diacnosis. Broad Whiteclifia with teleoconch of four or five whorls; mean
spire angle of about 65°; whorl profile strongly convex; last whorl often rather
swollen; collabral sculpture normally of up to sixteen strong, rather angular, costae
which becomes less frequent and either die out or are replaced by broad flexures on
pre-penultimate or penultimate whorl; spirally multistriate; subsutural collar
obsolete; suture becoming slightly canaliculate on last whorl or so; outer lip rarely
internally ribbed.
DEscripTIon. None of the specimens examined is complete, and the form of the
aperture is seen clearly only in one paratype. Whorls are enveloped just below the
periphery. The whorl profile is more strongly convex in the early whorls before
the collabral ribbing dies away. The whorl profileis almost horizontal at the adapical
suture, and usually becomes grooved by the third or fourth whorl. The subsutural
platform reaches a maximum width of 0-5 mm but seldom exceeds 0-3 mm wide; it
is not separated from the rest of the whorl-side by a rib. The teleoconch starts
with numerous weak, slightly opisthocline, collabral ribs. In the next quarter whorl
these increase in strength and gradually adopt the orthocline attitude of the collabral
ribs of the later teleoconch. There are normally about sixteen rather prominent,
angular, collabral ribs on the first whorls of the teleoconch. They are weakest just
below the suture and are of maximum strength from the shoulder to the adapical
suture. On the third teleoconch whorl they weaken considerably; they may be
just discerned on the fourth as faint flexures, which, on the body whorl, can be
seen to die away below the periphery. GG 419724 is unusual. On the second
whorl of its teleoconch about ten slight collabral crenulations occur for nearly half a
whorl. This is strongly reminiscent of the start of collabral ribbing in other speci-
mens, but dies away instead of developing further. On this specimen no other colla-
bral sculptural elements exist except for growth lines. In the other specimens the
spiral sculpture is first noticed within half a whorl of the collabral sculpture appearing.
It consists of between six and ten irregularly spaced bands. Usually they can be
seen to be ribs separated by interspaces wider than themselves. Often, however,
and particularly on the last whorl, this separation is reduced and the shell becomes
spirally multi-grooved. Bands of secondary strength may sometimes be seen.
Spiral sculpture is either weak or absent from the periphery of the body whorl.
Between seven and ten fairly prominent cords are present on the neck region.
Internal ribbing of the outer lip is seen clearly only on one specimen (Cu) which has
fifteen ribs extending back from the aperture for at least 45°. Irregular collabral
colour bands up to about 1 mm broad separated by gaps of similar size may be seen
on the last whorl of GG 19722/3 and one specimen (Cz); neither of these specimens
shows spiral colour banding. GG 19722/6, however, shows spiral but not collabral
colour banding. There isa dark band about 0-5 mm wide on the subsutural platform
208 SOME ENGLISH
with a slightly narrower band just below the platform, below this there are indications
of two more narrower bands well above the periphery.
DIMENSIONS. I 2 3 8 10 II
GG 19722/1 HT 12°9 8-0 707] 9+ I-61 1-67
GG 19722/2 PT 5:0 3:2 371 6+ 1°56 1°61
GG 19722/4 PT 6-7 4:0 3°9 8+ 1-67 1-72
GG 19722/5 PT 75 4:7 4:0 9+ I-60 1°88
GG 19722/7 PT 10:2 6-4 6-9 9+ 1°59 1-73
(see p. 192 for key)
REMARKS. This species closely resembles W. suturosa in shape and size. In
the latter, collabral sculpture is almost non-existent, spiral sculpture though similar
is weaker, and grooving of the suture occurs earlier and is more noticeable. As
indicated, GG 19724 has unusual sculpture. It seems advisable, however, not to
describe a separate species on the basis of a single specimen.
COLWELLIA gen. nov.
(PL. 7)
TYPE SPECIES. Cominella flexuosa Edwards in Lowry, Etheridge and Edwards,
1866.
Diacnosis. Agrees with Pseudocominella except in the following respects:
seldom exceeds 20 mm height; protoconch of about two and a half smooth whorls,
small, about I mm high, initially slightly flattened, first whorl very small; teleoconch
sculpture developing gradually with collabral sculpture developing first; teleoconch
with spiral and collabral sculpture of varying strength; subsutural platform either
poorly developed or absent ; growth lines of fasciole broadly sinuous as in Desovinassa ;
outer lip often denticulate within.
OTHER SPECIES ASSIGNED. WNassa antiquata Gabb, 1864 (pl. 7, fig. 11) Buccinum
auversiense Deshayes, 1865 (pl. 7, figs 6, 7); Ancillaria bretzi, Weaver, 1912 (pl. 7,
figs 8, 9); *Nassa cretacea, Gabb, 1864; *Molopophorus tejonensis Dickerson, 1915).
(See p. 201 for explanation).
GEOLOGICAL RANGE. Middle Eocene (Auversian) to Upper Eocene (late Pria-
bonian), Europe. Middle Eocene (Domengine Formation, California and Umpqua
Formation, Oregon) to Upper Eocene (Cowlitz Formation, Washington and Tejon
Formation, California), U.S.A.
REMARKS. The growth lines of the species assigned above to Colwellia are of
the type found in Pseudocominella and all the other new genera described herein:
they are approximately orthocline but are bent in an adoral direction in the sutural
(adapical) region. In the lectotype (pl. 8, fig. ra, b) of the type species of Molopo-
phorus, Bullia (M.) striata Gabb, 1869 (ANSP 4249), the growth lines are bent away
from the aperture in the sutural region, which appears to be canaliculate, as in
Brachysphingus (pl. 8, figs 3, 4). The same feature is shown by U.Cal. 7182-30750
PALAEOGENE NASSARIIDAE 209
(pl. 8, fig. 2). However, another specimen figured as Molopophorus striatus by
Dickerson (1915 : 67, pl. 8, fig. 6, given as fig. 7 in explanation to the plate) seems
to be misidentified and is almost certainly a Colwellia (pl. 7, fig. 10).
The type of growth line found in Molopophorus (s.s.) and Brachysphingus is
also found in Cyllene, a rare living genus of Nassariidae. This feature is found in
other widely-separated prosobranch superfamilies and is often associated with a
gutter or canaliculate suture which accommodates a posterior filament as in Oliva
(Volutacea) (Tryon, 1883 pl. 3) and Terebellum (Strombacea) (Jung & Abbott
1967, p. 446, pl. 319).
Sections through the columella of Brachysphingus show that it lacks the terminal
plait found in all definite Nassariidae including Cyllene. Although no specimens
of Molopophorus s.s. were available for sectioning (the lectotype is the only specimen
in the ANSP collections—Dr H. G. Richards, personal communication), the external
similarities of the two genera are so great that it is doubtful if either belongs to the
Nassariidae. Their general shell features including the fasciole ridge and reverse
S-shaped growth lines suggest that they may best be placed in the very broad
Buccinidae (sensu Wenz, 1941-1943).
In fact, it seems that Molopophorus (Gabb, 1869 : 156) is a synonym—based
on immature material—of Brachysphingus (Gabb, 1869 : 155), which is here selected
as generic name of the taxon.
A large number of North American west coast Tertiary species ranging in age
from the Meganos Formation (lower Eocene) of California (Clark & Woodford,
1927 : 117) to the Miocene of Washington State (Weaver, 1942 : 463-471) have been
assigned to Molopophorus and the genus is also recorded from the Oligocene and
Miocene of Japan (Oyama, 1960 : 73-74). These species differ considerably in
form. Vokes, moreover, (1939 : 140-141) mentions that there were two or three
distinct types of nuclear whorls among the Californian species ascribed to Molopo-
phorus and suggested that they were unlikely to be congeneric. Most of these
species need assigning to new genera—a task beyond the scope of this work. It
would appear, however, that they are fairly closely related to the European forms
described herein and would also seem to belong to the Nassariidae.
Gabb (1864 : 97) described Nassa cretacea and Nassa antiquata both of which
have been regularly assigned to Molopophorus by American workers in this century.
Nevertheless, he himself still assigned them to Nassa (1869, 219) in the paper in
which he described Molopophorus. The difficulties of interpreting Molopophorus
correctly have hindered workers who have suspected a connection between it and
European species previously assigned to Cominella.
We have assigned to Colwellia only those American species which have a very
strong resemblance to C. flexuosa and C. auversiensis.
In Desorvinassa the ramp is convex and the collabral sculpture weak. In Colwellia
the ramp is concave and the collabral sculpture is much stronger on the earlier
whorls. D. williamsi (Ypresian) and C. auversiensis (Auversian) appear to form a
link between the two genera.
The differences in the protoconch, given in the diagnosis, are sufficient to distin-
guish Colwellia from Keepingia, which almost certainly arose from it.
Cc
210 SOME ENGLISH
Colwellia flexuosa (Edwards, 1866)
(Pl. 7, fig. 1-5)
1866 Cominella flexuosa Edwards in Lowry, Etheridge & Edwards, pl. 3.
1880 Cominella (Buccinum) flexuosa Edwards MS; Judd: 154.
1881 Cominella (Buccinum) flexuosa Edwards MS; Keeping & Tawney: 105, 116.
1889 Cominella flexuosa Lowry; Bristow: 290.
1891 Cominella flexuosa Edwards MS; Newton: 168.
1921 Cominella flexuosa Edwards; Wrigley: 139.
1926 Cominella flexuosa Edwards; Jackson: 356.
1963 Cominella flexuosa Edwards in Lowry; Glibert: 67.
LECTOTYPE (designated herein). Upper Eocene, late Priabonian, Middle Headon
Beds (presumably Venus Bed), Colwell Bay, I1.0.W., GG 19709/1 (Ed).
PARALECTOTYPES (designated herein). Same details, GG 19709/2-30.
MATERIAL STUDIED. Restricted to Middle Headon Beds. Venus Bed, Colwell
Bay, GG 19710/1-11 (Wr); GG 19711 (Nu); (Cu); (S); Sand. Mus. 4778, 748; Sedg.
Mus. C 28281-4 (K&T). Headon Hill, 1.0.W., GG 19712/1-3 (St]B); Sedg. Mus.
C 28645-9 (KGET). Venus Bed, Whitecliff Bay, 1.0.W., GG 19713 (Cu). Brocken-
hurst Bed, Brockenhurst, GG 19714 (Ed).
FURTHER OCCURRENCES. Unknown.
Diacnosis. Colwellia with narrow sloping subsutural collar and rather broad
concave subsutural ramp; mean spire angle between 55° and 70°; angular can-
cellate ribbing on early whorls, becoming almost smooth, particularly below peri-
phery of last whorl; outer lip often weakly denticulate within; obvious sinus in
growth lines of peripheral region, and in some specimens a posterior siphonal notch,
developed at maturity.
DESCRIPTION. The protoconch is worn in all specimens, the junction with the
teleoconch cannot be seen. The aperture is slightly more than two-thirds shell
height. The teleoconch consists of up to five whorls. The suture normally lies on
the periphery, sometimes on later whorls it lies just below as the sutural slope in-
creases slightly. Below the rounded subsutural band the very slightly concave
ramp slopes away to the shoulder. Three or four weak spiral ribs occur on and
below the shoulder of spire whorls; that on the shoulder is the strongest. These
ribs become noticeably weaker on the last whorl. Up to two more spiral ribs,
which are usually weaker, may occur on the ramp. Between five and ten rather
widely spaced spiral cords are present on the neck of the body whorl. The collabral
ribs are stronger than the spiral ones and are like crested-waves in cross-section.
There are about twenty-one collabral ribs on the spire whorls of the teleoconch.
On the first half of the body whorl there are about sixteen increasingly weaker ribs.
Collabral ribbing becomes virtually obsolete on the final half whorl. The main
indication of its presence is slight axial nodes on the spiral ribs. Similar nodes are
formed on the spire whorls where the collabral and spiral ribs intersect giving a
rather cancellate appearance. In some specimens the outer lip is smooth internally,
in others there are up to fifteen unevenly spaced weak denticles about 1 mm long
PALAEOGENE NASSARIIDAE 211
and set about I mm inside the aperture, and a few shells show another row of
denticles about 45° inside the aperture. As well as the broad sinus in the growth
lines of the peripheral region of mature specimens there is a slight posterior siphonal
notch developed in the outer lip where it meets the suture (pl. 7, fig. 2c). Its strength
seems related to the size of the shell and it is sometimes absent in smaller specimens.
It may be presumed to mark the reaching of full growth as there is no trace, except
occasionally on the last quarter whorl, of its previous existence on earlier growth
lines. In a few specimens the colour pattern is clearly visible under ultra-violet
light. A pair of spiral bands, about 0-5 mm wide, separated by about 0-2 mm lie on
the periphery (thus appearing just above the suture on spire whorls). Another pair
of bands, about half the breadth, occurs at the top of the neck.
DIMENSIONS. I 2 3 4 5 6 7 8 9 10 II
GG 19709/1 LT 14:2 05 QO 65 TAU 2 20 eeted| 6 5 1:49 1°58
GG 19713 — 5°4 3°3 i Lp s20) 5H Aneel 1-630 51-65
GG 19709/6 PLT 10:8 6:8 6-05 te 2222 rz 4 © £59 1-64
GG 19709/2 PLT 14°3 8-3 S:0mnn Pp it SNE 7) 6) 1-72 1-66
GG 19709/4 PLT 14°9 9°4 OR § 18 23; 15-+ 7 OME SOM O2
GG 19709/3 PLT 20°'I 13:0 It2 6 Oo. 25, 28 Io) RG) RON, StF)
(see p. 192 for key)
REMARKS. Following Wrigley’s (1921 : 139) reasoning, we ascribe the author-
ship of this species to F. E. Edwards. Its resemblance to the earlier C. auversiensis
(Deshayes) (pl. 7, figs 6, 7) is striking. Both species have teleoconchs so similar
to those of Keepingia that it is necessary to compare them with both K. gossardi
(Nyst) (pl. 8, figs 5-7) and K. cassidaria (Sandberger) (pl. 8, fig. 9). C. auversiensis
differs from both C. flexuosa and K. cassidaria in being less tumid, having a less
concave ramp and less marked shoulder whilst its collabral sculpture, though weaker
on early growth stages is more persistent and its growth lines less flexuous. Sand-
berger figures two varieties of K. cassidaria. The first (1863, pl. 20, figs I, 1a, 1d,
Ie) is characterized by about four strong spiral ribs on the spire whorls and about
ten on the body whorl. It lacks collabral sculpture on the last and sometimes on the
penultimate whorl. The second variety (1863, pl. 20, figs 1b, 1c) which he refers to
(p. 230) as var. cancellata, is less common in B.M.(N.H.) collections. It has a
cancellate appearance; the primary spiral ribs are separated by secondary ones and
numerous collabral rugae are developed. Both forms of K. casstdaria may be
readily distinguished from C. flexuosa by their sculpture. K. gossardi differs from
C. flexuosa by being larger and having a relatively smaller aperture, less sinuous
growth lines, and more definite subsutural collar with a tendency for the suture to
be canaliculate. The sculpture of K. gossavdi is much stronger and persists to
maturity. It has numerous spiral ribs and about fifteen collabral folds per whorl.
Two specimens in the F. E. Edwards Collection (GG 19725-6) were labelled
by Edwards as being from the Bracklesham Beds of Brook, Hampshire. The first
was correctly identified as flexuosa and appears from the matrix to be from the
Brockenhurst Bed of the Brockenhurst area. The second is likely to be from Brook
but is a young Strepsidura turgida (Solander). The sculpture on the spire whorls of
Ct
212 SOME ENGLISH
these two species is remarkably similar. Edwards’ label may have caused Keeping
& Tawney (1881 : 105, 116) to indicate that the species occurs in strata older than the
Headon Beds.
There is a remarkably small variation in size of the known specimens. Most of
those in B.M.(N.H.) collections lie between 12 and 16 mm in height. The largest
and smallest specimens whose dimensions are given here are exceptional.
KEEPINGIA gen. nov.
(Pi; 8, figs 5-9)
TYPE SPECIES. Buccinum gossardi Nyst, 1836.
Diacnosis. Agrees with Colwellia, except in the following respects; sometimes
exceeding 20 mm in height; protoconch initially slightly heterostrophic, of one and
a half smooth whorls, separated from teleoconch by a whorl on which there are about
20 decreasingly opisthocline collabral folds, with spiral sculpture developing on later
part.
OTHER SPECIES ASSIGNED. **Cominella annandalei Vredenburg 1925; **Cominella
aturensis Peyrot, 1927; Buccinum bolli Beyrich, 1854; Buccinum cassidaria Sand-
berger, 1863; **Cominella praecedens Peyrot 1927; Nassa tarbellica Grateloup,
1834; **Buccinum uniseriale Sandberger, 1863. (See p. 201 for explanation.)
GEOLOGICAL RANGE. Lower Oligocene (Late Lattorfian) to Lower Miocene
(Burdigalian).
REMARKS. The similarities of the teleoconch suggest that Keepingia is close to
Colwellia but its consistently different protoconch in all species examined justifies
its separation. It is the only genus described herein that survives into the Miocene.
The first occurrence is of the type species in the Upper Tongrian Henis Clay of
Belgium, the species persists into the Rupelian of Belgium and France. The last
definite occurrence of the genus is that of K. tarbellica in the Burdigalian of the
Bordeaux basin.
As indicated above. no specimens of four of the species listed were available for
study, so the stratigraphical and geographical range of the genus as shown on Table 1
can only be tentative. This is particularly unfortunate in the cases of C. annandalet
from the Nari (Oligocene) of the Indian sub-continent. No species are known
from England.
THANETINASSA gen. nov.
(Pl. 9, figs 1-7)
TYPE SPECIES. Buccinum bicorona Melleville, 1843.
Diacnosis. Phos-like, relatively tall, up to about 20 mm high; teleoconch of up
to six whorls; mean spire angle of about 45°; aperture about half shell height; strong,
PALAEOGENE NASSARIIDAE 213
slightly sloping, subsutural platform developed; both spiral and collabral ribbing
strong, with beads produced at intersections; outer lip notably convex, only slightly
produced anteriorly; growth lines markedly recurved in neck region immediately
prior to terminal aperture, to form a ‘stromboid-like’ notch, associated with particu-
larly strong spiral ribbing; columella bent to the right above a terminal plait; growth
lines of fasciole as in Desovinassa, columellar and parietal callus weakly developed.
OTHER SPECIES ASSIGNED. None.
GEOLOGICAL RANGE. Palaeocene (Thanetian).
REMARKS’ The strong ‘stromboid-like’ notch developed at the final aperture
immediately distinguishes this genus from the others described in this paper.
All these have a slight sinuosity in a similar position, but nothing so marked. In
living Strombus the notch accommodates the right eye peduncle.
Both Phos and Nassarius s.l1. have similar notches. It is always strong in Phos,
but is of variable strength in Nassarius. The eye tentacles of Phos are joined at
their base to form a Y-shaped structure (Tryon, 1881 : 215, pl. 83, fig. 484), in
Nassarius (Tryon, 1882, pls 7, 9, 12, 14, 15, 18) the arrangement is rather similar.
It seems possible that this structure as a whole, rather than a single tentacle, lay
underneath the notch.
It is hoped that the relationship between the Nassariidae and the Phos-like
genera which were assigned to the Buccinidae by Wenz (1941) will be discussed in
a future paper in greater detail than would be appropriate here. The curved
columella, the form of the columellar plait, with its strong posterior boundary ridge
and the broad fasciole of Thanetinassa are very similar to those of contemporary
Desovinassa, which differs in having very weak sculpture The later genus
Pseudocominella is strongly sculptured and P. solanderi is similar in shape and size
toT. bicorona. Any similarity in their sculpture, however, is superficial. Moreover,
the columellar and fasciole features of the two genera are very different.
Thanetinassa should be compared with Buccitriton and Tritiaria (both assigned
by Wenz, 1941 : 1177-1178, to the Buccinidae) and Sagenella (tentatively assigned
by Wenz, 1943 : 1224, to the Nassariidae) All three genera occur in the Lower
Tertiary of the South-Eastern United States. In all three the fasciole is bounded
posteriorly by a much weaker ridge than in Thanetinassa, and the stromboid notch
is either weak or absent and is never associated with strong spiral ribs. In Buccitriton
the columella is short, terminating well above the most anterior part of the shell,
which in this case is the outer lip. In Thanetinassa, on the other hand, the terminal
columellar plait forms the most anterior portion of the shell. In Trttiarza the fasciole
is much narrower than in Thanetinassa and is spirally striate.
Thanetinassa bicorona (Melleville, 1843)
(Pl. 9, figs 1-7)
1843 Buccinum bicorona Melleville: 12, 73, pl. 10, figs 4, 5.
1850 Buccinum bicorona d’Orbigny: 303.
1865 Buccinum quoesitum Deshayes: 503.
214 SOME ENGLISH
1866 Buccinum quoesitum Deshayes: pl. 93, figs 9-12.
1889 Cominella bicoronata (Melleville) nom. mut. Cossmann: 138.
1901 Cominella bicovona (Melleville); Cossmann: 150.
1911 Cominella bicoronata (Melleville); Cossmann & Pissaro, pl. 37, fig. 178-7.
1934 Cominella bicovonata (Melleville); Cooper: 7.
1963 Comunella bicorona (Melleville) ; Glibert: 66.
MATERIAL STUDIED. ENGLAND, Palaeocene, Thanetian, Upper Thanet Sands’
Corbula Bed. Herne Bay, Kent, G57073-8 (J. E. Cooper Colln.). Bishopstone’
Herne Bay, G 60746 (J. E. Cooper Colln.); GG 19741/1-2 (Wr); GG 19742 (Wr);
GG 19743 (A. G. Davis Colin.). Bishopstone Gap, GG 19744 (Ru). Arctica morrist
Bed, foreshore exposure approx. 400 yards E. of Bishopstone Gap, GG 19745 (Ru).
FRANCE, Palaeocene, Thanetian, Chalons-sur-Vesle, Marne, G 19357/1-4 (L. Staadt
Colln.); GG 19740 (Wr); 5 shells EMP (Deshayes Colln.).
FURTHER OCCURRENCES. FRANCE, Sable Inférieurs (Thanetian). TyPE LOCALITY
Villiers-Franqueux (Melleville, 1843 : 73). Chalons-sur-Vesle, Brimont, Jonchery
(Deshayes, 1865 : 503; Chalons-sur-Vesle (Briart & Cornet, 1871 : 31); Chalons-sur-
Vesle, Chenay, Jonchery (Glibert, 1963 : 66).
DiaGnosis. As for genus.
Description. All the specimens are fragile, abraded, and with their earlier
whorls either missing or decorticated. It is impossible therefore to separate the
protoconch and teleoconch. GG 19741/2, however, is 12-3 mm high and has eight
whorls. Neither definite ribbing nor the subsutural platform appear until the fourth
whorl, so the first three may well comprise a rather acutely-spired, naticoid, proto-
conch. Most of the spiral ribs are distinctly weaker than the collabral ones and the
preservation in all specimens is such that spiral ribs cannot be seen until the third
teleoconch whorl. Up to nine spirals ribs occur on the spire whorls and about twenty
on the body whorl. The two strongest, which are stronger than any of the collabral
ribs, are those forming the sloping subsutural platform and the shoulder. The
concave ramp is formed in the interspace between these two ribs. Sometimes two
or three other fairly strong ribs occur on the neck above the fasciole, marking a
backwardly-directed fold in the growth lines. Sometimes a weak rib is developed
above the main rib of the subsutural platform. The collabral ribs are more or less
orthocline and decrease in strength below the periphery; on some specimens they
may be seen to follow the above-mentioned fold on the neck. They are prominent,
steep-sided, and separated by interspaces nearly twice their width. Their number
remains fairly constant, throughout growth. On the last whorl, their spacing
becomes less regular. An irregular, varix-like, pre-terminal aperture is often
developed just after the commencement of the last whorl, being situated on the
ventral surface of the shell, slightly to the left of the final aperture. Similar varices
may well be developed later on the last whorl, GG 19744 having a total of five.
Two weak spiral ribs are occasionally developed on the fasciole. Spiral ribbing has
not been seen within the outer lip.
PALAEOGENE NASSARIIDAE 215
DIMENSIONS. I 2 3 4 5 6 7 8 9 10 Ev
18°5 — — — —_- — 9 — — — —
een rae hee 20:0 = 9:0 6 AS SES SS = 1-77
ee ee OL ONE te ac EON Ol ae | i — —
prorat: ea 35 —- — 6 22 —- —- — — — 2°31
GG 19741/2 123. — 60 5-6 15 20 420 8 — — 2:01
155 85 83 6 23) 124 LS 8 — — —
GG 19357/1 ee 160 — — —- —- — — ~ ~~ 188 = 1:93
GG 19357/2 146 7:0 83 6 TO} LO 2 8 — 2:08 1:76
GG 19357/3 12: Oo: Onn On) 6 22 Zien nO 7 — 1:70 I-61
Remarks. The English specimens are notably more strongly varicate on the
last whorl than those from France. There are, however, no other apparent grounds
for specific separation. This species strongly resembles Buccinum montense (pl. 9,
figs 8a, b) described by Briart & Cornet (1871 : 30-31) from the Montian ‘Calcaire
Grossier’ of Mons; they compare the two species in detail, and both were subsequently
assigned to Cominella by Cossmann (1901 : 150). Examination of the fasciole of
Buccinum montense from Mons (GG 71604—7, Chavan Colln.) show that this species is
not a member of the Nassariidae but is congeneric with European Eocene and Oligo-
cene species assigned to Pollia of the Buccinidae.
Cossmann (1889 : 138) placed B. quoesitwm Deshayes (1865) in the synonomy
of B. bicorona Melleville (1843) remarking that Deshayes had overlooked the earlier
name. The whereabouts of Melleville’s specimens is unknown but his figures (1843,
pl. ro, figs 4, 5) appear to agree with those of Deshayes (1866, pl. 93 figs 9-12) and
Deshayes’ surviving specimens all from Chalons-sur-Vesle (EMP). Melleville’s
locality was Villers Franquex, at which the only horizon exposed is the middle
fossiliferous zone of the Sables de Chalons-sur-Vesle (Fritel, 1910 : 332).
REFERENCES
Axssott, R. T. 1960. The genus Stvombus in the Indo-Pacific. Indo-Pacific Mollusca,
Philadelphia. 1 (2) :33-146, pls 11-117.
ALBRECHT, J. C. H. & VaLtx, W. 1943. Oligocine Invertebraten von Sud-Limburg. Meded.
geol. Sticht., Maastricht, Ser.C.IV. 1 : (3). 163 pp., 27 pls.
BELLARDI, L. 1882. I molluschi dei terveni terziavi del Piemonte e della Liguria, 3. 253 pp.,
12 pls. Turin.
Beyricu, H. E. 1854. Die Conchylien des norddeutschen Tertiargebirges. Pt. 2. Z. dt.
geol. Ges., Berlin, 6 : 408-500, pls. 9-14.
Borson, S. 1820. Saggio di orittografia Piemontese. Memorie R. Acad. Sci. Torino, 25 : 180-
220, pl. 5.
Bosguet, J. 1880. Fossiles du systéme Tongrien et de l étage inférieur Rupélien in DEWALQUE,
G. Pyrodrome d'une description géologique de la Belgique: 2nd. ed.: 463-473. Brussels.
Briart, A. & CorNET, F.-L. 1871. Description des fossiles du calcaire grossierde Mons. Pt. 1.
Mém. cour. Sav. étr. Acad. v. Sci. Belg., Brussels, 36 : i—viii, 1-76, pls 1-5.
216 SOME ENGLISH
Bristow, H. W. 1889. The geology of the Isle of Wight. 2nd. (revised by C. Reid & A.
Strahan) ed. Mem. geol. Surv. U.K., London, xiv+349 pp., 5 pls, 84 text-figs.
Burton, E. St. J. 1933. Faunal horizons of the Barton Beds in Hampshire. Pyroc. Geol.
Ass., London, 44 : 131-167.
Crark, B. L. & Wooprorp, A.O. 1927. The geology and paleontology of the type section of
the Meganos Formation (Lower Middle Eocene) of California. Univ. Calif. Publs geol. Sct.,
Berkeley, 17 : 63-142, pls 14-22, I map.
Cooper, J. E. 1934. Oldhaven and Thanet Sand Mollusca of Herne Bay. J. Conch. Lond.,
20 : 4-8.
Cossmann, M. 1889. Catalogue illustré des coquilles fossiles de 1’Eocéne des environs de
Paris. Amnnls Soc. v. malacol. Belg., Brussels, 24 : 385 pp., 12 pls.
— 1901. Essais de paléoconchologie comparée. 4, 293 pp., 10 pls. Paris.
1915. Essais de paléoconchologie comparée. 10, 292 pp., 12 pls. Paris.
— & Pissarro, G. 1911. Iconographie compléte des coquilles de Eocene des environs de
Paris. 2, 65 pls. Paris.
Curry, D. 1958. Lexique stvatigyvaphique international, Europe; Angleterre, Pays de Galle
et Ecosse, Paléogéne. 1 (3a), XII, 82 pp. Paris.
Dat, W. H. tI909. Contributions to the Tertiary paleontology of the Pacific coast. I.
The Miocene of Astoria and Coos Bay, Oregon. Prof. Pap. U.S. geol. Suvv., Washington,
no. 59, 278 pp., including 22 pls in pagination.
Davis, A. G. 1952. The Brockenhurst Beds at Victoria Tilery, Brockenhurst, Hampshire.
Proc. geol. Ass., London, 63 : 215-219.
DESHAYES, G.-P. 1824-1837. Description des coquilles fossiles des envivons de Paris. 2, 814 pp.;
atlas 28 + 51 pp., LXV + 106 pls. Paris. pp. 499-780 (1835); pls. 1-105 (1837).
—— 1864-1866. Description des animaux sans vertébves découverts dans le bassin de Paris. 3 :
I—200 (1864); 201-658 (1865). 2, atlas 107 pp., 107 pls. (1866). Paris.
DEWALQUE, G. see BosguEtT, J. 1880.
DIcKERSON,R.E. t915. Fauna ofthe type Tejon: its relation to the Cowlitz phase of the Tejon
Group of Washington. Proc. Calif. Acad. Sci., San Francisco, Ser. 4, 5 : 33-98, pls 1-11.
Drxon, F. see SowERBy, J. deC. 1850.
Farcuap, H. 1936. Etude du Thanetien (Landenien marin) du bassin de Paris. Mém. Soc.
geol. Fr., Paris, N.S. 13, mém. 30 : 1-103, pls 1-6.
FIsHER, O. 1862. On the Bracklesham Beds of the Isle of Wight basin. Q. Jl geol. Soc.
Lond., 18 : 65-94.
ForBES, E. 1856. On the Tertiary fluvio-marine formations of the Isle of Wight. Mém.
geol. Surv. U.K., London, 162 pp., Io pls., 20 text-figs.
FRETTER, V. & GRAHAM, A. 1962. British prosobvanch molluscs. xvi + 755 Pp., 317 figs.
London.
FRriTeEL, P. H. 1910. Guide géologique & paléontologique de la région Parisienne. 356 pp.,
12 pls (maps), 162 text-figs. Paris.
Furon, R. & Kourtatcuy, N. 1948. La faune Eocéne du Togo. Mém. Mus. natn. Hist.
nat. Paris, N.S. 27 : 95-114, pls 8, 9.
Gass, W. M. 1864. Description of the Cretaceous fossils in Paleontology of California, Phila-
delphia, 1 : 57-236, pls 9-32.
1869. Cretaceous and Tertiary fossils in Paleontology of California, Philadelphia, 2, 299
pp., 36 pls.
GaRDNER, J. S., KEEPING, H. & Monkton, H. W. 1888. The Upper Eocene, comprising the
Barton and Upper Bagshot formations. Q. Jl geol. Soc. Lond., 44 : 578-635.
GIEBEL, C. 1864. Die fauna dev Braunkohlenformation von Latdorf bei Bernberg. 93 pp., 4 pls.
Halle.
GLIBERT, M. 1957. Pélécypodes et gastropodes du Rupélien supérieur et du Chattien de la
Belgique. Mém. Inst. rv. Sci. nat. Belg., Brussels, 137, 98 pp., 6 pls.
1960. Les Volutacea fossiles du Cénozoique étranger des collections de l'Institut Royal des
Sciences Naturelles de Belgique. Mém. Inst. rv. Sci. nat. Belg., Brussels, ser. 2, 61, 109 pp.
PALAEOGENE NASSARIIDAE 217
GLIBERT, M. 1963. Les Muricacea et Buccinacea fossiles du Cénozoique étranger des collections
de l'Institut Royal des Sciences Naturelles de Belgique. Mém. Inst. v.Sci.nat. Belg., Brussels,
Ser. 2, 74, 179 pp.
—— & de HEINZELIN de BRAvucourT, J. 1954. L’Oligocéne inférieur belge in Volume jubilaire
Victoy van Straelen, 1 : 281-438, pls 1-7. Brussels.
GRATELOUP, J. P. S. DE. 1834. Tableau (suite du) des coquilles qu’on recontre dans les
terrains tertiares grossiers (faluns) du bassin géologique de l’Adour (Landes). Act. Soc.
linn. Bordeaux, 6 : 270-320.
Gray, J. E.in Gray, M.E. 1850. Figures of molluscous animals selected from various authors.
4,iv + 219 pp. London.
Horz1, O. 1958. Die Molluskenfauna des oberbayerischen Burdigals. Geologica bav.,
Munich, 38, 348 pp., 2 pls., 5 text-figs.
IREDALE, T. 1918. Molluscan nomenclatural problems and solutions—no. 1. Proc. malac.
Soc. Lond., 18 : 28-40.
Jackson, J. F. 1926. A catalogue of Eocene and Oligocene fossils in the Museum of Isle of
Wight Geology, the Free Library, Sandown. Pyroc. Isle Wight nat. Hist. archaeol. Soc.,
Newport, 1, (6 for 1925) : 340-373.
Jupp, J. W. 1880. On the Oligocene strata of the Hampshire Basin. Q. Jl geol. Soc. Lond.,
36 : 137-177.
1882. On the relations of the Eocene and Oligocene strata in the Hampshire Basin.
Q. Jl geol. Soc. Lond., 38 : 461-489.
June, P. & Assort, R. T. 1967. The genus Tevebellum (Gastropoda: Strombidae), Indo-
Pacific Mollusca, 1 (7) : 445-454, pls 318-327.
KEEPING, H. 1887. On the discovery of the Nummulina elegans zone at Whitecliff Bay, Isle
of Wight. Geol. Mag., London, N.S., Dec. 3, 4 : 70-72.
& TAwNeEy, E. B. 1881. On the beds at Headon Hill and Colwell Bay in the Isle of
Wight. Q. Jl geol. Soc. Lond., 37 : 85-127, pl. 5.
KOENEN, A. von. 1864. On the correlation of the Oligocene deposits of Belgium, Northern
Germany, and the South of England. Q. Jl geol. Soc. Lond., 20 : 97-102.
1865. Die Fauna der unter-oligocdnen Tertiarschichten von Helmstadt bei Braunschweig.
Z. dt. geol. Ges., Berlin, 17 : 459-534, pls 15, 16.
1867. Das marine Mittel-Oligocén Nord-Deutschlands und seine Mollusken-fauna.
Palaeontogvaphica, Stuttgart, 16 : 53-127, pls 6, 7.
1889. Das Norddeutsche Unter-Oligocaén und seine Mollusken-fauna. Part 1. Abh.
geol. SpecKarte preuss. thir. St., Berlin, 10 : 1-280, pls 1-23.
Lowry, J. W., ETHERIDGE, R. E. & Epwarps, F. E. 1866. Chart of the characteristic British
Tertiary fossils, stvatigvaphically avvanged. 4 pls, 800 figs. London.
Martini, E. 1969. Nannoplankton aus dem Latdorf (locus typicus) und weltweite Para-
llelisieurungen im oberen Eozan und unteren Oligozin. Senckenberg. leth., 50 : 117-159,
pls 1-4.
—— & MoorkeEns, T. 1969. The type-locality of the Sands of Grimmertingen and calcareous
nannoplankton from the Lower Tongrian. Bull. Soc. belge Géol. Paléont. Hydrol. Brussels.
78 : 111-130.
& Ritzkowski, S. 1968. Was ist das ‘Unter-Oligocan’? Eine Analyse der BEY RICH’-
schen und V. KOENEN’schen Fassung der Stufe mit Hilfe des fossilen Nannoplanktons
Nachr. Akad. Wiss. Gottingen, II, Mathphys-K1., 13 : 231-250.
MELLEVILLE, M. 1843. Mémoire sur les sables tertiaives du bassin de Paris. 88 pp., Io pls.
Paris.
Morettet, L. & J. 1948. Le Bartonien du bassin de Paris. Mém. Serv. Carte géol. dét. Fr.,
Paris, 437 pp.
Morris, J. 1843. A catalogue of British fossils. x + 222 pp., London.
Newton, R. B. 1891. Systematic list of the Frederick E. Edwards collection of British
Oligocene and Eocene Mollusca in the British Museum (Natural History). xiii + 365 pp.
London.
218 SOME ENGLISH
Nyst, P.-H. 1836. Recherches sur les coquilles fossiles de Hoesselt et de Kleyn-Spauwen (province
du Limbourg). iii + 40 pp., 4 pls. Ghent.
1843. Description des coquilles et des polypiers fossiles des tervains tertiaives de la Belgique.
atlas, 48 pls. Brussels.
1845. Description des coquilles et des polypiers fossiles des terrains tertiaires de la
Belgique. Mém. cour. Sav. éty. Acad. y. Sci. Belg., Brussels, 17 : 1-697, pls 1-15.
Orpicny, A. D’. 1850. Prodrome de paléontologie stvatigvaphique universelle des animaux
mollusques et vayonnés. 2, 428 pp. Paris.
Oyama, K., Mizuno, A. & Sakamoto, T. 1960. Illustvated handbook of Japanese Paleogene
molluscs. 241 pp., 71 pls. Tokyo.
Preyrot, M. A. 1928. Conchologie néogénique de l’Aquitaine. Act. Soc. linn. Bordeaux, 79
suppl. : 5-263, pls 5-14.
PuiLippl, R. A. 1846-1847. Verzeichniss der in der Gegend von Magdeburg aufgefundenen
Tertidrversteinerungen. Palaeontogyaphica, Stuttgart, 1 : 42-44 (1846); 45-90, pls 7-10,
10a (1847).
Pow_E LL, A. W. B. 1929. The recent and Tertiary species of the genus Buccinulum in New
Zealand, with a review of related genera and families. Tvans. N.Z. Inst., Auckland, 60 :
57-101, pls 1-4.
REEVE, L. 1853-1854. Monograph of the genus Nassa. Conchologica Iconica, 8, 29 pls.
ReIp, C. 1902. The geology of the country around Southampton. Mem. geol. Surv. U.K.,
London, 70 pp., 21 figs.
—— & STRAHAN, A. see Bristow, H. W. 1880.
SANDBERGER, C. L. F. 1863. Die Conchylien des Mainzer Tertidrbeckens. 458 pp., 1 chart,
35 pls. Wiesbaden.
SOLANDER, D.C.in BRANDER,G. 1766. Fossilia Hantoniensia collecta, et in Musaeo Britannica
deposita. vi + 43 pp., 9 pls. London.
SOWERBY, J. DEC. 1823-1825. The Mineral Conchology of Great Britain. 5 : 1-64, pls 408—
443 (1823); 65-138, pls 444-485 (1824); 139-171, pls 486-503 (1825). London.
1850. Notes and descriptions of new species in Dixon, F. 1850, The geology and fossils of
the Tertiary and Cretaceous formations of Sussex : 163-194, pls 2-9, London.
STEwarT, R. B. 1927. Gabb’s California fossil type gastropods. Pyvoc. Acad. nat. Sct.
Philad., 78 (for 1926) : 287-447, pls 20-32.
STINTON, F. C. 1971. Easter field meeting in the Isle of Wight, Thursday 26th March to
Tuesday 31st March 1970. Proc. geol. Ass., London, 82 : 403-410.
Tawney, E. B. 1883. On the outcrop of the Brockenhurst Bed near Lyndhurst. Geol.
Mag., London, Dec. 2, 10 : 157-160.
TRavuB, F. 1938. Geologische und palaontologische Bearbeitung der Kreide und des Tertiars
im Ostlichen Rupertiwinkel, nordlich von Salzburg. Palaeontogvaphica, Stuttgart, A,
88 : 1-114, pls 1-8.
Tryon, G. W. 1881-1883. Manual of Conchology, 3, 310 pp., 87 pls (1881); 4, 276 pp., 58 pls
(1882); 5, 276 pp., 63 pls (1883). Philadelphia.
VINCENT, G. 1886. Liste des coquilles du Tongrien inférieur du Limbourg belge.
nat. Hist. (Geol.)
Mus.
e*
=) Ep anaeges =
Bull. Br.
PAG NIE, 2
Fics 1-8
Pseudocominella solanderi (Cossmann) p. 198
Upper Eocene, early Priabonian, Lower Barton Beds, Barton-on-Sea, Hampshire.
Fics 1a—d. Neotype, GG 19693/1 (Ho). Horizon A3. x4. Fig. 1d. Oblique view to
show colour-bands separated by the spiral rib which marks the weak shoulder of the last whorl.
Fic. 2. GG 19694/1 (Le), same sample as figs 3, 6, 8, Horizon A3, relatively tumid specimen.
Be
Fic. 3. GG 19694/2, same sample as figs 2, 6, 8 showing unusually persistent axial ribbing
and colour-banding. 5.
Fic. 4. GG 19698/1 (Nw). Horizon A2, juvenile shell showing early sculpture, and colour-
banding on shoulder. x6.
Fic. 5. GG 19695/2 (S). Horizon A3, same sample as fig. 7, juvenile. 6.
Fic. 6. GG 19694/3, same sample as figs 2, 3, 8, adult shell with persistent strong axial
ornament, and colour-banding. x 3.
Fic. 7. GG 19695/1, same sample as fig. 5, mouth of gerontic shell. 4.
Fic. 8. GG 19694/4, same sample as figs 2, 3, 6, protoconch, the erect appearance of the
first whorl is due to weathering. x 24 (stereoscan).
Fics ga, b
Pseudocominella bullata (Philippi) pp. 189, 193, 198
Lower Oligocene, Lattorfian, Latdorf, Germany, GG 19736/1 (Ed).
Fic. ga. Showing lack of axial sculpture on early whorls. x 2.
Fic. 9b. Showing protoconch and development of spiral sculpture. 12.
nat. Hist.
Mus.
Bull. Br.
PLATE 3
Fics 1-6
Pseudocominella armata (J. de C. Sowerby) p- 193
Middle Eocene, Auversian, Upper Bracklesham Beds.
Fics ta, b. Neotype. GG 19681/1 (Ed.), Bracklesham Bay, Sussex. 2.
Fics 2-4. GG 19685/1-3 (Ed), Bramshaw, Hampshire.
Fic. 2a. GG 19685/3. x2.
Fic. 2b. GG 19685/3. Showing fasciole. x5.
Fic. 3. GG 19685/2. Oblique view showing development of strong sculpture and pre-
terminal apertures on last whorl. x 3.
Fic. 4. GG 19685/1. Early whorls. 12 (stereoscan).
Fic. 5. GG 19683/1 (Ed), Brook, Hampshire, oblique view showing relatively weak sculpture.
x3.
Fic. 6. GG 19684 (Ed), Huntingbridge, Hampshire. x 3:5.
Fics 7, 8
Pseudocominella armata (J. de C. Sowerby)
Paris Basin, referred to by Deshayes 1865 : 499 as Buccinum fusiforme Deshayes,
(Deshayes Colin., EMP).
Fic. 7. Specimen from either Le Fayel, Mary or Caumont, Auversian, oblique view showing
sculpture. 3.
Fic. 8. Specimen from either Vendrest, Auvers or Acy, all Auversian (see p. 194 for explana-
(HVA), YK Sf.
Fics 9-10
Pseudocominella semicostata sp. nov. p- 196
Upper Eocene, Late Priabonian, Middle Headon Beds, Royden, Hampshire (Ed).
Both specimens referred to as Stvepsiduva armata (J. de C. Sowerby) by all previous authors.
Fic. 9. GG 19686/5. Juvenile shell. x5.
Fics toa, b. GG 19686/3. Larger shell showing gradual development of peripheral spines.
er
ie)
ine)
a
o
rh
2S)
ws
ee)
~
S
“a
~~
SS
By.
Bull.
PLATE 4
FIGs 1-3
Pseudocominella semicostata sp. nov. p. 196
Upper Eocene, late Priabonian, Middle Headon Beds.
All specimens referred to as Stvepsiduva avmata (J. de C. Sowerby) by all previous authors.
Fics 1-2. Whitley Ridge railway cutting, Brockenhurst, Hampshire (Ed ex Kk).
Fics ta, b. Holotype, GG 19687/1. x2.
Fic. 2. Paratype, GG 19687/2. 2°5.
Fic. 3. GG 19688/14, Brockenhurst, Hampshire, broken paratype showing columella. » 3.
FIGs 4-7
Desorinassa desori (Deshayes) p. 202
Palaeocene, Thanetian.
Fics 4-6. Jonchery-sur-Vesle, France.
Fics 4a, b. GG 19438/1 (L. Staadt Colin). x3. Fig. 4b. Slightly oblique side view to
show labial profile, posterior flaring of outer lip and subsutural platform.
Fic. 5. GG 19438/2 (L. Staadt Colin), oblique view of younger shell. x 4.
Fic. 6. GG19705/1 (Wr). Oblique view of even younger shell with weaker spiral sculpture
and showing naticoid protoconch comparable with that of Pseudocominella. 4.
Fics 7a, b. GG 19704, Thanet Sand, Reculver, Kent. (Wy), possibly the specimen referred
to by Cooper (1934 : 8) (see remarks, p. 203). x2.
Fics 8, 9
Desorinassa lata (Deshayes) p. 201
Palaeocene, Thanetian, Abbecourt and Bracheux, France.
(Deshayes Colin, EMP) with no indication as to which specimen comes from which locality.
Fic. 8. Lectotype, selected herein.
Fic. 9. Paralectotype selected herein. Both 1-67.
Bull. Br. Mus. nat. Hist. (Geol.)
PLATE 5
FIGs 1-3
Desorinassa williamsi sp. nov. p- 203
Palaeocene, Ypresian, Bognor Rock, foreshore at Bognor, Sussex.
Fics 1a, b. Holotype. GG19706 (Williams Colin). 2.
Fic. 2. Paratype. GG 19707 (Rev. A. Fuller Colin). Oblique view showing details of fine
sculpture on last whorl. x4.
Fic. 3. Paratype. GG 19708/1, probably from Bognor, history unknown, internal mould.
X1°6.
Fic. 4
Desorinassa ovata (Deshayes) p. 201
Lower Eocene, Cuisian, Cuise, France.
(Deshayes Colin, EMP) probably referred to by Deshayes, 1866 : 497. 4.
Fic. 5
Desorinassa acies (Watelet) p. 201
Lower Eocene, Cuisian, Sapicourt, France.
IScRNB sample IG 105901, fig. spec. no. 5600 I.S.T., referred to by Glibert (1963 : 66). 3.
Fics 6-11
Whitecliffia suturosa (Nyst) p. 204
Lower and Middle Oligocene, Belgium.
Fics 6-8. (de Looz Colln., IRScNB) sample IG 5002, Lower Oligocene (Lattorfian = Lower
Tongrian) Grimmertingen specimens referred to by Glibert & de Heinzelin (1954 : 382).
Fic. 6. Fig. spec. no. 5602 I.S.T., tumid specimen. 2:7.
Fics 7a, b. Fig. spec. no. 5603 I.S.T., typical specimen. Fig. 7a, 2-7; Fig. 7b, oblique
view to show protoconch and spiral sculpture on ramp. X3°5.
Fics 8a, b. (IRScNB). Fig. spec. no. 4070 I.S.T., Grimmertingen, specimen referred to as
Babylonia (Peridipsaccus) avchambaulti (Meunier) by Glibert & de Heinzelin (1954 : 368, 382,
jl, Hy ste Oe KB
Fics 9-11. Middle Oligocene, Lower Rupelian.
Fic. 9. GGu19721. (de Koninck Colin), Klein Spauwen. x3.
Fies 10, 11. (Bosquet Colln., IRScNB) sample IG 4285, Berg. Specimens referred to by
Glibert & de Heinzelin (1954 : 392, 393).
Fic. 10. Fig. spec. no. 5604 1.8.T. 2:5.
Fic. 11. Fig. spec. no. 5605 1.8.T. 2:5.
Bull. By. Mus. nat. Hist. (Geol.) 23, 3 PLATE
5
PAA EG:
Fics 1-4
Whitecliffia suturosa (Nyst) p. 204
Fics 1a, b. GG 19717 (Ed). Upper Eocene, late Priabonian, Middle Headon Beds, Royden,
Hampshire, specimen referred to as Comnella ventvicosa Edwards MS by Judd (1880 : 154)
and by Keeping & Tawney (1881 : 116), and possibly as Buccinum suturosum (Nyst), by von
Koenen (1867 : 83). 3.
Fics 2-4. Lower Oligocene, Lattorfian, Latdorf, Germany (Ed).
Fics 2a, b. GG 19720/1. Specimen with relatively strong spiral grooving. 3.
Fic. 3. GG 19720/2. Rear view of specimen with very weak spiral sculpture. x 3.
Fic. 4. GG 19719/2. Oblique view showing protoconch, form of suture and sculptural
details. 4.
Fics 5-10
Whitecliffia tumida sp. nov. p. 206
Upper Eocene, late Priabonian, Middle Headon Beds (Venus Bed), Whitecliff Bay, Isle of Wight.
Fics 5a, b. Holotype. GG 19722/1 (Cu). x4.
Fics 6-10. Paratypes. (All (Cz) except fig. 9).
Fics 6a, b. GG 19722/3. Specimen with inflated body whorl, Fig. 6a x3; Fig. 6b, showing
collabral colour banding. 2:7.
Fic. 7. GG19722/6. Showing broad colour band on ramp, with narrow one belowit. 2-7.
Fics 8a, b. GG19724. Small individual lacking strong axial sculpture on early whorls,
Fig. 8a, 5:5; Fig. 8b, x5.
Fic. 9. (Sedg. Mus., K&T) C.29305. 3.
Fic. 10. GG19722/2. Juvenile shell showing paucispiral protoconch. 11 (stereoscan).
ine)
73},
(Geol.)
Hist
nat.
Mus.
Bull. Br.
PALIN, 7
Fics 1-5
Colwellia flexuosa (Edwards) p. 210
Upper Eocene, Late Priabonian, Middle Headon Beds, Isle of Wight.
Fics ta, b. Lectotype. GG 19709/1 (Ed). Colwell Bay. x4.
Fics 2a-e. Paralectotype. GG 19709/3 (Ed). Colwell Bay, exceptionally large specimen;
2a, <3; 2b, oblique view of fasciole, x 4; 2c, side view of outer lip, showing changes in growth
lines with development of siphonal notch and peripheral sinus (compared with fig. 4), x3;
2d, 2e, ultra-violet photographs showing colour banding. 2.
Fics 3a-c. Paralectotype. GG 19709/4 (Ed). Colwell Bay, 3a, x5; 3b, 24 (stereoscan) ;
3c, X27 (stereoscan).
Fic. 4. GG 19711 (Nu), Venus Bed, Colwell Bay, side view showing abrupt transition from
relatively orthocline to very flexuous growth lines (compare with fig. 2c). 3.
Fics 5a, 5b. GG 19713 (Cu), Venus Bed, Whitecliff Bay, juvenile specimen. 6.
FIGs 6-7
Colwellia auversiensis (Deshayes) p. 208
Middle Eocene, Auversian, Auvers, Paris Basin.
(Deshayes Colln, EMP), specimens probably belonging to the type series of Buccinum auversiense
Deshayes (1865 : 498; 1866, pl. 94, figs 1-3), both x 2:5.
Fics 8, 9
Colwellia bretzi (Weaver) p. 208
Upper Eocene, Lower Cowlitz Formation, Vader, Washington State, U.S.A.
Fig. 8. GG 12783/1. (Wr) x5.
Fic. 9. GG 12783/2 (Wr) 16 (stereoscan).
Fic. 10
?Colwellia sp. p. 209
(Bruce Martin Colln, CAS., Loc. 244). Upper Eocene, Tejon Group, Tejon Quadrangle,
Kern County, California, U.S.A., east bank of Live Oak Creek about three quarters of a mile from
its mouth or from the edge of the San Joaquin Valley, and about three miles due east of the
mouth of Grapevine Canyon. Specimen referred to and figured as Molopophorus striatus
(Gabb) by Dickerson (1915 : 65, pl. 8, fig. 6 given as fig. 7 in explanation thereof). x6.
(See also pl. 8.)
Fic. 11
Colwellia antiquata (Gabb) p. 208
(U.Cal. 15939, Loc.A-819), specimen described and figured as Molopophorus antiquatus
(Gabb) by Vokes (1939 : 142, pl. 19, fig. 2), oblique view showing protoconch. x6.
Bull. By. Mus. nat. Hist. (Geol.) 23, 3
IRE AN ID, 78)
FIGs I, 2
Molopophorus striatus (Gabb) p- 209
Type species of Molopophorus, Upper Eocene, Tejon Formation, Tejon, California.
Fic. ta, b. Lectotype. (ANSP 4249) selected Stewart (1926 : 389, pl. 29, fig. 14), Ia x9;
Ib x6
Fic. 2. (U.Cal. 7182) Showing sculpture on spire. 9.
FIGs 3, 4
Brachysphingus gibbosus Nelson p. 209
Palaeocene, Martinez Formation, Ventura County, California (ex U.Cal.).
Fic. 3. G72721. Oblique view showing growth lines reversed at suture. 1-7.
Fic. 4. GG 12834/1. Section showing absence of columellar plait. x2.
Fics 5-7
Keepingia gossardi (Nyst) p- 212
Middle Oligocene, Rupelian.
Fias 5a, b. GG 19739/1. Etampes, France. 2:5.
Fic. 6. (EMP) unlocalised young shell. 16 (stereoscan).
Fic. 7. GG19748/1. ?Klein Spauwen (type locality), Belgium. x3.
Fic. 8
Keepingia bolli (Beyrich) [Ds Bur
GG 19749/1 (Wr), Upper Oligocene, Chattian, Almatil near Cassel, Germany. x 2.
FIG. 9
Keepingia cassidaria (Sandberger) p. 212
GG 19733/1 (Wr), Upper Oligocene, Chattian, Cyrenmergel, Sulzheim, Germany. 4.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 3
PLATE 9
FIGs 1-7
Thanetinassa bicorona (Melleville) p. 213
Palaeocene, Thanetian.
Fics 1-4. Chalons-sur-Vesle, France. (L. Staadt Colin.)
iGai GGO357)/Th ase
Fic. 2. GG 19357/3. 4.
Fic. 3. GG 19357/2. Oblique view showing ‘stromboid-like’ notch in outer lip. 4.
Fic. 4. GG 13357/4. Section showing form of columella. x 3.
FIGs 5-7
Corbula Bed, Upper Thanet Sands, Bishopstone Gap, Herne Bay, Kent.
Fig. 5. GGu19741/1 (Wr). x4.
Fic. 6. GG19744 (Ru). x3.
Fics 7a, b. G 60746 (J. E. Cooper Colln.). 3.
Fics 8a, b
Buccinum montense Briart & Cornet Pp. 215
G 71604 (Chavan Colin.). Palaeocene, Montian, Belgium.
Fie. 8a. XQ.
Fic. 8b. Oblique view of fasciole. 15.
Fic. 9
Nassarius arcularius (Linné) p. 181
Type species of Nassarius, Recent, Indo-Pacific. BMZD.19716 (Hugh Cuming Colln)
I. of Ticao,,Philippines, on reef, figd. Reeve (1853), pl. 4, fig. 25a). 1-5.
Fic. 10
Strepsidura turgida (Solander) p. 190
Type species of Stvepsiduva, GG 19750 (St/B), Upper Eocene, Bartonian,
Barton Beds, Barton-on-Sea, Hampshire. x2.
Fics t1a, b
Cominella testudinea (Lamarck) p. 180
Type species of Cominella, BMZD.1844.7.29.36 New Zealand.
iG elas G2
Fic. 11b. Oblique view of fasciole. 5.
Bull. Br. Mus. Hist. (Geol.) 23, 3
A LIST OF SUPPLEMENTS
TO THE GEOLOGICAL SERIES
OF THE BULLETIN: OF
THE BRITISH MUSEUM (NATURAL HISTORY)
. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya.
Pp. 213; 30 Plates; 2 Text-figures. 1965. £6.
. Er-Nacear, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper
Cretaceous—Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley,
Egypt, U.A.R. Pp. 291; 23 Plates; 18 Text-figures. 1966. £10.
. Davey, R. J., DowniE, C., SARGEANT, W. A. S. & WitiiaMs, G. L. Studies on
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248; 28 Plates; 64 Text-
figures. 1966. £7.
. APPENDIX. Davey, R. J., Downtz, C., SARGEANT, W. A. S. & WILLIAMS, G. L.
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24.
1969. 8op.
. Ettiott, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of the
Middle East. Pp. 111; 24 Plates; 17 Text-figures. 1968. {5.124.
. Ruopes, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni-
ferous) Conodont faunas, and their value in local and continental correlation.
Pp. 315; 31 Plates; 92 Text-figures. 1969. £11.
. Cuitps, A. Upper Jurassic Rhynchonellid Brachiopods from Northwestern
Europe. Pp. 119; 12 Plates; 40 Text-figures. 1969. £4.75.
. Goopy, P. C. The relationships of certain Upper Cretaceous Teleosts with
special reference to the Myctophorids. Pp. 255; 102 Text-figures. 1969. £6.50.
. OWEN, H. G. Middle Albian Stratigraphy in the Paris Basin. Pp. 164;
3 Plates; 52 Text-figures. 1971. {6.
. Sippigur, Q. A. Early Tertiary Ostracoda of the family Trachyleberididae
from West Pakistan. Pp. 98; 42 Plates; 7 Text-figures. 1971. £8.
anlle
igh Printed in England by Staples Printers Limited at their Kettering, Northants establishment
ar
K
|
,
|
aay Hise
DEVONIAN “g
(NATURAL HISTORY)
| Vol. 23 No.
PALAEOZOIC CORAL FAUNAS FROM
VENEZUELA, II. DEVONIAN AND
CARBONIFEROUS CORALS FROM THE
SIERRA DE PERIJA
BY
COLIN I SCRUTION-
Department of Geology
The University
Newcastle upon Tyne
Pp. 221-281; 10 Plates; 12 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 23° No. 4
KONDON =: 1973
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, 1s
issued in five series corresponding to the Departments
of the Museum, and an Histoncal series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar ‘year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 23, No. 4 of the Geological
(Palaeontological) series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation
Bull. Br. Mus. nat. Hist. (Geol.)
© Trustees of the British Museum (Natural History), 1973
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 31 May, 1973 Price £4°35
PALDAEOZOIC CORAL FAUNAS FROM
DEVONIAN AND
VENEZUELA, IL
CARBONIFEROUS CORALS FROM THE
SIERRA DE PERIJA
By C, Lf. SCRUTLON
CONTENTS
INTRODUCTION
ACKNOWLEDGEMENTS
STRATIGRAPHY OF THE Momma SiR | DE earn :
AGES AND RELATIONSHIPS OF THE CORAL FAUNAS
(a) Rio Cachiri Group
(i) Cano Los Guineos Boemetion
(ii) Cano Grande Formation
(iii) Cano del Oeste Formation
(b) Rio Palmar Formation .
SYSTEMATIC DESCRIPTIONS
RUGOSA .
Family LINDSTROEMIIDAE Potts.
Genus Steveolasma Simpson
S. sp.
Genus Grapeophyihon NEG
2S. sp. :
Genus Syvingaxon Tendotroen :
S. sp.
Family HADROPHYLLIDAE iicholean
Genus Hadvophyllum Edwards & Haime
H. sp.
Family HAPSIPHYLLIDAE Grabau
Genus Amplexizaphrentis Vaughan
A. sutherland sp. nov.
Family STREPTELASMATIDAE Nicholson
Genus Bowenelasma nov.
B. typa sp. nov. . 2
B. breviseptata sp. nov.
Genus Briantelasma Oliver
B. oliveri sp. nov. 5
Genus Heterophrentis Billings .
H. (H.) venezuelensis (Weisbord)
H. (H.) simplex (Hall)
Family CYATHOPHYLLIDAE Dana
Genus Heliophyllum Hall :
H. haili Edwards & Haime
H. wellsi sp. nov.
Genus Cylindrophyllum Simpson
C. elongatum Simpson .
Page
224
226
226
229
230
230
231
232
232
232
233
233
233
233
234
235
236
236
237
237
237
239
239
240
242
242
243
245
247
248
251
252
254
255
256
257
261
262
263
224 PALAEOZOIC CORAL FAUNAS
Family DURHAMINIDAE Minato & Kato F : : 264
Genus Durhamina Wilson & Langenheim : : 2 265
?D. sp. é : : ; ; : é : 265
D. sp. nov. : : . : ° : : 266
Family CYSTIPHYLLIDAE Edwards & Haime 5 : 269
Genus Plasmophyllum Dybowski : : c 269
P. secundum americanum (Edwards & Haime) : 269
IZ, 80. : : : d : : é 271
TABULATA ‘ : ° : : : c : 3 PP
Family FAVOSITIDAE . ; : : s : 5 272
Genus Favosites . ‘ : : 5 d a ‘ 272
F.. venezuelensis (Weisbord) . : c : c 273
F. arbuscula Hall : : j : c 5 274
VI. REFERENCES . ; : é : : : : 5 275
VII. AppENDIX. COMPOSITION AND LOCATION OF SAMPLES . : : 280
SYNOPSIS
Rugose and tabulate corals of Middle Devonian, and Lower Pennsylvanian age are described
from localities in the northern part of the Sierra de Perija, western Venezuela. The Palaeozoic
stratigraphy of the area is briefly reviewed and the ages and relationships of the coral faunas are
discussed. The new taxa Bowenelasma typa gen. et sp. nov., B. breviseptata sp. nov., Amplexi-
zaphrentis sutherland sp. nov., Briantelasma oliveri sp. nov. and Heliophyllum wellsi sp. nov.,
as well as species of Steveolasma, ?Stewartophyllum, Syringaxon, Heterophrentis (H.), Cvlindro-
phyllum, Durhanuna, Plasmophyllum and Favosites are described.
I. INTRODUCTION
TuIs is the second of two papers describing rugose and tabulate corals from the
Palaeozoic rocks of the Venezuelan Andes. The first paper (Scrutton 1971) dealt
with Silurian and Permo-Carboniferous corals from the Mérida Andes. This paper
describes material of Devonian and Carboniferous ages from localities in the northern
Sierra de Perija (see Fig. 1), some 300 km to the north-west and close to the
Venezuelan-Colombian border.
The bulk of the material was collected during 1959 by Dr J. M. Bowen (then
Compania Shell de Venezuela) but the opportunity is taken here to revise in addition
the small samples of corals collected by C. W. Yeakel, P. W. McFarland and R. A.
Liddle in 1924 (described by Weisbord 1926) and by R. A. Liddle in 1942 (described
by Wells 1943) which came from the same or nearby sections. The richest fauna is
one of upper Onesquethaw age (considered equivalent to the early Middle Devonian)
in terms of the eastern North American stages (see Oliver 1968, fig I), represented
by 14 of Bowen’s samples and including most if not all of the older collections.
A further three of Bowen’s samples are also Devonian in age. The Devonian
corals, which show very strong affinities with coral faunas of the same age in eastern
North America, include sixteen species belonging to eleven genera; one genus and
four species are new. Three of Bowen’s samples contain corals which, with some
supporting evidence, indicate probable Carboniferous ages. Three species, two of
them new, belonging to two genera are involved. The material is housed in the
Department of Palaeontology, British Museum (Natural History).
FROM THE SIERRA DE PERIJA, VENEZUELA 225
Yee
rs
Ke
i?
CARBONIFEROUS
DEVONIAN
+ +
Granite
E
Fossiliferous Somple Loc
PRE
DEVONIAN
Bese
ia
ny yay
EZ YEX.
Were,
“ SO Kits es TG
~~ ey
Fic. 1. Geological map of the northern Sierra de Perij4 showing sample locations (for
further information see Appendix). The area covered in detail in Fig. 2 is outlined.
Based on maps supplied by J. M. Bowen.
226 PALAEOZOIC CORAL FAUNAS
Il ACKNOWLEDGEMENTS
The author is most grateful to Dr J. M. Bowen (Shell International Petroleum
Corporation), who collected most of the material, for discussions on the stratigraphy
of the northern Sierra de Perija and for permission to use information not yet pub-
lished. Dr Bowen kindly made available the maps upon which Text-figs 1 and 2
are based. Acknowledgement is due to Compafiia Shell de Venezuela for permission
to describe Bowen’s collection which they have presented to the British Museum
(Natural History), and to Dr K. V. W. Palmer (Paleontological Research Institution)
who has facilitated the loan of material in her care.
Dr W. J. Oliver Jr. (U.S. Geological Survey) has kindly read the systematic
section of the manuscript and has contributed valuable discussion and information
concerning eastern North American coral faunas of Devonian age. Dr J. G. Johnson
(Oregon State University) has made valuable comments on the Devonian brachiopod
faunas associated with the corals.
R. F. Wise and P. J. Green (British Museum (Natural History)) have respectively
prepared and photographed material for this paper. E. Lawson (Department of
Geology, University of Newcastle upon Tyne) drafted the originals of Text-figs 1-3,
5, 6, 8, 9 and II.
III. PALAEOZOIC STRATIGRAPHY OF THE
NORTHERN SIERRA DE PERIJA
The basic framework of the Palaeozoic stratigraphy of the northern Sierra de
Perija was first established by R. A. Liddle and his co-workers during expeditions
along the course of the Rio Cachiri in 1924 and 1942. The results were published in
some detail in Liddle, Harris & Wells (1943). Subsequently a number of papers
have been published referring to the area (including Sutton 1946, Hea & Whitman
1960 and Miller 1962) and knowledge of the Devonian sections was summarised by
Weisbord (1968). These contributions, however, did not substantially modify the
stratigraphical results of Liddle’s expeditions. The most significant advance in
the understanding of the Palaeozoic successions is due to J. M. Bowen’s work in the
northern Sierra de Perija during 1959 when the material described in this paper was
collected. Both the geological map (Figs I, 2) and the stratigraphic successions
(Fig. 3) reproduced here are based on the results of his work (Bowen 1972).
From the first descriptions of the rich macrofauna yielded by the oldest fossiliferous
rocks of the area, published by Weisbord (1926), the strong resemblance to faunas of
the upper Lower Devonian to lower Middle Devonian Onesquethaw interval in
eastern North America was recognized. Liddle, in Liddle, Harris & Wells (1943)
later distinguished three formations of Devonian age in the Rio Cachiri section, the
Cafio Grande Formation, the Cafio del Oeste Formation and the Campo Chico
Formation in order of decreasing age, grouped together asthe Rio Cachiri Series (Cachiri
Group of Sutton 1946 : 1634). The rich invertebrate fauna was confined to the
Cano Grande Formation and Liddle described both the other formations as un-
fossiliferous (Liddle, Harris & Wells 1943 : 286, 289), although as Weisbord (1968 :
FROM THE SIERRA DE PERIJA, VENEZUELA 227
221) pointed out, Liddle records in his list of samples (Liddle, Harris & Wells 1943 :
313, sample 37) Heliophyllum halli and ‘Heterophrentis venezuelense’ apparently in situ
on the Cafio del Oeste Formation outcrop in the Cafio del Oeste section. In view of
Liddle’s statement in the text and the fact that Bowen’s more extensive collections
from the Cano del Oeste Formation do not contain these corals, it seems likely that
sample 37 was in fact a displaced raft from the Cafio Grande Formation such as are
known to occur on this section. An even more anomalous situation, however,
surrounds the original collection of fossils described by Weisbord (1926) and collected
in 1924 by C. W. Yeakel in the company of P. W. McFarland and Liddle. According
to Liddle (Liddle, Harris & Wells 1943, map) and Weisbord (1968 : 217) the collection
was taken in the Cafio del Norte about 4-5 km north of the Rio Cachiri from the upper
part of the Cafio Grande Formation. That the fauna is of Cafio Grande aspect can
hardly be doubted, but Bowen (1972; Fig. 4) records no outcrop of this formation
in, or even close to the Cafio del Norte. In fact the difference between Liddle’s
and Bowen’s maps of this particular section is most striking. Weisbord (1926 : 223)
does record that ‘the majority of our fossils were collected from float’ and this may be
the explanation. Alternatively, it is possible, in this sort of country, that the site of
the collection was wrongly located (fide Bowen). Unfortunately no definite solution
ed
=—- ~~ ~ ~
F
t
Lt
|
t
l
et
l
t
Campo Chico Formation
Cano del Oeste Formation
Cano Grande Formation
Pre- Devonian
metamorphic rocks
on ; 0 100 200 300 400
©3149 Fossiliferous Sample Locality 01 ___1___i_
m
Fic. 2. Geological map of the upper reaches of the Rio Cachiri, Sierra de Perijé showing
sample locations (for further information see Appendix). The area covered here is
indicated in Fig. 1. Based on maps supplied by J. M. Bowen.
228 PALAEOZOIC CORAL FAUNAS
to this problem seems possible at the moment and in this paper Yeakel’s collection
is considered as most likely to have originated from a displaced raft or possibly
faulted slice of Caio Grande Formation in the Cafio del Norte.
Bowen (1972) has recognized a fourth division of the Devonian Rio Cachiri
Group in the Sierra de Perija. This, the Caio Los Guineos Formation of the Rio
Lajas area, he considers to be lower in the sequence than the Cafio Grande Formation
although the two formations do not crop out in the same sections. They have a
very similar macrofauna and both formations are dated as Lower Devonian (Emsian,
equivalent to the middle Onesquethaw) mainly on the basis of brachiopods although
an early Middle Devonian age (upper Onesquethaw) is considered more likely here
(see p. 230-31). Bowen has also collected a fossil suite dominated by brachiopods
from the Cafio del Oeste Formation, previously not satisfactorily dated, from which
he deduces a lower Middle Devonian age. Finally, the highest, Campo Chico For-
mation has yielded palynomorphs indicating a Middle to Upper Devonian (probably
Frasnian) age (Bowen 1972). The Devonian corals, discussed in detail in the
following sections, come mainly from the Caio Grande Formation, for which they
suggest an upper Onesquethaw age.
Liddle (Liddle, Harris & Wells 1943 : 290) assigned all the Permo-Carboniferous
‘red-bed’ sequence and overlying crinoidal limestones in the Rio Cachiri section to
the Palmarito Formation of Christ (1927). Bowen (1972) has considerably modi-
fied this succession, recognizing in it the lithological units described by Arnold (1966)
from the Permo-Carboniferous of the Mérida Andes, about 300 km to the south-east.
The basal development of red-beds Bowen assigns to the SabanetaGroup. Equivalent
lower and upper divisions are recognized by both Arnold and Bowen which the
RIO CACHIRI RIO PALMAR
RIO PALMAR FM.
CARBONIFEROUS woo CANO INDI
C. DEL NOROESTE FM. | CANO DEL NOROESTE FM.
Nec
CAMPO CHICO FM. CAMPO CHICO FM.
DEVONIAN
MIDDLE
eccal
U
PRE-DEVONIAN PERIJA FM GRANITE OF RIO LAJAS
Fic. 3. Devonian and Carboniferous successions in the northern Sierra de Perija
(after Bowen 1972). For comments see text.
FROM THE SIERRA DE PERIJA, VENEZUELA 229
latter erects as new formations. The Cano del Noroeste Formation below (equiva-
lent to Arnold’s lower clastic member) contains a fauna of fusulinids, brachiopods
and palynomorphs indicating a lower Middle Carboniferous Namurian to Lower
Westphalian age (Lower Pennsylvanian) according to Bowen (1972) whilst the
Cafio Indio Formation above is unfossiliferous. The age of the Cano Indio Formation
is fixed as low in the Pennsylvanian, however, by the dating of the overlying Rio
Palmar Formation. This contrasts with the as yet poorly dated Sabaneta Group
(Sabaneta Formation of Arnold 1966) of the Mérida Andes, the type area, where
fossiliferous marine horizons are not known. Here, Arnold (1966 : 2371-2372)
records a possible Permian age for the higher parts of the Sabaneta Group on the
evidence of spores.
The Rio Palmar Formation is a thick carbonate sequence conformably succeeding
the Sabaneta Group. It is dated late Middle Carboniferous (late Lr. Pennsylvanian)
on the basis of fusulinids and palynomorphs (Bowen 1972) and is the only level
in the Permo-Carboniferous from which corals are described in this paper. This
new formation is not represented in the Mérida Andes (Bowen 1972) where
Arnold (1966 : 2373, 2377) records the Sabaneta Group grading upwards through a
transitional zone into the Palmarito Formation. In the northern Sierra de Perija,
the typical facies of the Palmarito Formation, which is absent from the Rio Cachiri
area but crops out further south, comes in unconformably above the Rio Palmar
Formation. According to Bowen (1972), the whole of the Upper Pennsylvanian
is apparently missing here, with the Palmarito Formation yielding Wolfcampian
fusulinids. In the Mérida Andes, however, the rich marine fauna of the Palmarito
Formation suggests that the late Pennsylvanian as well as the early Permian may be
represented (Arnold 1966 : 2378; Scrutton 1971 : 189). Thus it would appear that
the distinctive carbonate facies of the Rio Palmar Formation is the lateral equivalent
of rocks of upper Sabaneta or, less likely, lower Palmarito type to the south-east.
Between the Palmarito Formation and the Lower Cretaceous Rio Negro Formation,
Bowen (1972) has described a succession of pyroclastics, lavas and terrestrial
sediments of ?Permian to Jurassic age. As no material from these younger rocks
is described in this paper, they are not considered further here.
IV. AGES AND RELATIONSHIPS OF THE CORAL FAUNAS
The extent of previous work on South American Palaeozoic coral faunas was
briefly reviewed by Scrutton (1971 : 190). Two papers refer to Devonian corals
from the northern Sierra de Perija, those of Weisbord (1926) and Wells (1943).
Weisbord described the material collected by C. W. Yeakel in 1924 and Wells revised
the corals on the basis of further collections made by R. A. Liddle in 1942. In both
collections, the corals were all from the Cafio Grande Formation of the Rio Cachiri area.
J. M. Bowen’s much more extensive collections from a wider area in the northern
Sierra de Perija have more than doubled the species list for the Cafio Grande For-
mation as well as yielding small coral faunas from three further horizons. The
compositions of the faunas are listed in Table 1 and are discussed below in ascending
stratigraphical order.
230 PALAEOZOIC CORAL FAUNAS
TABLE I
Composition of coral faunas from the Rio Cachiri Group (Devonian) and
Rio Palmar Formation (Carboniferous) of the Sierra de Perija.
Rio Cachiri Group
Cafio Los CafioGrande Cajio del Palmar
Species Guineos Fm. Fm. Oeste Fm.
Stereolasma sp.
Syvingaxon sp.
Heliophyllum hall
H. wellsi
Heterophrentis (H.) simplex x
H. (H.) venezuelensis
Plasmophyllum secundum americanum
P. sp.
Acinophyllum vermetum
Briantelasma oliveri
Bowenelasma typa
B. breviseptata
Favosites venezuelensis
F. arbuscula
Cylindrophyllum elongatum — ? —
?Stewartophyllum sp. x
Hadrophylium sp. x
Amplexizaphrentis sutherlandi x
?Durhamina sp. x
Durhemina sp. nov. ?
x Xx X
DS ON ON OM OS PM OS ESD AIDS OS OOM
(a) Rio Cachiri Group
(i) Cato Los Guineos Formation. A small fauna of four different corals only is
available from this formation. Its composition suggests affinities with Middle
Devonian coral faunas of eastern North America where Heliophyllum halli Edwards
& Haime in particular is characteristic of the Onondaga Limestone and Hamilton
Group of New York and their equivalents elsewhere. Steveolasma appears to have a
similar, though less well documented range and Heterophrentis (H.) simplex (Hall)
has been recorded so far only from horizons of Hamilton age in New York and the
Ohio Valley (see Oliver 1968, fig. 1 for a correlation chart of the eastern North
America Devonian).
Bowen (1972) lists other fossils, principally brachiopods, from the Cano Los
Guineos Formation which he takes to indicate an upper Lower Devonian (Emsian)
age. There is some doubt, however, as to the reliability of this brachiopod fauna
in distinguishing between a late Lower and an early Middle Devonian age (J. G.
Johnson, pers. comm.). This matter is discussed more fully in the following section
on the Cafio Grande Formation which has yielded similar but richer coral-brachiopod
faunas. In view of the ambiguity of the brachiopod evidence, the affinities of the
coral fauna suggest that an upper Onesquethaw age (early Middle Devonian) is more
likely than a late Lower Devonian age for this formation.
FROM THE SIERRA DE PERIJA, VENEZUELA 231
(ii) Cano Grande Formation. The principal Onondaga—Hamilton species
present in this much richer coral fauna are Heliophyllum halli, Heterophrentis (H.)
simplex, Plasmophyllum secundum americanum (Edwards & Haime) and Favosites
arbuscula Hall. In addition, Acinophyllum vermetum (Weisbord)! is considered to
agree in all its essential characteristics with A. stramineum (Billings) from the lower
part of the Onondaga Limestone of Ontario (Oliver, in preparation) and Cylindro-
phyllum elongatum Simpson, which comes either from the Cafio Grande Formation
or the overlying Cafio del Oeste Formation, is also found at the same horizon in New
York. This evidence would clearly suggest an upper Onesquethaw (lower Middle
Devonian) age for the Cafio Grande coral fauna. Also present, however, is a new
species of Briantelasma. This genus has previously been recorded from rocks of
Silurian (? Wenlock) and Lower Devonian (Helderberg) age in eastern North America
(Oliver 1960a, b, 1963) and is the only element in the coral fauna not previously
recorded from the Middle Devonian. The new genus Bowenelasma provides no
direct evidence but its general characteristics relate it more to Briantelasma and
Silurian streptelasmatids rather than to typical Middle Devonian members of the
family. It should be stressed, however, that there is no sign of typical eastern North
American middle Onesquethaw corals in the Venezuelan fauna.
Bowen (1972) lists a brachiopod fauna from the Cano Grande Formation, very
similar to that in the Cafio Los Guineos Formation, which he takes to indicate a
late Lower Devonian age. He infers that this is supported by palynomorphs.
As previously noted (p. 230), however, Dr J. G. Johnson (pers. comm.) doubts
that confidence can be placed in the brachiopod fauna at the moment to distinguish
between a middle and an upper Onesquethaw age (that is, late Lower Devonian
and early Middle Devonian as currently understood in North America). In particular,
Johnson notes that Eodevonaria has been collected from rocks of upper Onesquethaw
age, and he regards the upper limits of the range of Leptocoelia to be uncertain.
Both genera have been taken to indicate a Lower Devonian horizon. He also draws
attention to the occurrence of Tropidoleptus (Morales 1965) and common Pentagonia
and Spinulicosta, all more suggestive of an upper Onesquethaw than a Lower
Devonian age, in the Floresta fauna of Colombia, with which the Venezuelan faunas
have much in common. Johnson thinks it likely that these South American
occurrences represent an overlap of late Lower and early Middle Devonian forms.
Dr L. Nijssen (pers. comm.), who made the palynological determinations, also has
reservations about distinguishing between an Emsian and an early Eifelian age on
the basis of the flora.
Although much revision of the North American coral faunas is necessary, Oliver
(1968 : 741) clearly indicates that an important break in the sequence of coral
assemblages there occurs between the middle and upper Onesquethaw. In this light,
the strong affinities of the coral fauna with faunas of upper Onesquethaw age in eastern
North America suggests that the Cafio Grande Formation should be dated upper
Onesquethaw (early Middle Devonian) rather than middle Onesquethaw at the
present time.
1 As Dr W. J. Oliver Jr. has a review of this species in preparation it is not described in the systematic
section of this paper.
232 PALAEOZOIC CORAL FAUNAS
The affinities between the Venezuelan and eastern North American coral faunas
indicates that the two areas must have formed part of the same province in early
Middle Devonian time. The presence of Bowenelasma and some species endemic to
Venezuela suggests that north-south migration may have been partially restricted
in some way, although these corals may be recorded from eastern North America in
the future.
(ii) Cano del Oeste Formation. This formation appears to be poor in corals
although it has provided a fairly varied fauna of brachiopods and molluscs which are
listed by Bowen (1972). One specimen each of ?Stewartophyllum sp. and
Hadrophyllum sp. indicate a tentative Middle Devonian age, whilst the brachiopods
have been more precisely dated as early Middle Devonian.
Although the specimen of Cylindrophyllum elongatum, which comes from a slipped
block, is listed by Bowen (1972) with the Cafio del Oeste fauna, Bowen has in-
formed the writer that the block compared lithologically with the Cano Grande
Formation outcropping further up the Cafio Pescado and may well have come from
that formation. If the block did originate from the Cafio del Oeste Formation,
however, then it would be from near the base, whereas the rest of the fauna, pro-
viding evidence for the age, derives from the very top of the formation.
(b) Rio Palmar Formation
This formation has also yielded two corals among a rather limited invertebrate
fauna. Amplexizaphrentis sutherlandi sp. nov. and ?>Durhamina sp. indicate a general
Carboniferous age and compare most closely with North American cordilleran
species. The other faunal elements provide more precise information and indicate
a late Lower Pennsylvanian age.
Durhamina sp. nov. is known only from a loose block whose stratigraphic relation-
ships are obscure. Of the known stratigraphy, it is most likely to have come from
the Rio Palmar Formation which is where Bowen (1972) lists it as “durhaminido
gen. and sp. nov.’. Possible Mississippian brachiopods were reported from the
same block, however, and it may represent an horizon as yet not recognized in situ
in the stratigraphic sequence.
V. SYSTEMATIC DESCRIPTIONS
The terminology used in the following descriptions is that proposed by Smith
(1945 : 4-9) and Moore, Hill & Wells (1956) unless otherwise indicated.
All the material collected by Bowen is now housed in the Department of Palaeon-
tology, British Museum (Natural History). The registered numbers for these
specimens are prefixed R and in each case the number of the sample in which the
specimen was collected is given in parentheses after the registered number. The
locations of Bowen’s samples are plotted on Text-figs 1 and 2 and full details are
given in the Appendix.
FROM THE SIERRA DE PERIJA, VENEZUELA 233
The sources of other material described below are indicated by the following
abbreviations: PRI—Paleontological Research Institution, Ithaca, N.Y.; NYSM—
New York State Museum, Albany, N.Y.
Order RUGOSA Edwards & Haime 1850
Suborder STREPTELASMATINA Wedekind 1927
Superfamily CYATHAXONIICAE Edwards & Haime 1850
Family LINDSTROEMIIDAE Poéta 1902
1971 Lindstroemiidae; Scrutton: 192, cum syn.
Discussion. The family concept was reviewed by Scrutton (1971 : 192).
Genus STEREOLASMA Simpson 1900
1g00 Steveolasma Simpson: 205.
1941 Steveolasma Busch: 395.
1949 Steveoelasma Stumm: 7.
1962 Steveolasma Stumm: 234.
1965 Steveolasma Stumm: 14.
DiacGnosis. Small ceratoid to trochoid corals. Major septa fuse in the axis to
form, with or without additional sclerenchyme, a prominent axial pillar. Minor
septa short, usually contratingent. Peripheral stereozone narrow. Cardinal
fossula narrow, poorly developed. Tabulae complete or incomplete, strongly
arched with a flat or slightly depressed axial area. No dissepiments.
TYPE SPECIES (by original designation). Stveptelasma rectum Hall 1876, pl. 10,
figs I-13 in part ?=Strombodes? rectus Hall 1843 : 210, text-fig. 87, 5 on p. 209,
and no. 48, fig. 5 on p. 44 of tables. Middle Devonian, Hamilton Group; western
New York State, U.S.A.
Discussion. Reference should be made to Stumm (1949: 7) and Stumm &
Watkins (1961 : 445) for comments on and a redescription of Steveolasma rectum.
Stereolasma sp.
(Pl. 1, figs 6, 7)
MATERIAL. R45129 (3849). Cano Los Guineos; Caio Los Guineos Fm., early
Middle Devonian (upper Onesquethaw).
DESCRIPTION. Small ceratoid coral, incomplete.
In the late neanic stage the mean diameter is II mm with 25 major septa. The
lumen is completely infilled by laterally contiguous wedge-shaped major septa merging
in the axial area with a core of sclerenchyme 3 mm in diameter. This core may be
formed by irregularly intertwined septal ends but the structure is not clear. There
are signs of rudimentary minor septa as wedges about I mm long between the major
septa.
234 PALAEOZOIC CORAL FAUNAS
In the ephebic stage (Pl. 1, fig. 6) there are-25 major septa at a mean diameter
of 14:5 mm. The major septa, except the cardinal septum, are 0-6—0-75 mm thick
where they are very slightly waisted one-third of the way along their length from the
periphery. Thecardinalseptumis notably thinnerato-4mm. Theseptaare thickest
about two-thirds of their length to the axis where they become laterally contiguous.
From this point they taper smoothly into the axis in a slight counter clockwise vortex
maintaining contact with their neighbours. The septa are longest in the alar area
where they all but meet in the axis. The counter and cardinal septa are shorter,
however, each reaching only three-quarters of the radius, leaving a narrow axial
gap about 4:5 mm long and 0-5 mm wide in the otherwise solid oval core of fused
septal ends, 7 mm x 6 mm in size. The peripheral ends of the septa alternate
with contratingent minor septa in a peripheral stereozone I-1:2 mm thick. The
minor septa generally appear as short thorny projections from the stereozone mainly
in the counter quadrants where they are more free standing. The minor septa
flanking the counter septum are 2 mm long and they decrease in size round towards
the cardinal septum near which they are 1-5 mm long. The minor septa contra-
tingent on the major septa immediately flanking the cardinal septum, however, are
again 2 mm long. Septal microstructure consists of an irregular structureless
core embedded in calcite showing growth lamellae running at a very low angle to the
septal faces axially and inward towards the core.
In the longitudinal section (Pl. 1, fig. 7), cut between the two sections described
above, the lumen is infilled with septal material except for two small peripheral
areas. The septal material shows bowl-shaped laminations, notched in the axis,
which are interpreted as growth-lines. In the smaller of the two voids, part of a
tabula can be seen curving upward and axially from the peripheral stereozone.
There are no dissepiments.
The specimen contains circular borings, 0-8-1: mm in diameter, of unknown
origin.
Discussion. This specimen appears to compare most closely in size and structure
with the type species of Stereolasma, S. rectum (Hall) from the Middle Devonian
Hamilton Group of eastern North America as illustrated by Simpson (1900, figs
16-18) and Stumm & Watkins (1961, pl. 58, figs 1-16). S. rectum, however, has
much thinner skeletal elements and less well developed minor septa except for the
counter-lateral minor septa which are strongly accelerated. The Venezuelan
specimen may belong to a new species but further material is required to confirm
this.
Genus STEWARTOPHYLLUM Busch 1941
1941 Stewartophyllum Busch: 393.
1949 Stewartophyllum Stumm: 9.
?1965 Stewartophyllum Federowski: 344.
TYPE SPECIES (by original designation). Amplexus intermittens Hall 1876;
pl. 32, figs 8-15. Middle Devonian, Hamilton Group; Moscow, N.Y., U.S.A.
FROM THE SIERRA DE PERIJA, VENEZUELA 235
DiaGnosis. Small subcylindrical to trochoid corals. Major septa dilated and
extending to axis where they fuse in neanic stage, withdrawn from axis in ephebic
stage. Cardinal fossula present. Minor septa very short or absent. Tabulae
complete and incomplete, flat axially, variably orientated peripherally. No dis-
sepiments (based on Stumm 1949 : 9 and Stumm & Watkins 1961 : 447).
Discussion. The type specimens of S. intermittens (Hall) have been redescribed
by Stumm and Watkins (1961 : 447) who showed that three other species described
by Busch should also be referred to Hall’s species. The diagnosis for the genus
given above takes account of this redescription.
?Stewartophyllum sp.
(Pl. 1, figs 4, 5)
MATERIAL. R45110 (3243). Cafio Grande; Cafio del Oeste Fm., early Middle
Devonian.
DESCRIPTION. Small curved trochoid coral, about 25 mm high and 20 mm in
maximum diameter.
In the late neanic-early ephebic stage, the lumen is almost entirely infilled. The
septa occur in a single series up to 1-25 mm thick at the periphery and wedge-shaped,
reaching between three and four-fifths of the way to the axis. They are arranged
in an irregularly pinnate pattern and are mostly laterally contiguous with scattered
very narrow gaps in the peripheral area between some septa. The axial ends of the
septa merge into solid sclerenchyme completely infilling the axis, in which no struc-
tural details can be seen. The septal structure compares with that described as
pseudotrabecular by Kato (1963, text-fig. ge).
In higher sections a subtriangular cardinal fossula containing a short, thin cardinal
septum, 4 mm long, is present on the convex side of the coral. There are slightly
larger gaps between the septa and signs of tabulae can occasionally be seen. The
axis of the coral is filled by a subcircular core of sclerenchyme 7 mm in diameter.
As the subcalicular level is approached, the subtriangular cardinal fossula increases
in size at the expense of the cardinal half of the core. The pinnate septal symmetry
is also more regularly developed immediately below the calice. The diameter here
is 16 mm with about 40 septa.
No longitudinal section is available.
Discussion. This single specimen is tentatively assigned to Stewartophyllum.
It appears to have considerable structural similarity with S. intermittens (Hall) from
the Middle Devonian Hamilton Group of eastern North America (see Stumm &
Watkins 1961, pl. 58, figs 17-28, 37-39) although it is clearly not conspecific.
Apart from the much higher septal number in the Venezuelan specimen, the
large axial core of sclerenchyme makes even a congeneric assignment open to
question. More material is required to assess the variability and significance
of the axial core.
236 PALAEOZOIC CORAL FAUNAS
Genus SYRINGAXON Lindstrém 1882
1970 Syvingaxon Sutherland: 1125.
1971 Syvingaxon Scrutton: 194, cum syn.
Discussion. See Scrutton (1971 : 194) for a diagnosis and full discussion of the
status of this genus.
Syringaxon sp.
(Pl. 1, figs 1-3)
MATERIAL. R45130 (3849). Cano Los Guineos; Cafio Los Guineos Fm., early
Middle Devonian (upper Onesquethaw).
DESCRIPTION. Small conico-cylindrical coral with moderately developed septal
grooves.
Coral circular in cross-section with a variably corrugated epitheca and a peripheral
stereozone about 0-6 mm thick. In the mature stages the major septa are 0-13-
0-2 mm thick midway along their length where they are very slightly waisted; the
cardinal septum is the thinnest. They reach approximately seven-eighths of the
radius towards the axis. The axial ends of the major septa are rhopaloid causing
them to be laterally contiguous and to form, with a small amount of additional
sclerenchyme, an aulos 2-2-5 mm in external diameter. In a subcalicular section
(Pl. 1, fig. 1), the internal dimensions of the aulos are 1:2 x 0-g mm. Ina section
approximately 1-5 mm below this (Pl. 1, fig. 2), the aulos is solidly infilled, possibly
by sclerenchyme coating a tabula as the longitudinal section shows the aulos to be
open for at least another 2-5 mm below the level of this section. The minor septa
are approximately half the radius in length except for the counter-lateral minor septa
which are equal in length to the counter septum. They are closely contratingent
so that the lumen between a major septum and its contratingent minor may be
closed at sub-calicular levels.
The longitudinal section is two-thirds infilled by septal material due to its being
cut slightly off-centre. There is a very narrow open core in the distal part of the
aulos containing irregularly spaced flat tabulae averaging 0-25 mm apart. In the
interseptal loculi one tabula can be clearly seen, flat near the peripheral stereozone,
and curving upwards towards the aulos. There are faint indications of what may be
further tabulae but none can be positively identified as such.
The subcalicular section is about 6-8 mm mean diameter with 20 major septa and
the section I-5 mm below is 6-4 mm in mean diameter with 18 major septa.
Discussion. This single specimen, although slightly larger in size at the same
septal number, is quite similar to S. ayvnoldi Scrutton (1971 : 196) from rocks of
Ludlovian age in the Mérida Andes of Venezuela. Further material is necessary,
however, to determine the status of the Devonian specimen.
S. rudis (Girty 1897 : 299, pl. 2, figs 7, 8), from the Lower Devonian of North
America, is in need of redescription and cannot be compared with the present
material.
FROM THE SIERRA DE PERIJA, VENEZUELA 237
Family HADROPHYLLIDAE Nicholson 1889
1937 Palaeocyclidae Bassler: 189, pars.
1949 Hadrophyllidae Stumm: 4.
1955 Porpitidae Jeffords: 12, pars.
1956 MHadrophyllidae Hill: 262.
1961 Hadrophyllidae Fontaine: 69.
1969 Hadrophyllidae Sutherland & Haugh: 27.
Tyre GENUS. Hadvophyllum Edwards & Haime 1850 : 67.
DiaGNnosis. Small, simple, discoid, patellate, button shaped or depressed
turbinate corals. Major septa usually arranged in quadrants and cardinal and alar
fossulae often strongly developed. Minor septa short and contratingent. Hori-
zontal partitions (?tabulae) developed in some genera.
Discussion. A brief resumé of the classification of this group of corals is given
by Sutherland & Haugh (1969 : 28). For the moment, Hill’s (1956 : 262) inter-
pretation of the Hadrophyllidae is accepted and attention is drawn to the presence
of what are probably tabulae in the specimen described below as Hadrophyllum sp.
and in Gymnophyllum wardi Howell (see Sutherland & Haugh 1069 : 35).
Genus HADROPHYLLUM Edwards & Haime 1850
1850 Hadrophyllum Edwards & Haime: 67.
1851 Hadrophyllum Edwards & Haime: 357.
1937 Hadvophylium Bassler: 197.
1952 Hadrophyllum Le Maitre: 41.
1955 Hadrophyllum Jeffords: 8.
1961 Hadvophyllum Fontaine: 69, cum syn.
Dracnosis. Discoid to broadly trochoid. Cardinal septum in oval fossula; other
cardinal quadrant septa pinnate and sub-parallel, counter quadrant septa more or
less radially arranged with a long counter septum. Minor septa short. Horizontal
partitions (?tabulae) may be developed.
TYPE SPECIES (by original designation). Hadrophyllum orbignyt Edwards &
Haime 1851 : 357, pl. 6, figs 4, 4a. Devonian; Charleston Landing, Indiana and
Clark County above Louisville, Kentucky, U.S.A.
Discussion. The status of this genus has been discussed by Jeffords (1955 : 8).
An amended diagnosis is given here to cover the occurrence of tabula-like structures
in the specimen of Hadrophyllum described below.
Hadrophyllum sp.
(Pl. 1, fig. 8, Text-fig. 4)
MATERIAL. R49275 (3243). Cafio Grande; Cafio del Oeste Fm., early Middle
Devonian.
B
238 PALAEOZOIC CORAL FAUNAS
DESCRIPTION. Small, broadly trochoid coral with smooth, slightly worn exterior.
Calice not seen.
A single section at the ephebic stage shows the lumen largely infilled by thick
septa arranged with striking pinnate symmetry. The septa are mostly between
I and 1:4 mm in thickness peripherally. The counter and alar septa are longest,
all three meeting in the axis. In the counter quadrants, the counter-lateral and next
four metasepta are roughly equal in length and about three-quarters the length of
the counter septum. The fifth and subsequent metasepta in the counter quadrants
become progressively shorter and more wedge-shaped against the alar septa. On
the cardinal side of the alar septa the first two metasepta are equal in length to the
alar septa and the subsequent metasepta become progressively shorter and more
wedge-shaped round towards the cardinal septum. The wall of the cardinal fossula
appears to be the result of sclerenchyme coating the successive ends of the metasepta
in the cardinal quadrants and is not formed by extra long metasepta flanking the
short, 3:5 mm long, cardinal septum. Very narrow gaps, about o-I mm wide,
separate the major septa in the peripheral area of the corallum. In several places
the gaps are crossed by narrow bars that are probably tabulae. There are short
triangular minor septa, about I mm long, inserted between nearly all the major
septa. The septal formula is CoA8K7A8C at a mean diameter of 13 mm.
No longitudinal section is available.
Discussion. The general characters of this coral agree with those of the genus
Hadrvophyllum although it cannot be readily assigned to any described species.
Comparison is made difficult as virtually no other species of Hadrophyllum has been
Fic 4. Septal arrangement in Hadvophyllum sp. (R49275). x5.
C = cardinal septum; K = counter septum; A = alar septa.
FROM THE SIERRA DE PERIJA, VENEZUELA 239
studied with, or illustrated by, thin sections. The appearance of interseptal
structures like tabulae in cross-section has never previously been recorded in a species
of Hadvophyllum but this may also be due to a lack of thin section study. For
example, rare tabulae have recently been recorded in thin sections of another member
of the Hadrophyllidae, Gymnophyllum wardi Howell, by Sutherland and Haugh
(1969 : 35).
Family HAPSIPHYLLIDAE Grabau 1928
1956 Hapsiphyllidae Hill: 267.
1964 Hapsiphyllidae Rowett & Sutherland: 46.
Genus AMPLEXIZAPHRENTIS Vaughan 1906
1906 Ampblexizaphrentis Vaughan: 315.
1958 Amplexizaphrentis Sutherland: 44.
1962 Amplexizaphrentis Armstrong: 30.
1962 Enniskillenia Kabakovitch im Soshkina & Kabakovitch: 323.
1964 Amplexizaphrentis Rowett & Sutherland: 47.
DiaGnosis. Small, solitary, trochoid to ceratoid corals in which major septa
may or may not unite around a conspicuous cardinal fossula. Septa commonly
withdraw from axis in late stages of growth. Alar pseudofossulae usually well
developed in early stages, but may become inconspicuous in ephebic stage. Tabulae
usually complete, arched or flattened axially except where abruptly depressed in the
fossula. No dissepiments. (After Rowett & Sutherland 1964 : 47).
TYPE SPECIES (see Opinion 854, I.C.Z.N. 1968). Zaphrentis curvulena Thomson
1881 : 223, 236. Lower Limestone Group, Lower Carboniferous; Brockley, near
Lesmahagow, Lanarkshire, Scotland.
Discussion. The status of the genus Amplexizaphrentis has been discussed by
Sutherland (1958 : 44 et seq.) and his conclusions are accepted here. More recently
a case to stabilize the generic name and type species was submitted to the I.C.Z.N.
by Shrestha (1966), supported by Mitchell (1966) and approved by the Commission
in Opinion 854. This effectively answered the comments on the validity of
Amplexizaphrentis expressed by de Groot (1963 : 39).
According to Rowett & Sutherland (1964 : 51) Barytichisma (Moore & Jeffords
1945 : 131) is distinguished from Amplexizaphrentis ‘primarily on the basis of the
unusually wide peripheral stereozone’. They also note in the diagnosis for that
genus that the septa are amplexoid and reach the axis immediately above the
tabulae only. Weyer (1965 : 450) also laid emphasis on amplexoid septa in Bary-
tichisma in maintaining the genus distinct from Amplexizaphrentis. If this distinc-
tion is accepted, the species described below seems to fall more naturally into
Amplexizaphrentis than Barytichisma despite a tendency to amplexoid septa at all
stages of growth. Certainly A. sutherlandi sp. nov. does not develop an unusually
wide peripheral stereozone.
240 PALAEOZOIC CORAL FAUNAS
Amplexizaphrentis sutherlandi sp. nov.
(Pl. 1, figs 9-13, Text-fig. 5)
DERIVATION OF NAME. After Dr P. K. Sutherland (University of Oklahoma).
DiaGnosis. Curved ceratoid Amplexizaphrentis with cardinal fossula on concave
side of corallum. Major septa thick, tending to amplexoid throughout ontogeny;
34 present in ephebic stage of holotypeat 14mm diameter. Minor septa rudimentary.
Tabulae wide, flat with downturned edges, inclined strongly from the counter to
cardinal side.
HorotyPe. R45127 (3784). Cafio Colorado; Rio Palmar Formation, Lr.
Pennsylvanian.
PARATYPES. R4g291 (3784; ?distal end of holotype), R49292 (3784). Same
locality and horizon as holotype.
DESCRIPTION. Curved ceratoid corals up to 60-70 mm long and about 25 mm
maximum diameter (estimated because of crushing of the calice). The cardinal
fossula is on the concave side of the corallum. Septal grooves weak or absent on a
slightly rugate epitheca.
In the early neanic stage (Pl. 1, fig. 10) the septa are short and variably developed
with a wide, open lumen. On the cardinal side a tabula is cut, on the upper surface
of which the cardinal septum and several metasepta in the left cardinal quadrant are
well developed. They are approximately 1-3 mm long and 0-5 mm wide at their
bases, the cardinal septum being fractionally longer than the others. No septa
can be distinguished in the zone where the tabula is in the plane of section and
immediately below the tabula, in the alar areas, the septa are very short and thorn-
like. In the counter area the counter septum is short and thin, about 1:3 mm long
and 0-I mm wide expanding to 0-5 mm wide where it merges into the thin peripheral
stereozone. The flanking septa are similar in shape but become progressively
shorter, from a maximumof 0-8 mm long, towards the alar areas. The mean diameter
is 7 mm; Ig septa can be counted but there are probably 22 or 24 present at this
level.
In a later neanic section (Pl. 1, fig. 11), approximately 2 mm higher than the
first, the septa in the cardinal quadrants are much more strongly developed. The
alar septa appear to meet across the axis of the corallum in a continuous band
o-5—0-8 mm thick and the metasepta in the cardinal quadrants are arranged in strongly
pinnate groups. The ends of successive metasepta outline a large suboval cardinal
fossula bisected by a long waisted cardinal septum extending to the axis. The
metasepta in the cardinal quadrants are o-4—0-6 mm thick peripherally and taper
towards the axis. Where tabulae are sectioned, they and the adjacent septa are
thickened by about 0-2 mm towards the periphery. In the counter quadrants,
the septa are thin short spines as in the previous section. Measurements are
difficult as the peripheral stereozone has been removed around much of these
quadrants. The mean diameter is 7-5 mm and the septal formula is C6A4K4A6C.
In an ephebic stage section (Pl. 1, fig. 9) 14 mm higher (measured centre to centre)
than the previous section, the counter quadrants are fully developed. The counter
FROM THE SIERRA DE PERIJA, VENEZUELA 241
septum 1 mm thick, is thicker than all other septa, which range between 0-5—0-7 mm
thick, and reaches into the axial area to meet other septal material. The flanking
septa are long and pinnately arranged with their axial ends fused and coated with
sclerenchyme to form strong walls parallel to the alar septa and suggesting very
weak alar fossulae. The arrangement of the septa in the cardinal quadrants is
similar to that described in the previous section except that the alar septa are now of
similar aspect to the cardinal metasepta and meet in the axis with an angle of about
120°. The cardinal fossula is more pear shaped and the cardinal septum has
shortened to 3-4 mm long (its axial end is damaged). Most of the major septa
taper towards the axis but some are slightly rhopaloid and may have very thin
needle-like axial extensions from the club-shaped thickening. The peripheral
stereozone is I mm thick where it is undamaged and slight kinks in it on either side
of the counter septum suggest the presence of rudimentary minor septa. The mean
diameter is 14 mm and the septal formula is C6AgKgAO6C.
A section in the ephebic stage of another specimen (R49292) shows all the septa
withdrawn from the axis except the metasepta flanking the cardinal septum. These
with a considerable thickening of sclerenchyme, outline an oval cardinal fossula
bisected by a thin cardinal septum. The diameter is 15 mm with an estimated
38 septa (C18K?18C).
In longitudinal section the tabulae are largely complete with few subsidiary
plates and a rather irregular spacing. They slope strongly downwards from the
counter to the cardinal side with a flat or slightly undulating surface and down-
turned peripheral margins. Successive complete tabulae may be as much as 3-4 mm
20
d (mm)
Fic. 5. Number of major septa plotted against diameter for specimens of Amplexi-
zaphrentis sutherlandi. The dashed line joins asterisks representing different sections
of the holotype.
242 PALAEOZOIC CORAL FAUNAS
apart during the late neanic-early epebic stages but become much closer-spaced,
less than I mm at the subcalicular level. The tabulae are extensively crested with
septal material in the peripheral areas. This material shows fibres directed axially
and upwards at varying angles and growth lines steeply dipping at the periphery,
curving round to the horizontal in the axial area. The tabulae in the axial area are
coated with sclerenchyme showing a zigzag fibre pattern. Although in some places
this appears to be an alteration phenomenon based on the calcite cleavage pattern, in
others it appears to be original and could reflect lateral movements in the basal
ectoderm of the polyp during calcification (see also Sutherland 1958 : 49). There are
no dissepiments.
Discussion. This species seems to compare quite closely with Amplexizaphrentis
cassa Sutherland (1958 : 54, pl. 8, figs 1-5, pl. 10, figs 1-4) from the middle
Mississippian Kindle Formation of the Tetsa River area, and particularly closely
with A. sp. B (Sutherland 1958 : 57, pl. 11, figs I-5) from the Prophet Formation
(?early Mississippian) of the Prophet-Muskwa Rivers area, both British Columbia.
A. sutherland is distinguished by the thickness of its septa, its large size and a
slightly higher septal ratio than A. cassa. In these features, however, it approaches
very closely to A. sp. B of Sutherland although the cardinal fossula in the latter
may be larger in proportion. The age of the Rio Palmar Formation appears to be
lower Pennsylvanian and thus A. sutherlandi is somewhat younger than either of the
Canadian species.
Superfamily ZAPHRENTICAE Edwards & Haime 1850
Family STREPTELASMATIDAE Nicholson 1889
1971 Stveptelasmatidae Scrutton: 206.
Discussion. The writer has recently outlined his views on the composition of this
family (Scrutton 1971 : 206). The new genus described here stands very close to
existing members of this group and does not necessitate any reconsideration of the
family concept.
Genus BOWENELASMA nov.
DERIVATION OF NAME. After Dr J. M. Bowen (Shell International Petroleum
Corporation) who collected the material described here.
DiaGnosis. Curved ceratoid corals with the cardinal septum on the convex
side of the corallum. Septa dilated to close the lumen in early ontogeny; strongly
dilated and more or less coated with sclerenchyme in the cardinal quadrants at the
ephebic stage. Cardinal fossula narrow or poorly developed. Intertwined septal
elements in the axial area may form a low boss in the calice. Minor septa well
developed. Tabulae steeply sloping axially and upwardsat the periphery, undulating
or highly domed in the axial area. No dissepiments.
TYPE SPECIES. Bowenelasma typa sp. nov.
FROM THE SIERRA DE PERIJA, VENEZUELA 243
Discussion. Bowenelasma is recorded so far only from the early Middle Devonian
of Venezuela. The genus is distinguished first and foremost by the strong, consistent
dilatation and thickening of the cardinal quadrant septa in the later, but subcalicular,
ontogenic stages. This feature appears occasionally in other streptelasmatid corals
and it is also characteristic of the genus Pseudophaulactis Zaprudskaya in Ivanovskii
(1963 : 32, pl. 6, fig. 2) from the Llandovery of the Siberian Platform. Bowenelasma
and Pseudophaulactis are very similar in appearance, the latter differing only in the
very attenuate nature of the septa in the counter quadrants and the general lack of
twisted septal ends in the axial area (see Ivanovskii 1965, pls 5, 6 and 7, fig. x). It
should also be noted, however, that the two genera are widely separated strati-
graphically and geographically.
Species of other streptelasmatid genera occasionally show significantly prolonged
dilatation and thickening in the cardinal quadrants during ontogeny as a specific
character or a subspecific variant. They include Kiaerophyllum semilunatum
Scheffen (1933 : 21, pl. 2, figs 4-6) from the Upper Ordovician of the Oslo region
which is considered a variant of Grewingkia buceros according to Neuman (1969 : 36),
Grewingkia europaeum hosholmensis Kaljo (1961 : 58, pl. 3) from the Upper Ordo-
vician of Estonia and the specimen identified as Dalmanophyllum dalman by
Ivanovskii (1963 : 36, pl. 7, fig. 4) (= Ditoecholasma dalmani in Ivanovskii 1965,
pl. 1, fig. 6) from the Llandovery of the Siberian Platform. All these are considerably
older than the new Devonian genus. Bowenelasma is clearly not congeneric with
Grewingkia, which is principally distinguished by its wide spongy axial structure
(Neuman 1969 : 35). Bowenelasma also lacks the distinctive blade-like columella
so prominent in illustrations of the type species of Dalmanophyllum (Edwards &
Haime’s 1851, pl. 1, fig. 6; Minato 1961, pl. 11, figs 16, 3c, 5b).
Comparison with its contemporary streptelasmatids shows Bowenlasma to be
readily distinguishable from all of them. This genus appears to be a late and little
modified descendant of the primitive streptelasmatid structural pattern and as such
contrasts quite strongly with the more structurally divergent Heterophrentis/
Siphonophrentis group. Briantelasma, however, may be more closely related (see
discussion under Briantelasma).
Bowenelasma typa sp. nov.
(Pl. 2, figs 1-11, Text-fig. 6)
DERIVATION OF NAME. From typus (L.) = type, indicating the type species.
Diacnosis. Bowenelasma with 45 major septa at a mean subcalicular diameter of
27 mm (holotype). In ephebic stage, major septa four-fifths radius in length,
dilated, coated with sclerenchyme and laterally contiguous in cardinal quadrants
and slightly deflected counter-clockwise. Axial area with few septal ends inter-
twined to form low calicular boss. Minor septa up to one-quarter radius in length.
Cardinal septum short in narrow fossula. Tabulae simple, convex, steeply sloping
upwards and axially at periphery, obscured in axis but apparently gently undulating.
244 PALAEOZOIC CORAL FAUNAS
HoLotyPeE. R45094 (3149). Cano Grande; Cafio Grande Formation, early
Middle Devonian (upper Onesquethaw).
PARATYPES. R45075, R45077 (both 3121), R45100 (3157); same locality and
horizon as holotype. R45121 (3618); Cafio Colorado; same horizon as holotype.
OTHER MATERIAL. R45082, R49247-9, R4g9251 (all 3121), R49263 (3157);
same locality and horizon as holotype. R45118 (3618), Cafio Colorado; R45279
(3323), loose boulder on Cafio del Oeste; both same horizon as holotype.
DESCRIPTION. Curved ceratoid corals up to about 90 mm long and 27 mm mean
diameter at the subcalicular level. The cardinal septum is located on the convex
side of the corallum. The epitheca is lightly rugate with weak septal grooves.
In the neanic stage (PI. 2, fig. 4), the lumen is almost completely infilled by thick,
irregularly pinnate septa. The major septa are between 0-6—0-7 mm thick near the
periphery and taper towards the axis, contiguous with their neighbours over most of
their length. Major septa in the counter and alar sectors, 3-0-3-5 mm long, reach
into the axial area where they fuse and almost completely fill the lumen. The other
septa wedge against these. The cardinal septum is about 1-3 mm long. Minor
septa are present as very short wedges 0-7-I-0 mm long, nearly always with very
small openings in the lumen beyond their axial ends. The counter septum is flanked
by extra long (1-7 mm) minor septa. There are also small openings in the lumen
periaxially, beyond the axial end of the cardinal septum and on the counter side of
one of the alar septa. The septal formula is C345K5A4C at a mean diameter of
7 mm.
Sections in the late neanic stage of the holotype (R45094c-d, Pl. 2, fig. 2) show
major septa of very variable length twisted in a wide counter-clockwise axial vortex.
Most of the major septa are 0:3-0:4 mm wide and half the radius in length. Only a
few irregular septal ends cross the axial area where there may also be a few isolated
spots of septal tissue. The cardinal septum is short and heavily invested with
sclerenchyme. Some of the other septa in the cardinal quadrants also appear to
be thickened but there is no marked contrast with the septa in the counter quadrants
at this level. Minor septa cannot be easily distinguished in the wider parts of the
very irregularly developed peripheral stereozone. Some of them appear to reach
2-5 mm in length and the stereozone may be up to 4 mm wide. There are 38 septa
at a mean diameter of 17 mm.
In the ephebic stage (Pl. 2, fig. 1) there is a strong contrast between cardinal and
counter quadrants. In both the major septa extend four-fifths of the distance to
the axis and are twisted in a weak counter-clockwise vortex. Those in the cardinal
quadrants however are between I-I-I:4 mm thick and laterally contiguous for
most of their length, whilst those in the counter quadrants are only 0:25-0:35 mm
across and more or less parallel sided. The cardinal septum, also contiguous with
its neighbours, is 5-5 mm long and situated in a weak narrow fossula closed on the
axial side by a tabula and converging septa. The axial area of the coral, about a
fifth of the diameter across, is traversed by two or three septal strands and traces of
tabulae. There may also be a few isolated spots of septal tissue. In R45077b there
is evidence that this structure in the axial area may produce a low boss in the floor
FROM THE SIERRA DE PERIJA, VENEZUELA 245
of the calice. Minor septa are also well developed, as wedges of variable size up to
I-5 mm across at the base and 3:5 mm long in the cardinal quadrants and as thin
spines projecting up to 2 mm beyond the peripheral stereozone in the counter quad-
rants. The minor septa flanking the counter septum are markedly longer than their
neighbours but measurement is difficult as this part of the coral is nearly always
crushed. The peripheral stereozone in the counter quadrants is quite thin, only
about 1 mm thick. In the holotype, the septal formula is C1oA13KI10A8C at a
mean diameter of 27 mm.
At subcalicular levels, the septa in the cardinal quadrants thin and separate from
their neighbours first on the axial side of the minor septa. The cardinal septum
thins most rapidly leaving a narrow, parallel sided fossula which it bisects. It
maintains a length of about half the radius whilst the septa in the cardinal quadrants
are still somewhat thicker than those in the counter quadrants but at slightly higher
levels it rapidly withdraws to about a fifth of the radius in length. Ina paratype,
R45121a (Pl. 2, fig. 10), the minor septa flanking the counter septum are about
5°5 mm long at this level and the septal formula is C1ioA1z2K12A9C at a mean
diameter of 28 mm.
The longitudinal section is dominated by septal traces and the shape of the
tabulae is difficult to see. At both walls there are thick solid septal deposits, rather
irregularly developed but much more pronounced on the cardinal side. In the axial
area individual septal traces form a complex and irregular pattern breaking up and
obscuring the tabulae. The tabulae can be clearly seen in the peripheral areas,
slightly convex, moderately spaced and sloping axially and upward at 5~-20° to the
wall. In the axial area, the tabulae appear to be gently undulating and in one case
(Pl. 2, fig. 6) sloping distinctly downward from the cardinal to the counter side.
The angle of slope is difficult to assess but appears to be as much as 45° in some
cases.
Measurements on the holotype and paratypes are plotted in Text-fig. 6.
Discussion. The only species so far assigned to Bowenelasma are the two species
from the Sierra de Perija described here. The type species, B. typa, which is the
more abundant by a factor of three in Bowen’s collection, is compared with B.
breviseptata under the discussion of the latter species.
Bowenelasma breviseptata sp. nov.
(Pl. 3, figs 1-7; Text-fig. 6)
DERIVATION OF NAME. From brevis (L) = short + septum, in reference to the
characteristically short major septa.
Diacnosis. Bowenelasma with 43 major septa at a mean subcalicular diameter
of 33 mm (holotype). In ephebic stage, major septa half radius in length, cardinal
quadrants heavily coated with sclerenchyme peripherally. Cardinal septum very
slightly longer than adjacent metasepta; no fossula developed. Wide axial area
partially filled by thickened and twisted septal ends. Minor septa up to one-third
radius in length. Tabulae simple, convex, steeply sloping upwards and axially at
the periphery, domed in the axis or obscured by septal traces.
246 PALAEOZOIC CORAL FAUNAS
HowotyPe. R45123 (3619). Cafio Colorado; Cafio Grande Formation, early
Middle Devonian (upper Onesquethaw).
PARATYPE. R45125 (3619). Same locality and horizon as holotype.
OTHER MATERIAL. R45105 (3199), Cafio del Sur; ? R4g256 (3154), Cafio Grande;
both same horizon as holotype.
DeEscRIPTION. Large curved ceratoid corals with the cardinal septum located in
the convex side of the corallum. Estimated original length of holotype 80-90 mm.
The epitheca has weak septal grooves and is gently rugate.
The late neanic stages (PI. 3, fig. 3) are largely infilled by major septa up to 1-7 mm
thick and laterally contiguous along most of their length. The major septa are
two-thirds the radius in length leaving a wide axial zone mostly infilled by loosely
and irregularly twisted septal elements. The cardinal septum is only just over one-
third the radius in length with a gap in the lumen beyond its axial end: the other
protosepta cannot be distinguished. The minor septa are well developed wedges
up to 5 mm long and 1 mm or a little more across the base.
In the ephebic stage (Pl. 3, fig. 1) septal thickening is confined to the cardinal
quadrants. The septa are about half the radius in length with a slight anticlockwise
displacement. In the cardinal quadrants the peripheral half to three-quarters of
the septa are thickened to about 2 mm across bringing adjacent septa into contact.
50
40
20
0 10 20 30 40
d (mm)
---- Btypa + holotype —-—-— B. breviseptata 0 holotype & R45105
@ other material + other material
Fic. 6. Number of major septa plotted against diameter for specimens
of Bowenelasma typa and B. breviseptata.
FROM THE SIERRA DE PERIJA, VENEZUELA 247
Otherwise the septa are between 0-6 and 0-7 mm across and parallel sided in both
quadrants. The cardinal septum is slightly longer than adjacent metasepta with the
peripheral dilatation imperfect ; there is no cardinal fossula. In the axial half of the
diameter, septal elements are loosely and irregularly twisted together and thickened
in part. This structure forms a low boss in the floor of the calice. The minor
septa are very long, and regularly developed reaching 5 mm or a third of the radius
in length. Unfortunately those flanking the counter septum cannot be seen in
the holotype due to crushing in this area. The minor septa are wedge-shaped in the
cardinal quadrants and parallel sided to gently tapering, 0-5—1-0 mm across in the
counter quadrants. The peripheral stereozone in the counter quadrants appears to
be thin but is everywhere worn and cannot be measured.
R45105 (Pl. 3, figs 5-7) differs somewhat from the other specimens through its
septa being generally less thickened. The section of the neanic stage available
(Pl. 3, fig. 5) shows no lateral contiguity between septa. In the ephebic stage the
septa are only slightly thicker in the cardinal quadrants than the counter quadrants,
although the cardinal septum itself is notably thick at 1:2 mm across. The minor
septa are up to 4-4 mm or two-fifths the radius in length and slightly variable in
development. The septal formula is C6AgKgA5C at a diameter of 22 mm.
The longitudinal section of the holotype (Pl. 3, fig. 4) is dominated by septal
traces and the peripheral stereozone. The latter is particularly thick on the cardinal
side. Traces of tabulae can be seen in the peripheral areas, sloping steeply axially
and upwards but they cannot be identified in the axial area among the irregular
vertical sections of septal elements. In R45105d (PI. 3, fig. 7), however, the structure
is very clear. The tabulae consist of large curved plates, convex upwards and
outwards. They form a high axial dome and are almost vertical peripherally where
they form a narrow trough against the peripheral stereozone. They are irregularly
spaced between I and 2:5 mm vertically.
Measurements are plotted in Text-fig. 6.
Discussion. At present it is not certain how much weight should be given to
the differences between R45105 and the holotype. R45105 is considered conspecific
with Bowenelasma breviseptata here although with more material and a better
understanding of the variation it may prove to be subspecifically or even specifically
distinct.
B. breviseptata is distinguished from B. typa principally through the much shorter
major septa in the mature stages and the lower septal number at comparable
diameters. JB. breviseptata also has more strongly developed minor septa and lacks
any clear sign of a cardinal fossula.
Genus BRIANTELASMA Oliver 1960
1960a Briantelasma Oliver: 89.
1960b Briantelasma Oliver: 6.
1963 Briantelasma Oliver: 26.
Diacnosis. Trochoid to cylindrical corals with subpinnately arranged major
septa extending half way or more to the axis. Lumen partly or completely infilled
248 PALAEOZOIC CORAL FAUNAS
by laterally contiguous septa and axial sclerenchyme especially in early growth
stages. Cardinal fossula present. Minor septa usually well developed. Tabulae
strongly domed with axial depression, complete and closely spaced. (Based on
Oliver 1960b : 6.)
TYPE SPECIES (by original designation). Briantelasma americanum Oliver 10960 :
89, pl. 14, figs 1-4. Reef facies of Coeymans Limestone, Helderbergian, Lower
Devonian; Madison and Oneida Counties, New York.
Discussion. The writer follows Oliver (1960b : 6) in regarding Briantelasma as
a member of the Streptelasmatidae in which the structural elements are completely
invested in sclerenchyme during almost the full ontogeny. The ancestors of
Briantelasma could well belong to the same lineage that evolved Bowenelasma as
there are similarities in the basic structural patterns of the two genera. They differ
principally, and very obviously, in the manner and extent of the development of
sclerenchyme and also in the detailed appearance of the tabularium and the peri-
pheral stereozone. There is no doubt that they represent distinct and divergent
genera over their presently known range.
Briantelasma oliveri sp. nov
(Pl. 4, figs 1-7; Text-figs 7, 8)
DERIVATION OF NAME. After Dr W. A. Oliver, Jr. (United States Geological
Survey).
DiacGnosis. Ceratoid to trochoid Briantelasma of variable size up to 39 mm
diameter with 60 major septa in the holotype. Lumen completely infilled by septa
and sclerenchyme to immediately sub-calicular levels. Major septa withdraw
from axis to approximately half radius in length in ephebic stage. Axis plugged by
structureless to vermiform loops of sclerenchyme which persists as low boss in
calice. Minor septa up to one-quarter radius in length. Tabulae present in floor
of calice; not distinguished in sclerenchyme.
HoLotyPE. R45090 (3129). Cafio Grande; Cafio Grande Formation, early
Middle Devonian (upper Onesquethaw).
PARATYPES. R45091 (3129); same locality and horizon as holotype. R45119
(3618) ; Cafio Colorado, same horizon as holotype.
OTHER MATERIAL. ? R45099 (3157); same locality and horizon as holotype.
? R4g9290 (3618) ; Cafio Colorado, same horizon as holotype. ? PRI 24426 (= 24433);
? loose block in Cafio del Oeste, same horizon as holotype. ? PRI 24429 (= 24430A);
same locality and horizon as holotype. ? PRI 24430B; ? locality of holotype or
PRI 24426, same horizon as holotype.
DESCRIPTION. Ceratoid to trochoid, straight to slightly horn shaped corals up
to 40 mm diameter at the base of the calice and an estimated I00 mm long. The
holotype, which is the longest specimen, is incomplete and has a crushed calice.
The cardinal septum is on the convex side in curved coralla. Epitheca moderately
rugate and bearing weak septal grooves.
FROM THE SIERRA DE PERIJA, VENEZUELA 249
In the neanic section R45091a (Pl. 4, fig. 5) the lumen is completely or almost
completely infilled by laterally contiguous septa and axial sclerenchyme. The major
septa vary between half and the full radius in length and their axial ends merge into
the sclerenchyme almost completely infilling the axis. They are between 0-6 and
0-7 mm in width and contiguous with their neighbours except for occasional narrow
gaps between one-quarter and one-third the radius from the periphery, probably on
the axial side of wedge-shaped minor septa. These minor septa are extremely
difficult to distinguish but appear to be present only in the counter quadrants.
There is a slight pinnate septal symmetry but again difficult to distinguish mainly
due to the character of the preservation. The septal formula is C6A8KQA6C at a
mean diameter of Io mm.
In the ephebic stage (Pl. 4, figs 3, 6), the laterally contiguous major septa are very
evenly developed 0-5 to 0-6 of the radius in length. With the exception of a few
specific major septa, they all have open axial ends merging with a solid axial core of
sclerenchyme which is either more or less structureless or contains a large scale
vermiform pattern (R45119a, b). The septal pattern in the holotype is illustrated
in Text-fig. 7. The counter septum, which is fractionally wider and longer than the
average, I'5 mm X II mm in the holotype, is characteristically flanked by counter-
lateral septa slightly shorter than the average and with closed axial ends. Some
septa in the alar areas, particularly new metasepta appearing immediately on the
counter side of the alar septa are also less than average length and have closed axial
ends. The cardinal septum, which is difficult to distinguish, appears to be about
average length and is flanked by shorter metasepta with the next adjacent meta-
septa curved slightly towards the cardinal septum at their axial ends. Minor septa
are well developed wedges up to a maximum of one-quarter the radius in length.
Fic. 7. Septal arrangement in Briantelasma oliveri (R45090a). X2.
C = cardinal septum; K = counter septum; A = alar septa.
250 PALAEOZOIC CORAL FAUNAS
The lumen is totally infilled. The septal formula is C1ioA13K15A10C at a mean
diameter of 23 mm in R45091b
Close to the base of the calice (Pl. 4, fig. 1) an opening first appears on the axial
side of the cardinal septum which shortens rapidly at this level. A narrow elongate
or sub-oval fossula is formed. Subsequently gaps appear between the major septa
on the axial side of the minor septa and rapidly enlarge. The septa appear to thin
almost simultaneously in all quadrants; contrasted thickening in various quadrants
at this level is due to sections slightly angled to the growth lines. As the major
septa withdraw to the periphery, with gently tapered axial ends, the axial plug of
sclerenchyme is isolated briefly as a low mound in the calice base. At this level in
the holotype, there is a peripheral stereozone 4 mm wide at a mean diameter of
39 mm. The stereozone thins in the walls of the calice and the minor septa project
from it, thin and narrowly attenuate.
In longitudinal sections (Pl. 4, fig. 4), there is evidence of some gaps in between
the septa and sclerenchyme at the neanic stage and in the cardinal area at later
stages. Two such gaps in the cardinal area occur in the longitudinal section of the
holotype (R450g90e) and show signs of tabulae trough shaped against the peripheral
stereozone and capped by flat subsidiary tabulae. The septa and sclerenchyme
show a strongly developed grain shallowy arched across the axis and sharply down
turned towards the peripheral trough-shaped tabulae. This may reflect the shape of
70
60
50
30
20
d (mm)
- - -® -- B oliveri * holotype
B. americanum
=== — B. knoxboroense serteseeeseee B. mainense
Fic. 8. Number of major septa plotted against diameter for specimens of Briantelasma
oliveri. The polygons outline the n/d variation fields for other species of Briantelasma.
FROM THE SIERRA DE PERIJA, VENEZUELA 251
tabulae suppressed, or obscured in the sclerenchyme. Unfortunately no longitudinal
section is available through the base of the calice, where tabulae are known to be
developed, to support this.
Measurements on the holotype and paratypes are plotted in Text-fig. 8. There is
a considerable variation in size, basal calice diameters ranging from 15:5 mm to
39 mm.
Discussion. Previous records of Briantelasma come from Silurian (? Wenlock)
and Lower Devonian (Helderberg) rocks in eastern North America. Briantelasma
oliveri, from the early Middle Devonian of the Sierra de Perija, thus considerably
extends the known range of the genus. The new species can be distinguished from
all earlier described species of Briantelasma through the major septa withdrawing
from the axis in the ephebic stage. Also, over the ontogenetic range available,
B. oliveri lies wholly outside the fields of variation in septal number with diameter
shown by the other species (see Text-fig. 8). It has higher septal ratios at corres-
ponding diameters and reaches a higher maximum diameter than the North American
species.
Genus HETEROPHRENTIS Billings 1875
Discussion. Oliver (1958 : 817, 825) suggested that Compressiphyllum Stumm
(1949 : 13), which differs from Heterophrentis s.s. only through a pronounced com-
pression of the corallum at right angles to the counter-cardinal plane, was best
considered as a subgenus of Heterophrentis.
Subgenus HETEROPHRENTIS (HETEROPHRENTIS) Billings 1875
1875 Heterophrentis Billings: 235.
1938 Heterophrentis Stewart: 20.
1949 Heterophrentis Stumm: 11.
1958 Heterophrentis (Heterophrentis) Oliver: 825.
21964 Heterophrentis Webby: 7.
1965 Heterophrentis Stumm: 18.
1968 Hetevophrentis Altevogt: 762.
DiaGnosis. Ceratoid to trochoid corals, circular in cross-section. Septa thin
throughout ontogeny; major reach more or less to axis, minor up to one-third radius
in ephebic stage; peripheral stereozone variable in thickness. Cardinal fossula
prominent. Tabulae complete or incomplete, flat axially, arched peripherally. No
dissepiments.
TYPE SPECIES (see Miller 1889 : 193). Zaphrentis spatiosa Billings 1858 : 178
= Zaphrentis prolifica Billings 1858 :176 (according to O’Connell 1914 : 183).
Onondaga Limestone, early Middle Devonian; Rama’s Farm, near Port Colborne,
Ontario, Canada.
Discussion. Numerous species of Heterophrentis (H.) have been described and
the scope of the snbgenus is reasonably well defined despite the fact that, according
252 PALAEOZOIC CORAL FAUNAS
to Sutherland (1958 : 45), the original specimens of neither Z. spatiosa, nor Z.
prolifica have been sectioned. The traditional interpretation of the subgenus is
followed here and would surely need to be stabilised even if Z. spatiosa proved not to
have the characters attributed to Heterophrentis (H.).
Heterophrentis (H.) is common in the Lower and Middle Devonian of eastern
North America. The subgenus has also been recorded in the Middle Devonian of
Spain and north-west Africa and is confirmed here in the early Middle Devonian of
Venezuela.
Heterophrentis (Heterophrentis) venezuelensis (Weisbord 1926)
(PI. 5, DSS 1,.2)
1926 Cyathophyllum venezuelense Weisbord: 4 pars., pl. 1, fig. 5 only.
1943 Heterophrentis venezuelensis (Weisbord) Wells: 97 pars., pl. 10, fig. 7 only.
Diacnosis. Heterophrentis (H.) with major septa two-thirds radius in ephebic
stage, tapering axially; about 64 at 33 mm mean diameter in lectotype. Wide axial
area occupied by tabulae only. Minor septa confined to peripheral stereozone
one-quarter radius in width. Cardinal fossula present but weak. Tabulae mainly
complete, gently undulating to depressed axially, sharply downturned peripherally,
very close spaced beneath calice floor.
LeEcTOTYPE (here chosen). PRI 21594 (= PRI 24421, same specimen). Loose
block (?) in Cafio del Norte; Cafio Grande Fm., early Middle Devonian (upper
Onesquethaw).
DEscriPTIon. The lectotype is an incomplete curved conical coral 80 mm high
between a diameter of 23 mm and the top of the calice. The epitheca is lightly
rugate. The counter-cardinal plane is at right angles to the plane of curvature.
In the ephebic stage, 10 to 15 mm below the base of the calice, the major septa
are regularly developed, 9-10 mm or two-thirds the radius in length. They are
between I and 1-3 mm thick at the edge of the peripheral stereozone and taper gently
towards the axis. Their arrangement is more or less radial except near the cardinal
septum which is flanked on either side by three or four shorter metasepta and fronted
axially by the concave side of a tabula. The cardinal septum itself cannot be easily
distinguished but may be slightly longer than the flanking metasepta. The axial
area of the corallum contains sections of tabulae but no septal material. The
peripheral stereozone is between 3:5 and 4 mm wide which is also the length of the
wedge shaped minor septa. There are at least 60 and an estimated 64 major septa
at a mean diameter of about 33 mm and thus a septal ratio of about I-94.
In longitudinal section, the tabulae are mainly complete and spaced about I mm
or slightly less apart at the subcalicular level. Towards the proximal end, the
spacing becomes wider and less regular. The tabulae are gently undulating or very
slightly depressed across the axial area and are sharply downturned peripherally.
FROM THE SIERRA DE PERIJA, VENEZUELA 253
They appear to be inclined downwards from the convex to the concave side of the
corallum. Subsidiary tabulae occur mainly on the periphery of the wide axial
plateau formed by the complete tabulae. Evidence from the cross-section suggests
that the tabulae are depressed in the cardinal fossula. Many tabulae have signs of
septa strongly developed on their upper surfaces and some tabulae are thickly
coated with septal deposits. The closer spaced tabulae in the 6-7 mm below the
calice floor, however, are not affected in this way.
Discussion. The syntypes of Cyathophyllum venezuelense Weisbord and the
additional specimens assigned to the species by Wells when he redescribed it as
Heterophrentis venezuelensis have been re-examined with the aid of a number of new
slices from which acetate peels have been prepared. No less than five species and
four genera are thought to be represented. Only two of the original syntypes are
species of Heterophrentis and these are not conspecific. One, PRI 21594 (= PRI
24421), has been selected as the lectotype and only known example of H. (H.)
venezuelensis and is the specimen described above. The other, PRI 21593, is a speci-
men of H. (H.) simplex (Hall) and is described below. The only additional material
of either species among Bowen’s collection is a single specimen assigned to H. (H.)
simplex. Of the other syntypes PRI 21592 (Weisbord 1926, pl. 1, fig. 1) is probably
Bowenelasma typa, and PRI 21791 (Weisbord 1926, pl. I, fig. 4) is Heliophyllum hall.
Of the additional material studied by Wells, PRI 24426 (two thin sections cut from
PRI 24433 and figured by Wells 1943, pl. 10, figs 8, 9), PRI 24429 (cut from PRI
24430A) and PRI 24430B are all questionably assigned to Briantelasma oliveri.
Wells (1943 : 98) did observe that Heterophrentis venezuelensis as then understood
would probably prove, with the study of additional material, to contain more than
one species.
Heterophrentis (H.) venezuelensis is difficult to compare with the many species of
Heterophrentis (H.) named from the Devonian of North America as very few com-
prehensive illustrations and descriptions for these species exist. Furthermore, the
Venezuelan species is represented only by the lectotype so that its range of variation
is unknown. As far as can be determined at the moment, the major septa are
comparatively short in H. (H.) venezuelensis, with a weak cardinal fossula and minor
septa confined to a well developed peripheral stereozone. This appears to distin-
guish the species, at least from H. (H.) prolifica (Billings) and H. (H.) spissa (Hall),
the two species with which Wells (1943 : 98) chiefly compared H. (H.) venezuelensis.
Stumm (1965 : 20) considered H. (H.) sfissa conspecific with H. (H.) inflata (Hall)
which he suggested in turn might be a junior synonym of H. (H.) prolifica. All
these ‘species’ appear to be characteristic of the early Middle Devonian in eastern
North America. H. (H.) venezuelensis certainly appears to differ markedly from the
holotype of H. (H.) inflata illustrated by Stumm (1965, pl. 4, figs 11, 12). As noted
above the writer believes that H. (H.) simplex, a third species compared with H.
(H.) venezuelensis by Wells (1943 : 98), is actually represented by a single specimen
among the syntypes. This specimen and the holotype of H. (H.) simplex are
figured here (Pl. 5, figs 3-5) and described below. They can be seen to contrast
strongly with the lectotype of H. (H.) venezuelensis.
Cc
254 PALAEOZOIC CORAL FAUNAS
Heterophrentis (Heterophrentis) simplex (Hall) 1843
(Pl. 5, figs 3-5; Text-fig. 9)
1843 Strombodes simplex Hall: 210, text-fig. 87, 6 on p. 209, no. 48, fig. 6 on p. 44 of tables.
1876 Zaphrentis simplex (Hall) Hall pars, pl. 21, figs 5, 8-10 (non figs 6, 7, 11).
1926 Cyathophyllum venezuelense Weisbord: 4 pars, pl. 1, figs 2, 3 only.
?1938 Heterophrentis simplex (Hall) Stewart: 23, pl. 2, figs 5-7.
1965 Heterophrentis simplex (Hall); Stumm: 21, ?pars, pl. 11, fig. 19, pl. 14, figs 5, 6.
?1968 Heterophrentis simplex (Hall); Stumm: 38, pl. 1, figs 6, 7.
DiaGnosis. Ceratoid Heterophrentis (H.). Major septa three-quarters radius in
ephebic stage, thin and tapering axially with a very slight counter-clockwise deflection
in axis. Cardinal septum short in suboval fossula. Minor septa one quarter radius
in length, confined to or more commonly projecting from variably developed
peripheral stereozone.
HoLotyPe. NYSM 360. Middle Devonian, Hamilton Group; Moscow, New York.
OTHER MATERIAL. PRI 21593. Loose block (?) in Cato del Norte; Cafio Grande
Fm., early Middle Devonian (upper Onesquethaw). R49293 (3849). Camo Los
Guineos; Cafio Los Guineos Fm., early Middle Devonian (upper Onesquethaw).
DistTRIBUTION. Middle Devonian, Hamilton Group of New York, U.S.A. and
equivalent horizons elsewhere in eastern North America; early Middle Devonian of
Sierra de Perija, Venezuela.
DESCRIPTION. The Venezuelan specimens are slightly curved incomplete ceratoid
corals lacking the proximal end.
60
50
n 40
30
20
10 20 30
d (mm)
Fic. 9. Number of major septa plotted against diameter for specimens of Hetevophrentis
(H.) simplex. The holotype is indicated by an asterisk. The dashed lines join points
representing different sections of the same specimen.
FROM THE SIERRA DE PERIJA, VENEZUELA 255
In the neanic stage of PRI 21593 (PI. 5, fig. 4), the lumen is partially infilled by
major septa between 0-8 and 1-0 mm thick, parallel sided and with blunt axial ends.
These septa are mostly laterally contiguous along the axial third of their length
where they are involved in a weak counter-clockwise vortex. A small area in the
axis is free of septal material. The cardinal septum is short, 3-5 mm or half
the radius in length, and flanked by short curved metasepta concave towards the
cardinal septum. Towards the periphery the major septa are separated by narrow,
parallel sided, ovoid or gradually widening wedge-shaped gaps terminated by the
peripheral stereozone. The stereozone is rather variably developed up to 1:25 mm
thick and contains traces of rudimentary wedge-shaped minor septa. The septal
formula is C7A8K8A7C at a diameter of I5 mm.
In the ephebic stage of PRI 21593 (PI. 5, fig. 5), the major septa are very slim and
taper gently from the peripheral stereozone where they are about 0-4 mm thick to
within a quarter of the radius or slightly less of the axis. There is a weak counter
clockwise vortex. A few isolated bars of septal material may occur in the axis.
In the cardinal quadrants, three or four metasepta flanking the cardinal septum curve
to meet each other, or join sections of tabulae, across the axial end of the cardinal
septum, delimiting a strong suboval fossula. The fossula is bisected by the cardinal
septum, 4:5—-5 mm or two-fifths the radius in length. The peripheral stereozone is
variably developed from 3:3 mm thick where the minor septa are completely con-
fined to the stereozone, to 1 mm thick from which the minor septa may project up
to a further 2 mm towards the axis. There are 42 major septa at a diameter of
24 mm, a septal ratio of 1-75. Measurements are plotted in Text-fig. 9.
No longitudinal section is available.
Discussion. The Venezuelan specimens are considered conspecific with the holo-
type of H. (H.) simplex which is figured here on PI. 5, fig. 3. The polished ephebic
section of Hall’s specimen is slightly larger—30 mm diameter with 53 major septa—
but in other features the specimens are almost identical.
H. (H.) simplex has slimmer, longer major septa and a much more distinct cardinal
fossula than H. (H.) venezuelensis. Furthermore, the septal ratio in this the smaller
species (I-77 at 30 mm in the holotype) is lower than that in H. (H.) venezuelensis
(I-94 at 33 mm in the lectotype).
Family CYATHOPHYLLIDAE Dana 1846
1963 Cyathophyllidae Birenheide: 367.
1966 Cyathophyllidae Pedder: 182, pars.
1969 Phillipsastraeidae Jell: 62, pars.
Discussion. Following his valuable revision of the Cyathophyllidae, Birenheide
(1963 : 368) commented on the structural similarities between this family and the
Disphyllidae, and Pedder (1966 : 182) later united the two groups of corals as sub-
families within the Cyathophyllidae. Jell (1969; see particularly pp. 64, 69), who
on the other hand united the disphyllids with the phillipsastreids in the Phillip-
sastraeidae, has also discussed this relationship. Although disputing Pedder’s
250 PALAEOZOIC CORAL FAUNAS
classification, he did point out the close similarities of the Billingsastraeinae, a new
subfamily of his Phillipsastraeidae, to the cyathophyllids.
The writer is of the opinion that Bzllingsastraea (as widely interpreted) and
Cylindrophyllum belong together with Heliophyllum in the same suprageneric group.
This is essentially the Billingsastraeinae of Jell (1969 : 68) with the addition of
Heliophyllum. Birenheide (1963 : 368) included Heliophyllum in his undivided
Cyathophyllidae but the Heliophyllum—Billingsastraea group, although considered
here to be more closely related to the cyathophyllids than the disphyllids, deserves
to be distinguished from the former on the basis of septal characteristics (see Text-
fig. 11). Itis therefore proposed that the Billingsastraeinae and the Cyathophyllinae
be considered subfamilies of the Cyathophyllidae.
The Phillipsastraeidae is interpreted in the sense of Scrutton (1968) (= Phillip-
sastraeinae of Jell 1969 : 65) but admitting the Marisastrinae (sensu Jell 1969 : 66)
if the presence of rhipidacanths can be confirmed in Marisastrum sedgwicki. The
Disphyllidae (= Disphyllinae of Jell 1969 : 68 and ? Paradisphyllinae Jell 1969 : 67)
is also accorded full family status. Perhaps the affinities of the three families
Cyathophyllidae, Disphyllidae and Phillipsastraeidae would be best recognized at
the superfamily level.
Subfamily BILLINGSASTRAEINAE Jell 1969
Genus HELIOPHYLLUM Hall 1846
1846 Hehophyllum Hall in Dana: 356.
1938 Helophyllum Stewart: 35.
1938 Heliophyllum Fenton & Fenton: 209.
1945 Heliophyllum Smith: 25.
1947 Helhophyllum Le Maitre: 30.
1949 Helhophyllum Stumm: 21, pars.
non 1949 Heliophyllum Soshkina: 88.
1962 Heliophyllum Stumm & Tyler: 267.
1963 Heliophyllum Birenheide: 404.
Dracnosis. Solitary or colonial rugose corals with long slender major and minor
septa bearing well developed yard-arm carinae in the dissepimentarium. Septa
dilated to fill the lumen in early ontogeny; dilatation may persist in tabularium of
cardinal quadrants in ephebic stage. Cardinal fossula long, narrow and weak.
Dissepimentarium of several rows of small globose dissepiments, horizontal or
reflexed peripherally, steeply sloping downwards axially. Tabulae complete or
incomplete, usually low domes but may be flat or slightly sagging.
TYPE SPECIES (by monotypy). Strombodes helianthoides? Hall (non Phillips,
nec Goldfuss) 1843 : 209, fig. 87, 3 on p. 209, fig. 48, 3 on p. 44 of tables = Helio-
phyllum halli Edwards & Haime 1850 : 69. York, Livingstone Co., N.Y., U.S.A.;
Ludlowville Fm., Middle Devonian (Hamilton Group) (Wells 1937 : 9).
FROM THE SIERRA DE PERIJA, VENEZUELA 257
Discussion. Birenheide (1963) in a revision of cyathophyllids from the German
Devonian has confirmed the views of Wang (1948 : 11) and others that Keriophyllum
Wedekind is not congeneric with Heliophyllum. He (1963 : 392) assigned the type
species of Keriophyllum to Cyathophyllum (Peripaedium) turbinatum, type species of
Peripaedium Ehrenberg. Birenheide (1963 : 405) did regard Keriophyllum dahle-
mense Haller (1936 : 615, pl. 33, figs 1-2, pl. 34, fig. 1), however, as a valid species of
Heliophyllum.
Heliophyllum is particularly common in the Middle Devonian of eastern North
America but it has also been recorded from Europe, Asia, North Africa and South
America. Some of these records may be species of Cyathophyllum but the acceptance
of K. dahlemense as a species of Heliophyllum and the confirmation of records of
H. halli from Spain (Altevogt 1968 : 764), North Africa (Le Maitre 1947 : 31) and
Afghanistan (Brice 1971 : 263) supports this distribution in part. It is doubtful,
however, if true Heliophyllum occurs in Australia.
Heliophyllum halli Edwards & Haime 1850
(Pl. 5, figs 6-8; Pl. 6, figs 1-6; Text-figs 10, 11)
1850 Heliophyllum halli Edwards & Haime: 69.
1926 Cyathophyllum venezuelense Weisbord: 224 (4), pars., pl. 35 (1), fig. 4 only. (PRI 21791).
1937 Heliophyllum halli Edwards & Haime; Wells: 9, pl. 1, figs 1-15.
1943 Heliophyllum halli Edwards & Haime; Wells: 363 (95), pl. 36 (10), figs 1-2.
1947 Heliophyllum halli Edwards & Haime; Le Maitre: 31, pl. 1, figs 1-7, pl. 2, figs 1-6, pl. 3,
fig. I.
1965 Helophyllum halli Edwards & Haime; Stumm: 36, pl. 32, figs 5, 6, 20-23.
1968 Helophyllum hall Edwards & Haime; Altevogt: 764, pl. 2, figs 5a, b, c.
21969 ©=Hehophyllum hal Edwards & Haime; Kaplan: 28, pl. 3, fig. 1.
1970 Heliophyllum halli Edwards & Haime; Brice: 263, pl. 18, fig. 6.
Diacnosis. Usually curved, ceratoid to turbinate solitary corals but capable of
considerable shape variation and rare increase. Major septa long, uniformly attenu-
ate in dissepimentarium but may be thickened in tabularium of cardinal quadrants.
Cardinal septum slightly shorter than other metasepta in long narrow fossula, other
metasepta usually arranged with bilateral symmetry about counter-cardinal plane.
Minor septa one-third to one-half radius in length. Septa in dissepimentarium
with well developed yard-arm carinae. Dissepiments small, globose, horizontal or
inclined towards epitheca peripherally, steeply sloping towards the axis. Tabulae
wide low-domed complete plates with subsidiary domed plates in axis and small,
inclined subsidiary plates periaxially.
MATERIAL. R45076, R45078, R45080-1, R45084-6, R49250, R49252, R49296
(3121); R45087 (3124); R45092-3, R49253 (3145); R49254 (3149); R45096-7,
R49255, R49258, R49260 (3154); R49262 (3157); all Cafio Grande; Cafio Grande
Formation. R45104, R49268-9 (3199); R45107—-9, R49274 (3221); all Cafio del Sur;
Cafio Grande Formation. R45120, R45122, R49286, R49288-9 (3618); R45124
258 PALAEOZOIC CORAL FAUNAS
(3619) ; all Cafio Colorado; Cafio Grande Formation. R45111-5, R49277-8, R49281,
R49284 (3323); all from loose boulder in Cafio del Oeste; Cafio Grande Formation.
R45128, R49294-5 (3849); all Cafio Los Guineos; Cafio Los Guineos Formation.
All early Middle Devonian (upper Onesquethaw).
DISTRIBUTION. Widespread in Middle Devonian of eastern North America and
also present in the Middle Devonian of North Africa, Spain, Afghanistan and
Venezuela. Records of this species from other areas doubtful.
DESCRIPTION. Curved conical trochoid to turbinate corals attaining a maximum
mean diameter of 38 mm and a height of about 50 mm. Corallum shape not greatly
variable. Epitheca with very weak septal grooves and gently rugate. The cardinal
fossula is invariably located on the convex side of the corallum.
In neanic sections, the lumen is almost entirely closed by thickened septa. In
R45104c (Pl. 6, fig. 2), the major septa, o-8-1-1 mm thick extend almost to the
axis where their ends are coated with sclerenchyme which fills the axial area. Along
their length they are laterally contiguous in parts and separated elsewhere by a
narrow sub-parallel sided gap between 0-05 mm and 0-2 mm wide crossed at intervals
by the traces of tabulae. There is a distinct bilateral symmetry, the cardinal
septum being longer than all the other septa and flanked by pinnate groups of
metasepta. Minor septa appear mostly as short wedges up to 2:8 mm long and
0-7 mm wide between the peripheral ends of the major septa forming a solid septal
stereozone around the cardinal quadrants. In the counter qudrants, however, there
is a very narrow unthickened zone around the periphery, about half as wide as the
length of the minor septa, in which major and minor septa alike are 0-2—0-3 mm wide.
Signs of carinae are seen only rarely as incipient structures within the thickened
peripheral parts of some septa. R45104c is 14:5 mm in mean diameter with 35
major septa.
In higher sections (R45108a, R45111c), the unthickened peripheral zone in the
counter quadrants grows in width and gradually extends circumferentially across the
alar areas towards the cardinal septum. Yard-arm carinae appear, one or two per
septum, on the peripheral unthickened parts of the septa. In the axis, where there
may be a weak counter-clockwise vortex, the core of sclerenchyme disappears in
most cases and the ends of the septa, usually just short of the centre, may be gently
tapered in an axial zone about 0-2 of the diameter across. The cardinal septum
shortens and the arrangement of the flanking metasepta defines a long narrow cardinal
fossula, well developed in some sections but hardly distinguishable in others.
Septal thickening is developed on the crests of the innermost series of dissepiments
and the tabulae. It begins to diminish first in the counter quadrants and often
disappears here completely by the late neanic—early ephebic stage. The boundary
between the unthickened and thickened portions of the septa remains sharp until all
trace of thickening disappears. The cardinal quadrants, however, are usually
heavily thickened into and throughout much of the ephebic stage, often with the
septa laterally contiguous except in the long narrow cardinal fossula. The outer
edge of the thickened zone moves inwards towards and usually across the dis-
sepimentarium/tabularium boundary, and the zone itself may be as small as half the
FROM THE SIERRA DE PERIJA, VENEZUELA 259
diameter in width in some ephebic sections. On the cardinal septum and one or two
flanking metasepta, however, the thickening starts very much closer to the periphery
and serves to emphasise the cardinal fossula.
In ephebic sections the epitheca is about 0-25 mm thick and the peripheral parts
of the septa about 0-I mm across. The major septa, with the exception of the
cardinal septum and a few flanking metasepta, all reach the axial area usually with
tapered ends bilaterally arranged about the counter-cardinal axis. Nearly always
the major septa are slightly thickened in the inner part of the tabularium. Sections
such as R49278a (PI. 6, fig. 5) with no notable septal thickening at all are rare. One
specimen R45115a-b (PI. 5, fig. 6) shows an irregular network of septal material in an
axial area about a fifth the diameter across. Minor septa range from a third to just
over half the radius in length and those flanking the counter septum may be slightly
longer than their neighbours. Yard-arm carinae are developed on major and minor
septa in the dissepimentarium, rarely with a strong contrast between counter (better
developed) and cardinal quadrants. Variation from specimen to specimen is notable
with individual carinae between 0-4 and 0-65 mm in average width, 0-6 to 1:2 mm in
average spacing and between means of 3 to 6 perseptum. The intensity of carina-
tion tends to increase with increasing size and septal number and with decreasing
septal thickening. The appearance of the cardinal fossula is also variable with the
cardinal septum ranging between extremes of a quarter to two-thirds of the radius
in length in a narrow, parallel sided fossula closed axially by the incurving ends of
the adjacent metasepta. Dissepiments are intercepted on average 0-5 mm apart,
usually uniserial between adjacent septa. Up to three rows closest to the epitheca
are concave abaxially, the rest concave adaxially. They may in some cases loop
between carinae rather than from septum to septum. Traces of tabulae are wider
spaced and more diffuse.
In longitudinal section, the lower parts and much of the peripheral areas par-
ticularly on the cardinal side are infilled by sclerenchyme. The dissepimentarium
is composed of small globose dissepiments up to 1:2 mm high but normally fairly
evenly developed around 0-75 mm high. The width of the dissepimentarium
increases towards the calice and contains up to six rows of dissepiments. They are
horizontal to slightly reflexed peripherally and steeply sloping downwards axially.
The dissepimentarium is notably modified in the cardinal fossula (Pl. 6, fig. 6)
where the peripheral two or three rows of normal globose dissepiments are followed
axially after a sharp break by large axially and downward sloping vesicles. The
tabulae are complete and incomplete. The complete series are broad low domes
I-2 mm apart. The incomplete tabulae are mainly subsidiary broad low domes
across the axial four-fifths of the tabularium diameter with a few small flanking
vesicles. The sclerenchyme coating the septa is commonly seen cresting the tabulae
as well. Traces of carinae are confined to the dissepimentarium. Close to the
epitheca they slope inwards and upwards at no more than 10° to the wall, but
towards the axis they curve over to an angle of about 45° and occasionally as low as
30° to the horizontal.
Septal microstructure is only clear in parts, where uniserial monacanthine
trabeculae can be seen. The cross-bar carinae which cut, and definitely
260 PALAEOZOIC CORAL FAUNAS
precede the septa in formation have the same general appearance as septal
tissue but no details of their trabecular structure can be determined.
Measurements of septal number against diameter are plotted in Text-fig. ro.
Discussion. Although the Venezuelan specimens have a considerably smaller
range of adult diameters than topotypic Heliophyllum halli (see Text-fig. 10) there
appears to be no structural distinction of note between the two groups of specimens.
Size and size linked factors such as septal number and intensity of carination, and
possibly also the more prevalent septal thickening in the Venezuelan sample are
probably all of ecological rather than genetic importance. Furthermore, although
the legendary variability of H. halli in external form is well documented (Wells
1937), variation in internal structure is not so well known neither for this species nor
for the host of other North American ‘species’ of Heliophyllum (see for example
Stumm (1965 : 35-38)). For these reasons it seems preferable not to distinguish
the Venezuelan material in any way from H. halli.
60
50
40
30
0 10 20 30 40 50 60 70 80 90
d (mm)
———e —— Hholli (venezuela) -—-—-—- H. halli (New York State) including
———4——— H.wellsi (holotype) x H.halli OH-hallireflexum + H arachne
Fic. 10. Number of major septa plotted against diameter for specimens of Hehophyllum
hall. from Venezuela and New York State, and for specimens of H. wellst.
FROM THE SIERRA DE PERIJA, VENEZUELA 261
Heliophyllum wellsi sp. nov.
(PIG, figs-7, 8; Pll 7, figs 1, 2; Vext-figs ro; r1)
DERIVATION OF NAME. After Professor J. W. Wells (Cornell University, Ithaca,
NEY..; U.S:A.}.
Diacnosis. Large solitary Heliophyllum with dissepimentarium one-half to
three-fifths the radius. Attenuate major and minor septa densely carinate in
dissepimentarium. Carinae yard-arm, sometimes with branching ends; they may
form reticulate network with septa close to the epitheca. Axial core of reticulate
septal lamellae about one-fifth the diameter across. Dissepiments small, globose,
numerous; tabulae incomplete, tabularium horizontal or slightly domed.
HoLotyrPe. R45088 (3126). Cano Grande; Caio Grande Formation, early
Middle Devonian (upper Onesquethaw).
PARATYPE. R45089 (3126). Same locality and horizon as holotype.
DEscripTION. The specimens are large conico-cylindrical fragments of the mature
stages of two coralla. The epitheca is moderately rugate.
In cross-section the epitheca is 0-2 mm thick and only slightly penetrated by the
expanded and shallowly convex ends of the septa. The septa are extremely thin
across the broad dissepimentarium, measuring 0:05-0:10 mm across; the peripheral
expansion is variable but not more than 0:30 mm across and less than 1 mm long.
There is no sign of a cardinal fossula. In the dissepimentarium, the septa bear abun-
dant yard-arm carinae, averaging 22-24 per septum, spaced 0-7 mm apart and
reaching 2 mm in length near the periphery. The carinae towards the inner dis-
sepimentarium are shorter and closer spaced normal cross-bars but many of those
in the outer dissepimentarium have bifurcating or geniculate ends. Insome instances
septa and carinae may form an intricate reticulate network just inside the epitheca.
The minor septa are about as long as the dissepimentarium is wide, that is half to
three-fifths the radius. A slight but well defined septal thickening to 0-3-0-4 mm
in width takes place at the dissepimentarium-tabularium boundary, gently tapering
towards the axis over some 5~7 mm. _ Incipient carinae can be clearly seen embedded
in these thickened parts of the major septa in higher ephebic sections. The axial
ends of the major septa terminate against an axial area 10-20 mm across containing
a dense and rather irregular reticulate network of septal material which is occasionally
weakly carinate. The details of this axial structure show considerable variation in
the sections available.
In longitudinal section the dissepimentarium is composed of numerous small
globose dissepiments on average 0-7 mm high, horizontally arranged near the
periphery and curving over to slope steeply downward towards the axis. The
tabulae are incomplete longer, flatter, vesicular plates arranged horizontally or
slightly domed across the axis. In the axial three-fifths of the tabularium diameter,
the tabulae are disrupted by numerous, vertical and sub-parallel, wavy septal
lamellae. The dissepimentarium is crossed by the traces of carinae, very regularly
spaced 0:5-0-6 mm apart normal to their length. The carinae are inclined 25—30°
to the epitheca at the periphery, curving over gently through 40° to lie approximately
262 PALAEOZOIC CORAL FAUNAS
40° to the horizontal at the inner edge of the dissepimentarium. They may just
penetrate the tabularium, bending sharply upwards through about 20° as they do so.
Diameter and septal number in the holotype are plotted on Text-fig. 10. The
paratype, of comparable size, is too crushed to measure accurately.
The fine structure of the epitheca, dissepiments and tabulae can be clearly seen
to be fibronormal. The septa appear to be composed of unitrabecular monacanths
and the fine structure of the carinae is of septal type but is otherwise obscure.
Discussion. Heliophyllum wellsi is distinguished in general by its large size, the
intensity of the septal carination and the apparent lack of a cardinal fossula. The
most striking feature of the new species, is however, the axial core of reticulate
septal lamellae which appears to be unique to H. wellsi among species of
Heliophyllum.
Genus CYLINDROPHYLLUM Simpson 1900
1938 Cylindrophyllum Stewart: 43.
1949 Cylindrophyllum Stumm: 33.
1949 Cylindrophylium Ehlers & Stumm: 23.
Fic. 11. Comparative septal carination in (a) Cylindvophyllum elongatum (R45116a)
x25, (b) Heliophyllum halli (R45115a) x10, and (c) Heliophyllum wellsi (R45088a) x6.
FROM THE SIERRA DE PERIJA, VENEZUELA 263
Diacnosis. Dendroid or phaceloid rugose corals. Corallites with attenuate
major and minor septa and radial symmetry. Septa with variably developed yard-
arm carinae in dissepimentarium, the major septa irregular in course with occasional
lateral projections in tabularium and frequently withdrawn from axis. Dis-
sepiments small, globose, usually sloping steeply downwards towards the axis.
Tabulae mainly incomplete; tabularium surface usually bowl-shaped, occasionally
flat or more rarely arched.
TYPE SPECIES (by original designation). Cylindrophyllum elongatum Simpson
1900 : 217, text-fig. 42. Onondaga Limestone (Edgecliff Member), early Middle
Devonian; Clarksville, Albany Co., N.Y., U.S.A.
Discussion. Cylindrophyllum has been commonly placed in the Disphyllidae but
the septal characteristics in particular suggest that the genus is more likely to be
related to the Heliophyllum-Billingsastraea group (see discussion of the Cyatho-
phyllidae).
Cylindrophyllum elongatum Simpson 1900
(Pl. 7, figs 3-6; Text-fig. 11)
1900 Cylindrophyllum elongatum Simpson: 217, fig. 42.
DracGnosis. Phaceloid Cylindrophyllum with corallites up to about 20 mm
diameter. Major and minor septa almost equal in length, more or less confined to
dissepimentarium. Septa straight or zigzagged, carinae usually well developed but
may be variable to almost absent in some corallites. Dissepiments small, globose,
up to four rows deep, strongly sloping axially and downwards. Tabulae complete
or incomplete; tabularium surface flat or bowl-shaped, rarely arched.
MATERIAL. R45116 (3539). Slipped block in Cafio Pescado; Cano Grande
Formation or lower part of Cafio del Oeste Formation, early Middle Devonian
(upper Onesquethaw).
DISTRIBUTION. Onondaga Limestone; New York, U.S.A. Cafio Grande Fm.
or lower Cafio del Oeste Fm. Sierra de Perija, Venezuela. Both early Middle
Devonian.
DEscrRIPTION. The specimen is a portion, 120 mm xX I00 mm xX 70 mm, of a
phaceloid colony in which most of the corallites are partially crushed. There are
no signs of connecting processes between corallites in this colony.
In cross-section the corallites were nearly all originally circular with a regularly
developed fibronormal epitheca 0-1 mm thick. In a few cases, adjacent corallites
grow with short lengths of their epithecae cemented back to back. Major and
minor septa are uniformly attenuate, 0-03 mm thick on average, except in the
peripheral 1 mm or less in which they gradually thicken towards the epitheca. The
septa have very bluntly wedge-shaped peripheral ends reaching 0-25 mm in thickness
which may penetrate up to half the thickness of the epitheca. Major and minor
septa appear to be formed of unitrabecular monacanthine trabeculae and are dif-
ferentiated by no more than 0-3 to 0-4 mm in length. They reach between a half
264 PALAEOZOIC CORAL FAUNAS
and two-thirds the radius to the axis, the minor septa ending at the tabularium/
dissepimentarium boundary, the major septa just penetrating the tabularium. The
septa may be straight or zigzagged. A few corallites have scattered carinae on one
or two septa only but most corallites have moderately developed carinae on all or
nearly all septa. The carinae appear to be of the yard-arm type as developed in
Heliophyllum but many are geniculate with a central portion displacing the septum
obliquely and only the extremities orientated normal to the septal face, thus giving
the appearance of alternate carinae on zig-zag septa. Dissepiments usually appear
as a single series between septa of straight to slightly curved bars 0-3-0-5 mm
apart. There is a wide axial area between a third and a half the radius across free
of septa and crossed only by curved sections of tabulae. No signs of the type of
increase were observed. Because of the crushing, accurate measurements were
possible on only a few corallites. The largest individual was 10-5 mm in diameter
with 29 major septa.
In longitudinal section the dissepiments are small, globose and regularly developed
0-4-0-6 mm high. The dissepimentarium is usually four rows, or sometimes possibly
five rows of dissepiments deep. The surface slopes axially and downwards at 45°—
55° to the horizontal. The carinae crossing the dissepimentarium may be almost
parallel to the epitheca at the periphery but incline more and more towards the axis
when traced towards the inner edge of the dissepimentarium where they are 45°—50°
to the horizontal. The tabulae are complete and incomplete, arranged horizontally
or with a slight axial depression. The complete tabulae are either slightly arched
or more usually slightly sagging and are supplemented or partially replaced by large
vesicular plates.
Discussion. This specimen appears to be conspecific with photographs of the
holotype and other specimens from the Edgecliff member of the Onondaga Limestone
of New York kindly provided by Dr W. A. Oliver, Jr. (U.S. Geological Survey).
The only points of difference appear to be a tendency for the New York material to
have a higher percentage of complete tabulae and to reach a larger corallite diameter.
The diagnosis for the species given here has been constructed with the aid of these
photographs.
Family DURHAMINIDAE Minato & Kato 1965b
1965b Durhaminidae Minato & Kato: 28.
1967 Durhaminidae Stephens: 426.
1967 Durhaminidae Federowski: 25.
DracGnosis. Solitary, fasciculate, cerioid or plocoid rugose corals. Axial structure
with median counter-cardinal plate in early ontogeny, poorly developed or lost in
ephebic stage and replaced by few irregularly arranged septal ends or septal lamellae.
Minor septa variably developed and scattered third order septa may occur. Septa
usually thin, sinuous or zigzagged in dissepimentarium, slightly dilated in tabularium.
Lonsdaleoid dissepiments may be variably developed, the others concentric or
herringbone: all dip steeply axially and downwards. Tabulae mainly incomplete,
FROM THE SIERRA DE PERIJA, VENEZUELA 265
peripherally flat or trough-shaped, axially flat-topped or arched domes. Clinotabellae
usually present. Axial tabellae may or may not be clearly differentiated.
TYPE GENUS. Durhamina Wilson & Langenheim 1962 : 504.
Discussion. The specimen described below as Durhamina sp. nov. definitely
possesses poorly developed sporadic third order septa and an examination of Easton’s
(1960, text-fig. 18) illustration of the holotype of Lonsdaleia cordillerensis (lower
right quadrant of largest corallite) shows them to be present to a similar extent in
the type species of Durhamina itself. This serves to underline the extremely close
relationship between the Durhaminidae and Waagenophyllidae and is of particular
interest in view of Minato & Kato’s (1965a : 13) observations on tertiary and higher
orders of septa in the Waagenophyllidae. The diagnosis given above is emended
accordingly.
Members of the Durhaminidae occur in the Middle Carboniferous to Middle
Permian rocks of Eurasia, western North America and South America.
Genus DURHAMINA Wilson & Langenheim 1962
1962 Durhamina Wilson & Langenheim: 504.
1965b Durhamina Minato & Kato: 34.
Diacnosis. Fasciculate Durhaminidae with weakly constructed axial structure
and scattered, weakly developed third order septa. Lonsdaleoid dissepiments
sporadically developed.
TYPE SPECIES (by original designation). Lonsdaleia cordillerensis Easton 1960 :
580, Text-figs 17, 18. Lr. Permian; nr. Ruth, eastern Nevada, and other localities
in eastern Nevada and southern California, U.S.A.
Discussion. Wilson & Langenheim (1962 : 504) provided a very full comparison
of Durhamina with similar genera, supplemented by Minato & Kato (1965b : 34).
It is only necessary here to draw attention to the scattered third order septa (see
discussion of family) not hitherto recorded in the genus.
?Durhamina sp.
(Pl. 7, figs 7, 8)
MATERIAL. R49285 (3495). Cafio del Norte; Rio Palmar Formation, Lr.
Pennsylvanian.
DEscrIPTION. Solitary cylindrical fragment 9 mm maximum diameter.
In cross-section the epitheca is thin and smooth. The major septa are between
a half and the full radius in length and are very slightly thicker in the tabularium
than dissepimentarium. The longer septa meet in the axial area, the shorter septa
have free tapered ends. Minor septa are fairly evenly developed, up to a third the
radius in length. Dissepiments are concentric between them and their flanking
266 PALAEOZOIC CORAL FAUNAS
majors but weakly herringbone beyond their axial ends when they are particularly
short. There are no lonsdaleoid dissepiments. There are 21 major septa at a
diameter of 7-5 mm.
In longitudinal section the dissepimentarium is narrow, a quarter the radius in
width, and composed of steeply inclined vesicles averaging I mm long and 0-25—
o-3 mm high. The tabulae are complete and incomplete but mostly the latter.
The complete tabulae are flat or gently sagging across the axis and downturned
peripherally. The incomplete tabulae are undulating plates, inclined gently axially
and upwards to meet the vertical trace of a septum in the axis. There are scattered
clinotabellae.
Discussion. This fragment can be closely matched with corallites from species
of Durhamina in which the axial structure is weak, and is therefore assumed to be
part of a corallite detached from a fasciculate coral colony. In view of the limi-
tations of the single specimen, however, the identification must be treated with
reserve.
Durhamina sp. nov.
(Pl. 8, figs 1-10; Text-fig. 12)
MATERIAL. R49276 (3270). From a boulder in the Cano Grande; horizon un-
certain, ? Rio Palmar Fm., Lr. Pennsylvanian, but see p. 232.
DESCRIPTION. The specimen is a large fragment, some I50 x I50 x 100 mm
before cutting, of a phaceloid or in parts cerioid colony set in a tough fine-grained
reddish-brown calcareous matrix. No external details of the corallites can be
observed.
In cross-section, the corallites are circular when free and uncrushed. There are
areas in the colony where groups of three to eight corallites become laterally con-
tiguous and assume a cerioid habit. Young hysterocorallites have slim major
septa, three-fifths to the full radius in length from corallite to corallite but of fairly
uniform length within a single corallite. In the smallest corallite (Pl. 8, fig. 5),
4 mm in diameter, the major septa are about 0-07 mm thick near the periphery,
tapering towards but not reaching the axis. The counter septum, however, crosses
the axial area with a slim blade-like expansion centred on the axis, 0-8 mm long and
0-12 mm thick, which imparts a strong bilateral symmetry to the corallite. In
slightly larger corallites (Pl. 8, figs 6, 7) with longer major septa, several of them
meet in the axis often deflecting the end of the counter septum or obscuring its
identification and the bilateral symmetry is weakened or lost. The peripheral
ends of the septa are often, but not in all corallites, interrupted by a single series of
lonsdaleoid dissepiments. When the septa are not interrupted they are usually
somewhat irregular in course and thickness at the periphery. The dissepiments are
very variable in appearance. Among the lonsdaleoid series and the inner two or
three series of similar sized normal dissepiments are scattered very much smaller
FROM THE SIERRA DE PERIJA, VENEZUELA 267
and more highly globose dissepiments. The dissepiments are irregularly arranged
in a single series or weakly herringbone between adjacent major septa. Minor
septa are absent or sporadically developed as short spines on the dissepiments.
In mature corallites, most major septa extend from the epitheca to the axis with
occasional interruptions along their length. The epitheca is a very smoothly and
regularly developed fibronormal wall 0-12 mm thick. It shows no signs of septal
grooves. The septa which have slightly expanded, flat to shallowly convex or wedge
shaped peripheral ends show little or no penetration of the inner face of the epitheca.
The major septa are slim and are usually irregularly zigzagged or sinuous in the
dissepimentarium due to dissepimental interference. They are not frequently
interrupted, however, and there is no well defined series of lonsdaleoid dissepiments.
Most septa show a slight and often sharp dilatation at the inner edge of the dissepi-
mentarium associated with a slight thickening of the innermost series of dissepiments.
Major septa are between 0-1 and 0-2 mm thick at this point from which they taper
smoothly towards the axis. In any one corallite about half the major septa will meet
in the axis usually resting against an irregular median plate formed by the extended
end of what is assumed to be the counter septum. These septa may be slightly
thickened in the immediate axial area. The rest of the major septa taper to a point
just short of the axis. A few of the septa may be interrupted periaxially thus lending
definition to the axial structure but no additional septal lamellae are involved.
Also, tabulae are intercepted more frequently in the axial area. Minor septa are
extremely irregularly developed. In some corallites they extend to or just across
the dissepimentarium/tabularium boundary, or very rarely up to half way across the
tabularium to the axis. They are very thin and zigzagged in the dissepimentarium
or may be frequently interrupted. At the other extreme the minor septa may be
no more than scattered spines on the crests of a few dissepiments. The dissepi-
mentarium is between 0-3-0-4 of the radius in width and disordered in appearance.
An irregular herringbone pattern is developed either between major septa when the
minor septa are weak, or between major and minor septa when the minor septa are
strong. In the latter case, scattered third order septa are developed as very short
pegs on the epitheca and as short spines cresting dissepiments (PI. 8, fig. 2). Septal
structure, although clearly trabecular, cannot be distinguished in detail and the style
of increase cannot be determined in cross-section.
In longitudinal section the dissepiments are mixed larger, less inflated, rather
elongate vesicles and smaller, more globose vesicles. The longer vesicles may reach
up to 3-4 mm long and 1 mm high normal to the dissepimental surface, whilst the
smaller average about 0-75 mm long and 0-4 mm high. The dissepiments slope
steeply and fairly uniformly axially and downwards at about 50° to the horizontal.
The tabularium is wide, between 0-6 and 0-7 of the diameter and consists of complete
and incomplete plates. Complete tabulae are rare, fairly wide flat-topped domes
with the peripheral extremities sigmoidally curved downwards and meeting the
dissepimental wall horizontally. The incomplete tabulae are axially dome-shaped
and periaxially trough-shaped plates with occasional clinotabellae, combined to give
a broad axial plateau to the tabularium surface, which may be gently domed or
depressed across the axis. There is no sign of any great modification of the tabulae
268 PALAEOZOIC CORAL FAUNAS
in the axial area corresponding to the weak axial structure in cross-section. In
longitudinal section, increase appears to be peripheral non-parricidal or lateral.
Measurements of corallite diameter and septal number are plotted in Text-fig. 12.
Corallites reach a maximum of 18 mm diameter but data are restricted by the
partial crushing of many corallites.
Discussion. Durhamina sp. nov. compares most closely with D. cordillerensis
(Easton) from the Lr. Permian of Nevada and California although it can be distin-
guished from that species on a number of points. The most striking is the more
regular development and greater simplicity of the axial structure in the Venezuelan
species. In D. cordillerensis the axial structure is of quite variable density and may
rarely be absent. When it is well developed, however, the septal lamellae involved
are clearly distinguished from the major septa, in contrast to the very primitive
arrangement in Durhamina sp. nov. The latter also has a more strongly developed
dissepimentarium than D. cordillerensis and indeed most other species of Durhamina.
Finally the Venezuelan species has larger corallites than the North American species.
Mature corallites are about 14 mm mean diameter with 22-25 major septa, reaching a
maximum of 18 mm diameter in D. sp. nov., compared to mean diameter about
8-9 mm with 22 to 28 major septa (Easton 1960 : 580) or 20-22 major septa (Wilson
& Langenheim 1962 : 506) and maximum diameter about 15 mm in D. cordillerensis.
The Venezuelan species has not been formally named here as it is represented by
only a single specimen from a loose block of uncertain stratigraphic position.
30
20
di (mm)
Fic. 12. Number of major septa plotted against diameter for corallites in one colony
of Durhamina sp. nov.
FROM THE SIERRA DE PERIJA, VENEZUELA 269
Suborder CYSTIPHYLLINA Nicholson 1889
Family CYSTIPHYLLIDAE Edwards & Haime 1850
Subfamily PLASMOPHYLLINAE Dybowski 1873
1964 Plasmophyllinae; Birenheide ; 15.
Genus PLASMOPHYLLUM Dybowski 1873
1873 Plasmophyllum Dybowski : 340
1964 Plasmophylium (Plasmophylium); Birenheide: 16 cum syn.
Diacnosis. Solitary or weakly colonial corals in which the lumen is filled by
vesicular tissue which may or may not be thickened with sclerenchyme. Septa are
absent or represented by short irregularly developed spines. The vesicles in general
increase in size towards the axis and are arranged with a shallow bowl-shaped to
inverted sub-conical surface. Thickening when present is based on this surface
and may extend from epitheca to axis or be confined to a single concentric zone.
TYPE SPECIES (by subsequent designation of Schltiter 1885 : 10). Cyathophyllum
limbatum Quenstedt 1879 : 465, pl. 158, fig. 37 (non fig. 38) = Cyathophyllum goldfussi
Edwards & Haime 1851 (non Castelnau 1843). Gerolstein, Eifel, Germany; Ahrdorf-
Schichten, Middle Devonian (Eifelian).
Discussion. Birenheide (1964) has provided a most valuable review of the
Devonian cystimorph group of corals, placing them all, with a few exceptions, in the
subgenera Plasmophyllum (Plasmophyllum) and Plasmophyllum (Mesophyllum).
The writer feels, however, that these two groups deserve full generic status based on
the distinction in septal development between them and that their close relationship
is best indicated at the subfamily level.
Plasmophyllum secundum americanum (Edwards & Haime) 1851
(Pl. 9, figs 1-4)
1851 Cystiphyllum americanum Edwards & Haime: 464, pl. 13, figs 4, 4a.
1964 Plasmophyllum (Plasmophylium) secundum americanum (Edwards & Haime) Birenheide:
23, pl. 24, fig. 117 (cum syn.).
Diacnosis. Trochoid or turbinate tending to subcylindrical Plasmophyllum with
vesicles clearly separable into two concentric zones. Vesicles of the inner zone
slightly larger than in the outer zone and increasing in size towards the axis in both
zones. Short blunt septal spines developed, particularly in the inner half of the
outer zone. Usually sclerenchyme is heaviest in a broad concentric band or bands
coating or completely obscuring the vesicles in the outer part of the inner zone.
MATERIAL. R45008 (3157). Cafio Grande; Cafio Grande Formation, early
Middle Devonian (Upper Onesquethaw). R45106, R49270, R49272 (3221). Cafio
del Sur; Cafio Grande Formation, early Middle Devonian (upper Onesquethaw).
D
270 PALAEOZOIC CORAL FAUNAS
DIsTRIBUTION. Widespread in rocks of Middle Devonian age in North America.
Cano Grande Fm., early Middle Devonian; Sierra de Perija, Venezuela. Also
present in the Middle Devonian of Spain, U.S.S.R., south-east Asia and Australia
according to Birenheide (1964 : 24, synonymy).
DESCRIPTION. Turbinate corals up to 40 mm diameter and 70 mm height.
Calice a shallow bowl with a strongly everted rim. Epitheca moderately rugate.
In cross-section, the epitheca is fibronormal and of somewhat variable thickness,
0-I-0-3 mm thick in the small part preserved. The lumen is entirely filled by vesi-
cular tissue, fine in an outer zone one-third the radius across in smaller sections to
one-half the radius in the largest subcalicular section, and coarser in the axial zone.
In the peripheral area the vesicles are highly globose and often circular or sub-
circular in cross-section, but somewhat variable in shape and generally ranging in
diameter from 0-5 to 2mm. They may be thin walled or lightly coated with fibro-
normal material locally extended into short, blunt septal spines up to I mm long
and 0-5 mm across at the base (Pl. 9, fig. 3). The coating has a general concentric
disposition in the corallum but does not form obvious zones. Towards the axis the
vesicles become less and less globose and the circular sections disappear. In the
general region of the boundary between the zones of large and small vesicles the
thickening abruptly increases. It may form a single broad somewhat ill defined
concentric band extending well into the zone of larger vesicles and ringing a rela-
tively narrow axial zone of lighter thickening about one-fifth the diameter across.
In other sections this may grade into a series of relatively narrow concentric bands of
thickening in the zone of larger vesicles. The thickening partly obscures the septal
spines which are coarser here and are developed to within a short distance of the
axis. Often isolated spots of sectioned septal spines occur in the immediately
periaxial area. The larger vesicles average about 2 mm across periaxially and
increase in size towards the axis.
In longitudinal section, the relative width of the zone of smaller dissepiments and
the degree of eversion of its upper surface is seen to increase with increasing size.
The vesicles gradually increase in size towards the axis and the septal spines are best
developed just outside the zone of larger vesicles and heavy thickening. The spines
may reach I-7 mm in length. Their microstructure is not perfectly clear but they
appear to consist of localized broad fans in the fibronormal material. At the inner
edge of the zone of smaller vesicles the surface slope is about 50° to the horizontal.
The larger vesicles are much thickened and may be completely obscured in a dense
mass of sclerenchyme. Towards the edge of the zone the thickening can be seen as
separate cones on the surface of the vesicles but the cones tend to merge in the axial
area. The vesicles in the axial area do not exceed 3:5 mm across their bases, and are
arranged with a shallow bowl-shaped surface across the axis.
Discussion. Birenheide (1964 : 23), in his comprehensive revision of the Devonian
cystimorphs, made Cystiphyllum americanum a subspecies of the Middle Devonian
species Plasmophyllum secundum. In particular he noted that larger examples
of P. secundum americanum could scarcely be distinguished from the West European
subspecies P. secundum conistructum Quenstedt. To judge from Birenheide’s
FROM THE SIERRA DE PERIJA, VENEZUELA 271
figures, however, and from a general comparison of American and German material,
there would appear to be several points of difference. Plasmophyllum secundum
americanum has much more regularly developed vesicles, both in size and shape,
particularly in the peripheral areas of the corallum. Furthermore, the development
of short blunt wedge-shaped septal spines, most conspicuous in the inner part of the
zone of smaller vesicles, is a characteristic feature of P. secundum americanum but
does not appear to be present in P. secundum conistructum. On the other hand,
typical P. secundum americanum, for example R45008 figured here (PI. 9, figs 1-4),
compares remarkably well with Wedekind & Vollbrecht’s (1931, pl. 17, figs 1-2)
illustrations of Paralytophyllum crassum (Wedekind) from the Givetian of Germany.
In particular, the development of septal spines and the size are more or less identical,
as is the calice shape, although this appears to be more variable in P. secundum
americanum as a whole. Birenheide (1964 : 28) listed Parvalytophyllum crassum in
his synonymy for P. secundum pseudoseptatum (Schulz). The fact remains that but
for the geographical isolation, any sort of distinction between the American material
and European P. secundum would be difficult to maintain. Birenheide’s synonymy
for P. secundum americanum is accepted uncritically.
Plasmophyllum sp.
(PI. 9; figs 5= 7)
1943 Zonophyllum sp. Wells: 98, pl. ro, figs 5, 6.
MATERIAL. R45095, R49259 (3154), PRI 24425 (PRI 24427 cut from same
specimen). Cafio Grande; Cafio Grande Fm., early Middle Devonian (upper
Onesquethaw).
DESCRIPTION. Small subcylindrical fragments 12-16 mm in diameter. The
corallum may be strongly constricted at intervals as a result of rejuvenation.
In cross-section the vesicles appear rather flattened sub-semicircular and variable
in size from about 0-5 mm to 5 mm across their bases. The epitheca and two or
three concentric internal surfaces are heavily thickened with fibronormal scleren-
chyme. The external wall thus varies between 0-6 and 1-5 mm in thickness. When
it is thinner, short blunt septal spines can be distinguished extending inwards about
I mm from the periphery and approximately 0-6 mm apart. Thickening of the wall
tends to obscure these spines completely. Similar septal spines may or may not
occur associated with inner zones of thickening. The vesicles show a general
increase in size towards the axis but the gradation, although irregular, is more or
less continuous and inner and outer size zones cannot be distinguished.
In longitudinal section, the vesicles slope axially and downwards at the periphery
and flatten out across the axis giving a bowl-shaped calice. The upper surface of
the vesicles is frequently thickened. At intervals up the length of the corallite the
lumen is entirely filled by sclerenchyme, more heavily deposited at the periphery so
that vesicular tissue is seen only infrequently to reach the epitheca unobscured.
D*
272 PALAEOZOIC CORAL FAUNAS
Discussion. Wells (1943 : 98) originally described some of this material as
Zonophyllum sp. Following Birenheide’s (1964) revision of the cystimorphs, the
Venezuelan material clearly belongs to the group of Zonophyllum he regards as
congeneric with Plasmophyllum.
Wells (op. cit.) noted that the Venezuelan material was similar in structure to
Cystiphyllum contfollis Hall (1876, pl. 30, figs 3-9) as redescribed by Fenton & Fenton
(1938 : 232, fig. 20, pl. 23, figs 9, 10, pl. 24, figs 4-7). Unfortunately the Fentons
did not section and figure Hall’s type material, all of which looks from Hall’s figures
to be too highly cystose to be closely related to the Venezuelan species. On the
other hand, one specimen figured by Fenton & Fenton (1938, pl. 24, figs 5-7) from
the Hamilton Group (Moscow Fm.) of Leicester, N.Y. as ‘Cystiphyllum’ conifollis
looks extremely similar to the Venezuelan specimens and may indeed be conspecific
with them. It seems best therefore neither to use the name conzfollis nor to erect a
new species until Hall’s original material has been redescribed.
Order TABULATA Edwards & Haime 1850
Suborder FAVOSITINA Sokolov 1962
Family FAVOSITIDAE Dana 1846
Genus FAVOSITES Lamarck 1816
1816 Favosites Lamarck: 204.
1851 Emmonsia Edwards & Haime: 152, 245.
1939 Favosites Lecompte: 80, cum syn.
1953 Favosites Ross: 48, cum syn.
1960 Favosites Philip: 196.
1965 Favosites Stumm: 60.
1965 Emmonsia Stumm: 65.
DraGnosis. Massive, ramose, foliaceous or encrusting cerioid corals. Corallites
generally contiguous, polygonal, of similar diameter throughout their length.
Corallite walls unthickened internally, but may be variably dilated in a thin peripheral
zone. Mural pores confined to corallite wall faces, usually one or two but may be
three to a face, arranged in vertical series. Septa present in some species, may be
variably developed usually as spines in vertical rows but also as septal striae.
Squamulae also present in some species, with or without septal spines, and also may
be variable in development. Tabulae usually complete and horizontal, frequently
suspended in squamulate forms.
TYPE SPECIES (see Edwards & Haime 1850: 60). F. gothlandicus Lamarck
1816 : 205 = Specimen . . . corallinum Gothlandicum Linnaeus 1745 : 39, fig. 27 and
1749 : 106, pl. 4, fig. 27.
Discussion. Considerable discussion of the genus Favosites has been published.
It is sufficient here to state that the writer accepts Philip’s (1960 : 190) opinion on
the status of squamulae in Favosites and also considers that Thamnopora should be
maintained as a distinct genus.
FROM THE SIERRA DE PERIJA, VENEZUELA 273
Favosites venezuelensis (Weisbord) 1926
(Pl. ro, figs I, 2)
1926 Pleuvodictyum venezuelense Weisbord: 6, pl. 1, figs 8, 9.
1943 Thamnopora venezuelensis (Weisbord) Wells: 99, pl. Io, fig. ro.
Diacnosis. Favosites, apparently nodular in form, with monomorphic corallites,
up to 2 mm diameter, having walls unthickened within the colony. Corallite walls
heavily thickened in a subcalicular zone 5~7 mm deep so as to completely close the
lumen in most cases. Squamulae present, particularly well developed in thickened
zone. Mural pores uniserial, sporadically biserial, 0-3 mm diameter. Tabulae
complete, horizontal, 7 in 5 mm.
Hototyre. PRI 21597 (PRI 24428 same specimen). Loose block (?) in Cafio
del Norte; Caio Grande Fm., early Middle Devonian (upper Onesquethaw).
MATERIAL. R45079 (3121). Cafio Grande; same horizon as holotype. R45101
(3182). Cano del Sur; same horizon as holotype.
DEscrRIPTION. Nodular colonies; R45101 is partly crushed, 85 mm long and
about 50 mm maximum diameter.
Internally, the corallites are polygonal in cross-section, 3 to 4 sided in immature
stages and mainly 6 to 7 sided when mature. They range up to almost 2 mm dia-
meter when measured between the axial plates of opposite walls but mature corallites
average about I-4 mm diameter. The corallite walls are thin with the axial plate
about 0-01—0-02 mm thick coated on either side by fibronormal tissue 0-02 to 0-04 mm
thick in mid-wall positions. The fibronormal tissue thickens slightly into the corners
of the corallites which are slightly rounded. Mural pores appear sporadically in mid-
wall or slightly offset positions. No definite indications of septal spines or squamulae
are seen in cross-sections through the centre of the colony.
Tangential sections close to the colony surface show variable degrees of thickening
of the corallite walls (Pl. ro, fig. 2). Thickening first builds up in the angles of the
corallite walls so that the lumen becomes increasingly rounded. At the same level
increasing signs of squamulae appear, of variable size with truncate or bluntly
tapering ends. There are also very rare indications of what may be septal spines.
Towards the colony surface, wall thickening may build up to almost or completely
close the corallites. The mural pores however appear to be least affected by this
excess thickening and even maintain close to their usual diameter when the corallites
they connect are apparently closed. At the colony surface, the thickening rapidly
diminishes leaving calices rounded and separated by walls about 0-3 mm thick at
their midpoints.
In longitudinal section the tabulae are complete, flat or slightly arched or sagging,
and fairly wide spaced 7in5 mm. They either cross from wall to wall or link facing
squamulae. The squamulae are flat to very faintly trough-shaped plates directed
horizontally in most cases. They are up to 0-8 mm long, 0:5 mm wide and become
more strongly developed as wall thickening begins to build up where they average
0-6 mm apart vertically. Corallites are normal to the colony surface for about Io
mm or rather more of their length and the corallite walls start to thicken quite
274 PALAEOZOIC CORAL FAUNAS
rapidly about 5-7 mm from the surface. Associated with the thickening, both
squamulae and tabulae become more numerous, the former also becoming increasingly
thickened. Mural pores are sectioned more frequently due to the thicker walls.
They are large and circular about 0-3 mm diameter and usually 1 mm apart in single
vertical rows. Most corallite walls show occasional doubling up of the pores with
variable horizontal spacing. Many pores appear to be capped by squamulae but the
relationship is apparently not invariable. In longitudinal section, the calices
appear to be bow] or funnel shaped and I to 2 mm deep for the most part but there is
considerable variability. Most but not all of them are floored by a zone where the
infilling of the corallites is more or less complete.
Discussion. This description is based mainly on the new material collected by
Bowen, particularly R45101, and considerably supplements the previous descriptions.
In particular, the well developed squamulae in this species have not been recorded
before.
The species should clearly be placed in Favosites as that genus is now understood.
The character of the wall thickening in F’. venezuelensis although apparently unique
in its development is comparable with that seen in many other species of Favosites
(for example see Ross 1953, pl. 21, fig. 6; Philip 1960, pl. 24, fig. 4), and is essentially
a mature surface feature of the colony. In Thamnopora the corallites are more or
less rounded throughout the colony.
F.. venezuelensis shows some similarities with Emmonsia radiciformis (Stewart
1938 : 69, pl. 16, figs 8-10, non Rominger 1876 : 34, pl. 12) and to a lesser extent
with Emmonsia carmani Stewart (1938 : 67, pl. 15, figs 1-3). Both these species
from the Middle Devonian of Ohio, however, appear to be more intensely squamulate
and lack the strong thickening of the distal corallite walls.
Favosites arbuscula Hall 1876
(Pl. ro, figs 3-6)
1876 Favosites arbuscula Hall, pl. 36, figs 1-9.
1953 Favosites (Emmonsia) arbuscula (Hall); Ross: 54, pl. 14, figs 2-9.
1965 Emmonsia arbuscula (Hall); Stumm: 65, pl. 64, fig. 13.
Di1aGNosis. Ramose Favosites with branches averaging 14 mm diameter. Coral-
lites oblique or perpendicular to surface, 1:2 to 1°38 mm mean mature diameter.
Walls 0-1 mm thick with little or no thickening at aperture. Mural pores 0-2 mm
diameter, uniserial. Squamulae abundant, sometimes concentrated in distinct
zones. Tabulae thin, often suspended from squamulae, I-1-5 mm apart.
MATERIAL. R45102-3, R49264-5 (3182). Cano del Sur; Caio Grande Formation,
early Middle Devonian (upper Onesquethaw).
DisTRIBUTION. Middle Devonian (Hamilton Group and equivalent units) of
eastern North America. Cafio Grande Fm. (early Middle Devonian) of the Sierra de
Perija, Venezuela.
FROM THE SIERRA DE PERIAJ, VENEZUELA 275
DESCRIPTION. Colonies ramose with branches averaging 14 mm diameter, close
packed or diverging. Largest fragment 80 mm long branching twice in planes at
right angles to each other.
In cross-section the corallites are three or four sided when immature, six or seven
sided when mature. There is a tendency for mature and immature corallites to be
arranged in a regular pattern. Mature corallite diameters reach 1-8 mm and average
about 1-5 mm. Their walls are thin, the axial plate about 0-02 mm thick flanked on
either side by fibronormal tissue 0-04 mm thick. There is only slight rounding of
corallite anglesinternally. Mural pores are infrequently sectioned, always close to the
mid-wall position and signs of septal spines or squamulae are extremely rare.
At the colony surface, corallite walls are slightly thickened up to about 0-25 mm
from side to side and often irregularly scalloped or provided with very short stubby
spines. The corallites are nearly all polygonal with rounded corners and are only
rarely subcircular. Mural pores are sectioned more frequently and occasional signs
of squamulae are seen.
In longitudinal section, the corallites diverge at a low angle to the axis of the
branch and curve over to open at the surface somewhat obliquely. Internally
corallite walls show slight irregular thickening and thinning with a more marked
but only slightly thicker zone within 1 to 1-5 mm of the branch surface. Mural
pores are infrequently sectioned up to 0-2 mm diameter and of indeterminate vertical
spacing. Squamulae are present as generally unthickened plates of flat to shallow
trough-shaped section, horizontally or slightly upwardly directed and about 0-5 mm
long. They are concentrated about 0-25 mm vertically apart in distinct zones I to
2 mm deep and with a vertical frequency of 4 to 5 mm. The tabulae are complete,
unthickened, flat to gently undulating and mostly wide spaced I to 1-5 mm but
sometimes up to 4mm apart. They are often suspended from the squamulae.
Discussion. The Venezuelan material appears to agree in most respects with
Favosites arbuscula Hall as redescribed by Ross (1953 : 54). Whether or not
Favosites digitata Rominger (1876 : 39, pl. 15) is conspecific with and therefore a
senior synonym of F. arbuscula cannot be determined in the absence of a modern
description and illustration of Rominger’s species. For the present it seems
preferable to accept the well described F. arbuscula.
The Venezuelan material possesses a combination of characters within the range of
F. arbuscula but not matched exactly in any of Ross’ (1953 : 55) subspecies. No
new subspecies designation is felt to be justified, however, and the material is referred
to F. arbuscula sensu lato.
VI. REFERENCES
ALTEvocT, G. 1968. Devonian tetracorals from Spain and their relation to North American
species. In Oswatp, D. H. (Ed.) International Symposium on the Devonian System,
Vol. 2 : 755-769, 2 pls. Calgary.
ARMSTRONG, A. K. 1962. Stratigraphy and palaeontology of the Mississippian System in
southwestern New Mexico and adjacent southeastern Arizona. Mem. Inst. Min. Technol.
New Mex., Socorro, 8 : i-vi, 1-99, pls 1-12.
ARNOLD, H. C. 1966. Upper Palaeozoic Sabaneta-Palmarito sequence of Mérida Andes,
Venezuela. Bull. Am. Ass. Petrol. Geol., Tulsa, 50 : 2366-2387.
276 PALAEOZOIC CORAL FAUNAS
BASSLER, R.S. 1937. The Paleozoic rugose coral family Palaeocyclidae. J. Paleont., Tulsa,
11 : 189-201.
Biriincs, E. 1858. Report for the year 1857 of E. Billings, Esq., Palaeontologist, addressed
to Sir W. E. Logan, F.R.S., Director of the Geological Survey of Canada. Geol. Suv.
Canada, Report of Progress for the year 1857 : 147-192.
1875. On some new or little known fossils from the Silurian and Devonian rocks of
Ontario. Can. Nat., Montreal, N.S., 7, 4 : 230-240.
BIRENHEIDE, R. 1963. Cyathophyllum- und Dohmophyllum- Arten (Rugosa) aus dem Mittel-
devon der Eifel. Senckhenburg. leth., Frankfurt a.M., 44 : 363-458, pls 46-62.
1964. Die ‘Cystimorpha’ (Rugosa) aus dem Eifeler Devon. Abh. Senckenb. naturforsch.
Ges., Frankfurt a.m., 507 : 1-120, 28 pls.
Bowen, J. M. 10972. Estratigrafia del Precretaceo en la parte norte de la Sierra de Perija.
Boln Geol. Dir. Geol. Publ. Espec., Caracas, 5 : 729-761.
Brice, D. 1971. Etude paléontologique et stratigraphique du Dévonien de 1’Afghanistan.
Contribution 4 la connaissance des brachiopodes et des polypiers rugueux. Notes Mém.
Moyen-Orient, Paris, 11 : i-viii + 1-364, pls 1-20.
Buscu, D.A. 1941. An ontogenetic study of some rugose corals from the Hamilton of western
New York. J. Paleont., Tulsa, 15, 4 : 392-411, 73 text-figs.
Curist, P. 1927. La coupe géologique le long du chemin de Mucuchachi a Santa Barbara
dans le Andes Vénézueliennes. FEclog. geol. Helv., Lausanne, 20, 4 : 769-792.
Dana, J. D. 1846. Genera of fossil corals of the family Cyathophyllidae. Am. J. Sci. Arts.,
New Haven, (2), 1 : 178-189, text-figs 1-5.
DE Groot, G. E. 1963. Rugose corals from the Carboniferous of Northern Palencia (Spain).
Doctor. Thesis Wis. Naturk. Rijksuniv. Leiden, 1963 : 1-123, pls 1-26.
DysBowsk1, W. N. 1873-4. Monographie dey Zoantharia sclerodermata vugosa aus dey Silur-
formation Estlands, Norvd-Livlands und der Insel Gotland . . ., pp. 257-532, pls 1-5, Dorpat.
Easton, W. H. 1960. Permian corals from Nevada and California. J. Paleont., Tulsa, 34,
3: 570-583, 18 text-figs.
Epwarps, H. M. & Hatme, J. 1850. A Monograph of the British Fossil Corals, Part 1.
Introduction, etc.: i-Ixxxv + 1-71, pls 1-11. Palaeontogr. Soc. [Monogy.], London.
1851. Monographie des Polypiers fossiles des Terrains palaeozoiques. Avch. Mus. natn.
Hist. nat. Paris, 5 : 1-501, pls. 1-20.
Euters, G. M. & Stumm, E. C. 1949. Corals of the Devonian Traverse Group of Michigan.
Part II, Cylindvophyllum, Depasophyllum, Disphyllum, Endophyllum and Synaptophyllum.
Contr. Mus. Paleont. Univ. Mich., Ann Arbor, 8, 3 : 21-41, 8 pls.
FEDOROWSKI, J. 1965. Lindstroemiidae and Amplexocariniidae (Tetracoralla) from the
Middle Devonian of Skalty, Holy Cross Mountains, Poland. Acta palaeont. pol., Warszawa,
10 : 335-363, pls 1-6.
1967. The Lower Permian Tetracoralla and Tabulata from Treskelodden, Vestspits-
bergen. Norsk Polavinstitutt Skriftey, Oslo, 142 : 1-44, 7 pls.
Fenton, C. L. & Fenton, M. A. 1938. Heliophylium and ‘Cystiphyllum’ corals of Hall’s
‘Illustrations of Devonian corals’. Ann. Cavneg. Mus., Pittsburgh, 27 : 207-250, pls 1-24.
Fontaine, H. 1961. Les Madréporaires paléozoiques du Viét-Nam, du Laos et du Cambodge.
Archs géol. Viét-Nam, Saigon, 5 : 1-276. Atlas, pls 1-35.
Girty, G. H. 1897. A revision of the sponges and coelenterates of the Lower Helderberg
Group of New York. New York State Mus., Ann. Rept, Albany, 48, 2 : 259-322.
Hat, J. 1843. Geology of New York, pt. 4, comprising the survey of the Fourth geological
district, pp. i-xxvill + 1-683, tables pp. 1-68, pls 1-19, Albany, N.Y.
1876. Illustrations of Devonian fossils; corals of the upper Helderberg and Hamilton
groups, pp. 1-7, pls 1-39. Albany N.Y.
Haier, W. 1936. Einige biostratigraphische Untersuchungen in der Rohrer Mulde unter
besonderer Beriicksichtigung der Keriophyllen. Jb. preu8. geol. Landesanstalt, Berlin,
56 : 590-632, pls 32-43.
FROM THE SIERRA DE PERIJA, VENEZUELA 277
Hea, J. P. & Wuitman, A.B. 1960. Estratigrafia y petrologia de los sedimentos precretacios
de la parte norte-central de la Sierra de Perija, Estado Zulia, Venezuela. Boln Geol. Dir.
Geol. Publ. Espec., Caracas 3, I : 351-376.
Hitt, D. 1956. Rugosa. In Moore, R. C. (Ed.) Treatise on Invertebrate Paleontology, F :
233-324, text-figs 165-219. Lawrence, Kansas.
Ivanovskil, A. B. 1963. Rugozy ordovika i Silura Sibirskoi Platformy, pp. 1-160, 33 pls.
Nauka, Moskva. (In Russian).
1965. Dyrevneisie vugozy, pp. 1-152, 39 pls. Nauka, Moskva. (In Russian).
Jerrorps, R.M. 1955. Septal arrangement and ontogeny of some porpitid corals. Palaeont.
Contr. Univ. Kans., Lawrence, Coelent. Art 2 : 1-16.
JELL, J. S. 1969. Septal microstructure and classification of the Phillipsastraeidae. In
CAMPBELL, K. S. W. (Ed.) Stratigraphy and Palaeontology : Essays in honour of Dorothy Hill:
50-73, pls 7, 8. Canberra.
Karjo, D. L. 1961. Dopolneniya k izucheniyu streptelazmid ordovika Estonii. Geoloogia
Inst. Uurim., Tallinn, 6 : 51-67, pls 1-4.
Kapran, A. A. 1969. Chetyrekhluchevye korally zhivetskikh otlozhenii beyanaula, chingiza
ipredchingizya. Izv.vyssh.ucheb. Zaved. geol. vaz. Moscow, 12, 1 : 19-29, 4 pls. (In Russian.)
Kato, M. 1963. Fine skeletal structures in Rugosa. J. Fac. Sci. Hokkaido Univ. (4, geol.
min.) 11 : 571-630, 3 pls.
Lamarck, J. B. P. DE 1816. AHistoive naturelle des animaux sans vertébres, 2 : 1-568. Paris.
LrecompteE, M. 1939. Les tabulés du Dévonien moyen et supérieur du bord sud du bassin de
Dinant. Mém. Mus. r. Hist. nat. Belg., Bruxelles, 90 : 1-229, pls 1-23.
Le Mairre, D. 1947. Le récif coralligene de Ouihalane. Notes Mém. Serv. Mines Carte géol.
Maroc, 67 : 1-112, pls 1-24.
1952. La faune du Dévonien Inférieur et Moyen de la Saoura et des abords de l’Erg el
Djemel (Sud-Oranais). Matér. Carte géol. Algéy., Alger, 1. Paléont., 12 : 1-171, 22 pls.
Lippte, R. A., Harris, G. D., & WELLS, J. W. 1943. The Rio Cachiri section in the Sierra
de Perija, Venezuela. Bull. Am. Paleont., Ithaca, 27 : 269-368 (1-100), pls 27-36 (1-10).
Linnaeus, C. 1745. Dissertatio, Coralia Baltica adumbrans, Quam... submittit Henricus
Fougt, pp. viii + 1-40, 1 pl. Upsala.
1749. Corallia Baltica ... proposita ab Henrico Fougt. Amoen. Acad. Stockholm &
Leipzig, 4 : 74-106, pl. 4.
MILLER, J. B. 1962. Tectonic trends in Sierra de Perija and adjacent parts of Venezuela and
Columbia. Bull. Am. Assoc. Petyol. Geol., Tulsa 46 : 1565-1595, 15 figs.
Mititer, S. A. 1889-1897. North American geology and palaeontology .. . 3vd Edition,
pp. 1-664 (1889). First Appendix, pp. 665-718 (1892). Second Appendix, pp. 719-793
(1897). Cincinnati.
Mryato, M. 1961. Ontogenetic study of some Silurian corals of Gotland. Stockh. Contr.
Geol., 8, 4 : 37-100, pls I-22.
Minato, M. & Kato, M. 1965a. Waagenophyllidae. J. Fac. Sci. Hokkaido Univ. (4, geol.
min.), 12, 3-4 : 1-241, 20 pls.
1965b. Durhaminidae (Tetracoral). J. Fac. Sci. Hokkaido Univ. (4, geol. min.), 13,
I : 11-86, 5 pls.
MirtcHEeLL, M. 1966. Comment on the proposed designation of a type-species for Amplext-
zaphventis Vaughan, 1906 (Anthozoa). Z.N. (S.) 1669. Bull. zool. Nomencl. London,
23, 2-3 : 82-83.
Moore, R. C., Hirt, D. & WELLs, J. W. 1956. Glossary of morphological terms applied to
corals. In Moore, R.C. (Ed.) Treatise on Invertebrate Paleontology, F : 245-251. Lawrence,
Kansas.
Moorg, R. C. & JeFrorps, R. M. 1945. Description of Lower Pennsylvanian corals from
Texas and adjacent areas. Publs. Bur. econ. Geol. Univ. Tex. Austin, 4401 : 77-208, 214 figs.
Morates, P. A. 1965. A contribution to the knowledge of the Devonian faunas of Colombia.
Boln Geol. Fac. Petrol. Univ. ind. Santander 19 : 51-111, 9 pls.
278 PALAEOZOIC CORAL FAUNAS
NeEumAN, B. 1969. Upper Ordovician streptelasmatid corals from Scandinavia. Bull. geol.
Instn Univ. Uppsala, N.S. 1, 1 : 1-73, 59 figs.
O’ConNELL, M. 1914. Revision of the genus Zaphrentis. Ann. N.Y. Acad. Sci., 23 : 177-192.
OxtiverR, W. A., Jr. 1958. Significance of external form in some Onondagan rugose corals.
J. Paleont., Tulsa, 32, 5 : 815-837, pls 104-106.
1960a. Rugose corals from reef limestones in the Lower Devonian of New York. Jf.
Paleont., Tulsa, 34, 1 : 59-100, pls 13-19.
1960b. Devonian rugose corals from northern Maine. Bull. U.S. Geol. Surv., Washington,
1111A : 1-23, pls I-5.
1963. Anew Kodonophyllum and associated rugose corals from the Lake Matapedia area,
Quebec. Prof. Pap. U.S. geol. Suvv., Washington, 430-C : 21-31, pls 9-14.
1968. Succession of rugose coral faunas in the Lower and Middle Devonian of eastern
North America. In Oswatp, D. H. (Ed.) Inteynational Symposium on the Devonian
System, Vol. 2 : 733-744. Calgary.
OPINION 854. 1968. Amplexizaphrentis Vaughan, 1906 (Anthozoa) : designation ofa type-species
under the plenary powers. Bull. zool. Nomencl. London, 25, 2-3 : 82-83.
PEDDER, A. E. H. 1966. The Devonian tetracoral Haplothecia and new Australian phacello-
phyllids. Proc. Linn. Soc., N.S.W., Sydney, 90 : 181-189, pl. 6.
Puitip,G.M. 1960. The Middle Palaeozoic squamulate favositids of Victoria. Palaeontology,
London, 3 : 186-207, pls 30-34.
QUENSTEDT, F. A. von. 1878-81. Petvefactenkunde Deutschlands, 6. Die Rohren- und Stern-
korallen., pp. 1-144 (1878); 145-624 (1879); 625-912 (1880); 913-1094 (1881); and Atlas.
Leipzig.
ROMINGER, C. 1876. Palaeontology. Fossil Corals. Geol. Surv. Michigan, Lansing, 3,
2 : I-161, 55 pls.
Ross, M. H. 1953. The Favositidae of the Hamilton Group (Middle Devonian of New York).
Bull. Buffalo Soc. nat. Sct., 21 : 37-89, pls 12-27.
Rowett, C. L. & SUTHERLAND, P. K. 1964. Wapanucka Rugose Corals. Bull. Okla geol.
Suvv. Norman, 104 : 1-124, pls 1-9.
SCHEFFEN, W. 1933. Die Zoantharia Rugosa des Silurs auf Ringerike im Oslogebiet. Shr.
norske Vidensk-Akad. mat.-nat. Kl., Oslo, 1932, 5 : 1-64, 11 pls.
SCHLUTER, C. 1885. Ueber neue Korallen aus dem mitteldevon der Eifel. Sber. niederrhein.
Ges. Nat. u. Hetlk., Bonn, 1885 : 6-13.
Scrutton, C. T. 1968. Colonial Phillipsastraeidae from the Devonian of south-east Devon,
England. Bull. By. Mus. nat. Hist. (Geol.), 15, 5 : 181-281, 18 pls.
1971. Palaeozoic coral faunas from Venezuela, I. Silurian and Permo-Carboniferous
corals from the Mérida Andes. Bull. Br. Mus. nat. Hist., (Geol.), 20, 5 : 183-227, 5 pls.
SHRESTHA, C. L. 1966. Amplexizaphrentis Vaughan, 1906 (Anthozoa): proposed designation
of a type-species under the plenary powers. Z.N. (S.) 1669. Bull. zool. Nomencl. London,
22, 5-6 : 348-350.
Srupson, G. B. 1900. Preliminary descriptions of new genera of Paleozoic rugose corals.
Bull. N.Y. St. Mus. Albany, 8, 39 : 199-222.
SmitH, S. 1945. Upper Devonian corals of the Mackenzie River region, Canada. Spec.
Pap. geol. Soc. Am., New York, 59 : vill + 1-126, pls 1-35.
SosHKINA, E. D. 1949. Devonskie korally Rugosa Urala. Tyvudy paleont. Inst., Leningrad,
15 : 1-160, pls 1-58. (In Russian.)
SosHxina, E. D. & Kapaxovicu, N. V. 1962. Podotryad Streptelasmatina. In ORLov,
U. A. (Ed.) Osnovy paleontologii, 2. Gubki, Arkheotsiaty, Kishechnopolostnye, Chervi:
317-324. Moscow. (In Russian.)
STEPHENS, C. H. 1967. Leonardian (Permian) compound corals of Nevada. J. Paleont.,
Tulsa, 41, 2 : 423-431, pls 52-54.
STEWART, G. A. 1938. Middle Devonian corals of Ohio. Sp. Pap. Geol. Soc. Am., New York,
8 : vii + 1-120, 20 pls.
FROM THE SIERRA DE PERIJA, VENEZUELA 279
Stumm, E. C. 1949. Revision of the families and genera of the Devonian Tetracorals.
Mem. geol. Soc. Am., New York, 40 : viii + 1-92, pls 1-25.
1962. Corals of the Traverse Group of Michigan. Part VIII, Steveolasma and Hetero-
phrentis. Conty. Mus. Paleont. Univ. Mich., Ann Arbor, 17, 10 : 233-240, 2 pls.
1965. Silurian and Devonian corals of the Falls of the Ohio. Mem. geol. Soc. Am.,
New York, 93 : x + 1-184, 80 pls. (Dated 1964 but published January 1965, vide Dr W.
A. Oliver, Jr.).
1968. The corals of the Middle Devonian Tenmile Creek Dolomite of northwestern Ohio.
Contry. Mus. Paleont. Univ. Mich. Ann Arbor, 22, 3 : 37-44, 3 pls.
Stumm, E. C. & Tyrer, J. H. 1962. Corals of the Traverse Group of Michigan. Part IX,
Heliophyllum. Contr. Mus. Paleont. Univ. Mich., Ann Arbor, 17, 12 : 265-276, 3 pls.
Stumm, FE. C. & Watkins, J. L. 1961. The metriophylloid coral genera Steveolasma, Amplexi-
phyllum and Stewartophyllum from the Devonian Hamilton Group of New York. /.
Paleont., Tulsa, 35, 3 : 445-447, pl. 58.
SUTHERLAND, P. K. 1958. Carboniferous stratigraphy and rugose coral faunas of Northeastern
British Columbia. Mem. Geol. Surv. Can., Ottawa, 295, 1-177, pls 1-33.
1970. A redescription of the Silurian rugose coral Syvingavon siluriense (McCoy).
J. Paleont., Tulsa 44, 6 : 1125-1128, pl. 152.
SUTHERLAND, P. K. & HauGu, B. N. 1969. The discoid rugose coral Gymnophyllum : growth
form and morphology. Jn CampBeLL, K. S. W. (Ed.) Stratigyvaphy and Palaeontology :
Essays in honour of Dorothy Hill: 27-42, pls 3, 4. Canberra.
Sutton, F. A. 1946. Geology of Maricaibo Basin, Venezuela. Bull. Am. Ass. Petrol. Geol.,
Tulsa, 30 : 1621-1741, pls 1-9.
THomeson, J. 1881. Onthe genus Alveolites, Amplexus, and Zaphrentis, from the Carboniferous
System of Scotland. Proc. R. phil. Soc. Glasg., 13 : 194-237, 4 pls.
VauGHAN, A. 1906. An account of the faunal succession and correlation. In Mat Ley, C. A.
& VAUGHAN, A. The Carboniferous rocks at Rush (County Dublin). Q. Jl geol. Soc. Lond.,
71: 1-52.
Wane, H. C. 10948. The Middle Devonian rugose corals of eastern Yunnan. Contr. Geol.
Inst., Nat. Univ. Peking, 33 : 1-45, 5 pls.
Wessy, B.D. 1964. Devonian corals and brachiopods from the Brendon Hills, West Somer-
set. Palaeontology, London, 7, 1 : 1-22, pl. 1.
WEDEKIND, R. & VOLLBRECHT, E. 1931. Die Lytophyllidae des mittleren Mitteldevon der
Eifel. Palaeontographica, Stuttgart, 75 : 81-110, pls 15-46.
WEISBORD, N. E. 1926. Venezuelan Devonian Fossils. Bull. Am. Paleont., Ithaca, 11 :
221-256 (1-36), pls 35-41 (1-7).
1968. The Devonian System in Western Venezuela. Jn Oswatp, D. H. (Ed.) Inter-
national Symposium on the Devonian System, Vol. 2 : 215-226, 2 figs. Calgary.
We tts, J. W. 1937. Individual variation in the rugose coral species Heliophyllum halk
E. & H. Palaeontogr. Am., Ithaca, 2, 6 : 1-22, pl. 1.
1943. Anthozoa. Jn LippiE, R. A., Harris, G. D. & WELLS, J. W. The Rio Cachiri
section in the Sierra de Perija, Venezuela. Bull. Am. Paleont., Ithaca, 27 : 363-368 (95-
too), pl. 36 (10).
WeEYER, D. 1965. Uber Amplexus zaphrentiformis White 1876 (Pterocorallia; Oberkarbon,
Pennsylvanian). Geologie, Berlin, 14 : 449-463, 2 pls.
Witson, E. C. & LaNGENHEIM, R. L., Jr. 1962. MRugose and tabulate corals from Permian
rocks in the Ely Quadrangle, White Pine County, Nevada. /. Paleont., Tulsa, 36, 3:
495-520, pl. 86-89.
280 PALAEOZOIC CORAL FAUNAS
VII. APPENDIX—COMPOSITION AND LOCATION OF SAMPLES
The most accurate system for the location of samples available is by means of the
Maracaibo Cathedral co-ordinates employed by Compania Shell de Venezuela on
their I : 10,000 topographic maps. The co-ordinates are in km. All the localities
are in the northern part of the Sierra de Perija, Zulia State, Venezuela and the
locations of 7m situ samples are plotted in Text-figs I and 2.
Cano Los Guineos Formation
Caiio Los Guineos section:
3849 Heliophyllum hall Siik-32 W 97°58
Heterophrentis (H.) simplex
Stereolasma sp.
Syringaxon sp.
Cano Grande Formation
Canio Grande section:
3121 Bowenelasma typa N Io-11 W 89:36
Favosites venezuelensis
Heliophyllum hal
3124 Heliophyllum hall N 10-04 W 89:34
3126 Heliophyllum wellsi N 10-02 W 89:32
3129 Briantelasma oliveri N 9:99 W 89:28
3145 Heliophyllum hall N 9-81 W 89:11
3149 Bowenelasma typa N 9:70 W 89:10
Heliophyllum halli
3154 ? Bowenelasma breviseptata N
Heliophyllum hal
Plasmophyllum sp.
3157 Bowenelasma typa N 9:81 W 89:04
? Briantelasma oliveri
Heliophyllum hall
Plasmophyllum secundum americanum
9°79 W 86-06
Cano del Sur section:
3182 Favosites arbuscula N 9:56 W 88-78
F. venezuelensis
3199 Bowenelasma breviseptata N 9:29 W 89:25
Heliophyllum halla
3221 Briantelasma sp. N 9:05 W 89-79
Heliophyllum halla
Plasmophyllum secundum americanum
FROM THE SIERRA DE PERIJA, VENEZUELA
Cafio del Oeste section (boulder) :
3323 Bowenelasma typa
Brniantelasma sp.
Heliophylium hall
Cafio Colorado section:
3618 Bowenelasma typa
Briantelasma oliveri
Heliophyllum halli
3619 Bowenelasma breviseptata
Heliophyllum hall
Cano Grande Fm. or lower Cano de Oeste Fm.
Cafio Pescado section (slipped block):
3539 Cylindrophyllum elongatum
Cano del Oeste Formation
Cafio Grande section:
3243 Hadrophyllum sp.
? Stewartophyllum sp.
Rio Palmar Formation
Cano del Norte section:
3495 ? Durhamina sp.
Cano Colorado section:
3784 Amplexizaphrentis sutherlandi
? Rio Palmar Formation
Cafio Grande section (boulder) :
3270 Durhamina sp. nov.
DR CoLiIn T. SCRUTTON,
Department of Geology
THE UNIVERSITY
NEWCASTLE UPON TYNE.
NE 7RU.
E
N 12°83
N 6-01
N 6-01
N 6:32
N 9-69
N 16:97
Ne 3-5r
N 10:51
W 87-79
W 91:98
W o1-98
W 89:83
W 88-11
W 85:97
W 90°47
W 86-68
281
PLATE1
Syringaxon sp.
Fics 1, 2. Cross-sections (slides). R45130b, a. x6.
Fic. 3. Longitudinal section (slide). R45130k. x6.
Catio Los Guineos; Cafio Los Guineos Fm., early Middle Devonian.
?Stewartophyllum sp.
Fics 4, 5. Cross-sections (slides). R45110b, a. X2°5.
Cafio Grande; Cafio del Oeste Fm., early Middle Devonian.
Stereolasma sp.
Fic. 6. Cross-section (slide). R45129a. x3.
Fic. 7. Longitudinal section in counter-cardinal plane; C on left (slide). R45120g. x 3.
Cano Los Guineos; Cano Los Guineos Fm., early Middle Devonian.
Hadrophyllum sp.
Fic. 8. Cross-section (polished surface). R49275. 2:5.
Cano Grande; Cafio del Oeste Fm., early Middle Devonian.
Amplexizaphrentis sutherlandi sp. nov.
Fic. 9. Cross-section (peel), early ephebic stage. R45127b (taken from holotype). x3.
Fic. 10. Cross-section (slide), early neanic stage 16 mm below R45127b. R45127a (cut
from holotype). x3.
Fic. 11. Cross-section (peel), neanic stage 14 mm below R45127b. R45127d (taken from
holotype). 3.
Fic. 12. Longitudinal section in counter-cardinal plane; C on right (slide). R45127e (cut
from holotype). x3.
Fic. 13. Longitudinal section in counter-cardinal plane; C on left (slide). R4g291b. x2.
Both Cano Colorado; Rio Palmar Fm., Lr. Pennslyvanian.
All cross-sections orientated with the cardinal septum at top centre.
E*
Bull. Br. Mus. nat. Hist. (Geol.) 23, 4
IIE ANAS, Ft
PAE 2
Bowenelasma typa gen. et sp. nov.
Fic. 1. Cross-section (slide), ephebic stage. R45094b (cut from holotype). 2.
Fic. 2. Cross-section (peel), neanic stage. R45094d (taken from holotype). 2.
Fic. 3. Longitudinal section in counter-cardinal plane; C on left (slide). R45094e (cut
from holotype). x2.
Fic. 4. Cross-section (slide), early neanic stage. R45100b. 4.
Fic. 5. Cross-section (slide), ephebic stage. R45100a. 2.
Fic. 6. Longitudinal section in counter-cardinal plane; C on left (slide). R45100c. x2.
Fic. 7. Cross-section (slide), ephebic stage, to illustrate septal structure. R45094b (cut
from holotype). x6.
Both Cano Grande; Cano Grande Fm., early Middle Devonian.
Fics 8, 9. Cross-sections (peels), neanic stage. R45121f,e. x2.
Fic. 10. Cross-section (slide), sub-calicular level. R4512ta. x2.
Fic. 11. Longitudinal section in counter-cardinal plane; C on left (slide). R4512th. x2.
Cano Colorado; Cano Grande Fm., early Middle Devonian.
All cross-sections orientated with the cardinal septum at top centre.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 4 PLATE 2
E**
PLATE 3
Bowenelasma breviseptata sp. nov.
Fics 1, 2. Cross-section (slide), ephebic stage. R45123a (cut from holotype). Fig. 1
Kee ie, 2, 3<-
Fic. 3. Cross-section (slide), neanic stage. R45123b (cut from holotype). x2.
Fic. 4. Longitudinal section in counter-cardinal plane; C on left (slide). R45123d (cut from
holotype). x2.
Cano Colorado; Cano Grande Fm., early Middle Devonian.
Fic. 5. Cross-section (peel), neanic stage. R45105¢c. x2.
Fic. 6. Cross-section (slide), ephebic stage. R45105b. 2.
Fic. 7. Longitudinal section (slide). R45105d. 2.
Cafio del Sur; Caio Grande Fm., early Middle Devonian.
All cross-sections orientated with the cardinal septum at top centre.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 4 PLATE 3
PLATE 4
Briantelasma oliveri sp. nov.
Fics 1, 2. Cross-section (slide), sub-calicular. R45090b (cut from holotype). Fig. 1, «2;
figs 25 XxX Ae
Fic. 3. Cross-section (slide), ephebic stage. R45090a (cut from holotype). x2.
Fic. 4. Longitudinal section (slide) in counter-cardinal plane; C on right. R45o0g0e (cut
from holotype). x2.
Fic. 5. Cross-section (slide), neanic stage. R45091a. x3.
Fic. 6. Cross-section (slide), ephebic stage. R45091b. x2.
Both Cano Grande; Cano Grande Fm., early Middle Devonian.
Fic. 7. Cross-section (slide), sub-calicular. R49290a. 2.
Cano Colorado; Cano Grande Fm., early Middle Devonian.
All cross-sections orientated with the cardinal septum at top centre.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 4 PLATE 4
PLATE 5
Heterophrentis (H.) venezuelensis (Weisbord)
Fic. 1. Cross-section (peel), ephebic stage. PRI 21594/PRI 24421 (taken from lectotype).
XK 2.
Fic. 2. Longitudinal section (peel). PRI 24421 (taken from lectotype). 1-5.
?Loose boulder in Cano del Norte; Cano Grande Fm., early Middle Devonian.
Heterophrentis (H.) simplex (Hall)
Fic. 3. Cross-section (polished surface), ephebic stage. NYSM 360 (holotype). 1-5.
Moscow, New York; Middle Devonian (Hamilton Group).
Fic. 4. Cross-section (polished surface), neanic stage. PRI 21593. 1-5.
Fic. 5. Cross-section (polished surface), ephebic stage. PRI 21593. X1I°5.
?Loose boulder in Cano del Norte; Cano Grande Fm., early Middle Devonian.
Heliophyllum halli Edwards & Haime
Fic. 6. Cross-section (slide), ephebic stage. R45115a. 2.
Loose boulder in Cano del Oeste; Cano Grande Fm., early Middle Devonian.
Fic. 7. Cross-section (slide), ephebic stage. R49289a. x2.
Fic. 8. Longitudinal section in counter-cardinal plane; C on right (slide). R49289b. x2.
Cano Colorado; Cano Grande Fm., early Middle Devonian.
All cross-sections orientated with the cardinal septum at top centre.
PLATE 5
nat. Hist. (Geol.) 23, 4
Mus.
Bull. Br.
PLATE 6
Heliophyllum halli Edwards & Haime
Fic. 1. Cross-section (slide), ephebic stage. R45104a. 2.
Fic. 2. Cross-section (slide), neanic stage. R4510q4c. 2.
Cano del Sur; Cano Grande Fm., early Middle Devonian.
Fic. 3. Cross-section (slide), ephebic stage. R45092b. 2.
Fic. 4. Longitudinal section in counter-cardinal plane; C on left (slide). R45092Cc.
Cano Grande; Cano Grande Fm., early Middle Devonian.
Fic. 5. Cross-section (slide), ephebic stage. R49278a. 2.
Fic. 6. Longitudinal section in counter-cardinal plane; C on left (slide). R49278d.
Loose boulder in Cano del Oeste; Cano Grande Fm., early Middle Devonian.
All sections orientated with the cardinal septum at top centre.
Heliophyllum wellsi sp. nov.
Fic. 7. Cross-section (slide), ephebic stage. R45088b (cut from holotype). I.
Fic. 8. Longitudinal section (slide). R45088e (cut from holotype). 1-5.
Cano Grande; Cano Grande Fm., early Middle Devonian.
Mus. nat. Hist. (Geol.) 23, 4
Bull. Br.
PLATE 7
Heliophyllum wellsi sp. nov.
Fics 1, 2. Cross-sections (slides), illustrating character of axial structure in ephebic stage.
R45088a, b (both cut from holotype). x2.
Cano Grande; Cano Grande Fm., early Middle Devonian.
Cylindrophyllum elongatum Simpson
Fic. 3. Cross-section (slide). R45116a. 2°5.
Fics 4-6. Longitudinal sections (slides). R45116h, j, g. 2:5.
Loose block in Cano Pescado; either Cano Grande Fm. or low in Cano del Oeste Fm., early
Middle Devonian.
?Durhamina sp.
Fic. 7. Cross-section (slide). R49285a. x4.
Fic. 8. Longitudinal section (slide). R49285b. x 4.
Cano del Norte; Rio Palmar Fm., Lr. Pennsylvanian.
Bull. By. Mus. nat. Hist. (Geol.) 23, 4 PLATE
PLATE 8
Durhamina sp. noy.
Fic. 1. Cross-section (slide) of corallite arrangement in colony. R49276b. xt.
Fic. 2. Cross-section (slide) of corallite with third order septa. R49276b. x6.
Fics 3-4. Cross-sections (slides) showing variation in appearance of mature corallites.
R49276c, a. 2.
Fics 5-7. Cross-sections (slide) of immature corallites. R49276a. 4.
Fics 8-10. Longitudinal sections (slides), fig. 8 showing increase. R49276h, i,j. 2.
Loose block in Cano Grande; Carboniferous, (?Rio Palmar Fm., Lr. Pennsylvanian).
8
Dy
PEACE
Bull. Br. Mus. nat. Hist. (Geol.) 23, 4
PLATE 9
Plasmophyllum secundum americanum (Edwards & Haime)
Fic. 1, 2. Cross-sections (slides). R45098b, c. X1°5.
Fic. 3. Cross-section (slide) showing form and arrangement of septal spines. R45098b.
Fic. 4. Longitudinal section (slide). R45098d. x 1°5.
Cano Grande; Cano Grande Fm., early Middle Devonian.
Plasmophyllum sp.
Fics 5-6. Cross-sections (slides). R45095a, b. 3.
Fic. 7. Longitudinal section (slide). R45095c. 3.
Cano Grande; Cano Grande Fm., early Middle Devonian.
x6.
PLATE 9
nat. Hist. (Geol.) 23, 4
Mus.
Bull. Br.
PLATE to
Favosites venezuelensis (Weisbord)
Fic. r. Cross and longitudinal sections (slide). R45101b. 3.
Fic. 2. Tangential section (slide). Rq45rora. 3.
Cano del Sur; Catto Grande Fm., early Middle Devonian.
Favosites arbuscula Hall
Fic. 3. Tangential section (slide). R45103a. 3.
Fics 4-5. Cross-sections (slides). R49264a, R45103b. x3.
Fic. 6. Longitudinal section (slide). R49264b. x3.
Cano del Sur; Cano Grande Fm., early Middle Devonian.
Bull. By. Mus. nat. Hist. (Geol.) 23, 4 PLATE ro
>"
——_
>
A LIST OF SUPPLEMENTS
TO THE GEOLOGICAL SERIES
OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya.
Pp. 213; 30 Plates; 2 Text-figures. 1965. {£6.
. EL-NaGcear, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper
Cretaceous—Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley,
Egypt, U.A.R. Pp. 291; 23 Plates; 18 Text-figures. 1966. 10.
. Davey, R. J., DowntE, C., SARGEANT, W. A. S. & Wititams, G. L. Studies on
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248; 28 Plates; 64 Text-
figures. 1966. £7.
. APPENDIX. DAvEy, R. J., DowniE, C., SARGEANT, W. A. S. & WILLIAMS, G. L.
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24.
1969. 8op.
. Ettiott, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of the
Middle East. Pp. 111; 24 Plates; 17 Text-figures. 1968. £5.12}.
. Ruopes, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni-
ferous) Conodont faunas, and their value in local and continental correlation.
Pp. 315; 31 Plates; 92 Text-figures. 1969. {II.
. Cuitps, A. Upper Jurassic Rhynchonellid Brachiopods from Northwestern
Europe. Pp. 119; 12 Plates; 40 Text-figures. 1969. £4.75.
. Goopy, P. C. The relationships of certain Upper Cretaceous Teleosts with
special reference to the Myctophorids. Pp. 255; 102 Text-figures. 1969. £6.50.
. OwEN, H. G. Middle Albian Stratigraphy in the Paris Basin. Pp. 164;
3 Plates; 52 Text-figures. 1971. 6.
. Sippigul, Q. A. Early Tertiary Ostracoda of the family Trachyleberididae
from West Pakistan. Pp. 98; 42 Plates; 7 Text-figures. 1971. {8.
ae.
BCR Printed in England by Staples Printers Limited at their Kettering, Northants establishment
as
= ae See é . rf
wer eS ae ee * Se
ate
A are
Zo aw
Ae a
PRODEINOTHERIUM FROM GEBEL ZELTEN
LIBYA |
AEMWUSE I
few ee
= sae Of
ft) aN 4
i = 4X >!
4 S
\
\
BY
JOHN MICHAEL HARRIS ||
Centre for Prehistory and Palaeontology, Nairobi —
Pp. 283-348; 5 Plates, 16 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 23 No. 5
LONDON : 1973
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 10949, 1s
issued in five series, corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
veady. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 23, No. 5 of the Geological
(Palaeontological) series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation
Bull. Br. Mus. nat. Hist. (Geol.).
© Trustees of the British Museum (Natural History), 1973
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 12 July, 1973 Price £2.85
PRODEINOTHERIUM FROM GEBEL ZELTEN,
LIBYA
By J. M. HARRIS
CONTENTS
Page
I. ABBREVIATIONS AND ACKNOWLEDGEMENTS . . 9 . : 286
II. INTRODUCTION. : : : 5 : : e : c 286
III. SYSTEMATIC DESCRIPTIONS. : 3 : : 5 6 291
Order PROBOSCIDEA . : : : : : : 291
Suborder DEINOTHERIOIDEA : : é : C 291
Family DEINOTHERIIDAE . : : : : : 291
Genus Deinotherium . : : : F : ; 292
Genus Prodeinotherium . 6 : : : : 5 293
Prodeinotherium Roo : : a . : : 296
Skull. ; : : : : ; 296
OTHER DEINOTHERIID SKULLS . : 3 ; 5 305
Mandible. : : 311
COMPARISON WITH OTHER DEINOTHERE MANDIBLES. 312
Dentition 2 : : : 3 : 313
TOOTH FUNCTION IN DEINOTHERES . : : é 318
Axial skeleton : : : : : d c 319
Appendicular skeleton . ; 330
SKELETAL EVIDENCE ON THE LOCOMOTION OF DEINO-
THERES . c ¢ : 5 : ¢ é 341
IV. CoNCLUSIONS . 3 : : 5 : : : : : 344
V. REFERENCES . : : : . ; : , : . 346
SYNOPSIS
A large assemblage of vertebrate fossils has been collected from Miocene continental and
near-shore marine lithosomes at the Gebel Zelten, Libya. Deinotheres were not prolific but
are of interest in that they include the most complete skull yet discovered, plus the associated
skull, mandible and partial skeleton of an immature individual. These are described and
compared with other deinothere remains and with the postcranial skeleton of fossil and recent
elephantoid proboscideans.
Features of the skull, skeleton and dentition indicate that the family Deinotheriidae possessed
two genera and Ehik’s taxon Prodeinotherium is resurrected for the earlier and more primitive
genus to which the Gebel Zelten specimens belong. The facial region of the Libyan Prodeino-
therium skull has a downcurved rostrum that is modified to bear a proboscis. The occipital
region, basicranium and anterior cervical vertebrae are adapted to impart a powerful downward
thrust to the tusks. The cheek teeth are separated into an anterior crushing battery and a
posterior shearing battery.
The postcranial skeleton of Prodeinotherium and Deinotherium exhibit graviportal adaptations
for support of the body weight. The atlas and axis vertebrae of deinotheres are distinctive and
other differences in the postcranial skeleton from that of elephantoids may be detected. The
scapula and foot bones of Prodeinotherium may be readily recognized but the long bones resemble
those of Elephas. More advanced characters shown by Deinotherium include reduction of the
scapular spine, shortening of the femur, elongation of the manus and pes, and reduction of the
first digit thus leading to functional tetradactyly. These may all be interpreted as cursorial
adaptations.
286 PRODEINOTHERIUM
I ABBREVIATIONS AND ACKNOWLEDGEMENTS
Individual specimens from Gebel Zelten are referred to by their University of
Bristol accession numbers, e.g. 6404: 44, 6418: 20 (M.26665). Specimens of
Deinotherium giganteum from Howenegg, Germany, and currently at the Johannes
Wurtemberg Universitaat, Mainz, had not been given permanent accession numbers
at the time of writing and are referred to in tables of measurements as ‘Howenegg’.
Recent specimens of Loxodonta africana used for comparison in the section on the
axial skeleton include OM 2209 from the National Museum of Kenya and 1961.8.9.82
from the British Museum (Natural History). Comparison between the appendicular
skeleton of Prodeinotherium and Elephas maximus was based on the skeleton of an
Indian elephant in the University of Bristol Department of Geology (catalogue
number U.B.20189). Deinothere material with KNM catalogue numbers is from
the Centre for Prehistory and Palaeontology, Nairobi. The catalogue numbers of
mammalian specimens from the British Museum (Natural History) are prefaced by
B.M.(N.H.)M. or (in tables of measurements) M. In the tables, measurements
are given in cms. unless otherwise stated; * = approximate measurement,
-+- = incomplete specimen.
This paper formed part of a Ph.D. thesis from the University of Bristol that was
supervised by Dr R. J. G. Savage for whose help and encouragement I am very
grateful. The work was financed by a grant from N.E.R.C. Iam grateful also to
Mrs D. M. Leakey who read and criticised the manuscript.
Prodeinotherium skull M.26665 was prepared by Mr A. Rixon in the Department
of Palaeontology, British Museum (Natural History). Deinothere material from the
1964 and 1966 University of Bristol expeditions was prepared by Mr. M. White of
the Department of Geology, University of Bristol. The photographs were prepared
by Mr R. Godwin and line drawings of the postcranial material by Mrs M. Clapp.
Field work in Libya was made possible by generous co-operation from Esso
Standard Libya and Oasis Oil Company of Libya Inc.
I am indebted to the staff of the various institutions that have permitted me to
examine material in their collections, and especially the late Professor C. Arambourg
(Institut de Paleontologie, Paris), Professor H. Tobein (Johannes Wurtemberg
Universitaat, Mainz), Professor R. Dehm (Institut ftir Palaontologie, Munich),
Dr E. Aguirre (Museo de Sciencias Naturales, Madrid), Dr L. S. B. Leakey (Centre
for Prehistory and Palaeontology, Nairobi), and Dr A. Sutcliffe (Department of
Palaeontology, British Museum (Natural History)).
Il: INTRODUCTION
THE Gebel Zelten is an elongate mesa lying approximately 200 kms inland from the
Gulf of Sirte and is bounded by the co-ordinates 19}$°-203°E and 28-29°N. The
gebel extends for some 140 kms in a northwest-southwest direction. It attains a
height of some 40-60 metres at its western extremity but gradually decreases in
elevation towards the east until it merges with the Calenscio Serir. At the eastern
extremity of the gebel the northern and southern scarps are separated by a distance
of some 50 kms. The scarps converge towards the west and near the centre of the
gebel they are only 8 kms apart. West of this point the southern scarp diverges
FROM GEBEL ZELTEN, LIBYA 287
sharply from the northern before merging at the extreme western extremity (see
text-fig. 1). The Gebel Zelten is deeply dissected by wadis that often reach 3 kms
in length. Most of the vertebrate sites are located in branches of the wadi systems.
The rocks exposed at the Gebel Zelten are of Miocene age and were attributed by
Selley (1966) to the Marada Formation. The rocks represent the mixed fluvio-
marine facies of the southern part of the Sirte Basin. The succession at Gebel
Zelten is predominantly of sandstone, sandy shales and shales, the units dipping
northward at an angle of about six seconds (Doust, 1968). According to Doust, the
base of the section at the northern scarp consists of intertidal and fluviatile sands and
shales that pass upward into bioturbated calcarenites and calcsilicates representing
intertidal and subtidal facies. The top of the succession comprises calcareous
crossbedded sandstones from an estuarine channel facies. On the southern scarp a
mixed intertidal and fluviatile facies passes upwards into intertidal sediments that
are ultimately overlain by the estuarine channel facies. Detailed discussions of the
geology are given in Selley (1966, 1967, 1968), Doust (1968), and Magnier (1962, 1968).
whe
NB Y
C\ Wadi el'ldem
Transportation
Corner
Fic. 1. Vertebrate localities of the Gebel Zelten.
Sites D, E, F and G are located between sites C and H.
288 PRODEINOTHERIUM
The vertebrate fauna of the Gebel Zelten was discovered in 1958 by personnel of
the Compagnie des Petroles Total (Libye) during a regional reconnaissance of Libya
(Magnier, 1962). In December 1960 an expedition from the Museum National
d’Histoire Naturelle, Paris, visited Gebel Zelten to exploit the vertebrate remains
(Arambourg & Magnier, 1961). Only an area 20 km long at the centre of the northern
scarp of the Gebel was prospected in detail but vertebrate specimens were collected
from all exposures examined. The material retrieved is currently housed in the
Institut de Paléontologie, Paris. A faunal list was given by Arambourg and
Magnier (1961), and later revised by Arambourg (1963b). Of the elements listed
only Bunolistriodon massai has been described in detail (Arambourg, 1963a) but
preliminary descriptions of Afrocyon burollett, Prolibyther.um magniert, Mastodon
pygmaeus and an aeopyornithid tibia have been published (Arambourg Ig61a,
1g61b).
From 1964, expeditions from the University of Bristol under the leadership of
Dr R. J. G. Savage have systematically collected vertebrate material from Gebel
Zelten. Exploitation of the southern scarp was undertaken in 1964 and of the
northern scarp in 1966. The 1967 and 1968 expeditions expanded and consolidated
the results of earlier investigations. In all, well over a thousand specimens,
representing several tons of unprepared material, have been collected by University
of Bristol parties. This is in addition to the material collected by the 1960 French
expedition and to the specimens donated by oil company personnel to the British
Museum (Natural History) and American Museum of Natural History. A list of
the faunal elements collected was given in Savage and White (1965) and revised in
Savage (1967) and Selley (1968).
Most of the vertebrate fossils have been collected from channels and flood plains
of the fluviatile facies, though remains are not infrequent in the tidal flat facies.
At the time of writing 24 sites had been discovered and these were assigned letter
prefixes in alphabetical order of discovery (see fig. I). Permanent site numbers and
other pertinent data are now on file in the Department of Geology, University of
Bristol. All the Gebel Zelten sites occur near the base of the exposed section and
range in areal extent from one to ten acres. A table showing the distribution of the
Gebel Zelten mammal fauna by sites was given by Savage (in Selley, 1968).
Desio (1935) referred the rocks on the southside of the Gebel Zelten to the Aquitanian
stage and those on the north scarp to the Burdigalian. The vertebrate fauna was
allocated a Burdigalian age by Arambourg (1963b) although it is not clear whether
this age was based on the mammalian elements or on the marine invertebrates.
Savage and White (1965) confirmed a Burdigalian age for the mammals, and Savage
(in Selley, 1968) later refined the age of the fauna to early Burdigalian or even late
Aquitanian. Definite conclusions on the age of the Gebel Zelten assemblage must
necessarily be delayed until all the mammalian groups represented have been
thoroughly investigated. To date only the carnivores (Savage, 1973) and giraffoids
(Hamilton, 1973) have been completed.
The Gebel Zelten deinothere skulls are not only the earliest skulls, though not the
earliest deinothere specimens (which are from East Africa), but are also unusually
complete. The best preserved specimen (6418 : 20) was collected in 1966 and was
FROM GEBEL ZELTEN, LIBYA 289
deposited in the British Museum (Natural History). The accession number of the
specimen is M.26665. All the permanent teeth are erupted and show wear but the
zygomatic, squamosal, auditory and pterygoid regions are poorly preserved.
A virtually complete skull of an immature individual was collected by the writer
from Site D in 1967. This specimen (6404 : 44) was completely buried and was
only discovered during the excavation of deinothere limb material, for which purpose
site D had been revisited. The skull was lying on its side (the only deinothere skull
from Gebel Zelten that was not lying upside down) and had been slightly deformed
by compaction after burial. The bone was in an extremely fragile state and the
right zygomatic arch disintegrated during excavation. The occipital region frag-
mented and became partially deformed during transportation to Bristol. Permanent
P3_M1 are fully erupted but only M! shows wear. M2 is only partly erupted.
The squamosal, auditory and orbital regions are better preserved than in M.26665.
Several of the facial sutures are still unfused. The mandible (6404 : 45) and much
of the skeleton belonging to this individual (M.26667a-1) were excavated from the
adjacent part of site D.
An incomplete deinothere skull (6404 : 14) and much of a mandible (6404 : 13)
were collected in 1964 from within a few yards of skull 6404 : 44. Unfortunately
skull 6404 : 14 was broken in transit from the Gebel Zelten and only the premaxilla,
jugal and dentition are now in a reasonable state of repair. The teeth of the 1964
specimens are in different stages of eruption and wear and are not thought to belong
to one individual.
The 1968 Bristol expedition to Gebel Zelten and Dor el Talha discovered an in-
complete and partly crushed skull at Gebel Zelten site L. Pieces of this skull were
collected but all except a fragment of the inferior half of the occiput are still in
Libya. The ventral edges of the lateral wings of the squamosal are better preserved
on this specimen than on M.26665. Two other partly eroded skulls were discovered
at site R by the 1966 expedition. These were allocated field numbers (6418 : 4,
6418 : 19) but have yet to be collected.
In addition to the deinothere postcranial material from site D, a cuneiform and
isolated vertebrae have been collected from other Gebel Zelten sites. Gebel Zelten
deinothere material is appreciably smaller than, and differs morphologically from,
equivalent specimens of Mastodon angustidens from Gebel Zelten. In the ensuing
descriptions, comparisons with recent elephant remains are based on vertebrae of
Loxodonta africana from the British Museum (Natural History), the manus and pes
of L. africana from the Osteology Department of the Centre for Prehistory and
Palaeontology, Nairobi, and fore and hind limb material of Elephas maximus in the
University of Bristol collections. Comparative deinothere material includes the
partial skeleton of ‘Deinotherium bavaricum’ in the Naturhistorisches Museum of
Vienna, the partial skeleton of D. giganteum from Valladolid in the Museo de
Sciencias Naturales of Madrid, and carpals and tarsals of D. gigantewm from Howenegg
in the University of Mainz collections.
Conclusions on deinothere taxonomy, morphology and functional anatomy were
made after viewing specimens in the following institutions: British Museum (Natural
History), London; Centre for Prehistory and Palaeontology, Nairobi; Department
290 PRODEINOTHERIUM
of Geology Museum, University of Bologna; Museo de Sciencias Naturales, Madrid;
Institut de Paleontologie, Paris; Naturhistorisches Museum, Vienna; Institut fir
Palaontologie und Historisches Geologie, Munich; Hessisches Landesmuseum,
Darmstadt; Naturhistorisches Museum, Mainz; Institut fiir Palaontologie, Johannes
Wurtemberg Universitat, Mainz; Naturhistorisches Museum, Wiesbaden. Exten-
sive use has been made also of data given by Bergounioux & Crouzel (1962a,
1962b), Dehm (1963), de Pauw (1908), Graf (1957), McInnes (1942), Pacheco (1930),
Palmer (1924), Sahni & Tripathi (1957), Stefanescu (1897, 1907), and Tobein
(1962).
The results of these investigations suggest that some revision of deinothere
systematics is necessary. Diagnoses are given at the beginning of the section on
systematic description because the Gebel Zelten deinotheres do not appear to belong
to the type genus.
TABLE I
Deinothere specimens from Gebel Zelten
Field Accession Field Accession
number number Description number number Description
6401: I Left M8 6404: 47 M.266671 Distal end of left
6401: 4 Right maxilla with humerus
P3_M3 6405: 98 Left cuneiform
6404: 13 Mandible with tusks 6409: 41 M? fragment
and cheek teeth 6410 2 Deciduous Pa
6404: 14 Skull fragments with 6412: I0 Tuskless mandible
dentition with worn cheek
6404 : 30 M.26667a Distal end of fibula teeth
6404: 31 M.26667b Right ulna 6412: 39 Left Me
6404 : 32 M.26667c Proximal end of right as eo ae =
cf eae 6413: 18 Left M8
6404: 33 M.26667d Right femur 6416 : 120 Tusk tip
6404 : 34 M.26667e Left ulna 6416 : 130 Race
6404: 36 M.26667{ Right scapula 6418: 20 Adult skull
6404: 37 M.26667g Left innominate 6418: 22 Mandible with tusks
6404 : 38 M.26667h Left rib head and cheek teeth
6404 : 39 M.26667i Right neuropophysis 6419: 15 Atlas vertebra
of 3rdcervicalvertebra 6419: 16 Tusk alveoli
6404: 41 M.26667j Proximal and distal 6419: 17 Enamel fragment
end of left radius 6421 a 2 Left M2
6404: 44 Immature skull 6421: 15 Molar fragment
6404: 45 Fragmented 6423: 13 Enamel fragment
immature mandible 6423: 45 Enamel fragment
6404 : 46 M.26667k Distal end of right 6423: 56 Enamel fragment
humerus **6424 : 79 Axis vertebra
** — Site unknown
FROM GEBEL ZELTEN, LIBYA 291
III. SYSTEMATIC DESCRIPTIONS
Order PROBOSCIDEA Illiger, 1811
Suborder DEINOTHERIOIDEA, Osborn, 1921
Family DEINOTHERIIDAE Bonaparte, 1845
DiaGnosis. Large herbivorous graviportal mammals. Dental formula $03
for the deciduous teeth and 2°:2:* for the permanent dentition; DP} and P3 with
well developed external crest; DP¢ and Mi} trilophodont ; the remainder of the cheek
teeth are bilophodont. Horizontal tooth replacement not developed so that all
permanent teeth may be erupted at the same time (cf. elephantoids). Mandibular
symphysis and lower tusks curved downwards so that the tusk tips are vertically or
nearly vertically aligned. Skull low with deep rostral trough, retracted external
nares, low orbit, inclined occiput, high occipital condyles, elongate paroccipital
processes, and diploe.
MIOCENE PLIOCENE PLEISTOCENE rd
P. hobleyi >
mn
rad
.@)
>
P. bavaricum Cc
Pe)
O
me)
m™m
D.
P. pentapotamige >
(op)
>
D. indicum
Fic. 2. Temporal and geographic distribution of deinotheriid taxa.
292 PRODEINOTHERIUM
REMARKS. The family Deinotheriidae originated in Africa. The earliest and
most primitive forms are from East African localities of early Miocene age. During
the Burdigalian the early representatives of this family migrated into Asia and
Europe. By the middle of the Miocene epoch, larger and more advanced repre-
sentatives had appeared in Europe and Asia. The primitive and advanced forms
coexisted until the end of the Miocene in Asia and the end of the Pontian in Europe.
The more advanced forms lingered on until the middle Pliocene in Eurasia. Only
the early and primitive form is known from the Neogene of Africa; a larger and more
advanced form occurred in the Pleistocene but the two forms are not known to have
coexisted.
Deinotheriid species have been commonly allocated to a single genus—Deino-
therrum—erected by Kaup in 1829. The species were defined upon geographic
distribution and characters of the mandible and dentition. While the cheek teeth
exhibit a great deal of variation in cusp morphology from specimen to specimen,
these differences do not teadily lend themselves to species differentiation, especially
when the several species are interpreted as belonging to a single genus. Moreover
the concept of a single deinothere genus does not adequately express the fundamental
division of the family into an early and primitive group of species (D. hobleyi,
D. bavaricum and D. pentapotamiae) and a later, more advanced group (D. giganteum,
D. indicum and D. bozasi). The recent discoveries of deinotheriid skull and post-
cranial material, coupled with the re-examination of well documented specimens
confirms that the two groups of deinotheres may be separated morphologically. It
would appear therefore that division of the family into two genera is warranted.
This idea was originally put forward by Ehik (1930) but has subsequently been
ignored. On evidence currently available, however, Ehik’s premise is substantiated.
The two genera are defined below.
Genus Deinotherium Kaup, 1829
REVISED DIAGNOsIS. Large deinotheres. Dental formulae as for the family;
tendency for the development of subsidiary styles on P3~4 and for simplification of the
postmetaloph ornamentation of M2-3 when compared to Prodeinotherium. The
skull rostrum not parallel to the mandibular symphysis and nearly horizontally
aligned; rostral trough and external nares wide; preorbital swelling sited anteriorly
on the rostrum; skull roof short and narrow at the temporal fossae; occiput slopes
gently posteriorly; occipital condyles elevated above the level of the external
auditory meatus; paroccipital processes very elongate. Postcranial skeleton with
cursorial modifications to graviportal structure; scapular spine reduced with no
acromion or metacromion; carpals and tarsals narrow with dolichopodous meta-
podials exhibiting functional tetradactyly.
FROM GEBEL ZELTEN, LIBYA 293
REMARKS. Most of the diagnostic characters of Deznotherium concern the skull
and postcranial characters. The teeth, however, are the most commonly preserved
remains. Teeth that may be attributed to Deinotherium are almost always larger
than the equivalent teeth of Prodeinother1wm. Both genera exhibit a tendency to
increase in size through geologic time, and in Europe there is an overlap in absolute
size of the cheek teeth of the two genera. This is due to the greater numbers and
greater longevity of deinothere specimens from that continent, but at any one horizon
the teeth of Deinotheriuwm are always unmistakably larger than those of Prodeino-
therium. On the basis of specimens examined and of those reported by Graf (1957),
McInnes (1942) and Sahni & Tripathi (1957), the minimum parameters of Deino-
therium teeth would appear to be as follows:
Tooth Length (in mm) Breadth (in mm)
Se 63 55
p4 59 62
M1 71 64
M? 70 74
M? 70 74
Ps 51 43
P, 61 48
Mi 74 54
Me 73 63
Ms 75 65
These parameters agree well with those noted by Bergounioux & Crouzel (1962a)
who also define parameters for the deciduous teeth.
TYPE SPECIES. Deinotherium giganteum Kaup, 1829 (including D. gigantissimum
Stefanescu, 1897).
OTHER RECOGNISED SPECIES: D. indicum Falconer, 1845; D. bozast Arambourg,
1934.
A detailed synonymy of D. gigantewm was given by Graf (1957) and of D. indicum
by Sahni & Tripathi (1957).
Genus Prodeinotherium Ehik, 1930
REVISED DIAGNOSIS. Small deinotheres. Dental formulae as for the family;
M2-3 with well defined postmetaloph ornamentation. Skull rostrum turned down
parallel to the mandibular symphysis; rostral trough and external nares narrow;
preorbital swelling close to orbit ; external nares anteriorly sited and nasal bones with
anterior median projection; skull roof relatively longer and wider than in
Deinotherium; occiput more vertically inclined; occipital condyles sited more
ventrally than in Deinotherium and level with the Frankfurt Plane; paroccipital
294 PRODEINOTHERIUM
processes short. Postcranial skeleton graviportally adapted; scapula with well
defined spine and stout acromion and metacromion; tarsals and carpals narrow but
not dolichopodous.
ReMARKS. This genus was originally proposed by Ehik (1930) for a small species
of deinothere from Kotyhaza, Hungary. The Kotyhaza specimens comprised a
jaw, some teeth, anda few incomplete limbelements. These specimens were generally
smaller than is normal in D. bavaricum but were subsequently attributed to this
taxon without objection (Graf, 1957; Bergounioux & Crouzel, 1962a; etc.). I agree
that the taxa bavaricum and hungaricum are probably conspecific but propose to
allocate them to Prodeinotherium. Ehik’s generic taxon is therefore resurrected to
encompass the small, early and primitive species of deinothere from the Neogene of
Africa and Eurasia. The slight change in spelling from Prodinotherium (Ehik,
1930) to Prodeinotherium is, in view of Kaup’s spelling of the type genus of the family
Deinotheriidae, considered to be a justifiee emendation according to the International
Code of Zoological Nomenclature (Stoll et al, 1961) section 9g, article 50, para
graph (c).
Specimens that may be attributed to species of Prodeinotherium are observed to
increase in size through geologic time. The maximum parameters for the permanent
cheek teeth of Prodeinotherium, based on personal observation and data from Graf
(1957), McInnes (1942), and Sahni & Tripathi (1957) are as follows:
Tooth Length (in mm) Breadth (in mm)
Ps 63 60
p4 61 62
Mt 73 69
M? 73 75
M3 69 72
P3 51 43
Py 60 52
My 73 53
M2 75 70
M3 81 68
Similar figures are given by Bergounioux & Crouzel (1962a) who also define size
limits for the deciduous teeth.
TYPE SPECIES. Pyrodeinotherium bavaricum (von Meyer), 1831.
OTHER RECOGNIZED SPECIES. P. pentapotamiae (Falconer), 1868; P. hobleyi
(Andrews), IgII.
A detailed synonymy of P. bavaricum was given by Graf (1957) and of P. pent-
apotanuae by Sahni & Tripathi (1957).
Skull.
Dentition.
Skeleton.
FROM GEBEL ZELTEN, LIBYA 295
TABLE 2
Comparison of Prodeinotherium and Deinotherium
Prodeinotherium
rostrum ventrally flexed
rostrum narrow and deep
preorbital swelling close to external
nares
external nares almost as deep as wide
external nares surmounted by median
projection of nasals
orbit above P8
skull roof nearly flat and inclined
anteriorly
occiput steeply inclined
paroccipital processes longer than in
elephantoids
occipital condyles cut ventrally by
Frankfurt Plane
P3-4 usually lack mesostyles
M2-3 with well developed postmetaloph
ornamentation
tusks nearly vertical
scapula—stout spine, metacromion and
acromion; supraspinous fossa _ well
developed
humerus—lateral epicondyle tapers
proximally
radius—medial half of head larger than
lateral half; distal epiphysis more
massive than in Mastodon angustidens
lunar—radial facet covers most of
proximal surface; magnum facet is
concavo-convex
cuneiform—posterolateral process of
similar length to that of Elephas but
does not articulate with unciform;
unciform facet is concavo-convex
unciform—cuneiform facet roughly tri-
angular; largest distal facet is for McV
magnum—proximal surface has large
posteromedial projection
metacarpals—laterally compressed but
of similar size to Mastodon angustidens
manus more plantigrade
femur—similar length to Elephas maxi-
mus
astragalus—tibial facet equant and
convex; prominent posteromedial pro-
cess
Deinotherium
rostrum almost straight
rostrum wide and shallow
preorbital swelling more anteriorly
sited
external nares much wider than deep
no median projection of nasals
orbit above P4
skull roof shorter and narrower at the
temporal fossae
occiput gently inclined
paroccipital processes longer than in
Prodeinotherium
occipital condyles
Frankfurt Plane
P3-4 often possess mesostyles
M2-3 with reduced postmetaloph orna-
mentation
tusks longer and may be recurved
beneath symphysis
scapula—reduced spine and no meta-
cromion or acromion; supraspinous
fossa greatly reduced
humerus—lateral epicondyle does not
taper proximally
radius—medial half of head corres-
pondingly larger; distal epiphysis more
massive than in Prodeinotherium
lunar—radial facet extends less far
posteriorly; magnum facet almost flat
elevated above
cuneiform—posteriolateral process is
relatively longer and more ventrally
inclined than in Prodeinotherium; unci-
form facet is biconcave
unciform—cuneiform facet extends far-
ther posterolaterally and distally, and
tapers more abruptly posteriorly;
largest distal facet is for McIV
magnum—posteriomedial projection is
less pronounced
metacarpals—more compressed lateral-
ly and distinctly more elongate than in
Prodeinotherium
manus more digitigrade
femur—proportionately 30%
than in E. maximus
astragalus—tibial facet equant but
nearly flat posteromedial process more
reduced than in Prodeinotherium
shorter
296 PRODEINOTHERIUM
Teeth are the most common deinothere remains, but although there is a great deal
of variation in minor topographic features on the cheek teeth the basic tooth
morphology is conservative throughout the known history of the group. Indeed,
when only a single genus—Deinotheriwm—was accepted, tooth morphology was of
far less importance for specific differentiation than absolute size of the teeth or
geographic distribution of the specimens. In effect only two distinct species could
be recognized—D. gigantewm and D. bavaricum. However, with these two taxa
separated by generic rank, tooth morphology assumes greater diagnostic importance.
It is entirely possible that detailed revision of known material will furnish evidence
for distinguishing different lineages within the two genera incorporating variations
in cheek tooth morphology. Such a revision is, however, beyond the scope of this
paper.
Only two deinothere species are currently recognized from fossil mammal sites in
Africa—Prodeinotherium hobleyi of the Neogene and Deinotherium bozasi of the
Quaternary. The Gebel Zelten deinothere teeth are more advanced than specimens
of P. hobleyi from the earliest Miocene of East Africa but more primitive than those
from Ngorora (Pliocene) in Kenya. The observed differences are insufficient at
present to warrant the erection of a separate taxon for the Libyan material.
Prodeinotherium hobleyi (Andrews), 1911
1911 Deinotherium hobleyi Andrews: 943.
1919 Deinotherium cuviert Kaup; Brives: go.
1957 Deinotherium bavaricum v Meyer; Graf: 152.
1967 Deinotherium cuviert Kaup; Savage: 263.
Skull
(Text-figs 3-6; Plates 1-3)
MATERIAL REFERRED. 6404 : 14, fragmentary adult skull with only the dentition,
right premaxilla, left jugal and occipital condyles well preserved; 6404 : 44, complete
but slightly distorted immature skull; M.26665, virtually complete skull but with
zygomatic arches, squamosal, auditory and pterygoid regions poorly preserved;
6418 : 4 and 6418 : 19, incomplete and uncollected skulls.
DescriPTION. The skull of Prodeinotherium hobleyi from Gebel Zelten is elongate
and low, the length of the cranium being twice that of the rostrum. The skull is
characterized by a relatively flat skull roof which ascends gently to the lambdoidal
crest, a small orbit that is open posteriorly, a broad and downturned rostrum that is
surmounted by massive preorbital swellings, high occipital condyles, and a back-
wards sloping occipital region. The rostrum is appreciably narrower than in
Deinotherium giganteum and is excavated to form a deep rostral trough anterior to
the external nares. The occipital region is very wide owing to the well developed
lateral wings of the squamosal bones. Much of the skull is formed from cancellous
bone, and where the surface bone has been eroded diplée similar to those of ele-
phantoid skulls may be detected.
Key to letters used in text-figures 3, 4, 5 & 6.
b = auditory bulla
en = externalnares
f = frontal
fo = foramen ovale
fr = foramen rotundum
ga = glenoid articulation surface
in = internal nares
io = infraorbital foramen
lc = lamboidal crest
ls = lateral wing of squamosal
m = maxilla
n = nasal
ns = nasal sinus
o = orbit
FROM GEBEL ZELTEN, LIBYA
—)
oc = occipital condyle
p = parietal
pf = posterior palatine foramen
pgf = postglenoid fossa
plf = posterior lacerate foramen
pm = premaxilla
po = postorbital process of frontal
pp = paroccipital process
ps = preorbital swelling of maxilla
pt = pterygoid process
rt = rostral trough
Ss = squamosal
tf = temporal fossa
zm = zygomatic process of maxilla
zs = zygomatic process of squamosal
Fic. 3. Prodeinotherium hobleyi skull, anterior view. Scale = 5 cm.
297
98
tf
Fic. 4.
PRODEINOTHERIUM
Prodeinotherium hobleyi skull, dorsal view. Scale = 5 cm.
FROM GEBEL ZELTEN, LIBYA
Fic. 5.
Prodeinotherium hobleyi skull, ventral view. Scale = 5 cm.
299
300 PRODEINOTHERIUM
Premaxilla. The dorsal surface of the deep rostral trough (text-figs 4, 5) anterior
to the external nares is lined by the premaxillae. The anterior portion of the
premaxilla forms a small flange projecting anteriorly from the upstanding rugose
walls of the rostral trough (Pl. 1a). The premaxillary-maxillary suture extends
from the lateral edge of this flange along the medial edge of the wall of the rostral
trough until it meets the anterior wing of the nasal (text-figs 3, 4). The suture has
completely fused in skull M.26665 but the premaxillary-maxillary contact is denoted
by the junction of smooth bone, typical of the medial walls and floor of the trough,
with the rugose bone of the lateral walls and dorsal surface. The suture is still
visible in skull 6404 : 44. The posterior limit of the premaxilla beneath the external
nares is not clear owing to the fusion of sutures in M.26665 and the cracked nature
of the bone in 6404 : 44. Perhaps the limit is marked by the anterior edge of the
cavernous nasal sinus (text-fig. 4, Pl. rb) that occurs on either side of the external
nares.
Viewed from the dorsal aspect, each premaxilla is roughly triangular with the
posterior edge of the bone forming the shortest side. In transverse section the distal
part of the premaxilla is L-shaped, the ascending arm lining the medial wall of the
rostral trough while the body forms the floor. Medial to the preorbital swelling of
the maxilla (text-figs 3, 4, 5; Pls. 1-2), the ascending process of the premaxilla
bulges medially in front of the nasal sinus, and here the surface of the bone is curved
sigmoidally. The facial process of the premaxilla ascends to contact both the nasal
and the anteriormost prolongation of the frontal bone (text-figs 3, 4), thereby ex-
cluding the maxilla from contact with the nasal. Ventrally the premaxilla is visible
only at the anterior third of the rostrum where the premaxillary flanges appear
above the diverging tips of the maxillae (text-fig. 5; Pl. 3b). The flanges of the
premaxillae converge ventrally.
Maxilla. This is the largest bone of the facial region, extending from the dental
alveoli up to the level of the external nares, and from the anterior tip of the rostrum
to behind the dentition. The facial process of the maxilla is contiguous with the
frontal, lacrimal and premaxilla. It continues forwards and downwards from the
orbit as the external wall of the rostral trough. Just anterior to the frontal-maxillary
suture the dorsal and dorsolateral parts of the rostral walls are swollen in skull
M.26665 to form a massive preorbital swelling. In 6404 : 44 the swelling is less
pronounced but this portion of the maxilla is considerably elevated above the
remainder of the dorsal rim of the rostral trough. The infraorbital foramen
(Pl. 2a and b) is large and the anterior opening lies above the anterior edge of the
third premolar. In skull M.26665 two deep grooves extend anteriorly from the infra-
orbital foramen. One runs for a short distance down the lateral surface of the
rostrum; the other extends parallel to the dentition to terminate on the dorsal
surface of the rostral trough immediately in front of the preorbital swelling. These
grooves mark the proboscideal branches of cranial nerves V and VII. In skull
6404 : 44 only the former groove is well developed. The jugal process of the maxilla
is stout, expanding posteriorly to define the posteroventral limit of the orbit (Pl. 2a).
The posterior opening of the infraorbital canal is sited above the base of the jugal
process of the maxilla and beneath the postorbital process of the frontal (Pl. 2b).
FROM GEBEL ZELTEN, LIBYA 301
In ventral aspect the maxillae diverge behind the anterior third of the rostrum,
revealing the margins of the premaxilla. Behind the anterior third of the rostrum
the intermaxillary suture extends posteriorly down the midline of the palate as
a cristiform ridge (Pl. 3b). The maxillary-palatine suture is fused in skull M.26665
but is presumed to have been level with the posterior root of the second molar and
to have incorporated the paired posterior palatine foramina. Shallow grooves
extend anteriorly from these foramina until level with the fourth premolars. The
lingual border of the alveolar region is continued forward from the anterior root of
the premolar as a broad, rounded ridge, converging on its fellow towards the tip of
the rostrum. On the medial side of this ridge a very weak crest extends anteriorly
to meet the ventral exposed portion of the premaxilla. The plate is vaulted in the
vicinity of the premolars and narrows as it continues forwards beneath the rostrum.
In the region of the molars the palate is convex.
Nasal. In skull M.26665 the nasal-frontal suture is fused but there is a pro-
nounced break of slope some 7 cms behind the external nares that probably indicates
the junction of these bones. The surface of the bone immediately above and
behind the external nares is much smoother than the adjoining bone. In 6404 : 44
the lateral wing of the nasal extends laterally until level with the front of the orbit
and the suture with the premaxilla is well defined. In dorsal plan the nasals are
crescentic (text-fig. 4). In M.26665 the nasals continue anteriorly at the intranasal
suture to form a well developed median projection (Pl. Ia and b) though this is less
well developed than that of the extant elephants. In 6404 : 44 the anterior portion
of the intra-nasal suture is marked by an indentation rather than by a projection;
this is attributed to the relative immaturity of the specimen. Apparently the nasal
bones do not contribute to the cavernous nasal sinuses on either side of the external
nares.
Lacrimal. Due to fusion of the sutures and to incomplete preservation the
lacrimal is not well delineated in skull M.26665 but in 6404 : 44 the lacrimal is seen
to form the anteroventral quadrant of the rim of the orbit. The lacrimal bone is
triangular with the base of the triangle elevated a short distance above the infra-
orbital canal. From the centre of the ventral half of the bone a lacrimal foramen
opens into a lacrimal canal that passes outwards to the anteroventral rim of the
orbit. Above the lacrimal foramen is a large traction epiphysis. The lacrimal is
contiguous only with the frontal and maxilla and does not form part of the infra-
orbital swelling of the maxilla (cf. Andrews, 1921; 530).
Jugal. This bone is missing from skull M.26665 but is known from 6404 : 44 and
from the incomplete skull 6404 : 14. The jugal joins the zygomatic process of the
maxilla at the posteroventral edge of the orbit, at which point the zygomatic process
of the maxilla attains its greatest height. From the vertical jugal-maxillary suture
the jugal increases in height posteriorly while simultaneously becoming flattened
from triangular to oval in transverse section. The posterior portion of the jugal
is overlapped by the zygomatic process of the squamosal. The posterior end of the
jugal forms a smooth convex curve and gives off a small conical process from the
posterior ventral surface. A jugal from skull 6404 : 44 is less massive and lacks
the conical process on the posterior ventral surface.
202 PRODEINOTHERIUM
Palatine. This is fused to the palatine process of the maxilla level with the
posterior root of M? and extends for some distance behind M3. The palatine is
pierced by two asymmetrically placed posterior palatine foramina. In M.26665 the
anteriormost part of the palatonarial border is sited some way behind M? and is
gently curved with the lateral, more posterior, portions missing. The internal
narial opening is almost circular.
Vomer. The vomer extends back behind the level of the anterior part of the
palatonarial border and forms a V-shaped crest along the roof of the internal nares.
Immediately above the posterior end of the vomer a small chip of bone is missing
in skull M.26665 thus revealing part of the cavernous ethmoidal fossa.
Frontal. In M.26665 the frontal bones occupy just over half of the skull roof
(text-fig. 4). The nasal-frontal suture is fused but the anterior edge of the frontals
may be determined as above. Part of the frontal-parietal suture is revealed in the
centre of the skull roof of M.26665 as a sinuous interdigitating groove but this cannot
be traced laterally. The frontals are slightly domed convexly in the midline and
over the large postorbital processes. There are no supratemporal ridges. Both the
frontal and parietal portions of the skull roof extend ventrally to contribute to the
temporal fossa. The postorbital processes of the frontals (text-fig. 4) are blunt
protruberances from each of which a ridge descends ventrally and posteriorly to
continue as the free end of the alisphenoid. In front of these processes the frontals
demarcate the upper rim of the orbit.
Parietal. The parietal bones form the posterior part of the skull roof. They
increase in width antetiorly from the lambdoidal crest. The lambdoidal crest forms
the most elevated portion of the skull roof and strongly delineates the latter from the
occipital region. The dorsal surface of the parietals is slightly concave in transverse
section and there is no sagittal crest.
Fusion of the sutures on the posterior half of the cranium makes the boundary of
the parietals indistinct. Even in the immature skull (6404 : 44) the junctions of the
parietals, squamosals, alisphenoids and frontals are not clear. Beneath the sharply
delineated rim of the temporal fossa the bone in M.26665 is sculpted by deep and
wide grooves denoting the point of attachment of the temporal muscle. Ventral to
the sculpted surface the bone is much cracked and broken. In 6404 : 44 the walls
of the temporal fossae are better preserved but the bone shows no temporal muscle
scars. No trace of an interparietal may be detected.
Supraoccipital. This contributes to the central and dorsal regions of the occiput
(Pl. 3a) and is concave both sagittally and transversely. The perimeter of this bone
is fused but it must have been roughly triangular; whether or not it formed the dorsal
border of the foramen magnum cannot be determined. The dorsal edge contributes
to the posterior side of the central portion of the lambdoidal crest. The lateral
junction with the squamosal may be indicated by a pronounced ridge that extends
from the paroccipital process to the lamboidal crest. The supraoccipital is excavated
in the midline by two deep triangular pits that form fossae for the nuchal ligament.
The dorsal edge of the nuchal fossa lies some 5 cms below the lambdoidal crest; the
ventral edge some 8 cms above the foramen magnum. As in elephantoid skulls, the
two halves of the nuchal fossa are separated by a narrow ridge of bone. The floor
FROM GEBEL ZELTEN, LIBYA 303
of each pit is comprised of sculpted bone, as is that region between the top of the
nuchal fossa and the lambdoidal crest.
Exoccipital. The paired exoccipital bones form the ventrolateral regions of the
occiput and support very large and prominent occipital condyles. From the
dorsolateral edge of the nuchal fossa of skull M.26665 a prominent crest curves
ventrally and laterally to terminate on the lateral edge of the paroccipital process.
This ridge is interpreted as separating the exoccipital from the lateral wing of the
squamosal and is continuous with the lateral limit of the supraoccipital. The
condyles are triangular and converge as they taper ventrally. The condyles are
limited ventrally by a deep groove (the condyloid fossa of Palmer, 1924) that is
also present, though less prominent, on the dorsal edge. The condyles are most
elevated from the rest of the exoccipital bone at their anteromedial edge at which
point they protrude almost at right angles from the plane of the occiput. The
paroccipital processes are incomplete on M.26665 but on 6404 : 44 are seen to extend
for some distance below the base of the occipital condyles, merging laterally with the
ventral edge of the lateral wing of the squamosal.
Basioccipital. Fusion of the sutures and incomplete preservation prevents delinea-
tion of the periphery of this bone in M.26665. The basioccipital descends steeply
from the foramen magnum and becomes progressively more narrow anteriorly
(text-fig. 5; Pl. 3b). The posterior part is deeply notched by the foramen magnum
and extends laterally to the medial edges of the occipital condyles. Anteriorly and
ventrally the body is keeled. The anteroventral junction with the basisphenoid is
probably level with the glenoid articulation surface. The basisphenoid passes
uninterruptedly from the basioccipital but the keel becomes less pronounced
anteriorly and disappears before reaching the internal narial opening.
The basicranium of 6404 : 44 has been distorted by crushing and the ventral
surface is now parallel with the left lateral side. Slightly anterior to, and on either
side of, the notch excavated by the foramen magnum is a large boss. This probably
served for the insertion of the rectus anticus muscles. The right boss is level with,
but somewhat below, the paroccipital process. The left boss is less well preserved.
From this point the basioccipital narrows appreciably as it continues anteriorly.
Some 5 cms in front of the anterior edge of the foramen magnum a pronounced
keel extends down the midline. This keel extends for at least 9 cms until level with
the eustachian opening of the auditory bulla but in front of this point the bone
is too badly displaced to allow further observation of the keel. On either side of the
posterior end of the keel are excavated large fossae for insertion of the rectus anticus
major muscles. An elongate notch, the posterior lacerate foramen, is sited anterior to
the paroccipital process and lateral to the basioccipital at the posterior end of the keel.
Sphenoid. The junctions of the various component parts of this bone with
each other and with contiguous elements are for the most part indistinct. The
anterior boundary of the alisphenoid forms part of the prominent cristiform outer
wall of a deep groove that extends from the pterygoid process to the postorbital
process of the frontal. The optic foramen, that probably perforated the orbito-
sphenoid, lies some distance above and anterior to the slit-like anterior lacerate
foramen, and the orbitosphenoid-alisphenoid suture undoubtedly lay somewhere
304 PRODEINOTHERIUM
between the two. Both foramina lie in a deep groove beneath the alisphenoid-
frontal ridge. From the optic foramen a shallow groove extends forwards to the
orbit, marking the course of the optic nerve. The foramen rotundum lies posterior
to the anterior lacerate foramen and opens from the alisphenoid flap into a large
alisphenoid canal. The mandibular branch of the fifth cranial nerve emerged from
the posterior opening of the alisphenoid canal while the facial and maxillary branches
emerged from the anterior opening. The position of the maxillary branch is marked
by a deep groove, partly covered by a flap of bone, that extends towards the ptery-
goid process. In skull M.26665 this groove extends almost vertically (Pls. 2a and b)
but in 6404 : 44 it passes anteriorly at an angle of 45°. Behind the posterior exit of
the alisphenoid canal, and above the anterior portion of the auditory bulla, is a large
foramen ovale (text-fig. 6).
The pterygoid region is very poorly preserved in skull M.26665. Ventrally and
medially it passes uninterruptedly into the presphenoid and posteriorly into the
alisphenoid. Nearly all of the ventral part that, together with the palatine, forms
the lateral border of the internal nares is missing. The posterior surface of the
pterygoid forms a smooth crest that. passes laterally to the anterior tip of the
petrosal. This region is also poorly preserved in skull 6404 : 44.
Squamosal. This is one of the largest bones of the skull and forms the postero-
ventral wall of the temporal fossa. It is only poorly preserved in M.26665 and in
many places the surface bone is missing to reveal diplée beneath. The squamosal
continues anteriorly and laterally from the supraoccipital to project as a vast lateral
wing forming the widest part of the skull (text-fig. 4; Pl. rb). The external edge of
this wing passes uninterruptedly from the lambdoidal crest into the paroccipital
process of the exoccipital. In lateral aspect the junction of the squamosal with
both the alisphenoid and the parietal is indistinct. The zygomatic process of the
squamosal is less stout than that of the maxilla, being almost quadrate in transverse
section near the point of origin but becoming more slender as it extends forward
Fic. 6. Pvrodeinotherium hobleyi right auditory bulla. Scale = 5 cm.
FROM GEBEL ZELTEN, LIBYA 305
to overlap the jugal. The auditory meatus opens onto the external surface of the
lateral wing immediately posterior to, and slightly above, the zygomatic process.
In M.26665 this region is poorly preserved but in 6404 : 44 the passage of the meatus
is clearly seen. The passage of the meatus is gently inclined medially. The
glenoid articulation surface is convex and elongate transversely. There is no
postglenoid process and behind the glenoid surface there is a shallow transverse
channel forming the floor of the auditory meatus as in Elephas.
Tympanic bulla. The tympanic bullae are incompletely preserved in M.26665
but are seen to be relatively larger than in the skull of D. gigantewm from Eppeslheim,
Germany. Much of the surface bone on the Gebel Zelten specimen is missing and
sufficient detail is preserved to show only the eustachian opening below and behind
the foramen ovale. In skull 6404 : 44 both bullae are present but the left bulla is
incomplete and crushed and the right bulla (text-fig. 6) is incomplete. The bullae
are bounded medially by the basioccipital and basisphenoid (contributing in part to
the posterior lacerate foramen), posteriorly by the paroccipital process, and dorsally
by the squamosal and alisphenoid. The bulla projects ventrolaterally as a semi-
circle of bone. In dorsal aspect it may be divided into two unequal portions—a
posterior concave surface and an anterior (slightly larger) convex area. The regions
are separated by a ridge that runs posteriorly and laterally from the foramen ovale
to merge with the lateral perimeter level with the posterior lacerate foramen. The
concave area thus enclosed presents an irregular surface pitted with numerous
small foramina. The anterior portion of the dorsal surface is convex and consists
mainly of smooth bone although it is pierced by several small foramina and canals
near the anterolateral edge. The eustachian opening occurs at the anteromedial tip.
On skull 6404 : 44 the ventral surface of both bullae is incomplete but sufficient
bone is preserved to permit a composite description. The ventral surface is divided
sagittally into two unequal portions by a crest that extends from the paroccipital
process to the anterolateral margin of the bulla. The portion of the bulla lateral
to the crest is the smaller. The ridge is most pronounced posteriorly where it
separates the posterior lacerate foramen from the large stylomastoid foramen, and
becomes progressively less distinct anteriorly. The bone anterior to the stylo-
mastoid foramen and lateral to the ridge is concave transversely and heavily
sculpted. This part of the bulla is equivalent to the tympanohyal pit. Medial to
the crest the surface of the bulla is smooth but cut by transverse shallow grooves.
This portion of the left bulla is crushed and fragmented; on the right bulla the surface
bone is missing from the anteromedial region to reveal the cavernous tympanum.
In comparison with that of the extant elephants, the tympanic bulla of Prodeino-
therium is less elongate sagittally and wider transversely with a larger stylomastoid
foramen.
OTHER DEINOTHERIID SKULLS. Prodeinotherium pentapotamiae. An incomplete
skull of P. pentapotanuae was collected in 1912 by Pilgrim from the Lower Chinji
Sandstone, four miles from Chinji in the Salt Range of India. Two portions of the
skull are preserved—a palate with cheek teeth (I.M. A.460) and a basicranium (I.M.
A.461). These are of similar size to those of the Gebel Zelten skulls and were
306 PRODEINOTHERIUM
described by Palmer (1924). The skull foramina that Palmer identified are sited
similarly to those of the Gebel Zelten skulls. Only the base of the vertical tapering
pillar that Palmer identified as the hyoid is still preserved but the basal tympanohyal
portion would have fitted well into the elongate and narrow tympanohyal pit
anterior to the stylomastoid foramen in Gebel Zelten skull 6404: 44. Unlike
Palmer, I do not believe that the right paroccipital process of the Indian skull can
be far from its original position. Palmer had correctly identified the tympanic
bullae, unlike Andrews (1921), but these are now only poorly preserved on the
Indian specimen.
Eppelsheim skull. The skull of Deinotherium gigantewm from Eppelsheim,
Germany, was originally described by Klipstein and Kaup (1836) and the most
recent description of this specimen was given by Andrews (1921). In the light of
the information provided by the Gebel Zelten Prodeinotherium material it is now
necessary to revise some of Andrews’ interpretations of the Epplesheim specimen.
For example, on page 527 Andrews remarks:
. de Blainville suggests that possibly the occipital surface had been
crushed down towards the floor of the skull. Careful examination of the
specimen, however, does not seem to support this idea and it seems probable
that . . . it represents the condition of the living animal.’
TABLE 3
Prodeinotherium hobleyi skull measurements
Parameter M.26665 6404: 44
Total length of skull . ue e.O4 73
Width anterior tip of rostrum. Be Bees IO-l
Length rostrum to external nares eae 34
Width skull at external nares : 56 38
Width rostral trough at external nares. 17:8 118
Length external nares to lambdoidal crest 40 31
Minimum width skull roof . , ae Ae 28
Width lambdoidal crest ; 59°6 35
Height foramen magnum to lambdoidal
crest : é . A 2554 =
Height nuchal fossa : ; : 2» 12-3 12°6
Width nuchal fossa. : ; E59 EL
Width occipital condyles. : D272 —
Height foramen magnum . : ; 6-4
Width foramen magnum. Fi] —
Height molar alveoli to Jambdoidal crest 41-9 oi
Width glenoid condyles : 13°7 —
Length palatonarial border to rostrum tip 56 84
Width palate at P? ; : : 73 —
Width palate at M!_ .. : : if OTEO —
Width palate at M3. : : ; 83 —
FROM GEBEL ZELTEN, LIBYA 307
The cracked condition of the bone and asymmetry of the left and right lateral
wings of the squamosals reveal that the Eppelsheim skull has undergone at least a
modicum of postmortem distortion, either during burial or after excavation.
Differences between the cast of the ventral side and the specimen itself indicate that
some post-excavation restoration has been necessary. Indeed, much of the
squamosal region is now plaster of paris. Certain anomalous features of the
Eppelsheim skull are discussed later and indicate that restoration has not always been
skilful or accurate.
On page 528 Andrews stated ‘In Deinotheriwm no development of cellular bone
seems to have taken place.’ This is clearly wrong. In the Eppelsheim skull, as in
the Gebel Zelten skulls, portions of the surface bone are missing to reveal cellular
bone beneath. This phenomenon was noted in de Blainville’s earlier description
of the skull (de Blainville, 1837) and was confirmed by Palmer (1924) in his descrip-
tion of the skull of P. pentapotamiae. Skull 6404: 14 from Gebel Zelten was
unfortunately shattered in transit to Bristol but the fragments salvaged leave no
doubt that much of the skull was composed of cellular bone.
On page 529 Andrews noted that ‘in front of the orbit the maxilla forms a great
mass of bone terminating in a rugose, somewhat concave surface for the attachment
of muscles’. These concavities are preservation anomalies and not original mor-
phological features. That on the left side is much smaller than the right concavity.
In corollary the preorbital swellings of the Eppelsheim skull are unequal, that on the
left being much larger. During restoration a large piece of bone has been cemented
on to the anterior part of the left swelling thus increasing its total size. This piece of
bone is the same length as the concavity on the right side and, if it were to be
transferred there, the preorbital swellings would be (a) symmetrical, (b) of similar
size and (c) convex.
Further, on page 530, Andrews declared:
‘The point of union of the premaxilla (with the maxilla) is probably marked
by the sudden narrowing of the snout about 40 cms in front of the orbit.’
This sudden narrowing marks the anterior termination of the preorbital swelling.
The ‘suture’ to which Andrews refers in the next sentence occurs on both the dorsal
and ventral surface of the rostrum and bisects both the premaxilla and maxilla. It
appears likely that the anterior part of the rostrum was once detached and the
‘suture’ marks the point of breakage. The actual suture between the premaxilla
and maxilla is well seen on Gebel Zelten skull 6404 : 44 and there is no reason to
believe that the geometry of these bones differed appreciably on the Epplesheim
skull.
The anterodorsal surface of the external nares appears to be complete and is
similar in morphology to the equivalent portion of a skull fragment of Deinotherium
giganteum from Palencia, Spain. There is no anterior median process in contrast
to the condition in Gebel Zelten skull M.26665. The lacrimal bone cannot be
observed in the Eppelsheim skull but was more likely to have occupied a similar
position to that in 6404 : 44 than to have formed part of the preorbital swelling
(cf. Andrews, Ig2I : 530), especially as the latter feature is found farther forward
308 PRODEINOTHERIUM
than in the Gebel Zelten skulls. The ‘irregularly arranged pits’ at the anterior end
of the premaxilla (Andrews, Ig2I : 531) are not, as Andrews suggested, rudimentary
tusk alveoli but diploe revealed by erosion of the external surface of the bone at this
point.
Any comparison made between the occipital regions of the Eppelsheim and Gebel
Zelten skulls is liable to be suspect for the roof of the Eppelsheim specimen is
incomplete posteriorly. The lambdoidal crest appears to be missing and much of
the bone between the foramen magnum and the lambdoidal crest is either absent or
distorted (cf. Andrews, 1921 : 527). The shape of the occipital surface of the
Eppelsheim skull appears to be more oval than in the Gebel Zelten skulls and the
nuchal fossa is shaped differently. The nuchal fossa of the Eppelsheim skull may
be divided into two portions—an anterior deep and wide oval pit (long axis trans-
versely orientated) and a posterior triangular trough that is shallower and tapers
ventrally. The latter trough may be all that remains of the true nuchal fossa on the
Eppelsheim skull.
The ‘large post-tympanic flanges’ (Andrews, Ig2I : 527) are equivalent to the
paroccipital processes of other mammals and are, as Andrews described, formed
partly of the exoccipitals and partly of the squamosals. The paroccipital processes
are smaller in the Gebel Zelten skulls. The ‘tongue-like process of bone’ (Andrews,
1921 : 527) that is attached to the left paroccipital process was incorrectly identified
by Andrews as the paroceipital process itself. This piece of bone is seen only on the
left side of the skull and is more likely to represent part of the hyoid bone (cf. Palmer,
1924: 4). The large convex bulbs that Andrews (1921 : 530; fig. 3) interpreted as
tympanic bullae are seen also on Gebel Zelten skull 6404 : 44 where I have interpreted
them as bosses for the insertion of the rectus anticus muscles. The tympanic
bullae of the Epplesheim skull are more likely to be represented by a strong crest of
bone that terminates posteriorly at the hyoid and extends forward apparently to
merge with the pterygoid flange. A petrosal of D. gigantewm was described by
Claudius (1865) but the original specimen is now lost and the accuracy of its identi-
fication must be regarded as questionable (pers. comm. H. Tobein).
Munich skull. Stromer (1938) described the remains of a skull, mandible and
partial skeleton of D. gigantewm from the Sarmatian Flinzsande of Munich. Un-
fortunately all except the cast of the palate were subsequently destroyed and only
Stromer’s description and diagrams are now available for study. The skull was
originally incomplete, lacking the lower part of the occiput, the basicranium, the
ventral walls of the temporal fossa and part of the skull roof. However the frag-
ments that were collected are of importance in that they are less distorted than some
portions of the Eppelsheim skull.
The rostral flanges of the premaxilla and maxilla are separated in the midline as
in the Eppelsheim skull, and are flexed at a similar angle, i.e. less steeply than
Prodeinotherium. The preorbital swellings were either not developed or have not
been preserved. The surface of the skull roof was relatively flat and, as restored
by Stromer, may have been more horizontal posteriorly than in the Gebel Zelten
skulls. Only the upper portion of the occiput was preserved and was described as
FROM GEBEL ZELTEN, LIBYA 309
strongly inclined posteriorly, though this cannot be detected from the illustrations
given by Stromer. The lateral wing of the squamosal was more vertically aligned
than in the Eppelsheim skull and strongly resembles that of Prodeinotherium,
indicating that the Eppelsheim skull was badly deformed in this region.
Palencia skull. The skull roof and palate of D. giganteum from Castrillo de Villa-
vega, Palencia, Spain, is housed in the Madrid Natural History Museum. The nasal
bones are represented but are incompletely preserved on the right side. They
surmount an external narial opening that is greatly compressed dorsoventrally.
The lateral sinuses, which are large on the Gebel Zelten Prodeinotherium skulls, are
much less prominent in the Palencia specimen. The nasal-frontal suture is fused
but the nasals are demarcated as a flat surface projecting anteriorly at an angle from
the remainder of the skull roof. The intra-nasal suture is fused and there is no
anterior median projection of the nasals over the external nares. The skull roof
forms a hollow behind the posterior edge of the nasals and in front of the lambdoidal
crest. The lambdoidal crest is incomplete on the right side but on the left extends
down the lateral wing of the squamosal to contribute to the widest part of the skull.
The postorbital process of the frontal is very large and the anterior portion is located
in front of the external nares. Behind the postorbital process the lateral surface
of the temporal fossa is strongly delineated from the skull roof, the angle between
the two surfaces being in the order of 45°.
Only the dorsal edge of the occipital surface is preserved and this had been in-
completely prepared. It is not therefore possible to determine at present whether
the occiput was inclined as in the Eppelsheim skull or the Gebel Zelten specimens.
The nuchal fossa is incomplete. The shape indicated by the upper part of the fossa
is approximately triangular and the dorsal edge of the fossa sited only some 5 cms
below the lambdoidal crest. Although the orbit is not represented on this specimen
the position of the postorbital processes suggests that the orbit was sited in advance
of the external nares in contrast to the situation in the Gebel Zelten specimens.
Discussion. Comparison of the skulls of Prodeinotherium and Deinotherium, as
represented by the Gebel Zelten, Eppelsheim and Palencia specimens, yields evidence
of both positive and negative allometric changes. Using the overall skull length as
a basis for comparison, parameters that remain the same in both genera are skull
width at the external nares, width at the lambdoid crest, and length of the rostrum.
The only major increase in width is in the rostral trough of Deinotheriwm and this is
exhibited in both the Palencia and Eppelsheim skulls. In both these skulls the
external nares are also sited farther posteriorly than in Prodeinotherium and are
considerably less deep than wide—converging on the elephantoids in both position
and shape. The length and width of the skull roof are proportionately smaller in
Deinotherium than in Prodeinotherium. The facial/cranial region ratio remains
constant and the decrease in skull roof length in Deinotherium is directly propor-
tional to the increase in area of the occipital region due to the anterior inclination of
the latter. The shorter skull roof of Deinotheriwm is also correspondingly narrower.
This is a compensatory feature to permit a large area of origin of the temporal
muscles despite a shorter skull roof.
310 PRODEINOTHERIUM
The majority of the differences between the cranial morphology of Deinotherium
and Prodeinotherium involve only relatively minor allometric changes and some of
these are foreshadowed by the differences between the immature (6404 : 44) and
mature (M.26665) Prodeinotherium skulls. Thus the rostrum is more steeply down-
turned in M.26665 than in Deinotheriuwm and the rostral flexure of 6404 : 44 is even
more pronounced. Decrease in rostrum curvature in Deinotherium, together with
increase of rostral width and retraction of the external nares, suggests the develop-
ment of a more powerful and effective proboscis in the later genus. The length
measurements of 6404 : 44 and M.26665 are very similar but the skull width para-
meters are relatively smaller in the immature specimen.
In Deinotherium the elevation of the occipital condyles, elongation of the paroc-
cipital processes and increase in inclination of the occiput are associated with
functional changes enhancing the downward movement of the head (and thereby
the tusks). The greater inclination of the occiput has contributed to a decrease
in the length of the skull roof, although this is also affected by the more posterior
siting of the external nares. The shorter skull roof limits the potential area of
attachment of the temporal muscles but this is compensated for by excavation of the
temporal fossa and decreased width of the skull roof. In this way major changes in
cranial morphology in the Deinotheriidae are associated with proboscis development
and with increase in potential movement of the skull on the neck, both factors
suggesting changes in feeding methods and habits.
TABLE 4
Deinotherium skull measurements
Parameter Eppelsheim Munich Palencia
Total skull length 109 ad =
Width anterior tip rostrum 34°7 — —
Length rostrum to external nares 53°5 — —
Width skull at external nares 79 — 69
Width rostral trough at external nares 49°8 as 32*
Length external nares to lambdoidal crest 29 33 34
Minimum width skull roof 48 40 45
Width at lambdoidal crest 98 — —
Height lambdoidal crest to foramen magnum 47°73 — —
Height lambdoidal crest to nuchal fossa 9°4 — —
Height nuchal fossa 19°5 — —
Width nuchal fossa ice) ~- —
Width occipital condyles 30°5 — —_
Height foramen magnum 5°6 — —=
Width foramen magnum 14-1 -— —
Height lambdoidal crest to molar alveoli 50* — —
Width glenoid condyles 27 21 —
Length palatonarial border to rostrum tip 68 —- —
Width palate at P38 15°8 77 a
Width palate at M! 18-7 = ——
Width palate at M3 05 10 --
FROM GEBEL ZELTEN, LIBYA 311
Mandible
(Plates 4-5)
MATERIAL REFERRED. 6404 : 13, incomplete horizontal rami with tusk fragments
and cheek teeth; 6404 : 45, fragmented immature mandible with tusks and dentition ;
6412 : 10, tuskless mandible with worn cheek teeth; B.M.(N.H.)M.26666, mandible
with tusks and cheek teeth; 6419 : 16, symphysis fragment with tusk alveoli.
DESCRIPTION. Four nearly complete mandibles and a number of mandibular
fragments have been collected from various sites at Gebel Zelten. None of the
specimens is complete but sufficient information can be derived from the specimens
to provide the following composite description.
The horizontal ramus is sigmoid, rising gently in front of Mg and flexing ventrally
at M, until at the tusk alveolus it subtends an angle of 115° to the alveolar border.
The cheek teeth are erupted in almost parallel rows on the median dorsal surface
of the horizontal ramus. The alveolar border is produced anteriorly to form a
strong ridge extending from the third premolar to the tusk alveolus. These ridges
converge anteriorly and enclose a gutter-like suprasymphysial depression similar to
that of the elephantoids. The symphysis itself is broad and extends anteriorly
from the vicinity of the posterior root of the third premolar. In one specimen
(6412 : 10) a wide and shallow pit is preserved on the.ventral and medial side of the
horizontal ramus beneath the posterior root of Ps. The pit may represent the point
of insertion of the anterior belly of the digastric muscle. On each ramus there are
three mental foramina. In mandible 6404 : 13 the posterior foramen is slightly
anterior to the front root of P; in 6412 : Io it is level with the hind root of this
tooth. In both specimens the anterior mental foramen is sited about 8 cms in
front of the posterior mental foramen, and the intermediate foramen varies in location
between the two.
Beneath the third premolar the horizontal ramus is oval in transverse section,
being deeper than wide and slightly more convex laterally than medially. Beneath
Ms the horizontal ramus has flat and parallel lateral and medial sides and develops a
small keel on the ventral surface. The dorsal surface of the horizontal ramus
increases in width posteriorly and the lateral surface becomes the leading edge of the
ascending ramus when level with M3, from which it is separated by a wide, oval and
concave platform.
The leading edge of the ascending ramus is nearly at right angles to the horizontal
ramus. The coronoid process is small and directed anteriorly. The mandibular
condyle is elevated only a short distance above the coronoid process and is a little
wider than the latter is long. The condyle is almost cylindrical. The inferior and
posterior borders of the ascending ramus are appreciably thicker than the adjoining
bone. There is a pronounced notch beneath the dental foramen on the inferior
border of mandible 6412 : 10 that may mark the insertion of the sternomandibular
muscle. The medial surface of the ascending ramus is divided unequally by a ridge
extending from the base of the mandibular condyle to the alveolar border beneath
Ms. A second ridge descends posteriorly from the anteromedial face of the coronoid
process and joins the first beneath the hind edge of the coronoid process. The
312 PRODEINOTHERIUM
triangular area thus enclosed serves for the insertion of the temporal muscle (which
also occupies a depressed area of similar size on the lateral face of the ascending
ramus). The masseter muscle inserts on the lateral surface of the angle of the jaw.
The internal pterygoid muscle inserts similarly on the medial surface of the angle and
the external pterygoid muscle inserts above the dental foramen and between the
mandibular condyle and coronoid process. The digastric muscle inserts on the
ventromedial surface of the mandible beneath and in front of the coronoid process.
COMPARISON WITH OTHER DEINOTHERE MANDIBLES. The degree of curvature
and length of the mandibular symphysis varies appreciably with the various deino-
there taxa. The mandibular symphysis of Deinotherium giganteum is longer and
has a more vertically aligned symphysial region than that of Prodeinotherium
bavaricum. D. bozasi of the African Quaternary has a relatively short mandibular
symphysis but this is flexed at right angles and therefore more abruptly than in any
other deinotheriid taxon. The symphysis of D. levius (here interpreted as synony-
mous with D. giganteum) is intermediate in length and degree of flexure between
D. giganteum (sensu stricto) and P. bavaricum (Graf, 1957). Arambourg (1934)
believed the apparently more acute flexure of D. Jevius separated this taxon from
D. giganteum, and Laskarev (1944) deduced that D. levius had African rather than
European affinities. The mandibular symphysis of P. hobleyi from Gebel Zelten is
similar to that of P. bavaricum.
Sahni & Tripathi (1957) separated the two Asian deinotheriid taxa on the basis
of the transverse section of the horizontal ramus level with M3. In D. indicum the
jaw bulges on either side of Mg, the lateral bulge being more pronounced and giving
rise to a flat platform on the lateral side of this tooth and in front of the ascending
ramus. In contrast, the jaw of P. pentapotamiae has a narrow and elliptical cross
section. In this respect P. hobley: from Gebel Zelten is similar to D. indicum.
Graf (1957) and Bergounioux & Crouzel (1962a) showed that the mental foramina
of the mandible varied greatly in position. Those of D. giganteum and D. bozasi
tended, however, to be sited more anteriorly than those of P. bavaricum. The mental
foramina of the Gebel Zelten Prodeinotherium are also sited farther forward than in
the examples of P. bavaricum quoted by Graf (1957 : 168).
TABLE 5
Prodeinotherium hobleyi mandible measurements
Left horizontal ramus (6412 : IO)
Total length . : : ‘ : =< 163
Height at tusk alveolu : : , : 127
Height at Ps. : : : : : 13°53
Height at M,_ . : : F ; E I2°3
Height at Ms... : . : : 3 Lit
Width at Ps. ; : : : : 7°3
Widthat M; : 7 d ! ‘ 7h |
Width at M3_«tj ; : : ; : 10-7
FROM GEBEL ZELTEN, LIBYA 313
Left ascending ramus (6412 : IO)
Length . : ; ; : a i22"6
Height at coronoid process : : 5 26
Height at mandibular condyle : ‘ 206
Mandibular condyle ee =
Length ©. : ; : ‘ 4°7
Width ; : : : ; ; ‘ 103
Dentition
(Text-fig. 7; Plates 4-5)
MATERIAL REFERRED. 6401 : 4, right maxilla with P?-M3; 6404 : 13, mandible
with tusks and cheek teeth; 6404 : 44, immature skull with cheek teeth; 6404 : 45,
fragmented immature mandible with tusks and cheek teeth; 6412 : 10, tuskless
mandible with worn cheek teeth; M.26665, adult skull with cheek teeth; M.26666,
mandible with tusks and cheek teeth; 6401 : 1, LM3; 6404 : 14, LP?-M? and RP3-4;
6409 : 41, M2 fragment; 6410 : 2, RP4; 6412 : 39, LM2; 6412 : 40, LMj; 6412 : 54,
LP4; 6413 : 18, LM®; 6416 : 120, tusk tip; 6416 : 130, tusk; 6421 : 2, LM2; 6404 : 28,
6405 : 48, 6419 : 17, 6421 : 15, 6423 : 13, 6423 : 45, 6423 : 56, enamel fragments.
DEscripTIon. Teeth are the most commonly preserved deinotheriid remains and
differ sufficiently in size and morphology from other Cenozoic mammals to permit
ready identification even on incomplete specimens. The terminology employed in
previous descriptions of deinothere teeth has varied greatly from author to author.
The teeth are basically lophodont, the lophs originating through coalescence of
adjacent cusps in a bunodont ancestral stock. Terms used here are based on the
standard cusp nomenclature (Osborn, 1907) and those used by Osborn for probos-
cidean teeth (Osborn, 1942). The terms are here used only to express topographic
equivalence with the dental characters of other mammals and have no genetic
significance.
Superior dentition. Downturning of the rostral region involved the posterior
retreat of the external nares, accompanied by withdrawal of the premaxillae to line
the floor of the rostral trough. The incisors, canines and anterior cheek teeth were
lost during, if not before, the hindward migration of the premaxillae. All lophs
and cingula on the upper teeth are rugose until worn. The lingual edges of the
transverse lophs are taller than the labial edges and show more wear. A well
developed facet is formed on the anterior surface of each transverse loph and extends
for the entire width of the loph. Parallel facets are seen also on the anterior and
posterior cingula when the tooth is worn sufficiently for these to occlude with the
lower dentition.
The third premolar is equant and possesses two internal cusps and an ectoloph
(text-fig. 7A). The protocone is the larger cusp and is separated from the hypocone
by a deep but very narrow valley. The protocone extends labially and anteriorly
to the base of the ectoloph, thus forming a low, incipient protoloph. The hypocone
bears a small mesostyle on its anterolingual face. The hypocone is elongate labially
but does not forma metaloph. The strong ectoloph is formed by the coalition of the
paracone, metacone, and an intermediate cusp that is taller than either. The
314 PRODEINOTHERIUM
posterior end of the ectoloph is continuous lingually with the posterior cingulum that
terminates lingually on the posterolingual face of the hypocone. The median valley
is wide and open posteriorly ; it is deepest between the labial bases of the protocone
and hypocone. The anterior cingulum connects the base of the protocone to that
of the ectoloph. There are no lingual or labia cingulla.
The fourth premolar is bilophodont and subequant, being longer than broad.
The ectoloph is less strong than in the third premolar and the intermediate cusp is
reduced to a swelling on the anterior face of the metacone. The transverse lophs are
not straight, the protoloph being convex anteriorly and the metaloph convex
posteriorly. Both lophs are connected labially with the ectoloph. From the
posterior lingual face of the protocone a ridge extends posterolabially to the centre
of the median valley and a similar ridge extends down the anterior face of the
A
Fic. 7. Pyrodeinotherium hobleyi upper teeth. A = left P?, B = right M!; Not to scale.
ac = anterior cingulum, h = hypocone, me = mesostyle, p = protocone, T = tritoloph,
E = ectoloph, M = metaloph, P = protoloph, pe = posterior cingulum.
FROM GEBEL ZELTEN, LIBYA 315
hypocone. The metaloph of the fourth premolar is the only loph in the upper
dentition to bear a posteriorly directed wear facet. The median valley is widest
labially next to the ectoloph and narrows, opening lingually between the protoloph
and metaloph. It is almost, but not quite, dammed by a small mesostyle on the
anterior face of the hypocone. The anterior and posterior cingula are well developed.
There is a minute internal cingulum.
The anterior molar is trilophodont and subrectangular, being longer than wide and
narrowest posteriorly (text-fig. 7B). All three lophs bear ridges directed towards
the midline of the tooth from their posterolingual and posterolabial edges. The
tritoloph (posterior loph) is much less wide than either the protoloph or metaloph
and the ridges from the posterolingual and posterolabial edges of the loph are less
pronounced than in the protoloph or metaloph. Between the protoloph and meta-
loph the anterior median valley is blocked labially by a small bune between the
paracone and metacone, and lingually by a short spur connecting the protocone
to the hypocone. This spur bears a small mesostyle. The posterior median valley,
between the metaloph and tritoloph, is much less deep. It is blocked lingually by a
short spur connecting the lingual edges of the metaloph and tritoloph, and labially by
the junction of sharp ridges extending down the posterior face of the metaloph and
anterior face of the tritoloph. The anterior cingulum is strong but the posterior
cingulum is less well developed than in the premolars.
The second molar is bilophodont and slightly longer than wide. The protoloph
is similar to, but taller than, the metaloph. The centrally directed ridges on the
hind face of the protoloph are well developed and stronger than in M1. The ridge
from the lingual edge of the loph extends beyond the midline of the tooth; that from
the labial end of the loph is more posteriorly directed and fuses with a similar ridge
extending from the crown of the metacone. The median valley is open labially
but is bounded lingually by a short spur joining the protoloph to the metaloph.
The postmetaloph ornamentation is rather complex. A strong ridge extends from
behind the hypocone to join the lingual edge of the posterior cingulum. A more
ventrally directed ridge connects that from the hypocone to a posterolabially
directed ridge from the metacone, thus forming a subsidiary loph behind the meta-
loph. The posterior cingulum bears a large style on its labial extremity.
The third molar is subequant, and tapers posteriorly. Like the second molar it is
bilophodont. The topography of the protoloph is similar to that of M2. The
metaloph is less wide than the protoloph, and the postmetaloph ornamentation is
simpler. A ridge descends from the hypocone to the lingual termination of the
posterior cingulum. The metacone gives off a strongly developed ridge that
descends posteriorly and then centrally to the midline of the tooth to connect with
the posterior cingulum. In some teeth the metacone ridge is also connected to the
hypocone ridge and thus the postmetaloph ornamentation of M3 is more variable
than in M2. There is no style on the posterior cingulum.
Inferior dentition. Tusks are present in mandibles 6404 : 13, M.26666, and
6404: 45. The tusks are oval in transverse section with the long axis aligned
anteroposteriorly. Enamel was preserved only on the tusks of the immature
mandible (6404 : 45) where it covered the entire erupted surface of the tooth. The
c
316 PRODEINOTHERIUM
tusks in 6404 : 45 are short and nearly straight but in mandible M.26666 they
curve downwards until the tip is nearly vertical. The dentine shows no ‘engine
turning’ structure as in Mastodon but forms bands parallel to the periphery of the
tooth.
As in the superior cheek teeth, all lophs, cingula and isolated cusps of the lower
molars and premolars are rugose until worn. Except on Py, all wear facets on the
transverse lophids occur on the posterior surface. The labial edges of the lophids
are less high but show more wear than the lingual edges.
The third premolar is triangular and tapers anteriorly. The protoconid is the
largest and tallest cusp. It is elongate sagittally and is sited in the midline of the
tooth. The protoconid is continued forwards as a high narrow ridge extending to
the lingual edge of a very abbreviated anterior cingulum. It is also connected
posterolingually to a tall metaconid. From the junction of the protoconid and meta-
conid, a narrow and less elevated ridge proceeds posteriorly to merge with an
elongate hypoconid. This in turn connects with an elongate entoconid via a weakly
developed, low, posterior cingulum. A C-shaped hypoconulid may or may not
connect with the posterolabial edge of the entoconid. The protoconid and outer
face of the ectolophid became worn more rapidly than the remainder of the tooth.
The fourth premolar is bilophodont and represents a wider and more elongate
version of Ps. The protoconid and metaconid are set farther apart and are con-
nected by a protolophid that is curved with the convex face posteriorly. The
anterolingual extension of the protoconid extends more laterally than in P3. The
protoconid is also joined to the hypoconid by a low ectolophid. There is no hypo-
conulid but the hypoconid is connected to the entoconid by a hypolophid that is
straighter and less tall than the protolophid. There are no well defined wear facets
on the hypolophid but the lateral extremities become worn faster than the central
portion and small wear facets occur on both sides of the lophid. The hypoconid is
connected by a short spur to the strong posterior cingulum.
The first molar is trilophodont, the protolophid and hypolophid being wider than
the tritolophid. A strong anterolingual ridge on the anterior face of the labial edge
of the protolophid projects towards, but does not join, a weaker anterolabial ridge
from the lingual extremity of the lophid. Similar but less strongly developed ridges
may be traced on the anterior faces of the hypolophid and tritolophid. The anterior
and posterior median valleys are open at both sides. The anterior cingulum is
less well developed than the short, centrally placed posterior cingulum that stands
proud at the rear of the tooth.
The second and third molars are bilophodont and are differentiated only by the
posterior taper on the Mg and the better developed posterior cingulum on that tooth.
From the labial edge of the protolophid a strong medial ridge extends postero-
ventrally to meet the anterior cingulum. A similar but less strong ridge extends
from the lingual extremity of the lophid. These ridges are duplicated in a less
pronounced fashion on the anterior face of the hypolophid. The median valley
is open at bothends. As in Mj, the wear facets on the lophids face posteriorly. The
posterior cingulum of Mg is less wide than that of Mg but is taller, more curved,
and extends farther posteriorly.
p4
M2
M3
My
Mg
ap
prot
trit
*
FROM GEBEL ZELTEN, LIBYA
TABLE 6
Prodeinotherium hobleyi tooth measurements (in mm.)
Upper premolars
6401: 6404: 6404: 6412: M.26665(1)
+ 14(I) 14(r) 55
ap 47°5 564 Ceieiae == 40°5
prot — 55:0 — 511 —
met 50°1 55°60 — — 47°5
ap Aare 47°1 = Te 47°9
prot — 62:7 64:0 — 5855
met — 60°3 = — 56-7
Upper molars
6401: 6401: 6404: 6413: M.26665(1)
I 4 14(l) 18
ap —_ 68-6 76:6 — 70°5
prot — 58-8 60-7 —— 62°5
met — 56-7 60-6 — 61°5
eet 47°7 49°2 = =
ap — 67°8 732 — 70:6
prot — 67°9 69:0-+ —_— —
met — 66:5 64:0-+ — —
apy 9 075°7 67-2 = 59°3 728+
prot 77:2 67°9 —_ 62-0 77:6
met 66-9 62:6 — — 739
Lower premolars
6404: 6404: 6404: 6404: 6410: 6412
13(1) 13(r) 45(1) 45(r) 2 10(1)
ap 43°3 45°5 491 5°°7 a 39°6
tr 32°3 35°0 40°7 39°6 a 31°7
ap = 544 531 = 58-9 == 44°3
prot 43-0 43°9 = 46'9 38°4 38-6
hyp 45°6 43°2 a 43°3 34°9 =
Lower molars
6404: 6404: 6404: 6404: 6412: 6412 6412:
13(I) 13(r) 45(1) 45(r) ro(1) 10(r) 39
ap = 70°0 65°3 82-7* = 55°0 55°3 —
prot 48-8 47:0 47°4* —_— 38°8 41°8 —
hyp 51:0 49°1 48-2* = 39°5 39°8 =
trit 45:9 44°3 38-3 = 33°8 34°2 =
apo 71-9 om 76°3 57°0 54°8 =
prot 65°5 61-6 — 61-1 503 50°4 52°8
yp 59°0 = 62:2 48-0 49°5 =
ap — — a= 84-0 66-9 67:8 _—
prot — = = 63°4 541 54°9 =
hyp == = = 59°3 47°6 46°4 =
= greatest sagittal length: tr = transverse width;
= width at protoloph(id) ;
= width at tritoloph(id) ;
= approximate measurement.
met = width at metaloph(id) ;
hyp = width at hypolophid;
M.26665(r)
43°7
46°1
44:9
58°4
57°6
M.26665(r)
715
62:9
317
318 PRODEINOTHERIUM
TOOTH FUNCTION IN DEINOTHERES. The function of deinothere tusks has yet to
be satisfactorily explained. Conventional interpretations include pulling branches
down to mouth level and digging for food. It is unlikely that deinothere tusks were
used for digging for the following reasons. The tusks would fail to reach the ground
unless the animal was either kneeling or standing at a lower elevation than the
surface at which digging occurred. Such wear facets as have been observed are not
consistent with a digging function. Food excavated from below ground surface
would inevitably be contaminated with soil which would result in rapid attrition
of the cheek teeth; no compensatory increase in hypsodonty of the deinothere cheek
teeth has been observed.
The tusks of immature deinotheres are covered by enamel. Enamel production
apparently ceases shortly after the eruption of the tusks. The absence of enamel on
the tusks of mature deinotheres suggests that the enamel-bearing portion was subse-
quently removed by wear. I have observed wear facets in two places on deinothere
tusks—on the posterior surface of the tusk tip (P. hobleyz) and on the anteromedial
surface of the tusk tip (P. hobleyi, P. bavaricum, D. bozasi). The anteromedial
facets may be attributed to constant abrasion by the proboscis but might conceivably
have been produced during the action of stripping bark or branches from vegetation.
A detailed description of the cheek tooth function in deinotheres is in the process
of preparation. A summary of the preliminary conclusions is given below.
Deinothere cheek teeth occlude orthally and may be functionally divided into anterior
crushing teeth and posterior shearing teeth. In the premolars some shearing is
effected during occlusion of the ectoloph and ectolophid. The transverse crests on
the premolars serve to prevent anterior dislocation of the jaw during the occlusal
stroke of the mandible and perform a crushing function during the recovery stroke.
The transverse crests of the deciduous fourth premolars and of the molars have well
developed shearing facets—on the anterior surface of the lophs and rear of the lophids.
These facets are maintained until the transverse crests have been almost completely
removed by wear. In unworn teeth the facets are aligned almost vertically but
become progressively more horizontal with wear. The change in angle of the shear
facet is seen not only between individuals of different maturity but also within single
tooth rows, the facets on the posterior end of the tooth row being oriented more
vertically than on the anterior molars.
The anterior crushing battery and posterior shearing battery are reconstituted
three times during the life of the individual as follows:
anterior battery posterior battery
Juvenile animal DP2-3 DP4
Immature adult P3-4 M1
Mature adult P3-4, Mr M2-3
The anterior trilophodont molar performs different functions at different stages of
wear. Initially it serves as a shearing tooth in contrast to the crushing premolars.
By the time the second and third molars are fully functioning the anterior molar is
often too worn to provide an efficient shearing action and becomes incorporated into
the anterior crushing battery.
FROM GEBEL ZELTEN, LIBYA 319
The width of the anterior molar, its attitude in the alveolus, and its alignment in
the tooth row all suggest that it functions primarily as a posterior premolar rather
than as an anterior molar. The shearing action of this tooth is of limited duration
but sufficient to cope during the transition between deciduous and permanent
dentition. The bifunctional nature of the tooth row may well explain why deino-
theres failed to develop horizontal tooth replacement as in the elephantoids.
Axial skeleton
(Text-figs 8-13)
MATERIAL REFERRED. M.26667h, left rib head; M.266671, right neuropophysis
of third cervical vertebra; 6419 : 15, atlas vertebra; 6424 : 79, axis vertebra.
DescripTIon. Adlas vertebra. The atlas vertebra of Prodeinotherium hobleyi
(text-fig. 8, 6419 : 15) is smaller than that of Mastodon angustidens (6412 : 185)
from Gebel Zelten. Although the transverse processes of the mastodont atlas are
incomplete the Prodeinotherium atlas was probably wider. The facets for articula-
tion with the occipital condyles are comma-shaped in both specimens but the masto-
dont facets are larger, relatively taller, less wide and apparently more concave.
Differences on the dorsal edge of the neural arch are especially pronounced. The
neural spine of the mastodont atlas forms an elongate transverse ridge with large
and widely separated scars for the insertion of the rectus muscles. The Prodeino-
thertum atlas spine is a shorter, blunt protruberance that is bounded by rectus
muscle scars that are smaller and face more ventrally than in Mastodon. The
transverse processes of the mastodont specimens are incomplete but were apparently
more horizontal than those of Prodeinotherium and are sited higher on the neuro-
pophysis, extending for some 4 cms above the dorsal edge of the axis facet. The
odontoid fossa of the mastodont atlas is also incomplete but appears to rise more
steeply anteriorly and to be more concave than that of Prodeinotherium.
The atlas attributed to Prodeinotheriwm does not articulate well with a Prodeino-
therium axis vertebra (6424 : 79) from an unknown locality at Gebel Zelten, mainly
due to differences in the articular facets. The general proportions of the occluding
facets and odontoid fossa and processes are however sufficiently similar for the
poor articulation to be attributed to variation between individual specimens.
TABLE 7
Atlas vertebva measurements
Parameter Prodeinotherium Mastodon Loxodonta
hobleyi angustidens africana
6418315 M.26656 1961.8.9.82
Maximum length 10°4 10.7-+-* 78
Maximum height 17-2 20:0 17'9
Width at transverse process 28-7 24°3+ 30°74
Width at occipital facets 18°4 20°7 Pita)
Width neural canal Or 7°3 8:0
Height neural canal 5:0 33) 3°9
Width odontoid fossa 56 6°5 Ps
Height odontoid fossa 6-2 56 6:2
Width at atlas facets 16°7 16-3* 17°7
PRODEINOTHERIUM
Fic. 8. Prodeinotherium hobleyi atlas vertebra.
5 cm.
anterior view, B = posterior view. Scale =
A=
FROM GEBEL ZELTEN, LIBYA 321
————uc“K«
Sa
22
Fic. 9. Lowxodonta africana atlas vertebra.
A = anterior view, B = posterior view. Scale = 5 cm.
322 PRODEINOTHERIUM
Axis vertebra. The sole deinotheriid axis from Gebel Zelten (6424 : 79) is incomplete
dorsally and ventrally (text-fig. 10). In overall size it is slightly smaller than the
axis vertebra of the extant African elephant (text-fig. 11) but the odontoid process,
although eroded, is longer and stouter in Prodeinotheriwm. The transverse processes
are broken at their bases and the entire ventral half of the centrum is missing. The
cephalic articular surfaces are oval, extending posteroventrally from above the odon-
toid process. It is not clear whether, as in Loxodonta, the articular facets extend to
meet below the odontoid process because they are incomplete. The neural canal
is circular in transverse section. The neural archisnotched anteriorly and posteriorly
for the passage of the second and third cervical spinal nerves, the posterior notch
being the deeper.
v/
('
lll
\
Fic. 10. Pyrodeinotherium hobleyi axis vertebra.
= anterior view, B = posterior view, C = right lateral view. Scale = 5 cm.
FROM GEBEL ZELTEN, LIBYA 323
The spine is the most diagnostic feature of the Prodeinotheriwm axis. In the
extant elephants ‘the neuropophyses blend together above and develop a thick
bifurcate spine before coalescing with the centrum’ (Owen, 1866 : 547). In the
Gebel Zelten axis the neuropophyses are less tall but longer anteroposteriorly than in
Loxodonta. The bifurcation of the spine is extreme, beginning at the anterior tip
of the spine and diverging at an angle of 55° (cf. 30° in Loxodonta). The twin crests
Fic. 11. Lowxodonta africana axis vertebra.
A = anterior view, B = posterior view, C = left lateral view. Scale = 5 cm.
324 PRODEINOTHERIUM
thus formed ascend backwards less steeply than in the elephant but reach farther
posteriorly and are separated by a U-shaped valley that widens posteriorly and
descends at an angle of nearly 30° from the horizontal. In Loxodonta this valley
appears only at the posterior portion of the spine and is horizontal. The post-
zygapophyses of the elephant axis are convex; the greater part of the articular
surface faces ventrolaterally but the dorsomedial corner faces posteriorly. Much
of the articular surfaces of the zygopophyses of the Gebel Zelten Prodeinotherium
axis are eroded but they are seen to be larger than in the elephant, to be concave
and to face ventrolaterally. A small triangular facet may be discerned along the
dorsomedial edge of, and at right angles to, the main articular surface but this may
have served for the attachment of muscles or ligament rather than for articulation.
Above the postzygapophyses much of the bone is missing but this region of the
spine must originally have been greatly swollen.
Although the neuropophyses are appreciably wider and longer than those of the
elephant, the centrum is only slightly longer. It is not until the relative areas of the
neural canals are taken into consideration that the axis of Prodeinotherium is seen
to be more massive and appreciably longer than that of the African elephant. The
larger surface of the neural spine affords a greater area of attachment for the rectus
capitus posterior major muscle, obliquus capitus inferior muscle, senuspinalis capitus
and semispinalis cervicus muscles. All four muscles serve to extend the head on the
neck or, acting unilaterally, to rotate or flex the head on the neck. The narrow
width of each diverging supraspinous crest suggests that the nuchal ligament was
inserted only on the posterior portion of the dorsal surface of the supraspinous crest.
The great width of the spine reflects the great width of the nuchal ligament that is in
turn reflected by the wider fossae for the nuchal ligament on the occiput of Prodeino-
therium than in elephantoids. The relatively low dorsoventral height of the spine
may reflect the elevated condition of the occipital condyles and/or the less elevated
head of deinotheres.
TABLE 8
Axis vertebva measurements
Parameter Prodeinotherium Loxodonia
. hobleyi africana
6424:79 1961.8.9.82
Maximum length 17°1 13°4
Maximum width 17°5-+ 17°5
Maximum height 17-7 24°6
Width at atlas facets 176+ 17-2
Width neural canal 5:2 6-2
Height neural canal 5:0 5°8
Width odontoid process 379+ 3°4
Height odontoid process 4°7 36
Maximum length neuropophysis 15:2 I1°9
Minimum length neuropophysis 5°5 36
Minimum width neuropophysis 15°3 TF,
Height neuropophysis above neural canal 76+ 8-1
Length postzygopophysis 52+ 5°5
Width postzygopophysis 5255 Si
FROM GEBEL ZELTEN, LIBYA 325
Third cervical vertebva. An incomplete right neuropophysis of a third cervical
vertebra M.266671 was associated with immature Prodeinotherium skeletal elements
from Gebel Zelten site D. The specimen (text-fig. 12) is not well preserved but
incorporates the region between the neural spine and the vertebrarterial canal.
The neural spine is very low and could have been elevated only slightly above the
valley separating the two wings of the axis spine. The neural canal is compressed
dorsoventrally. The neural arch is longer anteroposteriorly but less stout than in
Loxodonia. The prezygapophysis is convex and faces anterolaterally. The post-
zygapophysis is concave and faces ventrally. Both zygapophyses are aligned more
horizontally than in Loxodonta. The neural arch is notched anteriorly and posteriorly
for the passage of the spinal nerves, the posterior notch being deeper and extending
-
YA
(6
\
Wy
We
tll
Ww
Fic. 12. Prodeinotheyium hobleyi third cervical vertebra.
A = anterior view, B = posterior view, C = lateral view. Scale = 5 cm.
326 PRODEINOTHERIUM
to the dorsal edge of the vertebrarterial canal. The surface of the bone is not well
preserved and muscle attachment areas cannot be detected.
It is interesting that the articular surfaces of the zygapophyses have reversed
curvature when compared to the elephantoids in which the postzygapophysis and not
the prezygapophysis is convex. This is believed to have some bearing on the
direction of movement of the neck in elephantoids as opposed to deinotheres. The
centra of the cervical vertebrae of the Gebel Zelten deinothere shows some flattening
anteroposteriorly as in the elephantoids but the neuropophyses are considerably more
massive. Taking into consideration the diameter of the neural canal, the cervical
vertebrae of the Gebel Zelten Prodeinotherium were more elongate than those of
Loxodonta.
TABLE 9
Measurements of third cervical vertebra
Parameter Prodeinotherium Loxodonta
hobleyi afvicana
M.266671 1961.8.9.82
Maximum length neuropophysis WeWse Fal
Length neuropophysis at neural arch 371 2:8
Width transverse process above vertebrarterial canal 2:8 Ig
Width neural canal 70+ 6:8
Height neural canal 4°4 5°7
Height neural arch above centrum 5:9 9:0
Fic. 13. Loxodonta africana third cervical vertebra.
A = anterior view, B = posterior view. Scale = 5 cm.
FROM GEBEL ZELTEN, LIBYA 327
Rib One left rib was collected with the immature Prodeinotherium skeleton
from site D. Other rib fragments were noted but were too incomplete to warrant
collection. The head of the rib M.26667h is large, rounded and divided by a deep
transverse groove into two convex oval facets for articulation with two thoracic
vertebrae. The transverse groove is widest medially, and the cranial articulation
facet is larger than the caudal. The neck is short and a little wider than the head.
On the dorsal edge of the neck is an oval, slightly convex tubercule that articulates
above the caudal facet of the head. On the caudolateral side of the tubercule is a
rounded prominence, one fourth the size of the tubercle, that may have served for
attachment of the longissimus dorsi muscle. The shaft of the rib extends ventro-
laterally from the neck but flexes downwards at nearly 40° at the angle. The dorsal
surface of the angle bears an elongate prominence for the attachment of the
iliocostal muscle. On the cranial surface of the rib, between the head and the
angle, there are two elongate fossae. On the caudal surface a deep groove appears
midway between the head and angle near the lateral border of the shaft. A trans-
verse section of the shaft proximal to the angle resembles the shape of an hourglass;
distal to the angle it is concavoconvex (concave caudally). Distal to the caudal
depression the cranial face turns laterally and the transverse section becomes oval
and progressively flattened towards the end of the rib. Extending over the cranial
surface of the shaft from the angle are a number of irregular prominences that serve
for the attachment of various parts of the serratus posticus muscles.
The bicipital head of the rib suggests that it was articulated towards the front
of the rib cage.
TABLE I0
Measurement of Prodeinotherium vib M.26667h
Parameter Dimension
Length head . ; ; é . 2 j 53
Width head. j : é : : : 5:0
Width neck. : ; : : : : 5:2
Length neck . F ; : s : 3 Br7
Length neck to angle 4 ; b 5 : 10°7
Length at angle ‘ : é : ‘ . 6:8
Breadth at angle. c é : : : 2°
Discussion. In mammals generally, only one muscle is attached to the anterior
face of the arch of the atlas vertebra. This muscle, variously recorded as the rectus
capitis dorsalis minor (Sisson & Grossman, 1947), rectus posterior minor (Jayne, 1898),
and rectus capitis lateralis posterior minor (Shindo & Mori, 1957) inserts on the
occiput lateral to the nuchal fossa and serves to extend the head on the neck. The
reduced width of the neural spine of the deinothere atlas compared to that of
elephantoids is due, at least partly, to the fact that the sites of origin of the rectus
muscles are closer together and face upwards and outwards rather than forwards.
The line of action of the muscles is thus less horizontal and more vertically aligned
than in Mastodon or the elephants. The change in line of action of the musculature
328 PRODEINOTHERIUM
reflects the different geometry of the occiput of Prodeinotherium which has a more
dorsally placed nuchal fossa than has that of the elephantoids. The more posteriorly
inclined transverse processes may be partly due to a more vertical alignment of the
muscles extending from the transverse process to the occiput, or may reflect the
importance of muscles originating from the ventral surface of the transverse process
to insert on the basicranium and paroccipital process.
The Gebel Zelten Prodeinotherium axis differs in many features from that of the
elephantoids, the more striking differences being associated with the development of
the neuropophyses. The muscles attaching thereto serve to move both the neck on
the thorax and the head on the neck. The cross sectional area of the anterior
opening of the neural canal of the axis vertebra of Loxodonta (ca. 4000 mm?) is half
as large again as that of the Gebel Zelten deinothere axis (ca. 2600 mm2) and if this
area may be taken to suggest the relative size of these two mammals (cf. Radinsky,
1967) then the Gebel Zelten deinothere, although smaller than the African elephant,
has much larger and longer neuropophyses than the living animal. It is obvious
therefore that the axis of Prodeinotherium plays a more important role in the move,
ment of the head and neck than that of Loxodonta.
Five major muscles are attached to the axis spine. The rectus capitis posterior
major, obliquus capitis inferior, and semispinalis capitalis originate on the axis and
move the head about the atlantooccipital joint; the scalenus and semispinalis
cervicus insert on the axis and move the head on the thorax. The lamellar portion
of the nuchal ligament also inserts on the spine of the axis. Three muscles possess
considerably larger attachment areas on the spine of the Prodeinotherium axis than
on that of the elephant. These are the obliquus capitis inferior, the rectus capitis
posterior minor and the semispinalis cervicus. The former rotates, extends and
fixes the atlantooccipital joint (and thereby the head), while the rectus capitis
extends the head and the semispinalis cervicus extends the neck. All three are
attached to the upper part of the neuropophysis. That the spine is less elevated
than in Loxodonta suggests that its function as a lever for moving the head on the
thorax is less important than its capacity to move the head on the neck. The
external walls of the spine are orientated more laterally than in Loxodonta thus
affording a more nearly perpendicular line of action for the muscles moving the head
and thereby enhancing the mechanical efficiency of these muscles. The sagittal
length and divergent nature of the neural spine of the axis also affords a greater
area of attachment for the digitation of the lamellar portion of the nuchal
ligament.
The postzygapophyses of the axis and zygapophyses of the succeeding cervical
vertebrae of deinotheres are more horizontally aligned than in the elephantoids.
The curvature differs also—the postzygapophyses of deinotheres are concave and the
prezygapophyses convex in direct contrast to the condition normally found in the
elephantoids. Thus while the zygapophyses of the anterior cervical vertebrae of
elephantoids permit an upward flexure of the neck, those of Deinotherium and
Prodeinotherium flex in the opposite direction. The differences in direction of
flexure may be related to the different ways in which the tusks function in deino-
theres and elephantoids.
FROM GEBEL ZELTEN, LIPYA 329
The centrum of the axis is slightly longer in Prodeinotherium than in Loxodonta
and is appreciably longer when the area of the neural canal is used as a basis for
comparison. The neural arch of the third cervical vertebra from Gebel Zelten is also
longer than its equivalent in Loxodonta and it is probable also that this was true of
the remaining vertebrae of the neck. Increase in the length of the neck amounts to
a decrease in the lever arm of the musculature supporting the head on the neck and
moving the head on the thorax. The decrease in lever arm of these muscles reduces
their mechanical efficiency, and a larger and stronger musculature is therefore
required to support the head on the neck. This is reflected by the large area of
insertion of the semispinalis cervicus muscles in Prodeinotherium. Without access
to the cervical vertebrae of Deinotherium one cannot deduce whether the relatively
long neck of the Gebel Zelten deinotheres is a primitive feature, the neck becoming
shorter and mechanically more efficient in later forms, or whether it was a functional
adaptation to permit the head to be nearer the ground and thereby enhancing the
potential use of the tusks for feeding.
The thoracic vertebrae of Deinotheriwm are incompletely known, the anterior
thoracics being best known from the skeleton of D. ‘gigantissimum’ (= D. giganteum)
from the Pliocene of Manzati, Rumania. The neural spines of these vertebrae are
inclined steeply posteriorly. Slyjper (1946) pointed out that the neural spines act
as levers to transmit muscular force to the centra. The force applied determines
the height and direction of the spines. Increase in total body size is accompanied
by a relative increase in muscle strength, the size and shape of the spines being
therefore dependent also on body size, weight of the head, and length of the neck.
The most favourable direction of the neural spine is perpendicular to the muscular
force acting on the spine, but intermediate directions are achieved when two or
more forces act in different directions upon one spine. In the elephant three main
forces are applied to the neural spine of the anterior thoracic vertebrae by the
nuchal ligament, the sfinalis dorsi, and the multifidus spinae muscles. Tension in
the nuchal ligament exerts a force pulling the spines anteriorly whilst contraction of
the muscles pulls the spines posteroventrally.
The funicular portion of the nuchal ligament exerts most of its force on the anterior
thoracic vertebrae. The anterior thoracic neural spines of Loxodonta are more
perpendicular to the centra than those of Elephas, although in both animals the
spines slope backwards. This indicates that the nuchal ligament of Loxodonta
is more horizontally aligned and, conversely, that the head of Elephas is more
elevated with regard to the anterior thoracic vertebrae. Because the nuchal ligament
functions most effectively when the lever arm (neural spine) is perpendicular to the
line of action of the ligament one may deduce that mammals whose heads are
sited above the withers will have backward pointing spines whereas those mammals
whose skulls are on the same horizontal plane as the withers will have upright
spines. The few mammals with skulls held below the level of the withers will also
have vertical neural spines but because the line of action of the ligament will not be
perpendicular to the spines the lever arm of the ligament must increase to compensate
for reduction in efficiency. The neural spines of these mammals will therefore be
330 PRODEINOTHERIUM
correspondingly longer than those of mammals in which the skull and shoulders are
level.
The vertebral column of D. ‘gigantissimum’ was restored and mounted by
de Pauw (Stefanescu, 1907; Pl. 1). The neural spines of thoracic vertebrae 2-5 are
far more steeply inclined than in those of either of the extant elephants. The
original specimens from Manzati, Rumania, were incomplete and their exact length
cannot be determined. Backward inclination of these neural spines may be inter-
preted in two ways: either the head of the deinothere was carried above the shoulders
(the backward inclination being influenced by the line of action of the nuchal liga-
ment) or the vertebral column was rigid (the position of the spines being determined
by the multifidus muscles). Total length as well as the angle of inclination of the
neural spines must be considered before either alternative can be fully investigated.
With regard to the rigidity of the vertebral column, none of the extant graviportal
mammals can flex their lumbar and thoracic vertebrae. In view of the large
size of the deinotheres it seems likely that the vertebral column would have been
similarly modified and perhaps the extreme spinal inclination suggests extreme
inflexibility.
Appendicular skeleton
(Text-figs 14-16)
MATERIAL REFERRED. M.26667a, distal end of fibula; M.26667b, right ulna;
M.26667c, proximal end of right humerus; M.26667d, right femur; M.26667e, left
ulna; M.26667f, scapula; M.26667¢, left innominate; M.26667j, proximal and distal
end of left radius; M.26667k, distal end of right humerus; M.26667]1, distal end of left
humerus; 6405 : 98, left cuneiform.
DESCRIPTION. Scapula. An almost complete right scapula (M.26667f, text-fig.
14) was associated with the immature skull from Gebel Zelten site D. The left
scapula was also present but was partly overlain by skull 6404 : 44 and was destroyed
during excavation of the latter. The fragility of scapula M.26667f resulted in the
collapse of the vertebral border and cranial and caudal angles during transportation
from the site but these portions were subsequently restored with the aid of photo-
graphs taken at the site.
The articular fossa is broad and concave. The scapular tuberosity is poorly
developed but the coronoid process is well defined. The scapular notch is deeply
indented. The spine divides the lateral surface of the scapula into supraspinous
and infraspinous fossae of similar proportions to those of Elephas maximus. The
spine attains its highest elevation in its ventral third at the point of divergence of
the acromion and metacromion. From here it descends gradually to the neck and
caudal angle. Near the neck the cranial surface of the spine is concave but becomes
convex near the cranial angle. The reverse is true of the caudal surface.
Although the scapula is similar in size to that of the Indian elephant, the caudal
angle is sited dorsal to the point of divergence of the acromion and metacromion.
In the Indian elephant the base of the acromion is level with the caudal angle. The
FROM GEBEL ZELTEN, LIBYA 331
B
Fic. 14. Right scapula of (A) Pyodeinotherium hobleyi and (B) Elephas maximus. Scale = 5 cm.
D
332 PRODEINOTHERIUM
head of the Prodeinotherium scapula is wider and less concave than that of the
elephant and the spine ascends more gently from the neck than in Elephas. The
scapular notch is deeper and the scapular tuberosity less pronounced than in
Elephas.
TABLE II
Scapula measurements
Parameter Prodeinotherium Elephas
hobley1 maximus
M.26667j U.B.20189
Length head to vertebral angle 52+ 62
Transverse width at caudal angle 35 42
Length caudal angle above head 25 15
Length caudal angle behind spine 26 30
Length spine 40 44
Length acromion 9 17
Length metacromion 6 16
Maximum width spine behind metacromion 5 3-7,
Width head (straight) 1533 1453
Width head (curved) 17 16
Depth head 10* 8-3
Humerus. The distal extremities of a right and left humerus M.26667k and
M.266671) were associated with the immature skull 6404 : 44. They are complete
from the condyles to the deltoid tuberosity and proximal end of the lateral supra-
condyle ridge. This part of the bone is slightly shorter proximodistally than the
equivalent part of the humerus of the Indian elephant but is appreciably wider and
deeper suggesting a larger area of origin of the many flexor and extensor muscles
that arise from this portion of the humerus.
In cranial aspect the distal end of the musculospiral groove is seen to curve
laterally towards the proximal end of the lateral supracondylar ridge. This is a
strong ridge, straighter and more salient laterally than its equivalent in the elephant.
Asin Elephas the lateral epicondyle does not project very far laterally but it is stronger
cranially in Prodeinotherium. Medial to the lateral epicondyle there is a small but
deep fossa separated by a low ridge from the very wide but shallow coronoid fossa.
The median epicondyle of the trochlea is larger than the lateral, the disparity in size
being more pronounced than in Elephas.
In caudal aspect the lateral and medial epicondyloid ridges are considerably wider
than their counterparts in Elephas, as are also the epicondyles. Thus, despite the
great overall width of the distal portion of this bone, the olecranon fossa is con-
siderably narrower than in Elephas. In Elephas the epiphysial suture between the
median epicondyle and the condyle is aligned diagonally whereas in Prodeinotherium
it runs medially, thereby increasing the area of the condyle. The anconeal process
of the Prvodeinotherium humerus is narrower but no more salient than that of
Elephas.
FROM GEBEL ZELTEN, LIBYA 333
Fic.15 Fic. 16
Fic. 15. Radius of Prodeinotherium hobleyi and Mastodon angustidens.
A = P. hobleyi left radius (M.15429), cranial view
B= M. angustidens right radius (6412 : 166), cranial view
C = P. hobleyi head of left radius (6404 : 41), dorsal view
D = P. hobleyi head of left radius (6404 : 41), caudal view
Fic. 16. Radius of Prodeinotherium hobleyi and Mastodon angustidens.
A = P. hobleyi left radius (M.15429), caudal view
B = P. hobleyi left radius (M.15429), distal view
C = M. angustidens right radius (6412 : 166), caudal view
D = M. angustidens right radius (6412 : 166), distal view
Scale = 5 cm.
Dt
334 PRODEINOTHERIUM
TABLE 12
Humerus measurements
Parameter Prodeinotherium Elephas
hobley1 maximus
M.26667k M.266671 U.B.20189
Width at dorsal edge of lateral epicondyle 18-5 183 14°6
Length from lateral epicondyle to distal end of condyle 19°7 19°7 14:6
Width at median epicondyle 20°5 20:9 19:0
Length from median epicondyle to distal end of
median condyle 14'0 13°4 10°5
Width of articular surfaces on condyles 16:2 16-7 1593
Depth at median epicondyle 14:2 14°8 13°4
Minimum width shaft above lateral epicondyle 974 1O'L 77
Minimum depth shaft above lateral epicondyle 9:2 9:3 7:6
Depth of articular surface of median condyle 12:0 10:2 I2'I
Depth of articular surface of lateral condyle 10-2 10:2 II‘O
Radius. Only two proboscidean radii have been collected from Gebel Zelten—
the head and neck of a left radius of P. hobleyi (text-fig. 15C and D; M.26667j) and a
nearly complete right radius (M.26654; text-figs 15 and 16) of Mastodon angustidens.
Two left proboscidean radii from East African Miocene localities are present in the
collections of the British Museum (Natural History). That from Rusinga (B.M.
(N.H.)M.21872) was attributed to P. hobleyi by MacInnes (1942) but belongs to
Mastodon angustidens. The other specimen (B.M.(N.H.)M.15429) is from Maboko
Island and belongs to Prodeinotherium (text-figs 15 and 16).
The proximal articular surface of the radius of Mastodon angustidens is trapezoidal
in outline, with the medial side longer than the lateral. The capitular depression
for articulation with the humerus is not large and extends through the centre of the
articular surface parallel with the caudal and cranial edges. At right angles and
distal to the capitular depression, articulation facets for the ulna occur on the medial
and caudal sides, the medial facet being the smaller. Beneath the neck and caudal
edge of the median articulation facet for the ulna, the radius is swollen to produce
a radial tuberosity for the insertion of the biceps brachii muscle. This tuberosity
is more pronounced than in Prodeinotherium or Elephas. Immediately below the
radial tuberosity the shaft is approximately rectangular in transverse section, the
caudal and cranial edges being the longest and the lateral edge being slightly shorter
than the medial. Distally the medial and lateral edges diverge and the shaft is
compressed craniocaudally but whereas the shaft of M. angustidens retains this
shape throughout its length, that of Prodeinotherium becomes more equant in its
distal half.
Halfway down the shaft of M. angustidens the medial edge of the cranial surface
is flattened for the insertion of the pronator teres muscle. The interosseous crest
extends down the medial side of the caudal face from beneath the radial tuberosity.
Distal to the insertion of the pronator teres, and at the proximal end of the distal
third of the shaft, the interosseous crest diverges, a strong ridge extending towards
the medial edge of the epiphysis and a weaker ridge towards the lateral. Lateral
FROM GEBEL ZELTEN, LIBYA 335
to the interosseous crest is a deep depression extending from the radial tuberosity
to the distal fourth of the bone.
The radius of Prodeinotherium is smaller than that of M. angustidens and is less
straight proximodistally and craniocaudally. The radial tuberosity is comparatively
less well developed and the interosseous ridge diverges halfway down the shaft.
The distal extremity is less wide and deeper craniocaudally than in M. angustidens.
The radii of both M. angustidens and P. hobleyi are of similar length to the radius of
Elephas maximus but the shafts are much wider and the entire bone stouter.
TABLE 13
Radius measurements
Parameter Prodeinotherium Mastodon Elephas
hobleyi angustidens maximus
M.26667} M.15429 M.26654 M.22872 U.B.20189
Length head to
distal epiphysis — 48 63 59 43
Width proximal
epiphysis 5°8 50+ 7:0 6:5 4°6
Depth proximal
epiphysis 9:0+ 8-5 10°3 85+ 8-5
Width at distal
epiphysis a 10-6 13°7 T4°5 Io-l
Depth at distal
epiphysis = 8-3+ Viet 6-9 6°5
Width at centre
of shaft == 54+ 1 Te 43
Depth at centre
of shaft — 471 375 3°7 371
Ulna. Right and left ulnae were associated with skull 6404 : 44 but the shaft of
the left ulna (M.26667e) is shattered and only the right ulna M.26667b could be
prepared completely. The right ulna lacks the distal and olecranon epiphyses and
also the proximal portion of the semilunar notch including the anconeal process.
The bone is approximately equal in length to that of the extant Indian elephant.
The proximal extremity is the stoutest part of the bone and the olecranon, even with-
out the epiphysis, is larger than the olecranon plus epiphysis of the Indian elephant.
On the posterolateral side of the semilunar notch is a large scar marking the insertion
of the anconeous muscle. A coronoid process projects cranially on either side of the
radial notch, the medial process being the larger. The ulnar tuberosity, located on
the craniomedial side of the ulna and separated from the medial coronoid process
by a large depression, is less well developed for the insertion of the brachialis muscle
than is the equivalent tuberosity in the elephant.
The middle third of the ulna shaft is triangular in transverse section with cranial,
medial and lateral faces but reverts to a pentagonal cross-section at the distal
epiphysis. The interosseous ridge extends distally along the medial edge beneath
the ulnar tuberosity but dies out at the proximal edge of the distal third of the
shaft. Flat surfaces on the caudal aspect of the bone mark the areas of attachment
336 PRODEINOTHERIUM
of the external pollicis longus, extensor communis and flexor digitorum profundus
muscles and the ligament humeroantibrachialis. These are all developed much as
in the Indian elephant. The shaft is of similar length to that of the Indian elephant
but is considerably deeper and wider.
TABLE 14
Ulna measurements
Parameter Prodeinotherium Elephas
hobleyi maximus
M.26667b M.26667e U.B.20189
Length from olecranon to distal epiphysis 61* 53* 63
Width olecranon 13°6 12-2 15:2
Depth olecranon below semilunar notch HOF 12-2 13-2
Width semilunar notch 7-0 = 6-7
Width coronoid processes 19°4 a 15°4
Width distal end of shaft I1°8 10:9 12:2
Depth distal end of shaft 13°7 I4'7 II‘g
Width midpoint of shaft 8-4 = 673
Depth midpoint of shaft 7:0 — 6-3
Carpal bones. A small number of carpal bones have been collected from Gebel
Zelten sites. MacInnes (1942) reported several carpals from Rusinga Island, Kenya,
and these he attributed to P. hobleyi. Examination and comparison of the East
African and Gebel Zelten material has revealed that only three of MacInnes’ speci-
mens may definitely be attributed to Prodeinotherium—an incomplete left cuneiform
(M.21875), a right magnum (M.21877) and a left unciform (M.21882). These are
redescribed with the Gebel Zelten material.
Cuneiform. The left cuneiform from Gebel Zelten (6405 : 98) is better preserved
than that from Rusinga. The proximal surface that articulates with the ulna bears
two articulation facets (a median rectangular facet and a lateral triangular facet)
separated by a sagittal ridge. In Elephas and Mastodon the proximal articular
surface of the cuneiform is saddle-shaped with a transverse depression through the
centre. The distal surface of the deinothere cuneiform bears a triangular concavo-
convex facet for articulation with the unciform, much as in the elephant or mastodon.
On the caudal border there is a triangular facet for articulation with the pisiform,
the hypoteneuse of the triangle being bounded by a deep groove extending down the
large posterolateral process from the caudal junction of the proximal facets. The
distal surface of the posterolateral process of the Gebel Zelten cuneiform is eroded
but, as in Elephas, it bears a facet for articulation with the fifth metacarpal. The
medial surface bears a cranial and (larger) distal facet for articulation with the
lunar.
FROM GEBEL ZELTEN, LIBYA 337
TABLE 15
Cuneiform measurements
Parameter P. hobleyi D. giganteum = M. angustidens E. maximus
6405 :08 (Howenegg) M.21874 U.B.20189
Maximum length 8:5 15°6 8-6 7-9
Maximum width Tez 19°3 Iro+ II‘
Depth cranial face 48 9°3 5°0 4°6
Depth caudal face 5:0 8-1 5°3 5°1*
Length proximal facet 673 I1'9 5:9 5°5*
Width proximal facet or 14-7 84+ 77
Length distal facet 71 14'8 8-2 6:5
Width distal facet 75 13°6 8-0 75
Length posterolateral process 36 7:0 4:0 3:9
Width posterolateral
process 5°0 6-3 — 3°5
Depth posterolateral
process 76 9°5 — 7a
Lunar. An incomplete lunar (M.21876), identified by MacInnes (1942) as the left
lunar of Prodeinotherium hobleyi is almost certainly mastodont. However the right
lunar of both P. hobleyi and Mastodon angustidens are present in the collections of the
Centre for Prehistory and Palaeontology, Nairobi. The Prodeinotherium lunar
(KNM-RU 12091) is triangular and almost complete although the lateral edges are
somewhat irregularly preserved. The facet articulating with the radius occupies
nearly the whole of the proximal surface and extends nearly to the caudal edge of the
bone. In contrast, that of Mastodon terminates more cranially. The radial facet
is convex in the anteroexternal corner but concave over the remainder of the cranial
surface. The ulnar facet at the anteroexternal corner is rhomboid and not triangular
as in Mastodon. The magnum facet on the distal surface is convex cranially and
concave caudally.
TABLE 16
Lunar measurements
Parameter P. hobleyi D. giganteum M.angustidens E. maximus
KNM-RU1i291 (H6wenegg) KNM-RU 1260 U.B. 20189
Maximum length 8-7 16:0 10°3 8-1
Maximum width 8-7 15°4 10°8 8-1
Maximum depth 5°8 10:8 71 53
Length radial facet 73 11-4 75 6-7
Width radial facet 73 13°6 8-5 6:0
Length magnum facet 707 139 9:0 71
Width magnum facet 8-0 1374 8-6 Tez
Length ulnar facet 4-2 7:0 5°4 3°2
Width ulnar facet 3°55 4°3 4:0 2:9
Magnum. The right Prodeinotherium magnum from Rusinga (M.21877) is
incompletely preserved but is more complete than a left magnum of Mastodon
angustidens from Gebel Zelten (M.26652) that provides a basis for comparison.
In Prodeinotheriwm the proximal surface of the magnum articulates with the lunar
338 PRODEINOTHERIUM
and is L-shaped, a postero-medial projection (the foot of the L) standing proud. In
Mastodon or Elephas this projection merges gradually with the body and is relatively
less distinct. The proximal facet of the deinothere magnum is divided into two
portions—an almost flat cranial surface for articulation with the cranial part of the
lunar and a convex caudal region that occludes with the caudal part of the lunar
and scaphoid. The scaphoid articulates with the caudomedial projection of the
magnum. The distal articular facet is pear-shaped (stem posterior) and concave
for articulation with the third metacarpal. Four facets may be discerned on the
medial side. The cranioproximal, craniodistal and caudoproximal facets are tri-
angular and denote points of contact with the trapezoid. A mediodistal facet
extends anteriorly beneath the craniodistal facet to articulate with the second
metacarpal. The lateral side of the magnum adjoins the unciform and four facets
may be observed. Two large proximal facets are separated by a central groove in
contrast to the situation in the elephantoids in which the facets are contiguous.
The distal unciform facets on the magnum are also separated, in contrast to the
situation in Elephas, but the craniodistal facet is poorly developed as in Mastodon.
TABLE 17
Magnum measurements
Parameter P. hobleyi D. giganteum = M.angustidens E.maximus
M.21877 (H6wenegg) M.26652 U.B.20189
Maximum length 8°5 15°3 II‘2 8-7
Width cranial edge 5°7 10:2 6-9 For
Width caudal edge 6:8 10:0 6:8 71
Depth cranial edge 57 10°4 6-7 6-2
Depth caudal edge TRG) 13°4 oo 78
Length proximal facet 78 14°7 9:2 76
Width anterior edge
proximal facet 4°7 9-9 6:6 7X
Width posterior edge
proximal facet 6-6 10-0 6:8 71
Length distal facet 6:8 14°0 — 6-7
Width distal facet 4°5 6:2 53+ 4°5
Unciform. Three unciforms of Mastodon angustidens M.26653, M.26655, M.26657)
are known from Gebel Zelten and contrast strongly in several features with the left
Prodeinotherium unciform (M.21882); described by MacInnes (1942). In Prodeino-
thertum, the dorsal facet for articulation with the cuneiform is roughly triangular
inoutline. It reaches its maximum width at the anterior edge and tapers posteriorly.
The cranial and caudal portions are convex but are separated by a transverse concave
depression. The facet reaches its highest elevation at the caudomedial edge.
The medial surface of the Prodeinotherium unciform bears a large proximal and two
smaller distal facets for articulation with the magnum. Similar facets are also
exhibited by Elepbhas and Mastodon. Three facets are apparent on the distal
surface of the unciform. The medial facet is elongate, faces distomedially, is divided
into two by a concave transverse groove and supports the lateral edge of the third
metacarpal. The central facet is triangular (tapering posteriorly), is concave and
FROM GEBEL ZELTEN, LIBYA 339
articulates with the fourth metacarpal. It is separated from a larger rectangular,
convex facet for Metacarpal V by a sagittal groove that is deepest posteriorly.
Adjacent to the caudolateral edge of the McV facet but extending onto the proximal
surface of the bone is a small, triangular, concave facet for the pisiform.
In the Gebel Zelten mastodont unciforms the cuneiform facet approximates to a
rhomb and is longer than broad, the maximum breadth being achieved near the rear
of the bone. In Elephas this facet is appreciably wider than long, the caudal
edge being wider than the cranial, although the latter is relatively wider than in
Mastodon. The distal facets of the Gebel Zelten mastodont unciforms are arranged
much as in Prodeinotherium but are more elongate anteroposteriorly. These facets
are of similar length in Prodeinotherium and the Indian elephant but in the latter the
facets for McIV and McV are subequal in size and that for McIV is almost flat.
TABLE 18
Unciform measurements
Parameter P. hobleyi D. giganteum = M.angustidens E. maximus
M.21882 (Howenegg) M.26655 U.B.20189
Maximum length 9:2 16-4 10-6 8-4
Maximum width 10°3 17°5 11-6 973
Maximum height 8-9 10:9 8:9 6:6
Length cuneiform facet 7:6 14°8 8-3 6°5
Width cuneiform facet 9:0 16:2 8:5 Ql
Length McV facet 6:8 8-9 6:5 5°9
Width McV facet 4:2 53 5:0 39
Length McIV facet 6:7 12:2 7:0 5:2
Width McIV facet 4°7 8-7 6°5 54
Length McIII facet 51 1371 5°5 59
Width MclIII facet 1-6 3°6 I°3 2:0
Pelvis. One half of a pelvis (M.26667g) was associated with the immature deino-
there skeleton from Gebel Zelten site D. It may have belonged with the juvenile
skull (6404 : 44), the adult skull (6404 : 14) or neither. Unfortunately the pelvis
was shattered during transportation from the site.
Femur. A right Prodeinotheriwm femur (M.26667d) was associated with skull
6404: 44. It lacks both the head and the distal epiphysis but the shaft is approxi-
mately the same length as that of the extant Indian elephant. The shaft is com-
pressed craniocaudally and is virtually straight. The greater trochanter was
developed much asin Mastodon or Elephas. The lesser trochanter is poorly preserved.
The distal half of the cranial surface of the shaft bears a large ridge that becomes
more swollen distally as it extends towards the patellar surface. This ridge is
comparatively larger than in Elephas and suggests a larger area of attachment for
the vastus intermedius muscle. The caudal surface of the Prodeinotherium femur,
where preserved, strongly resembles that of the Indian elephant.
340 PRODEINOTHERIUM
TABLE I9
Femur measurements
Parameter Prodeinotherium Elephas
hobleytz maximus
M.26667d U.B.20189
Length shaft 78-7 80
Width shaft at proximal epiphysis 2575+ 22:9
Depth shaft at proximal epiphysis 12-24 10°7
Width shaft at distal epiphysis 20°8+ 16-4
Depth shaft at distal epiphysis 176+ 13°0
Width at centre of shaft II3 8-6
Depth at centre of shaft 9°6 65
Fibula. The distal third of the shaft of a Prodeinotheriuwm fibula (M.26667a) was
associated with skull 6404 : 44. The anterointernal border or interosseous ridge is
separated from the anteroexternal border in the proximal half of the specimen by a
narrow and flat anterior surface. The two borders converge in the distal half and
the external border fades out before reaching the distal extremity. The posterior
surface is round and broad but forms a sharp ridge immediately above the distal
extremity. The lateral face is convex outwards and presumably served for the
attachment of the peroneous brevis muscle. The medial face is concave and afforded
attachment for the flexor longus hallucis muscle. From the distal epiphysis the bone
decreases in width proximally, the change being gradual along the anterior edge but
with a fairly sharp genuflexion along the posterior edge just below the dorsal limit of
the specimen. From the shape of the posterior edge of the Gebel Zelten specimen
it would seem that the bone was originally about as long as that of the Indian
elephant. It differs from the latter, however, in that the distal extremity of the
Gebel Zelten specimen is half as wide again.
Tarsal bones. MacInnes (1942) described an astragalus, two naviculars and an
external cuneiform from Rusinga and attributed them to the taxon Prodeinotherium
hobleyi. Comparison of the astragalus (M.21880) with mastodont astragali from
Gebel Zelten and East Africa and with the astragalus facets of mastodont and
deinothere tibiae confirms MacInnes’ identification. The external cuneiform and
navicular differ both in size and in some morphological features from the equivalent
bones of the Indian elephant but are likely to have belonged to Mastodon angustidens.
Astragalus. The deinothere astragalus is more equant than that of Mastodon.
The dorsal and medial surfaces bear a rectangular convex facet for articulation with
the tibia. This facet is as long but less wide and more convex than that of
M. angustidens. The anterior border of the Prodeinotherium astragalus from Rusinga
is broken but the missing portion was probably of similar appearance to that of
Mastodon except that the groove on the craniomedial surface, separating the tibial
facet from the navicular facet, was less deep and less wide than in Mastodon. There
are three facets on the distal surface—a triangular and convex navicular facet and
two calcaneum facets. In Prodeinotherium the navicular facet extends to the
craniomedial border of the astragalus, the medial edge of the facet being concave.
The medial tip of the navicular facet of Mastodon is also concave but extends less
FROM GEBEL ZELTEN, LIBYA 341
far medially. The groove separating the navicular facet from the lateral calcaneal
facet is deeper but less wide than in Mastodon. The lateral calcaneal facet is tri-
angular and is flatter than that of Mastodon. The caudomedial calcaneum facet is
oval and concave. It is separated from the lateral facet by the continuation of the
same groove that separates the latter from the navicular facet. The medial calcaneal
facet is separated from the navicular facet only by a very shallow groove and extends
posteriorly onto the under surface of the massive posteromedial projection. On the
lateral surface of the astragalus the facet for articulation with the fibula appears
to be as long as, but much wider than, that of Mastodon.
TABLE 20
Astragalus measurements
Parameter P. hobleyi D. giganteum = M. angustidens E. maximus
M.21880 (H6wenegg) M.26658 U.B. 20189
Length lateral edge 79 — 74 7°4
Length medial edge I1l‘o 16:0 7:84 10:2
Maximum width Io:9+ 17°7 10-0* II°5
Depth medial edge pas 9°5 7X 7:9
Depth lateral edge 4°4 7:4 BZ 4°4
Length tibial facet 6-9 — 73+ 6-9
Width tibial facet FoR — 8-3 8-3
SKELETAL EVIDENCE ON THE LOCOMOTION OF DEINOTHERES The limb material
of the Gebel Zelten deinotheres is too fragmentary to permit a detailed functional
analysis. Nevertheless, comparison of these specimens with other Prodeinotheriwm
and Deinotherium remains affords some information about the locomotion of these
animals. The most important skeletal elements in this respect would appear to be
the scapula, femur, manus and pes.
Smith and Savage (1956) determined that in graviportal mammals the scapula
tends to be high and broad to afford sufficient surface area for the serratus and other
shoulder muscles. They noted that the scapula blade of graviportal mammals
is frequently expanded posteriorly to increase the mechanical advantage of the main
retractor muscle of the forelimb (the teres major) and that the spine is usually heavy
and may bear a posteriorly projecting flap in the region overlying the infraspinous
fossa (Rhinoceros, Brontops). Alternatively the spine may end in a large acromion
(Loxodonta, Uintatherium). The trapezius muscle originates on the spine and
metacromion, the deltoid on the acromion. Both these muscles aid in abducting
the forelimb.
Compared with the scapula of Elepbhas maximus, the scapula of Prodeinotherium
hobleyi (text-fig. 14) is less tall (51 v 62 cm) and less broad (25 v 42cm). The caudal
angle is sited more dorsally above the head of the scapula (25 v 15 cm) and is less
elongate posteroventrally from the spine (26 v 30 cm). Because the caudal angle of
the scapula is sited more dorsally in Prodeinotherium than in Elephas, the mechanical
advantage of the teres muscle is thereby decreased. Smith and Savage (1956)
have shown that the mechanical advantage of the teres muscle is equivalent to the
length of the moment arm of the teres muscle divided by the perpendicular distance
342 PRODEINOTHERIUM
from the glenoid to the ground. The mechanical advantage of the teres muscle of
the Indian elephant is nearly 1/13 but that of P. hobleyi is 1/16.
The supraspinous fossa of Prodeinotherium hobleyi is relatively smaller than that of
Elephas, and the scapular spine rises less abruptly and more dorsally. An acromion
process is developed on the spine of the P. hobley: scapula and is sited more dorsally
than in Elephas. A metacromion is also present, sited level with the base of the
acromion and forming a flap overhanging the infraspinous fossa. Both acromion
and metacromion are reduced in comparison with those of the Indian elephant.
Behind the metacromion the spine of P. hobleyi is broader and more robust than in
the Indian elephant.
The supraspinous fossa of D. gigantewm from Valladolid, Spain, is even more
reduced than that of the Gebel Zelten deinothere. The caudal angle of the Deino-
therium scapula is less posteroventrally produced than in the extant elephants and,
significantly, there are no processes diverging from the relatively slender spine.
The absence of acromion and metacromion was also noted by Stefanescu (1899) on
the scapula of D. ‘gigantissimum’ from Manzati, Rumania. Reduction of the spine
and supraspinous fossa, and the absence of acromion and metacromion would in
artiodactyls be regarded as evidence of cursorial modification (Smith & Savage, 1956).
The retention of a rigid body axis is essential in animals of elephantine bulk in
order to support the weight of the body. The method of rapid locomotion is thus
limited to a fast amble or trot. The more rapid action of the teres muscle of deino-
theres when taken in conjunction with other features of the scapula does, however,
suggest that Deinotherium was potentially capable of more rapid locomotion than the
elephantoids. Such an interpretation is supported by other features of the appendi-
cular skeleton and tallies with Tobein’s (1962) interpretation of the function of the
Deinotherium manus and pes. Prodeinotheriwm, although more typically graviportal
than Deinotherium, shows some signs of cursorial adaptation.
The humerus of the Gebel Zelten Prodeinotherium must have been slightly shorter
than that of the Indian elephant but was still appreciably longer than the ulna.
The shaft of the humerus is compressed laterally and elongated anteroposteriorly in
direct contrast to the condition in Elephas. The lateral epicondyle is shorter than in
Elephas but projects farther laterally. The condyles of the distal epiphysis have a
greater degree of curvature in Prodeinotherium than in Elephas suggesting that
the ulna of P. hobleyi may have been capable of being flexed through a greater angle.
The ulna of the Gebel Zelten deinothere is of similar length to that of the Indian
elephant but is proportionately stouter. As in the elephantoids the nearly vertical
alignment of the forelimb requires an olecranon process that diverges at right
angles from the shaft. There is no evidence to suggest that the ulna was fused to
the radius at its distal end although fusion at this point would seem likely. The distal
epiphyses of both the ulna and radius were proportionately more massive in Deino-
therium than in Prodeinotherium.
The radius of Prodeinotherium is best known from Maboko Island, Kenya, that
from Gebel Zelten being represented only by the head. Prodeinotheriwm radius
M.15429 is again about the size of that of Elephas maximus. It is less straight
proximo-distally and craniocaudally than in the elephantoid proboscideans. The
FROM GEBEL ZELTEN, LIBYA 343
radial tuberosity is less well developed than in Mastodon angustidens and the interos-
seous ridge diverges halfway down the shaft. The radius shaft of both Mastodon
angustidens and P. hobleyi is relatively wide laterally and compressed craniocaudally,
but, whereas the mastodont shaft retains this shape throughout its length, that of
Prodeinotherium becomes progressively more equant distally.
Only the large carpal bones of Prodeinotheriwm have been examined. These are
serially arranged and are smaller but similar in shape to those of D. gigantewm from
Hoéwenegg, Germany (Tobein, 1962). The morphology of the proximal surface
of the cuneiform suggests that the medial malleolus of the ulna extended farther
distally in deinotheres than in elephantoids. No metacarpals have been collected
from Gebel Zelten but a limited amount of information about the metacarpals may
be gleaned from the distal row of carpal bones. In the unciform the facet for
McV is proportionately larger and less laterally orientated than in the Howenegg
Deinotherium specimen and the facet for McII1issmaller. The magnum of Prodeino-
therium bears a large facet for McIII and a smaller and more laterally aligned facet
for McII.
Ehik (1930) described metacarpal fragments of P. bacaricwm from Kotyhaza,
Hungary. These were laterally compressed, but less so than the metacarpals of
Deinotherium giganteum from Pikermi, Greece, described by Dietrich (1916). The
metacarpals of P. bavaricum are less elongate than those of D. gigantewm and resemble
more closely those of Elebhas. Whereas Dietrich (1916) and Tobein (1962) stated
that the first metacarpal of D. giganteum is sufficiently reduced to suggest functional
tetradactyly of the manus, Ehik alleges that in P. bavaricum the fifth metacarpal
shows the greatest reduction. The distal articulation surface of the fourth meta-
carpal of D. giganteum is concave but that of P. bavaricum is concavoconvex. Ehik
(1930 : 12) implies that the manus of P. bavaricum was more plantigrade than that of
D. giganteum.
The metacarpals of D. gigantewm are dolichopodous (Tobein, 1962). Whereas
the proximal epiphysis of the fourth metacarpal of D. gigantewm is orientated at
right angles to the longitudinal axis of the shaft of the bone, the proximal epiphysis
of McIV of P. bavaricum is aligned obliquely to the longitudinal axis of the shaft.
The digits of P. bavaricum must therefore have spread farther laterally, were probably
more uniform in size and are likely to have been less digitigrade than those of
D. gigantewm. The trapezium and trapezoid bones of Prodeinotherium have not been
examined and it is therefore impossible to say if the first metacarpal was reduced as
in D. giganteum.
The long bones of the hind limb of the deinotheres are similar in many respects
to those of the elephantoids. The head of the femur is vertically aligned and, as in
other graviportal mammals, the femur is longer than the tibia. Osborn (1929 : fig. 670)
formulated the general rule that, relative to the length of the tibia, the femur
of cursorial mammals shortens with increase in potential speed while the pes elements
become more elongate. The femur and tibia of Prodeinotherium are of comparable
lengths to those of Elephas maximus but the femur of D. gigantewm from Valladolid
is relatively shorter with respect to the tibial length. The tibia/femur ratio of
E. maximus is 0:55 whereas that of D. gigantewm is 0-71, indicating that the femur of
344 PRODEINOTHERIUM
D. giganteum is about 30% shorter than that of the extant Indian elephant.
Reduction of the first metatarsal in Deinotherium was noted by de Pauw (1908),
and the narrow elongate pes of D. gigantewm in conjunction with the dolichopody
of the digits convinced Tobein (1962) that D. gigantewm was capable of swifter
locomotion than its contemporary elephantoids. The reduction in length of the
Deinotherium femur would appear to support Tobein’s interpretation.
The equant tibial facet of the astragalus of all deinothere taxa from which this
bone is known is a distinctive character that reflects the comparatively narrow width
of the pes of deinotheres compared to that of elephantoids. The tibial facet of
Prodeinotherium astragali is more convex than in astragali of its contemporary
elephantoids but that of Deinotherium is flatter. The large posteromedial projection
from the Prodeinotheriwm astragalus is not present in either Mastodon or Elephas
and is reduced in Detnotherium. The surfaces of the Prodeinotherium astragalus
that articulate with the fibula and medial malleolus of the tibia are larger and more
vertically orientated than in Mastodon angustidens.
IV. CONCLUSIONS
The family Deinotheriidae is known from Africa, Europe and western Asia. All
representatives of this family are large. Features of the skull and dentition serve
to distinguish the deinotheres from the elephantoids and distinctive characters may
also be seen in the postcranial skeleton. On their dental characters the deinotheres
appear to be most closely related to the barytheres and moeritheres and may have
been derived from ancestral Barytherium stock during the early Tertiary.
The family Deinotheriidae is represented by two genera, the primitive Prodeino-
therium and the later and larger Deinotherium. The Gebel Zelten specimens are
assigned to Prodeinotherium hobleyi. They include the earliest deinothere skulls
yet recovered and the fine condition of preservation of the skulls permits their
use for reinterpreting the morphology of less well preserved Deinotheriwm skulls.
The Gebel Zelten specimens exhibit a number of primitive deinothere features
including a steeply downturned rostrum, narrow and more anterior external nares,
a skull roof that is proportionately wider and longer, less elevated occipital condyles,
shorter paroccipital processes and a more vertically inclined occiput.
The facial region of P. hobleyi was modified for the attachment of a proboscis as
was that of Deinotherium. The cranial region was adapted for greater movement of
the skull on and with the neck than in elephantoids. The most important modi-
fications enlarge the paroccipital processes and elevate the occipital condyles to
increase the downward thrust of the mandibular tusks. Reversed curvature of the
zygapophyses of the cervical vertebrae (compared to that of elephantoids) also aids
in the downward movement of the head.
In detail the dental characters render the deinotheres unique. Loss of the superior
pre-cheek teeth and retention of a single pair of downturned inferior tusks is a combi-
nation not exhibited by any other mammal. The main function of the tusks may
have been for self defence and feeding, both involving a violent downward thrust of
FROM GEBEL ZELTEN, LIBYA 345
the head. It would seem unlikely that the tusks were used for digging and their
main function in feeding was probably for clearing and stripping vegetation. Wear
facets on the anteromedial surface of the tusk tip in a number of deinothere tusks
would appear to support this interpretation. Lophodont cheek teeth have been
evolved in several mammalian groups but the trilophodont anterior molars of the
deinotheres are unique. The deciduous fourth premolar is also trilophodont and
served as a shearing tooth in contrast to the crushing function of the second and
third milk premolars. The anterior permanent molar initially serves as a shearing
tooth but after the posterior molars have been erupted it becomes part of the anterior
crushing battery. The second and third molars are bilophodont and persist as
shearing teeth until the lophs are removed by wear. The angle of shear changes from
nearly vertical in newly erupted teeth to almost horizontal in greatly worn teeth.
The shearing surface is maintained by thegosis. There is no increase in hypsodonty
or brachyodonty throughout the recorded history of the deinotheres, and the
length versus width parameters of the teeth of all deinotheriid taxa remain constant.
Deinothere vertebrae are unfortunately rare but the axis and atlas vertebrae are
known. They may readily be distinguished from those of the elephantoids and are
modified to provide large attachment areas for the muscles of the neck. The
zygapophyses of the cervical vertebrae have reversed curvature relative to those of
the elephantoids. The neural spines of the anterior thoracic vertebrae were ap-
parently longer and more posteriorly inclined than those of contemporary elephan-
toids. Differences from elephantoid structure may also be seen in the appendicular
skeleton, the scapula and foot bones being most distinctive, the long bones less so.
Advanced characters shown by Deinotherium include reduction of the scapular
spine, elongation of the carpals, tarsals and metapodials, elevation in posture of the
manus and pes and reduction of the first digit leading to the functional tetradactyly
of the feet. Many of the limb modifications point to a cursorial adaptation of
Deinotherium, but Prodeinotherium resembles Elephas in the size and proportions of
the long bones.
Prodeinotherium and Deinotherium are separated on the basis of size, minor modi-
fications of the dentition and characters of the skull and postcranial skeleton. Six
species are recognized: P. hobleyi, P. bavaricum, P. pentapotamiae, D. giganteum,
D. indicum and D. bozast. Each species is based primarily on geographic distribu-
tion. The conservative dental characters of deinotheres are of little use for specific
differentiation but the skulls and skeletal elements may prove more diagnostic
when these are better known.
Deinotheres must have originated in Africa during the early Tertiary but are
first known from the early Miocene. During the early Miocene Prodeinotherium
migrated from Africa via the Middle East (and perhaps the western Mediterranean)
into Eurasia. By the late Miocene Deinotheriumhadappeared. Possibly two species
evolved separately, D. giganteuwm from P. bavaricum in Western Europe and D.indicum
from P. pentapotamiae in western Asia. More probably Deinotheriwm was derived
from P. bavaricum and then spread eastwards. Deinotherium and Prodeinotherium
coexisted in Eurasia during the late Miocene. Pvodeinotheriwm had become extinct
in Eurasia by the end of the early Pliocene and Deinotheriwm by the middle Pliocene.
346 PRODEINOTHERIUM
P. hobleyi was present in East and North Africa during the early Miocene but was
thereafter restricted to East Africa. It survived into the Pliocene but is not known
to have coexisted with D. bozasit which had appeared by the beginning of the Pleisto-
cene. Whether D. bozasi evolved in situ or represented the southerly migration of
the Eurasian deinotheres cannot yet be determined but the former seems more likely
in view of the limited distribution of African deinotheres after the early Miocene.
Both Prodeinotherium and Deinotherium were browsing mammals that used their
tusks and probosces to gather food. The proboscis of Prodeinotherium was smaller,
and the skeleton lacked many of the cursorial adaptations of Deinotheriwm.
Prodeinotherium probably inhabited more densely vegetated regions while Deinotherium
frequented more open territory. There is no obvious successor to Deinotherium in
the middle or late Pleistocene of Africa. Whether it failed to adapt to changing
environmental conditions or was supplanted through competition from a group or
groups of smaller herbivorous mammals must remain a matter for conjecture.
V. REFERENCES
ANDREWS, C. W. i191t. On a new species of Dinotherium from British East Africa. Proc.
zool. Soc. London, Dec. 1911 : 943-945.
1921. A note on the skull of Dinotherium giganteum in the British Museum. Tb7d., Sept.
1921 : 525-534.
ARAMBOURG, C. 1934. Le Dinotherium des Gisements de l’'Omo. C. R. Soc. géol. Fr. Paris,
1934 : 86-87.
1961a. Prolibytherium magniert, un Vellericorne nouveau du Burdigalien de Libye.
Ibid, 1961, no. 3 : 61-62.
1961b. Note preliminaire sur quelques vertébres nouveaux du Burdigalien de Libye.
Ibid, 1961, no. 4 : 107-108.
1963a. Le genre Bunolistriodon Arambourg 1933. Bull. Soc. géol. Fr. Paris, (7) 5 : 903-
git.
1963b. Continental vertebrate faunas of the Tertiary of North Africa. In Howell, F. C. &
F. Bourliere (Edit) African ecology and human evolution. Chicago, 1963 : 55-64.
—— & P. MacnierR, 1961. Gisements de vertébres dans le bassin Tertaire de Syrte (Libye).
C. R. Acad. Sci. Paris, 252 : 1181-1183.
BERGouNIOUx, F. M. & F. CrouzELt. 1962a. Les Deinotherides d’Europe. Avnls. Paléont.
Paris, 48 : 13-56.
1962b. Les Deinotherides d’Espagne. Bull. Soc. géol. Fy. Paris, (7) 4 : 394-404.
1963. Migrations des Mastodontes Miocenes. C.R. Congr. nat. Soc. Sav., Sect. Sct.,
Paris, 87 : 867-872.
De BLAINVILLE, H. M. D. 1837. Note sur la téte de Dinotherium giganteum actuellement a
Paris. C.R. Acad. Sci. Paris, 4, no. 12 : 421-426.
Brives, A. 1919. Sur la decouvert d’une dent de Dinotherium dans la Sabliere du Djebel
Kouif pres Tebessa. Bull. Soc. Hist. nat. Afy. N. Alger., 10 : 90-93.
Craupius, M. 1865. Das Gehorlabyrinth von Dinotherium giganteum nebst Nemerkungen
iiber der Werth der Labyrinthformen fiir die Systematik der Saugerthiere. Paldeonto-
gvaphica, Stiittgart 13 : 65-74.
Deum, R. 1963. Paldontologische und geologische Unterschuchungen im Tertiar von
Pakistan: 3. Dinotherium in der Chinji-Stufe der Unteren Siwalik Schichten. Bayer.
Akad. Wiss. Wien, (n.s.) 114 : 1-34.
DietricH, W. O. 1916. Uber die Hand und Fuss von Dinotherium. Zeitschr. dt. geol. Ges.
Berlin, 68 : 1-44.
FROM GEBEL ZELTEN, LIBYA 347
Desio, A. 1935. Studi geologio sulla Cyrenaica, sul Deserto Libico, sulla Tripolitania e sul
Fezzan Orientali. Mission Scient. R. Acc. d'Italia a Cufra (1931). Roma, 1.
Doust, H. 1968. Palaeoenvivonment studies in the Miocene of Libya. Vol. 1 of unpub.
Ph.D. thesis, Imperial College, London.
Eutk, J. 1930. Pvodinotherium hungaricum N.g., N.sp., Geol. Hungarica Ser. Palaeont., fasc.
6, Jan. ’25 : I-24.
FaLconerR, H. 1845. Description of some fossil remains of Dinotherium, Giraffe and other
Mammalia, from the Gulf of Cambay, western coast of India. Q. Jl geol. Soc. Lond. 1 : 356—
37/2:
Graf, I. E. 1957. Die Prinzipien der Artbestimmung bei Dinotherium. Palaeontographica,
Stuttgart, Abt.A, 108 : 131-185.
Hamitton, W. R. 1973. The Lower Miocene ruminants of Gebel Zelten, Libya. Bull. Br.
Mus. nat. Hist. (Geol.), 21, 3 : 73-150
JAYNE, H. 1898. Mammalian anatomy part 1: the skeleton of the cat. Philadelphia, 816 pp.
Kaup, J. J. 1829. Deinotherium giganteum. Isis (XXII), Jena, 4 : 401-404.
Kuipstein, H. & Kaup, J. J. 1836. Beschreibung und Abbildungen von dem in Rheinhessen
aufgefundenen colossalen Schedel des Dinotherii gigantet mit geognostischen Mittheilungen uber
die knochenfuhrenden Bildungen des mittelrheinischen Tertiarbeckens. Darmstadt, 38 pp.
LasKAREV, V. 1944. Uber die Dinotherien reste aus Serbien. Neues Jb. Min. Monatsch,
Stuttgart, 1944, 2/3, abt.B : 67-77.
Macinnes, D. G. 1942. Miocene and post-Miocene Proboscidea from East Africa. Tvans.
zool. Soc. London, 25, part 2 : 33-106.
Maenigr, P. 1962. Etude géologique du gisement de vertébres du Gebel Zelten (Libye).
C.R. Soc. géol. Fy. Paris, 1962, fasc. 2 : 55-57.
1968. Etude lithostratigraphique du Miocene inferieure du Bassin de Syrte (Libye).
Proc. Comm. Medt. Neog. Strat; Geol. Bologna, 35 : 119-130.
von MEYER, H. 1833. Beitrage zur Petrefaktenkunde. Das Dinotherium Bavaricum, mit Ruck-
sicht auf die riesenmassigste fossile Thiergattung der Dinotherien uberhaupt, und auf die
Struktur der Mahlzahne in den Tapiren. Nova Acta Acad. Leop. Carol., Halle, 16 : 485-516.
Osporn, H. F. 1907. Evolution of mammalian molar teeth to and from the triangular type.
New York. 250 pp.
1929. The titanotheres of ancient Wyoming, Dakota and Nebraska. U.S. geol. Su.
Washington, Monograph 55, 2 : 703-953.
1942. Proboscidea, volume 2. American Museum of Natural History: 803-1675.
OwEN, R. 1866. On the anatomy of vertebrates, vol. II. London: 437-444.
PACHECO-HERNANDEZ, F. H. 1930. Fisiografia, geologia y paleontologia del territorio de
Valladolid. Comm. Invest. pal. y prehist., no. 37 (ser. pal. no. 9) : 1-192.
PALMER, R. W. 1924. On an incomplete skull of Dinotherium, with notes on the Indian
forms. Mem. geol. Surv. India, [Paleont. Indica], (n.s.) 7, mem. no. 4 : I-13.
DE Pauw, L. F. 1908. Trois mois au Musée de Bucharest ou quelques considerations sur le
Dinotherium de Manzati. Brussels. 35 pp.
Rapinsky, L. 1967. Relative brain size; a new measure. Science, Feb. 1967, 155, no. 3674 :
836-838.
SAHNI, M. R. & Tripatui, C. 1957. A new classification of the Indian deinotheres and
description of D. ovlovii sp. nov. Mem. geol. Surv. India, [Paleont. Indica], 33, no. 4 : 1-33.
SavaGE, R. J. G. 1967. Early Miocene mammal faunas of the Tethyan region. Syst. Assoc.
Publ. no. 7 : 247-282.
1973. Megistotherium, gigantic hyaenodont from Miocene of Gebel Zelten, Libya. Bull.
Br. Mus. nat. Hist. (Geol.), 22, 7 : 483-511
— & Hamitton, W.R. 1973. Introduction to the Miocene mammal faunas of Gebel Zelten.
Libya. Bull. Br. Mus. nat. Hist. (Geol.), 22, 8 : 513-527
—— & WuitE, M.E. 1965. Two mammal faunas from the early Tertiary of central Libya.
Proc. geol. Soc. Lond., no. 1623 : 89-01.
348 PRODEINOTHERIUM
SELLEY, R. C. 1966. The Miocene rocks of the Marada and Jebel Zelten area, central Libya;
a study of shoreline sedimentation. Guidebook Petr. Expl. Soc. Libya. 1966. 30 pp.
—— 1967. Paleocurrents and sediment transport in near shore sediments of the Sirte Basin,
Libya. J. Geol., Chicago, 75; no. 2 : 215-223.
1968. Near shore marine and continental sediments of the Sirte Basin, Libya. Proc.
geol. Soc. Lond., no. 1648 : 81-90.
SuHinpbo, T. & Mori, M. 1957. Musculature of the Indian Elephant Part III: Musculature of
the trunk, neck and head. Ak. Fol. Anat. Jap., 28 : 18-39.
Sisson, S. & GRossMAN, J. D. 1947. The anatomy of the domestic animals. 3rd edit. revised.
New York. 972 pp.
SLIJPER, E. J. 1946. Comparative biological-anatomical investigations on the vertebral
column and spinal musculature of mammals. Kon. Ned. Akad. Wet. Verh. (Tweedie
Sectie), 42, no. 5 : 1-128.
SmitH, J. MAYNARD & SAvaGE, R. J. G. 1956. Some locomotory adaptations in mammals.
J. Linn. Soc. London (zoo), 62, no. 288 : 603-622.
STEFANESCU, G. 1897. Dinotherium gigantissimum Stef. trouve a Munztac. Cong. geol. Int.
(7th session), 1897, St. Petersburg: cxc—cxci.
1907. Quelques mots sur le Dinotherium gigantissimum Stef. trouve a Munztac. Cong.
geol. Int. (10th session), 1906, Mexico, 1 : 427-430.
Stott, N. R. ef al. 1961. International Code of Zoological Nomenclature adopted by the
XVth International Congress of Zoology. London. 176 pp.
STROMER, E. 1938. Huftier-Reste aus dem unterspliocanen Flinzsande Munchens. Bayer.
Akad. Wiss. Wien, Abh. A, 44 : 1-39.
ToBEIN, H. 1962. Uber Carpus und Tarsus von Dionotherium giganteum Kaup. Palaeont.
Z., H. Schmidt-Festband, Nov. 1962 : 231-238.
J. M. Harris, B.Sc., Ph.D.
CENTRE FOR PREHISTORY
NAIROBI
East AFRICA
PLATE 1
Fic. a. Prodeinotherium hobleyi skull (M.26665), anterior view.
Fic. b. Prodeinotherium hobleyi skull (M.26665), dorsal view.
Bull. By. Mus, nat. Hist. (Geol.) 23, 5 PLATE 1
Ta
tb
IGN, 2
Fic. a. Prodeinotherium hobleyi skull (M.26665), left lateral view.
Fic. b. Prodeinotherium hobleyi skull (M.26665), right lateral view.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 5 IPILAN IIE 2
2b
E*
PLATE 3
Fic. a. Prodeinotherium hobleyi skull (M.26665), posterior view.
Fic. b. Prodeinotherium hobleyi skull (M.26665), ventral view.
Bull. By. Mus. nat. Hist. (Geol.) 23, 5
PLAT
Ie
3
PLATE 4
Fic. a. Prodeinotherium hobleyi mandible (6412 : 10), right lateral view.
Fic. b. Prodeinotherium hobleyi mandible (6412 : 10), dorsal view.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 5 PLATE 4
4a
TINIE, 5
Fic. a. Prodeinotherium hobleyi mandible (6404 : 13), dorsal view.
Fic. b. Prodeinotherium hobleyi right maxilla (6401 : 4) ventral view.
Bull. By. Mus. nat. Hist. (Geol.) 23, 5 PLATE 5
A LIST OF SUPPLEMENTS
TO THE GEOLOGICAL SERIES
OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya.
Pp. 213; 30 Plates; 2 Text-figures. 1965. 6.
. EL-Naccar, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper
Cretaceous—Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley,
Egypt, U.A.R. Pp. 291; 23 Plates; 18 Text-figures. 1966. {r10.
. Davey, R. J., Downtz, C., SARGEANT, W. A. S. & Wittiams, G. L. Studies on
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248; 28 Plates; 64 Text-
figures. 1966. £7.
. APPENDIX. Davey, R. J., Downig, C., SARGEANT, W. A. S. & WiLLIaMs, G. L.
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24.
1969. 8op.
. Evtiotr, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of the
Middle East. Pp. 111; 24 Plates; 17 Text-figures. 1968. £5.12.
. Ruopes, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni-
ferous) Conodont faunas, and their value in local and continental correlation.
Pp. 315; 31 Plates; 92 Text-figures. 1969. {11.
. Caitps, A. Upper Jurassic Rhynchonellid Brachiopods from Northwestern
Europe. Pp. 119; 12 Plates; 40 Text-figures. 1969. £4.75.
. Goopy, P. C. The relationships of certain Upper Cretaceous Teleosts with
special reference to the Myctophorids. Pp. 255; 102 Text-figures. 1969. £6.50.
. Owen, H. G. Middle Albian Stratigraphy in the Paris Basin. Pp. 164;
3 Plates; 52 Text-figures. 1971. {6.
. Sippigui1, Q. A. Early Tertiary Ostracoda of the family Trachyleberididae
from West Pakistan. Pp. 98; 42 Plates; 7 Text-figures. 1971. £8.
«
BC Printed in England by Staples Printers Limited at their Kettering, Northants establishment
_ CIRRIPEDES FROM THE UPPER -*°""..
CRETACEOUS OF ALABAMA — Lem
“REGION, USS.A.
I. PALAEONTOLOGY
| J. S. H. COLLINS
>) GEOLOGY
eee 'B, MELLEN
Vol. 23 No. 6
LONDON : 1973
GULE REGION, ..U.S.A.
IL PALAEONTOLOGY
BY
JOSEPH STEPHEN HENRY COLLINS
Dulwich, London
Pp 349-380; 5 Plates, 3 Text-figures
Il. GEOLOGY
BY
FREDERIC FRANCIS MELLEN
Jackson, Mississippi, Seen el
Pp 381-388; 2 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 23 No. 6
LONDON : 1973
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY) instituted in 1949, 1s
issued tn five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 23 No. 6 of the Geological
(Palaeontological) series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation
Bull. Br. Mus. nat. Hist. (Geol.).
© Trustees of the British Museum (Natural History), 1973
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 13 July, 1973 Price £2.25
GIRRIPEDES FROM THE UPPER CRETACEOUS
OF ALABAMA AND MISSISSIPPI, EASTERN
GULF REGION, U.S.A.
PART I. PALAEONTOLOGY
By J. S. H. COLLINS
CONTENTS
Page
INTRODUCTION é : : : : 5 ‘ : . = 351
HISTORICAL REVIEW . 5 : : 5 : : 6 : 3 352
SYSTEMATIC DESCRIPTIONS : ‘ A : . ° : . 354
ACKNOWLEDGEMENTS ‘ : : ‘ : : 5 > 5 386
REFERENCES . 5 . : : : 5 . . . ° 387
SYNOPSIS
From the Upper Cretaceous Selma Chalk and Ripley Formation of Mississippi and Selma
Chalk of Alabama, U.S.A. valves of nine new species and one new subspecies of cirripedes are
described and figured: Cretiscalpellum harnedi, C. macrum, C. vallum, C. venustum, Arcoscal-
pellum bakeri, A. campus, A. hubrichti, A. witherst, Brachylepas angulosa, Virgiscalpellum gabbi
apertus. Further valves belonging to Virgiscalpellum gabbi (Pilsbry) and two other Virgiscal-
pellum valves are described and figured from the Ripley Formation. They occur in silty, finely
sandy and highly foraminiferal and ostracodal horizons within the ‘Selma’ which represents an
‘outer neritic’ environment of the continental shelf. Three biostratigraphic horizons are
recognized: (1) zone of Vivgiscalpellum of Ripley and basal Prairie Bluff (Maastrichtian) age;
(2) zone of Cretiscalpellum vallum of basal Annona or upper Coffee (Campanian) age; and
(3) zone of Arcoscalpellum hubrichti of middle Mooreville (Santonian) age. These three zones
supplement known faunal zones within the Upper Cretaceous.
INTRODUCTION
BETWEEN 1935 and 1938 F. F. Mellen corresponded with T. H. Withers of the
British Museum (Natural History) and sent numerous small collections of cirripede
valves to him for study and for description in a paper Withers had planned. The
new forms recognized were received too late for inclusion in the monographic
Catalogue of Fossil Cirripedia, Vol. 2, Cretaceous, 1935. The holotypes were to be
retained in the British Museum (N.H.) and paratypes were to be returned to the
Department of Geology and Geography at Mississippi State University.
The intervention of World War II, the retirement of Withers and his death in 1958,
prevented the completion of the study of the Mississippi barnacle collections which,
according to present tabulation, numbered 146 valves from seven localities (see
below).
Bull. By. Mus. nat. Hist. (Geol.) 23, 6
352 CRETACEOUS CIRRIPEDES
Between 1967-1969 many valves were secured from previous collections and
additional hundreds were collected by F. F. Mellen and others, not only from those
of the earlier sites that still existed, but from new localities scattered over several
Counties in Alabama and Mississippi. A further group of 36 valves, from Greene
County, Alabama, were lent to J. S. H. Collins by the Paleontological Research
Institute, Ithaca, New York.
HISTORICAL REVIEW
The present material contains only one, Virgiscalpellum gabbi (Pilsbry), of the
eight species of cirripedes previously recorded from the Cretaceous of the United
States of America. Of the other seven species, the generic position of one, ‘Scalpel-
lum’ inaequiplicatum Shumard 1862, from the Upper Senonian of Texas, was con-
sidered by both Shumard and Withers (1935) as provisional, owing to the poor pre-
servation of the specimen. The inadequate description of Scalpellum sp. listed by
Adkins 1928, from the Upper Austin Chalk of Texas, was also considered to be
equivocal by Withers (1935). Of the remaining species, Loriculina ?texanum Withers
is recorded from the Middle Albian of Texas; Calantica (Titanolepas) martini Withers,
Stramentum haworthi (Williston) and Squama spissa Logan are known from the
Senonian of Kansas. Stvamentum canadensis (Whiteaves) (Withers, 1935) has been
described from the Fort Benton group of Manitoba and recently Russell (1967)
added a further Canadian species, ?Calantica saskatchewanensis, from the Bearpaw
Formation of southwestern Saskatchewan. Arcoscalpellum conradi (Gabb, 1876)
from the Vincentown Formation of New Jersey, must now be regarded as a
Palaeocene form (Richards, 1958), which substantiates Withers’ (1935) opinion that
the scutum and carina of this resembled certain Tertiary species.
Students and collectors of the rich fauna of Upper Cretaceous sediments of Missis-
sippi, Alabama and Tennessee made great use of Wade (1926) in the identification of
fossils, despite its limitation to basal Ripley strata of Coon Creek, McNairy County,
Tennessee. It was soon recognized that Virgiscalpellum gabbi, first recorded from
Coon Creek, was present in the Ripley strata of Oktibbeha County, Mississippi, and
that other barnacle types were also present.
The collections to be considered afford an aggregate of some 1493 valves ; from these
46 different valves have been distinguished and are included in the following species:
Cretiscalpellum harnedt sp. nov.
C. macrum sp. nov.
C. vallum sp. nov.
C. venustum sp. nov.
Arcoscalpellum bakeri sp. nov.
A. campus sp. nov.
A. hubrichti sp. nov.
A. withersi sp. nov.
Virgiscalpellum gablt gabbi (Pilsbry)
V. gabbi apertus ssp. nov.
V. sp.
V. sp.
Brachylepas angulosa sp. nov.
ALABAMA & MISSISSIPPI 353
This material not only allows nine new species and one new subspecies to be des-
cribed, but also includes the first member of the sub-order Brachylepadomorpha, to
be recorded from the Western Hemisphere. The new material of V. gabbi allows
us to add the scutum, tergum, carinal and rostral latera to the known valves of that
species. Moreover, these valves show that V. g. gabbi is related to the Maastrichtian
species V. hagenowianum (Bosquet) which occurs only at Maastricht.
Both Pilsbry (1933) and Withers expressed doubts whether or not the upper latus,
figured by Withers (1935, p. 39, fig. 2, erroneously called a scutum by Wade, 1926)
should be ascribed to V. gabbi. In his notes on the valves present in the original
collections sent to the British Museum (Nat. Hist.), Withers had considered that this
valve really belonged to the species here named A. withers: but in view of the
additional material in the present collections, which bear a striking similarity to the
upper latus of V. darwinianum (Bosquet) figured by Withers (1935, pl. 38, fig. I)
there seems little to support this opinion. Until such time as more material becomes
available, these particular upper latera are here retained, with reservation, in
V.g. gabbi. The present material from the Ripley Formation of Mississippi shows
that the carina of V. gabbi is very variable in structure and carinae similar to the
one Withers set aside in his notes as a new species are here regarded as a subspecies
of V. gabbi. Also, a distinct scutum and tergum ostensibly from the Ripley Forma-
tion and Annona chalk respectively are described, but not named. These anomalies
indicate that much remains to be learned of the genus Virgiscalpellum, particularly
from the American Cretaceous. It is felt that extensive new collections are desirable
to allow a re-study of the genus to be made.
The valves of Arcoscalpellum hubrichti have characters in common with both of
the European Groups of A. maximum (J. de C. Sowerby) and A. fossula (Darwin) ;
both A. campus and A. withersi have affinities with the Group of A. fossula.
Of the four species of Cretiscalpellum present, C. vallum is the best known; not only
is it represented by numerous carinae, but also by eight other different valves. C.
macrum and C. venustum, the geologically oldest species of the genus in the present
collection, are known from only fifteen and four valves respectively, while C. harnedi,
the youngest of the genus in this study, is represented by thirteen valves.
Withers (1927, 1935, 1953) and Cheetham (1963) have fully discussed the mor-
phology and classification of fossil lepadomorph cirripedes and the latter has also
provided a comprehensive survey of North American Tertiary species.
Following the practice adopted by Darwin, a carina, when present, has been
selected as holotype.
ABBREVIATIONS
The following abbreviations relating to Museum and other collections have been
used in the text: BMNH, British Museum (Natural History); MSU, Mississippi State
University; PRI, Paleontological Research Institution.
354 CRETACEOUS CIRRIPEDES
SYSTEMATIC DESCRIPTIONS
Series CIRRIPEDIA Burmeister, 1834
Order THORACICA Darwin, 1851
Suborder LEPADOMORPHA Pilsbry, 1907
Family SCALPELLIDAE Pilsbry, 1916
Genus CRETISCALPELLUM Withers
1922 Cretiscalpellum Withers : (9), 9, 374-
1935 Cretiscalpellum Withers : 2, 144.
Diacnosis: Scalpellidae with seventeen valves including four pairs of large and
little differentiated, much overlapping lower latera; subcarina much larger than
rostrum ; carina with umbo apical.
TYPE SPECIES: C. unguis (J. de C. Sowerby).
RANGE: Aptian (Lower Greensand) to Maastrichtian.
Cretiscalpellum macrum sp. nov.
(Plate 1, Figs 1-6)
Diacnosis: Carina thin, moderately arched with fine apico-basal ridge, acute
basal angle. Tergum and scutum thin, tergum with carinal margin divided into
upper and lower portions.
HoLotypPe: A carina. BMNH In.64414. (PI. 1, fig. 1), Upper Senonian, Moore-
ville Chalk; 3 miles E. of West Greene and 3 miles W. of Clinton, Greene County,
Alabama.
MATERIAL: 15 valves:
British MusEUM
In.64414. Carina. Holotype. Pl. 1, fig. 1. 385
ACKNOWLEDGEMENTS : c . : : 2 - 5 : 386
REFERENCES . : 5 : . : : . : o 387
STRATIGRAPHY
Hitherto, cirripedes from the ‘Selma chalk’ have all but been ignored stratigraphi-
cally in favour of other more easily recognized macroscopic and abundant micro-
scopic index fossils. It was thought that cirripedes might be of biostratigraphical
importance from the taxonomic study of the Mississippi/Alabama collections. The
horizons of the various species were tabulated and it was seen that they are ap-
parently naturally grouped with no inter-horizon overlapping.
The cirripedes of the ‘Selma chalk’ may be summarized as follows:
Ripley Formation—Navarroan (Maastrichtian) :
C. harnedi, A. bakeri, A. campus, A. withersi, V. g. gabbi, V. g. apertus,
B. angulosa.
Basal Annona—Tayloran (Campanian):
C. vallum.
Middle Mooreville—Austinian (Santonian):
C. macrum, C. venustum, A. hubrichiz.
The genus Virgiscalpellum is restricted to the Ripley and the basal part of the over-
lying Prairie Bluff. This is overlain almost directly by deposits which contain
Diploschiza mellent, a small bivalve used as an index fossil in the Prairie Bluff. Two
new species of Arcoscalpellum and one of Cretiscalpellum found with Virgiscalpellum
also appear to be restricted to this horizon. The Ripley Formation of Oktibbeha
County, Miss., seems to be the most prolific cirripede unit yet described from North
America. An attempt has been made to zone this unit on the basis of its cirripede
fauna, but other than the one specimen of B. angulosa, from the mid-Ripley Barr
Pasture Locality (M4) there seems to be no significant variation, apart from an
apparent increase in abundance of all forms in the middle and upper parts of this
unit.
C. vallum appears to have a restricted vertical range. It is found in deposits of
outer neritic sediments at the base of the Annona chalk (which is the equivalent of
the upper part of the Coffee Sands) some 20-30 ft below the lowest range of Diplo-
schiza cretacea, an important zonal indicator.
A. hubrichti and two new species of Cretiscalpellum seem to be restricted to a zone
which lies near the mid portion of the 250 ft thick Mooreville chalk (marl).
Bull. Br. Mus. nat. Hist. (Geol.) 23, 6
Cc
382 CRETACEOUS CIRRIPEDES
Approximately 250 ft of chalk, and chalky marl separates the zone containing
Virgiscalpellum from that with C. vallum, while about 200 ft of chalky marl separates
the zone containing C. vallum with that of A. hubrichti.
From these observations it is clear that there are three distinct cirripede biostrati-
graphic ‘zones’ within the ‘Selma’. These, designated in descending order, are:
Zone of Virgiscalpellum (Ripley and basal Prairie Bluff—Maastrichtian).
Zone of Cretiscalpellum vallum (basal Annona or upper Coffee—Campanian).
Zone of Arcoscalpellum hubrichti (middle Mooreville—Santonian).
Electrical Log
MABEN
OIE WATER WELL
BIOSTRATIGRAPHIC
(PALEOCENE)
TERTIARY
DIPLOSCHIZA MELLE
NAVARRO aie
GROUP
ONE OF
V/RGISCALPELLUM
MAASTRICHT/IAN
EXOGYRA
CANCELLATA
TAYLOR | coonewax 1
GROUP BEC * |$DIPLOSGHIZA GRETACEA
‘ANNONA nf
=
NS
=
Q
=
x
G
<4} CRETISCALPELLUM
VALLUM
MOORE- ARCOSCALPELLUM
AUSTIN VILLE HUBRICHTI
SANTONIAN
Fic. 4. Correlation and biostratigraphic zones in ‘Selma chalk’ represented on a selected
electrical log, Maben Water Well, SE. 4+ of SW. 4 of NW. 4 of Section 31, T. 20 N., R.12E.,
Oktibbeha County, Mississippi. Elevation 466 feet. Position of Zone of Cvretiscalpellum
vallum is interpreted at 1000 feet (—534). This Zone crops out at elevation 200 feet on south
valley wall of Tibbee Creek, NE. } of SE. 4 of Sec. 6, T. 19 N., R. 16 E., Clay County, 244
miles to the east. The westerly dip of 734 feet is at an average rate of 30 feet per mile,
slightly less than the rate of ‘true’ dip, which is west-southwesterly, about 35 feet per mile.
ALABAMA & MISSISSIPPI 383
GEOGRAPHICAL RANGE: The zone of Virgiscalpellum has been established over an
outcrop distance of 155 miles from Coon Creek, McNairy County, Tennessee, to south
central Noxubee County, Mississippi. Although most, if not all, of the six species
recorded appear possibly to be restricted to this zone, the designation is chosen
because V. gabbi (Pilsbry) was first reported (by Wade) from the lower Ripley Coon
Creek beds of Tennessee; also, this species is the dominant one throughout the Ripley
of Oktibbeha County, Mississippi.
The zone of Cretiscalpellum vallum has been established over an outcrop distance
of 564 miles from Tibbee Creek, Clay County, Miss., to one mile west of Gainesville,
Sumter County, Alabama.
The zone of Arcoscalpellum hubrichti has been established at many localities in an
outcrop belt about two miles wide in Greene County, Alabama; in 1970, Dr E. E.
Russell (Mississippi State University) located the species 4 miles west of Columbus,
Lowndes County, Miss. (M12), in the Mooreville, in a position estimated to be 50 ft
above its base, thus extending the lateral outcrop over a distance of 65 miles between
EUROPEAN TEXAS MISSISSIPPI ALABAMA
STAGES
PRAIRIE BLUFF PRAIRIE BLUFF
maastricurian| NAVARRO ee
GROUP
McNAIRY RIPLEY RIPLEY
GOON CREEK
CAMPANIAN NCOR DEMOPOLIS DEMOPOLIS
GROUP
= = ARCOLA ARCOLA
ANTONIAN
Aafecsre MOOREVILLE MOOREVILLE
CONIACIAN RODE EUTAW
EAGLE FORD EAGLE FORD
TER OMIAS GROUP (LOWER EUTA
=" U. TUSCALOOSA GORDO
WOODBINE |
CENOMANIAN
GROUP |M. TUSCALOOSA POKER
EOLINE
L. TUSCALOOSA! ¢ ot ToNDALE
Fic. 5. Correlation chart showing European stages, Texas Group terminology and Upper
Cretaceous stratigraphic nomenclature in Mississippi and Alabama. No attempt was made
to show range of hiatuses in Texas or Alabama or in the Mississippi subsurface.
GAS ROCK
=
x
=
is}
=
Ly
ry
AUSTIN
384 CRETACEOUS CIRRIPEDES
the States. Barnacle valves were also found at Cochrane, Pickens County, Alabama,
some 15 miles distant from West Green. Recent search has not re-discovered this
particular locality, but as Cochrane is on Mooreville terrain, it is probable that the
valves recorded were of A. hubrichtt. Valves of A. hubrichti outnumber those of the
accompanying species of Cretiscalpellum by a ratio of 200 : I.
A detailed examination of cores and electrical loggings are needed for greater
refinement in the stratigraphical placing of the zones of C. vallum and A. hubrichtz.
Cirripedes of the ‘Selma chalk’ may well be found in sediments of widely varying
bathymetric depths, but the present collection seems to have been preserved best in
deposits consisting of thin-shelled molluscs, comminuted shell fragments, fine sands
and silts. It is suggested that these deposits were formed on outer neritic bottoms
of the continental shelf—in depths of 200-600 ft. A notable exception is V. gabbi
from highly glauconitic and extremely fossiliferous marls of the basal Ripley, Coon
Creek beds of McNairy County, Tennessee, which indicates very shallow marine
conditions.
The various formations composing the ‘Selma chalk’ are tabulated for Alabama,
Mississippi and Texas, and correlated with the European Stages (Fig. 5). Below
the ‘Selma’ the lower and upper Eutaw are shallow marine and estuarine sediments
and would probably yield a cirripede fauna with intensive collecting. Beneath, the
sediments of Tuscaloosa Age are almost entirely continental except in the deepest
sub-surface.
CRETACEOUS DEPOSITION
All the present cirripede collections were made from deposits on a platform area
lying north of a tectonic hinge which marks the northern limit of the Mississippi
Interior Salt Basin. The structure of these deposits indicate a massive step by step
transgression of the Mesozoic seas from the south. During the late Upper Cretaceous
the area of north Mississippi, west Tennessee, west Kentucky, southeast Missouri
and northeast Arkansas was covered by marine water for the first time since the
retreat of the shallow Upper Carboniferous (Pennsylvanian) seas, Alleghany Revolu-
tion (Rodgers, 1967).
The Appalachian Mountain Chain to the north and northeast and the Nashville-
Ozark and Ouachita Uplifts to the north and northwest were probably the proven-
ances of the sediments.
The initial dip (Paleoslope) of the strata of ‘Selma’ age was in a south easterly
direction, except in central Alabama and eastwards, although a period of intense
volcanism reached its climax during late Taylor or early Navarro times in west-
central Mississippi, southeast Arkansas and northeast Louisiana. This produced
the highly uplifted and truncated Jackson Dome and the Sharkey-Monroe Uplift
from which sediments and igneous detritus have been eroded. Scattered biotite
flakes, pyroclastic granules and pebbles, and bentonite deposits in the shallow water
Tuscaloosa, Eutaw, Coffee and Ripley Formations which became increasingly
abundant near the volcanic uplifts, would seem to be evidence of the importance of
contributions of the late Mesozoic volcanism to the geological history of the area.
ALABAMA & MISSISSIPPI 385
It is further suggested that the axis of the Mississippi Embayment and its lateral
Desha Basin which extends west into southeast Arkansas were a subsidence caused
by the initial uplift of the Jackson Dome and the Sharkey-Monroe Uplift to the west.
LOCALITIES AND COLLECTIONS
MISSISSIPPI
‘Original Collections’
Cirripede valves, from seven localities and totalling 146, were from the Ripley Formation of
Oktibbeha County (140) and from the Tayloran chalk of Clay County (6). Of these, 81 were
collected by H. H. Harned, Jr., and 65 by Mellen. At the time these collections were made
there were no good maps of the area, consequently the collections were designated:
OC.1. 2 miles E. of State College.
(Probably NW of NW}, Sec. 5, T. 18 N., R. 15 E., Oktibbeha County.)
OC.2. } mile E. of State College (H.H.H., Jr.)
(Probably Bardwell Pasture, S.W. corner NEi of NW}, Sec. 6, T. 18 N., R. 15 E.,
Oktibbeha County.)
OC.3. 4 mile E. of State College.
(Probably near Centre W4 of NW} Sec. 6, T. 18 N., R. 15 E., Oktibbeha County.)
OC.4. Catalpa Creek.
(NW34 of NE#t of Sec. 25, T. 18 N., R. 14 E., Oktibbeha County: basal Prairie Bluff.)
OC.5. 5 miles NW. of State College (Stoney Point).
(SW4 of SWH of Sec. 16, T. 19 N., R. 14 E., Oktibbeha County: basal Prairie Bluff
and Upper Ripley.)
OC.6. Tibbee Creek (44 miles S. of West Point).
(NE# of SEH, Sec. 6, T. 19 N., R. 16 E., Clay County: basal Annona or upper Coffee.)
OC.7. 14 miles E. of State College—Barr Pasture (H.H.H. Jr.)
(Probably N.W. corner of NE+ of NW} of Sec. 6, T. 18 N., R. 15 E., and other
outcrops lying to the northwest and northeast.)
Serial numbers have been included to simplify locality recordings in the text.
Other collections made by Harned during the 1930’s were included in material sent directly to
Collins later, are marked ‘Barr and Bardwell Pastures’, ‘Barr’, and ‘Sand Creek and Barr’. It
would be imprecise to attempt to assign more specific location descriptions to the old collections
due to the fact that some of the old chalk gullies have been completely filled and obliterated in
recent years.
MISSISSIPPI
‘Recent Collections’
Mr. ‘Oktibbeha County, from Dunn Seiler Museum’—exact localities unknown, but all
undoubtedly from the Ripley of the MSU campus area.
Mz. Barr & Bardwell Pastures: NE} of NW} Sec. 6, T. 18 N., R. 15 E., Oktibbeha County
(H. H. Harned, Jr., Coll.) (middle Ripley).
M3. Barr Pasture: N.W. Corner of NE} of NW} of Sec. 6, T. 18 N., R. 15 E., Oktibbeha
County (H. H. Harned, Jr. Coll.) (middle Ripley).
M4. Barr Pasture: cf. ‘M3’. (middle Ripley)
Ms. Bardwell Pasture: S.W. Corner of NE} of NW} of Sec. 6, T. 18 N., R. 15 E., Oktib-
beha County (H. H. Harned, Jr. Coll.). These gullies have been filled and com-
pletely obliterated. (middle Ripley)
Ato.
AIl.
CRETACEOUS CIRRIPEDES
Sand Creek & Barr: Probably SW Sec. 31, T. 19 N., R. 15 E. & N.W. Corner of NE}
of NW Sec. 6, T. 18 N., R. 15 E. Oktibbeha County (H. H. Harned, Jr. Coll.).
(middle & lower Ripley)
6 Miles N.W. of Starkville (H. H. Harned, Jr. Coll.) Locality indeterminable, but prob-
ably upper Ripley.
N. of Evans Hall: NW} of NW3 Sec. 1, T. 18 N., R. 14 E., Oktibbeha County.
Chapel Hill Church locality of S. valley wall Catalpa Creek, 5 miles S. & 1-75 miles E.
of Barr Pasture locality, NE} of NE Sec. 32, T. 18 N., R. 15 E., Oktibbeha County.
Trim Cane—Josey Creeks confluence; SW of SW} Sec. 20, T. 19 N., R. 14 E., Oktib-
beha County.
4% miles S. of West Point, S. valley wall of Tibbee Creek, E. of U.S. Highway, 45—W.:
NE of SE} Sec. 6, T. 19 N., R. 16 E., Clay County. (basal Annona or upper
Coffee.)
Cuts in Highway 82 bypass, c. 4 miles W. of Columbus, Secs. 26 & 27, T. 19 N., R. 17E.
Lowndes County.
ALABAMA
Approx. S.W. corner SE} of NE} Sec. 27, T. 23 N., R. 1 W., on public road (through
W. M. Steele land), 14 miles N. of West Greene, Greene County.
c. 1 mile N.E. of ‘Ar’, c. SW of SEH, Sec. 23, T. 23 N., R. 1 W., Greene County.
c. 14 miles N.W. of ‘Ar’, near centre NW3 Sec. 22, T. 23 N., R. 1 W., Greene County.
2 miles E. of West Greene, 4 miles N. of Clinton, near Centre SW Sec. 36, T. 23 N.,
R. 1 W., Greene County.
3 miles E. of West Greene & 3 miles W. of Clinton, c. miles N.E. of church, c. Centre
SW3 Sec. 31, T. 23 N., R. 1 E. Greene County.
Mt. Olive Church, around the common quarter corner, Secs. 30 & 31, T.22 N.,R.1E.,
Greene County.
1:2 miles due E. of Mt. Olive Church and in SW} of SW} Sec. 32, T. 22 N., R.TE.,
Greene County.
I mile S.E. of Mt. Olive Church locality; 14 miles N.E. of Mt. Olive Church, north
of Highway, Greene County.
Shallow washes in brushy area W. of paved road 44 miles S.S.W. of Eutaw in SW}
of NE# of Sec. 20, T. 21, N., R. 2 E., Greene County.
1 mile W. of Gainesville, NE}? Sec. 10, T. 21 N., R. 2 W. Sumter County.
‘Bonanza’, SW of SW}, Sec. 32, T. 22 N., R. 1 E., Greene County.
ACKNOWLEDGEMENTS
The writers are greatly obligated to many people for assistance in the collecting,
preservation and study of the cirripedes covered in the present work: to Professors
Troy J. Laswell, Ernest E. Russell and Donald M. Keady of the Department of
Geology and Geography at Mississippi State University for assistance in collecting
and encouragement of student participation, and for affording a repository for barn-
acle collections and literature in the Dunn-Seiler Geology Museum; to Andrew
Martin, Terry Christian, William Honey and John Kline for contributed speci-
mens; to Stephen Hook who collected with one of us (F.F.M.) and made numerous
other contributions during 1968; to Horace H. Harned, Jr., and Wentworth V.
Harned, who, as students of geology at Mississippi State University, collected
and supplied considerable material from the Ripley of Oktibbeha County: to
ALABAMA & MISSISSIPPI 387
Leslie Hubricht, of Meridian, Miss., collector of the type material of Arcoscalpellum
hubrichti and to Dr K. V. W. Palmer for kindly allowing access to this material; to the
Keeper of Palaeontology, British Museum (Natural History), for making available
Withers’ notes and the original material; to Dr W. T. Dean, R. Baker (retired),
S. F. Morris and W. R. Smith, also of the British Museum (Natural History) for kindly
allowing access to specimens in their charge, and to C. W. Wright, of London, for
allowing us to examine specimens in his collection and for much valuable advice.
We wish to record our warmest thanks also to Dr James W. Ward of the Anatomy
Department of the University of Mississippi School of Medicine and Drs Gordon
Gunter and J. Y. Christmas of Gulf Coast Research Laboratory, who furnished
specimens of recent cirripedes and expressed much interest in this work; to Dr
Wayne E. Moore, Division Paleontologist of Chevron Oil Company, for his critical
review of the geological parts of this paper; to the Photographic Department, British
Museum (Natural History) and to E. Kentish, of London, who prepared the photo-
graphs.
Support for extensive field work and other geological studies by Mellen has been
afforded by Germany-Gulmon Oil Company; the Mississippi Geological Survey;
and by the Mississippi Research and Development Center under its Grant Number
1-68-70. Also, many palaeontologists and stratigraphers were of assistance and
encouragement during the investigation.
REFERENCES
Apxins, W. S. 1928. Handbook of Texas Cretaceous Fossils. Univ. Texas Bull. 2838,
1-304, pls., I-37.
BRAUNSTEIN, J. 1950, 1959. Subsurface Stratigraphy of the Upper Cretaceous in Mississippi.
In Cretaceous of Mississippi and South Tennessee: Eighth Field Trip Guidebook, Mississippi
Geol. Soc.; In Upper Cretaceous Outcrops Northeast Mississippi and West Central Alabama:
Fourteenth Field Trip Guidebook, Mississippi Geol. Soc., Jackson.
Capitan, W. M. 1954. Subsurface Geology and Oil and Gas Possibilities of Northwestern
Arkansas; Bull. geol. Surv. Avk., Little Rock, 20.
CHEETHAM, A. H. 1963. Gooseneck Barnacles in the Gulf Coast Tertiary. J. Paleont.,
Tulsa, Oklahoma, 37 : 393-400, pl. 46.
Coiiins, J. S. H. 1965. Arcoscalpellum comptum (Withers), a cirripede new to the Gault.
Palaeontology, Lond., 8 : 629-633, pl. go.
Darwin, C. R. 1851. A Monograph on the Fossil Lepadidae, or Pedunculated Cirripedes of
Great Britain. vi—88 pp., 5 pls., Palaeontogr. Soc. Monogy., London.
Fisk, H. N. 1944. Geological Investigation of the Alluvial Valley of the Lower Mississippi
River. Mississippi River Comm.
Grouskorr, J. G. 1955. Subsurface Geology of the Mississippi Embayment of Southeast
Missoun: Rep. Mo. geol. Surv. Wat. Res., Rolla, 37, Sec. Ser.
Hoek, P. P.C. 1907. The Cirripedia of the Siboga Expedition: Siboga Exped. Rept., 314,
Cirripedia Pedunculata, 127 pp., Io pls.
MELLEN, F. F. 1958. Cretaceous Shelf Sediments of Mississippi, Bull. Miss. geol. Surv.,
Mississippi, 85.
Monroe, W.H. 1941. Notes on Deposits of Selma and Tipley Age in Alabama. Bull. geol.
Suvv. Ala., Montgomery, 78.
388 CRETACEOUS CIRRIPEDES
PirsBry, H. A. 1916. The Sessile Barnacles (Cirripedia) contained in the Collection of the
U.S. National Museum; including a Monograph of the American Species. Bull. U.S. nat.
Mus., Washington, 93 : xii—366 pp., 76 pls., 99 text figs.
1933. An unusual Cretaceous Cirriped. Science, 77 : 1994, 283-284.
Pirssry, H. A. & Orssen, A.A. 1951. Tertiary and Cretaceous Cirripedia from Northwestern
South America. Pyvoc. Acad nat. Sci. Philad., 103 : 197-210, pls. 6-11.
Pryor, W. A. 1960. Cretaceous Sedimentation in Upper Mississippi Embayment. Bull.
Am. Ass. Petrol. Geol., Chicago 44 : 9.
RAINWATER, E. H. 1963. Geological History and Oil and Gas Possibilities of Mississippi; In
Mississippi Geologic Research Papers—1962, Bull. Miss. geol. Surv., 97.
Ricuarps, H. G. 1958. The Cretaceous Fossils of New Jersey. Bur. Geol. and Topogr.,
Trenton, N.J., Pt. 1., vi-266, 46 pls.
RODGERS, J. 1967. Chronology of Tectonic movements in the Appalachian Region of Eastern
North America, in Symposium on the Chronology of Tectonic movements in the United
States. Amer. Jour. Sct., New Haven, 265, no. 5, pp. 408-27, table.
RUSSELL, L. S. 1967. A Pedunculate Cirripede from Upper Cretaceous Rocks of Saskat-
chewan. J. Paleont., Tulsa, Oklahoma, 41 : 1544-1547, 5 text figs.
SHUMARD, B. F. 1862. Descriptions of new Cretaceous fossils from Texas. Pyvoc. Boston
Soc. Nat. Hist., 8 : 188-205.
STEPHENSON, L. W. & Monror, W.H. 1940. The Upper Cretaceous Deposits (of Mississippi).
Bull. Miss. geol. Surv., 40.
STEPHENSON, etal. 1942. Cretaceous Correlation. Bull. geol. Soc. Amey., New York 53 : 435-
448.
Wave, B. 1926. The fauna of the Ripley Formation on Coon Creek, Tennessee. U.S. Geol.
Surv., Prof. Paper, 137, 1-272, 62 pls.
WitTHeErRS, T. H. 1928-1953. Catalogue of Fossil Civripedia in the Department of Geology,
British Museum (Natural History) : 1928 Triassic and Jurassic. xii-154, 12 pls.; 1935
Cretaceous. xvi-434, 50 pls.; 1953 Tertiary. xv—396, 64 pls. London, British Museum
(Natural History).
1945. New Cretaceous Cirripedes and Crab. Ann. Mag. nat. Hist., London, Ser. 2,
5 : 552-561, pl. 2.
J. S. H. CoLiins FREDERIC F. MELLEN
63, OAKHURST GROVE 1202 STANDARD LIFE BUILDING
E. DuLwicH JACKSON
Lonpon, S.E.22. MISSISSIPPI 39201
DESsAr
PLATE 1
Cretiscalpellum macrumi sp. nov. Page 354
Upper Senonian, Mooreville Chalk.
Fic. 1. Carina. Outer view. Holotype. In.64414. A5 : Greene County, Alabama.
Fic. 2. Carina. Outer view. A8. Paratype. MSU 1311.
Fic. 3. Scutum (right). Outer view. Ai. Paratype. In.64415.
Fic. 4. Tergum (right). Outer view. A4. Paratype. In.64416.
Fic. 5. Tergum (left). Outer view. A8. Paratype. MSU 1312.
Fic. 6. Tergum (left). Outer view. A8. Paratype. In.64417.
Fics. 1-6 x2 diam.
Cretiscalpellum venustum sp. nov. Page 356
Lower or middle Mooreville Chalk, Upper Senonian.
Fic. 7. Carina. a, outer view; b, side view. Holotype. In.64418. A7 : Greene County,
Alabama.
Fic. 8. Carina. Outer view. A8. Paratype. MSU 1313.
Fic. 9. Tergum (left). Outer view. A8. Paratype. In.64419.
Fics. 7-9 x2 diam.
Cretiscalpellum harnedi sp. nov. Page 357
Maastrichtian, Middle Ripley Formation.
Fic. 10. Scutum (right). a, outer view; b, inner view. M8 Paratype. In. 64420.
Fic. 11. Carina. a, side view; b, outer view; c, inner view. Holotype. In.64421. M3:
Oktibbeha County, Mississippi.
Fic. 12. Tergum (right). a, outer view; b, inner view. M8. Paratype. In.64422.
Fics. 10-12 x4 diam.
Cretiscalpellum venustum sp. nov. Page 350
Fic. 13. Transverse section of carina, MSU 1313 at apex, c. x8 diam.
Cretiscalpellum vallum sp. nov. Page 358
Upper Senonian, basal Annona or upper Coffee Formation.
Fic. 14. Carina. a, outer view; b, inner view, x2 diam. Mir. Paratype. MSU 1316.
Fic. 15. Carina. a, outer view; b, side view, x2 diam. Holotype. In.64423. Murr:
Clay County, Mississippi.
Fic. 16. Carina, with small bivalve on the outer surface, x2 diam. Mur. Paratype.
MSU 1355.
Fic. 17. Transverse sections of carina, In.64423, at a, c. 7-0 mm and J, c. 14-0 mm from apex.
c. x5 diam.
Fic. 18. Tergum (right). Outer view, x2diam. M11. Paratype. In.64426.
Fic. 19. Scutum (left). Outer view, x4 diam. Mir. Paratype. MSU 1317.
Fic. 20. Scutum (left). a@, outer view; 6, inner view, x2 diam. M11. Paratype. In.
04425.
Fic. 21. Upperlatus. Outer view, x4diam. M11. Paratype. MSU 13109.
Fic. 22. Subcarina. a, inner view; b, outer view, x2 diam. Mrr. Paratype. In.64424.
Bull. By. Mus. nat. Hist. (Geol.) 23, 6
PEATE 1
10b
12b
Ila ie) Ile
| a
15a
17b
20a 20b
D*
22a
22b
PLEAD E, 2
Cretiscalpellum vallum sp. nov. Page 358
Fic. 1. Subcarina. Outer view, x4 diam. M1r. Paratype. MSU 1320.
Fic. 2. Tergum (left). a, inner view; b, outer view, x2diam. Mir. Paratype. In.64428.
Fic. 3. Tergum (left), with small bivalve on the outer surface, x2 diam. Mir. Paratype.
In.64435.
Fic. 4. Tergum (left). Outer view, x4diam. M11. Paratype. MSU 1318.
Fics 5-6. Carinal latera (left & right). Outer views, x4 diam. M11. Paratypes. In.
64433-34.
Fic. 7. Rostral latus (left). Outer view, x2 diam. Mir. Paratype. In.64429.
Fic. 8. Subcarinal latus (right). Outer view x4 diam. Mir. Paratype. In.64430.
Fic.9. Inframedian latus. Outer view, x4diam. Ato. Paratype. In.64431.
Fic. 10. Upper latus. Outer view of juvenile valve, x6 diam. OC.6. Paratype. In.
64432.
Fic. 11. Upperlatus. Outer view, x4 diam. Mii. Paratype. In.64427.
Fic. 12. Subrostrum. Outer view, x4 diam. Mr1r. Paratype. I[n.64483.
Arcoscalpellum hubrichti sp. nov. Page 363
Upper Senonian, Mooreville Chalk.
Fic. 13. Carina. a, outer view; b, side view; c, inner view, x2 diam. At : Greene County,
Alabama. Holotype. - 6072PRI.
Fic. 14. Carina. a, outer view; b, side view, x2 diam. A6. I[n.64438.
Fic. 15. Carina and associated left tergum which has become orientated through 180°.
Outer view, natural size. Az. MSU 1358.
Fic. 16. Scutum (right). a@, outer view; b, inner view, x2 diam. A7. Paratype. In.
64436.
Fic. 17. Scutum (left). Outer view, x2 diam. A7. Paratype. In.64437.
Fic. 18. Scutum (left). Outer view, x2 diam. A7. Paratype. MSU 1325.
Fic. 19. Upper latus (right). Outer view, x4diam. A3. Paratype. MSU 1359.
Fic. 20. Transverse sections of carinae; at a, 1o-o mm & b, 15-0 mm from the apex, c. x 4
diam. Paratypes. 6082a—6082bPRI.
Bull. By. Mus. nat. Hist. (Geol.) 23, 6 ILA WIS,
l6a-b
20a-b
PLATE 3
Arcoscalpellum hubrichti sp. nov. Page 363
Fic. 1. Tergum (right). Outer view, x2diam. Ar. Paratype. 6074PRI.
Fic. 2. Tergum (right). a, outer view; 6, inner view, X2diam. A3. MSU 1324.
Fic. 3. Tergum (right). Outer view, x2 diam. Ar. In.64439.
Fic. 4. Upper latus (left). Outer view, showing well developed growth beyond the umbo,
x2 diam. A7. Paratype. In.64443.
Fic. 5. Upper latus (right). Outer view, x2 diam. A7. Paratype. In.64444.
Fic. 6. Carinal Jatus (right). Outer view, x2diam. A7. Paratype. In.64440.
Fic. 7. Carina. Side view of apical part, x2diam. Az. Paratype. 60873aPRI.
Fic. 8. Carinal latus (right). a, outer view; b, inner view, x2 diam. A7. Paratype.
In.64441.
Fic. 9. Carinal latus (left). Outer view, x2diam. A7. Paratype. In.64442.
Arcoscalpellum bakeri sp. nov. Page 367
Maastrichtian, Ripley Formation.
Fic. 10. Carina. a, outer view; b, side view; c, inner view, x2 diam. My4 : Oktibbeha
County, Mississippi. Holotype. In.64446.
Fic. 11. Transverse sections of carina taken at, a, c. 5-o mm and Bb, c. 18-0 mm from the apex
x5 diam. Paratype. In.64448.
Fic. 12. Carina. a, outer view; b, inner view, x4 diam. M3. Paratype. In.64447.
Fic. 13. Tergum (right). Outer view, x4 diam. Mz. Paratype. In.64449.
Arcoscalpellum campus sp. nov. Page 369
Maastrichtian, Middle Ripley Formation.
Fic. 14 Carina. a, outer view; b, side view, x2 diam. M5 : Oktibbeha County, Missis-
sippl. Holotype. In.64452.
Fic. 15. Scutum (left). a, outer view; b, inner view, x4 diam. M5. Paratype. MSU
1328.
Fic. 16. Carinal latus (right). a@, outer view; b, inner view, x4 diam. M4. Paratype
In.64453-
Arcoscalpellum withersi sp. nov. Page 371
Maastrichtian, Ripley Formation.
Fics 17-18. Rostral latera (left & right). Outer view, x4 diam. M5. Paratypes.
In.64454-64455.
Bull. By. Mus. nat. Hist. (Geol.) 23, 6 PAs 3
PLATE 4
Arcoscalpellum campus sp. nov. Page 369
Fic. 1. Scutum (left), Outer view, x4 diam. M5. Paratype. In.64450.
Fic. 2. Tergum (left). a, inner view; b, outer view, x4 diam. M5. Paratype. In.64451.
Fic. 3. Tergum (left). Outer view, x4 diam. M4. Paratype. MSU 1329.
Fic. 4. Transverse section of carina, In.64452, at c. 4.0 mm from the apex, x6 diam.
Arcoscalpellum withersi sp. nov. Page 371
Fic. 5. Carina. Outer view, x2 diam. OC.7. Paratype. MSU 1330.
Fic. 6. Carina. a, outer view; b, side view; c, inner view, x2 diam. M3: Oktibbeha
County, Mississippi. Holotype. In.64456.
Fic. 7. Transverse sections of carina, at, a, c. 6-0 mm and 8, c. 11-0 mm from the apex,
x5 diam. M3. Paratype. In.64463.
Fic. 8. Scutum (right). Outer view, x2 diam. M4. Paratype. In.64459.
Fic. 9. Scutum (right). a, outer view; b, inner view, x2 diam. M3, Paratype. MSU
1331.
Fic. 10. Tergum (right). Outer view, x2diam. M2. Paratype. MSU 1332.
Fic. 11. Tergum (left). Outer view, x2 diam. M3. Paratype. In.64457.
Fic. 12. Tergum (left). Outer view, x2 diam. Mz. Paratype. MSU 1425.
Fic. 13. Scutum (left). Outer view, x2 diam. M4. Paratype. MSU 1426.
Fic. 14. Carinal latus (right). a, outer view; b, inner view, x4 diam. M4. Paratype.
In.64461.
Fic. 15. Upper latus (right). Outer view, x4diam. M4. Paratype. In.64462.
Virgiscalpellum gabbi gabbi (Pilsbry) Page 374
Maastrichtian, Ripley and basal Prairie Bluff Formations.
Fic. 16. Upper latus (right). Outer view, x4 diam. OC.1. In.64474.
Fic. 17. Carina. a, outer view; b, side view, x2diam. M8. MSU 1334.
Fic. 18. Carina. a, side view; b, inner view, x2diam. OC.1. In.64464.
Bull. By. Mus. nat. Hist. (Geol.) 23, 6 PLATE 4
PLATE 5
Virgiscalpellum gabbi gabbi (Pilsbry) Page 374
Fic. 1. Tergum (left). a@, outer view; b, inner view, x2 diam. M2. In.64472.
Fic. 2. Tergum (left). Outer view, x2 diam. Mz. MSU 1335.
Fic. 3. Scutum (right). a, outer view; b, inner view, x2diam. Mz. Paratype. In.64467.
Fic. 4. Tergum (left). Outer view, x2diam. OC.2. In.64469.
Fic.5. MRostrallatus (left). Outer view, x4diam. M3. Paratype. MSU 1337.
Fic. 6. Rostral latus (left). Outer view, x2 diam. M3. Paratype. In.64475.
Fic. 7. Scutum (left). Outer view, x2 diam. Mz. Paratype. In.64470.
Fic. 8. Carinal latus (right). Outer view, x4 diam. OC.1. Paratype. In.64473.
Fic. 9. Transverse sections of carinae: a, open lower limb 4:0 mm from umbo, In.64465;
infilled lower limb 8-0 mm from umbo, In.64466. 5 diam. M2.
Fic. 10. Carina. a, side view; b, inner view, x2 diam. With the umbo and part of the
lower limb infilled. M8. MSU 1416.
Fic. 11. Scutum (right). Outer view, x2 diam. Mz. MSU 1336.
Virgiscalpellum sp. Page 379
Upper Senonian, basal Annona or upper Coffee Formation.
Fic. 12. Scutum (right). Outer view, «x4 diam. OC.6: Clay County, Mississippi. In.
64476.
Virgiscalpellum sp. Page 379
Maastrichtian, Upper Ripley Formation.
Fic. 13. Tergum (left). Outer view, x4 diam. OC.3 : Oktibbeha County, Mississippi.
In.64477.
Virgiscalpellum gabbi gabbi (Pilsbry) Page 374
Fic. 14. Carina. a, outer view; b, side view; c, inner view, x2 diam. With thickened
lateral margins. OC.1. In.64487.
Virgiscalpellum gabbi apertus sub-sp. nov. Page 378
Maastrichtian, Ripley and basal Prairie Bluff Chalk.
Fic. 15. Carina. a, outer view; b, side view; c, inner view, x2 diam. OC.4 : Oktibbeha
County, Mississippi. Holotype. In.64478.
Brachylepas angulosa sp. nov. Page 380
Maastrichtian, Ripley Formation.
Fic. 16. Rostrum. a, outer view; b, side view; c, inner view, x4 diam. M4 : Oktibbeha
County, Mississippi. Holotype. In.64479.
Bull. By. Mus. nat. Hist. (Geol.) 23, 6 PLATE 5
A LIST OF SUPPLEMENTS
TO THE GEOLOGICAL SERIES
OF THE: BULLETIN. OF
THE BRITISH MUSEUM (NATURAL HISTORY)
. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya.
Pp. 213; 30 Plates; 2 Text-figures. 1965. {6.
. Et-NaceGar, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper
Cretaceous—Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley,
Egypt, U.A.R. Pp. 291; 23 Plates; 18 Text-figures. 1966. fro.
. Davey, R. J., Downtz, C., SARGEANT, W. A. S. & Witttams, G. L. Studies on
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248; 28 Plates; 64 Text-
figures. 1966. £7.
. APPENDIX. DAVEY, R. J., Downtg, C., SARGEANT, W. A. S. & WILLIAMS, G. L.
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24.
1969. 8op. ie Se
. Ettiott, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of the -
Middle East. Pp. 111; 24 Plates; 17 Text-figures. 1968. {5.12%.
. Ruopes, F. H. T., Austin, R. L. & Druce, E, C. British Avonian (Carboni-
ferous) Conodont faunas, and their value in local and continental correlation.
Pp. 315; 31 Plates; 92 Text-figures. 1969. {1I1.
. Cuitps, A. Upper Jurassic Rhynchonellid Brachiopods from Northwestern ,
Europe. Pp. 119; 12 Plates; 40 Text-figures. 1969. £4.75.
. Goopy, P. C. The relationships of certain Upper Cretaceous Teleosts with —
special reference to the Myctophorids. Pp. 255; 102 Text-figures. 1969. 6.50.
. OwEN, H. G. Middle Albian Stratigraphy in the Paris Basin. Pp. 164;
3 Plates; 52 Text-figures. 1971. {6.
. Sippigui, Q. A. Early Tertiary Ostracoda of the family Trachyleberididae
from West Pakistan. Pp. 98; 42 Plates; 7 Text-figures. 1971. £8.
0-05) between the
spacing of carinal nodes on the two kinds of branches.
Cross-sections of some heavily encrusted branches showed that although the
reverse surface may be smoothly rounded and without striae, secondary laminae
FROM FERMANAGH, IRELAND 487
within the branch become progressively more corrugated towards the axial region.
This tendency reaches its maximum at the contact with the strongly ribbed under
side of the basal plate, a part of the primary skeleton. The ribbing seen in cross-
sections is merely a reflection of the striated or longitudinally ridged condition in
solid specimens, and it therefore seems that although the presence or absence of
‘striae’ on the reverse may be an indication of the age, or the state of weathering of a
specimen, it is unlikely to be of taxonomic value.
Genus SEPTOPORA Prout, 1859
DiaGnosis. Zoarium a fenestrate, flabellate or leaf-like expansion; primary
branches numerous; increasing by bifurcation or interpolation; the lateral branches
unite with those from the adjacent primary branches; apertures in two rows on
primary and lateral branches; reverse usually with fine striae and scattered
dimorphic pores (Nickles & Bassler 1900: 41).
TYPE SPECIES. S. cestviensis Prout 1859. Mississippian, Illinois.
Septopora hibernica sp. nov.
(Pl. 26, figs 1-8)
MATERIAL. Only ten specimens were available. They are not in such good
condition as those of other species, being encrusted with secondary tissue to an
unusual degree, so that surface features are commonly obscured. Some branches
lack the entire apertural face, presumably owing to partial replacement, so that only
the base and sides remain. It is possible that all specimens belonged to a single
colony, and this should not be overlooked in using the following data for comparative
purposes. The largest fragment was 42 x 17mm. PD.4885-91, PD.5336-8.
Measurements (N = 10):
B € D
LAE I-20-1°50 1-400 0-100 7-14
F.w. I*50-2:00 1-640 0-156 g°51
L.ap.s. 0:35-0:40 0°375 O-O1I 3:00
B.w. 0:72-0:87 0-770 0-043 5°58
(primary)
B.w. 0:45-0:67 0-515 0:070 13°59
(secondary)
Note: Data for zooecial apertures are not included as these structures are com-
monly broken, abraded, or so reduced by coarse grained silicification that original
dimensions are not evident.
Micrometric formula: 6—8/7—9//12—14/none.
Apertures per fenestrule.
Range of specimen modes: 3
Distribution of specimen modes: 10
DiacGnosis. Septopora of unknown zoarial shape, lacking carinal nodes and
‘dimorphic pores’. With well developed keel on primary, but not secondary
branches. The latter have a variable growth habit, causing fenestrule shapes to be
G
488 VISEAN FENESTRATE BRYOZOA
irregular. Primary branches may show a Ptylopora-like pattern and bear two rows
of small, closely-spaced zooecial apertures. Chamber base shapes triangular or
hemi-hexagonal with short lateral walls. Structural data as above.
TYPE SPECIMENS. Holotype: PD.4885. Paratypes: PD.4886 to PD.48q0.
DESCRIPTION. There is no certain evidence of zoarial shape though, judging
from the structure of available fragments, it was probably a more or less upright,
foliaceous or fan-shaped growth. Two of the larger fragments have a pronounced
Ptylopora-like branch pattern and are only distinguishable from that genus by the
cross-bars (secondary branches) which bear zooecia.
Primary branches are straight or gently flexed. They are stout and somewhat
flattened on the obverse, which bears a prominent, rib-like median keel. In shallow,
longitudinal grooves on either side of this are two rows of zooecial apertures. The
reverse is broadly rounded and longitudinally striated, the striae serving to separate
a number of ridges, each of which bears on its crest a row of minute, rounded papillae.
These are about 0-01 mm in diameter and spaced about 0:04 mm apart. From eight
to ten parallel ridges are visible on the reverse of a branch.
Secondary branches (corresponding to dissepiments in the Fenestellidae) are
variably developed, and their original shapes are commonly obscured by heavy
secondary deposits. Where two unite in the chevron pattern characteristic of
Septopora, they leave the main branches at angles of about 50°. This arrangement
is uncommon in the present material in which, owing to space restriction and
irregularities of growth, others predominate. Close to the junction of primary
branches secondaries may be short and straight, rather like stout fenestellid dissepi-
ments. In other cases a secondary appears to have developed independently of its
partner, forming an oblique cross bar between adjacent primary branches. Bow-
shapes are of fairly common occurrence, the angular junction between components
having been rounded by later schlerenchymal growth. In other examples irregular
shapes have resulted from asymmetrical unions between opposing secondary
branches. Well developed secondaries have two rows of alternating apertures, but
in many cases the arrangement is irregular. Some appear to lack apertures alto-
gether, but this is probably due to sealing by schlerenchymal overgrowth rather
than to a genuine absence of zooecia. Secondary branches do not have a median
keel and the cross-section is approximately circular. The reverse shows the same
characteristics as in primaries.
Fenestrules in Septopora are typically chevron-shaped with rounded extremities,
but these are rare in the present specimens. The irregular growth of secondary
branches inevitably results in many variations of fenestrule shape, which may be
round, oval, rectangular or even triangular. Variable amounts of secondary
encrustation cause still further diversity and commonly convert regular to irregular
shapes. The relatively small size of fenestrules and their variable configuration
might almost be said to characterize the specimens examined.
Zooecial apertures are in two alternating rows close to the keel and away from
branch margins. They may be circular, with a diameter of about 0-15 mm, or (more
commonly) oval, measuring about 0-225 by 0-125 mm. Some are relatively small
FROM FERMANAGH, IRELAND 489
and more than their own diameter apart; others are much larger and separated by
only a thin bar. Much depends on the amount of secondary encrustation present,
for there is no doubt that this has in many cases reduced the size of apertures or even
sealed them altogether. On the other hand, many have been enlarged by breakage
or incomplete replacement of their rims. Apertures on both kinds of branches
were probably of about the same size.
Few zooecial base shapes are distinguishable in the specimens, but it is possible
that there is a slight difference in this respect between primary and secondary
branches. Those of the former seem to have hemi-hexagonal shapes with a length
of about 0-32 mm and maximum width of 0-2mm._ The short lateral walls are about
0-05 mm long. On secondary branches chambers appear to be of similar size but
triangular, with the maximum length (parallel to the branch margin) about 0-3 mm
and maximum width about 0:25 mm. Only three chamber bases of each kind were
seen, however, and it is uncertain whether the observed difference is general or not.
Discussion. This is the first positive record of Septopora in the British Isles,
though Whidborne (1895, p. 183) mistakenly assigned a poorly preserved specimen
(probably of Ptiloporella) to the genus. Nor has Septopora been found in other
European countries, with the exception of Russia where a number of species have
been reported in recent years (Nikiforova 1938; Shishova 1952, 1957; Morozova
1955). The genus was introduced by Prout in the United States, and several species
were described by Ulrich (1890). A possible reason for the previous absence of
Septopora from faunal lists is that, due to lines of weakness provided by the angular
junctions between secondary branches, the colonies were particularly liable to
breakage. The resultant fragments typically consist of a primary branch with a
number of secondaries on each side, all broken at the junction angle. Such fragments
are exceedingly difficult to distinguish from Penniretepora stipes (always abundant in
Carboniferous bryozoan faunas) and could easily have been described as such in the
literature.
Among described forms of Septopora the present specimens are closest to the type
species, which is from approximately the same horizon (lower Chester) in Illinois.
Common features include a clear median keel on the obverse, about three zooecial
apertures per fenestrule and the absence of small accessory apertures (‘dimorphic
pores’). There are also important differences, however, S. cestriensis having a
rather finer mesh (ten or eleven branches and fenestrules in 10 mm, according to
Prout), carinal nodes and apertural peristomes. Ulrich (1890; 628) recorded the
species from the lower and middle Chester of Illinois and Kentucky, but his speci-
mens had an even finer meshwork than those of Prout. He also mentioned the
presence of accessory pores on both obverse and reverse, a feature not shown by
Prout’s specimens or the present ones. The average branch width of Ulrich’s
material was 0-5 mm, considerably less than that of the Irish specimens, though this
might be accounted for by the thick secondary encrustation of the latter. Shishova
(1952; 162) found, in the Dinantian of the Moscow region, specimens that she
assigned to S. cestriensis. She stated their formula as: 14/103—114//22-223, which
differs widely from that of the Fermanagh specimens. Also her material had
490 VISEAN FENESTRATE BRYOZOA
lozenge-shaped (rhomboidal) chamber bases, and there were accessory pores on the
reverse.
The only other described species which the present specimens resemble is S.
subquadrans Ulrich, from the upper Chester. This differs from S. cestriensis mainly
in fenestrule shape, and has the formula: 7-12/10}—12//21. The number of apertures
per 5 mm is greatly in excess of that shown by the Fermanagh specimens, however,
and there are accessory apertures on both surfaces (Ulrich 1890; 6209).
In view of the lack of correspondence with existing species it is clearly necessary to
introduce a new name for the present material. S. hibernica seems appropriate for
the first recorded occurrence of the genus in Ireland.
Other Genera
Note: In addition to the above forms, three further fenestrate bryozoans belonging
to the Carrick Lough fauna have been described separately elsewhere by the author.
In each case the account given follows the basic pattern used here. The additional
species and location of their descriptions are:
1. Ptilofenestella carrickensis gen. et sp. nov. In 1965, Palaeontology 8 : 478-401.
2. Muinilya nodulosa (Phillips). In 1965, Geol. Mag. 102 : 135-142.
3. Polypora stenostoma sp. nov. In 1971, Palaeontology 14 : 178-187.
“REFERENCES
Bass_erR, R. S. 1935. In: Fosstlium Catalogus, 1: animalia. Ed. W. Quenstedt. Pars 67:
Bryozoa, 1-229, s’Gravenhage.
1953. Bryozoa: in Tveatise on Invertebvate Paleontology, ed. Moore: i-xili, G1—253.
Kansas.
BELL, W. A. 1929. The Horton—Windsor District, Nova Scotia. Mem. geol. Surv. Can. 155 :
25-84, pl. 12-13.
Brunton, C. H. C. 1966. Silicified productoids from the Visean of County Fermanagh.
Bull. Br. Mus. nat. Hist. (Geol.), London 12 : 173-243, pls 1-109.
CAMPBELL, K.S.W. 1961. Carboniferous fossils from the Kuttung rocks of New South Wales.
Palaeontology, London 4 : 428-474, pls 53-63.
Canu, F. 1900. Révision des bryozoaires du Crétacé figurés par d’Orbigny. Bull. géol. Soc.
Fr., Paris (3), 28 : 334-463.
CuRonIic, J. 1949. In: Newell, N. D., Chronic, J., and Roberts, T. G., The Upper Paleozoic
of Peru. Mem. geol. Soc. Am., Washington 58 : 1-241, 44 pls.
Conpra, G. E. & Exias, M. K. 1944b. Study and revision of Archimedes (Hall). Geol. Soc.
Am. Spec. Pap. 53 : i-vili, 1-243, 41 pls.
Cumincs, E. R. 1904. Development of some Paleozoic bryozoa. Am. J. Sci., New Haven
17 : 49-78.
1906. Description of the bryozoa of the Salem Limestone of southern Indiana. Rep.
Dep. Geol. Nat. Res. Indiana, Indianapolis 30 : 1274-1296, pls 27-40.
Dawson, J. W. 1878. Acadian Geology (3rd edition). i—xxviii, 1-797, London.
EICHWALD, E. 1860. Lethaea Rossica; Paléontologie dela Russe: Ancienne Periode. Bryozoa :
355-494, pls 23-32, Moscow. In French.
Exias, M. K. 1937a. Depth of deposition of the Big Blue (late Paleozoic) sediments in Kansas.
Bull, geol. Soc. Am., Rochester, N.Y. 48 : 403-432.
—— 1937b. Stratigraphic significance of some late Paleozoic fenestrate bryozoans. J. Paleont.,
Tulsa, Okla. 11 : 306-336.
FROM FERMANAGH, IRELAND 491
Exias, M. K. 1950. List of identified species in Hunterian collection of Carboniferous bryozoa, with
brief notes on thety most characteristic chavactervs. Unpublished manuscript: 1-4.
1956. A Revision of Fenestella subantiqua and related Silurian fenestellids. J. Paleont.,
Tulsa, Okla. 30 : 314-332, pl. 43.
1964. Stratigraphy and paleoecology of some Carboniferous bryozoans. C.7., Cinquiéme
Congr. internatn. de stratigvaphie et de la géologie du Carvbonifére : 375-382, pls 1-5.
Extras, M. K., and Conpra, G. E. 1957. Fenestella from the Permian of West Texas. Mem.
geol. Soc. Am., Washington 70 : IX and 158; 23 pls.
Forrste, A. F. 1887. Flint Ridge bryozoa. Bull. Denison Umiv. Sci. Lab. 2 : 71-88, pl. 7.
GerorGE, T. N., and OswaLp, D. H. 1957. The Carboniferous rocks of the Donegal syncline.
Q. Jl geol. Soc. Lond., London 113 : 137-183.
GoLtprFuss, A. 1826-44. Petrefacta Musei Universitatis Regiae Borussicae Rhenanae Bonnensis,
I. 252pp. Diisseldorf.
Hatt, J. 1881. Bryozoans of the Upper Helderberg and Hamilton groups. Absty. Trans.
Albany Inst. 10 : 1-26.
Hatt, J. & Simpson, G. B. 1887. Palaeontology of New York, 6: Corals and bryozoa, i-xxvi,
1-298, 66 pls, Albany, N.Y.
HaArRMER, Sir S. F. 1934. The polyzoa of the Siboga Expedition. Pt. III Cheilostomata, Asco-
phora. Family Reteporidae, i—viii, 503-637, pls 25-41, Leiden.
Hinks, T. 1880. A history of the British marine polyzoa 1: i-cxli, 1-601, Vol. 2, 83 pls, London.
HIsINGER, W. 1837. Lethaea Svecica seu Petrificata Sueciae Iconibus et Chavacteribus Illustrata.
124 pp., 38 pls, Stockholm.
Kaisin, F. 1942. Les bryozoaires fenestrellinidés et acanthocladiidés du Tournaisian de la
Belgique. Mém. Inst. Géol. Louvain, 13 : 93-141, pls 1-8. In French.
Kine, W. 1850. The Permian fossils of England. Palaeontographical Society i-xxxvii,
1-258, 28 pls, London.
KoeEnic, J. W. 1958. Fenestrate bryozoa in the Chouteau Group of central Missouri. /.
Paleont., Tulsa, Okla. 32 : 126-143, pls 21-22.
Lacaalj, R. & GAUTIER, Y. V. 1965. Bryozoan assemblages from marine sediments of the
Rhone delta, France. Micropaleontology, New York 11 : 39-58.
LikHAREV, B. 1934. Upper Carboniferous of Ferghana. Bull. geol. Soc. China, Peking
13 : 155-172.
LonsDALE, W. 1839. In Murchison, R. I., The Silurian System 2 : 675-677. London.
LowenstaM, H. A. 1950. Niagaran reefs of the Great Lakes area. J. Geol., Chicago, etc.
58 : 430-480.
MapDGEFRAU, K. 1933. Zur entstehung der mitteldeutchen Zechstein-Riffe. Centralbl.
Mineralogie, Jahrg. 1933 (Abt. B, 11) : 621-624, Berlin. In German.
M’Coy, SirF. 1844. A synopsis of the character of the Carboniferous Limestone fossils of Iveland :
i-vill, I-207, pls 1-29, Dublin.
McFarian, A. C. 1942. Chester bryozoa of Illinois and western Kentucky. J. Paleont.,
Tulsa, Okla. 16 : 437-458, pls 65-68.
MitterR, T. G. 1961. Type specimens of the genus Fenestella from the Lower Carboniferous
of Great Britain. Palaeontology, London 4 : 221-242, pls 24-27.
1962a. North American species of Fenestella from the Carboniferous of Great Britain and
Ireland. J. Paleont., Tulsa, Okla. 36 : 120-125, pls 23-24.
1962b. On Hemitrypa hibernica M’Coy. Geol. Mag. London, Hertford 99 : 313-321.
1963. The bryozoan genus Polypova M’Coy. Palaeontology, London 6 : 166-171, pls
23-24.
Moore, R.C. 1929. A bryozoan faunule from the Upper Graham Formation, Pennsylvanian,
of north central Texas. J. Paleont., Tulsa, Okla. 3 : 1-27, pls 1-3.
NEKHOROSHEY, B. P. 1926. Lower Carboniferous bryozoa from the Kuznetsk Basin. Bull.
Com. géol. Leningrad 43 : 1237-1290, pls 19-20. In Russian, with English summary.
1956. Nizhni Kamennougolnye Mshanki Altaia y Sibiri. Tvudj vses. nauchno-issled.
Inst. Geol.; Min. Geol. Och. Nedr. S.S.S.R. (n.s.) 13 : 1-419, pl. 57, Moscow. In Russian.
492 VISEAN FENESTRATE BRYOZOA
NEWELL, N. D. and others. 1953. The Peymian reef complex of the Guadalupe Mountains
vegion, Texas and New Mexico. i—xix, 1-236, 32 pls, San Francisco.
NickLes, J. M. & Bassler, R. S. tIg00. A synopsis of American fossil bryozoa. Bull. U.S.
geol. Surv., Washington 173 : 1-663.
Nixirorova, A. I. 1926. Lower Carboniferous bryozoa of Turkestan. Bull. Com. géol.
Leningrad 45 : 175-192, pls 4-5. In Russian, with English summary.
1927. Lower Carboniferous bryozoa from the Donetz Basin. Bull. Com. géol. Leningrad
46 : 245-268, pls 12-14. In Russian, with English summary.
1933a. Contribution to the knowledge of the Lower Carboniferous bryozoa of Turkestan.
Trudy uses. geol-vazv. Obed. NKTP Leningrad 207 : 1-26, pl. 12. In Russian, with
English summary.
1933b. Stratigraphic range of the Carboniferous bryozoa of the U.S.S.R. Tvrudy vses.
geol.-vazv. Obed. NKTP Leningrad 268 : 1-32. In Russian, with English summary.
1938a. Types of Carboniferous bryozoa in the European part of the U.S.S.R. Akad.
Nauk. S.S.S.R., Inst. Paleontology, Paleontology U.S.S.R. 4 : 1-290, 55 pls. In Russian,
with English summary.
1938b. Stratigraficheskoe raspredelnie mshanok v neftenosnykh rifovykh isvestnyakakh
Ishimbaevskogo raiona. Leningr. vses. nauchno-issled. geol.-vazv. Inst. (Ser. A) 101 : 76-89.
In Russian.
Oakey, K. P. 1948. In Muir-Wood, H. M.: Malayan Lower Carboniferous fossils. Brit.
Mus. (Nat. Hist.) : 1-118, 17 pls, London.
D’ORBIGNY, A. 1850. Pyvodvome de paleontologie stratigraphique, 3 vols : LX and 394; 1-427;
1-196, Paris. In French.
OswaLp, D.H. 1955. The Carboniferous rocks between the Ox Mountains and Donegal Bay.
Q. Jl geol. Soc. Lond., London 111 : 167-186.
PHILLIPS, J. 1836. Illustrations of the geology of Yorkshive, Pt II : XII and 185, 14 pls,
London.
PRANTL, F. M. 1934. Carboniferous bryozoa from Dobsina (Slovakia). Bull. int. Acad.
Sci. Prague 35 : 225-243, pls 1-2.
Prout, H. A. 1859. Third series of descriptions of bryozoa from the Paleozoic rocks of the
western states and territories. Tvans. Acad. Sci. St Louis, St Louis, Mo. 1 : 443-452,
pls 17-18.
Ross, J. P. 1961. Ordovician, Silurian and Devonian bryozoa of Australia. Bull. Miner.
Resour. Surv. Aust., Canberra 50 : 1-171, pls 1-28.
SARLE, C. J. Ig01. Reef structures in Clinton and Niagara strata of western New York.
Am. Geol., Minneapolis 28 : 282-299.
SCHWARZACHER, W. 1961. Petrology and structure of some Lower Carboniferous reefs in
north-western Ireland. Bull. Am. Ass. petrol. Geol., Chicago, etc. 45 : 1481-1503.
— 1963. Orientation of crinoids by current action. J. sedim. Petrol., Tulsa, Okla. 33 :
580-586.
SuisHova, N. A. 1952. Podmoskovnye i dono-medveditzkie kamennougol’nye mshanki
roda Septoporva. Trudy paleont. Inst., Moskva 40, 159-176, pls I-V.
SHRUBSOLE, G. W. 1879. A review of the British Carboniferous Fenestellidae. Q. Jl geol.
Soc. Lond., London 35 : 275-281.
1880. A review and description of the various species of British Upper Silurian Fenestelli-
dae. Q. Jl geol. Soc. Lond., London 36 : 241-254.
1881. Further notes on the Carboniferous Fenestellidae. Q. Jl geol. Soc. Lond., London
37 : 178-189.
SHULGA-NESTERENKO, M. I. 1941. Lower Permian bryozoa of the Urals. Akad. Nauk. S.S.S.R.,
Paleont. Inst., Paleont. U.S.S.R. 5 : 1-226, 67 pls. In Russian, with English summary.
1951. Kamennougolnye fenestellidy Russkoi platformy. Akad. Nauk. S.S.S.R., Paleont.
Inst. 32 : 1-61, 34 pls. In Russian.
1955. Kamennougolnye mshanki Russkoi platformy. Akad. Nauk. S.S.S.R., Paleont.
Inst. 57 : 1-207, 32 pls. In Russian.
FROM FERMANAGH, IRELAND 493
Simpson, G. B. 1895. A handbook of the genera of the North American Paleozoic bryozoa.
A. Rep. State Geol. (N.Y.) for 1894 : 407-608.
Simpson, I. M. 1954. The Lower Carboniferous stratigraphy of the Omagh syncline, Northern
Ireland. Q. Jl geol. Soc. Lond., London 110 : 391-408.
SPJELDNAES, N. 1957. The name Fenestella. J. Paleont., Tulsa, Okla. 31 : 675-676.
Stacy, L. W. 1936. Correlation of zoarial form with habitat. J. Geol., Chicago, etc. 44 :
60-65.
STUCKENBERG, A. 1895. Korallen and bryozoen der steinkohleablagerungen des Urals USW.
Mém. Com. géol. St Petersburg 10 : 1-244, 24 pls. In German.
TAVENER-SMITH, R. 1965. A new fenestrate bryozoan from the Lower Carboniferous of
County Fermanagh. Palaeontology, London 8 : 478-491, pl. 66.
—— 1966a. Ovicells in fenestrate cryptostomes of Visean age. J. Paleont., Tulsa, Okla.
40 : 190-198, pl. 25.
1966b. The micrometric formula and the classification of fenestrate cryptostomes.
Palaeontology, London 9 : 413-425.
1969. Skeletal structure and growth in the Fenestellidae (Bryozoa). Palaeontology,
London 12 : 281-309, pls 52-56.
1971. Morphology and significance of unusual frontal features in the bryozoan Polypora
stenostoma. Palaeontology, London 14 : 178-187, pl. 25.
TuHeEopoRIvicH, G.I. 1941. Migration of Upper Paleozoic reefs in south Bashkiria. Akad.
Nauk. S.S.S.R. Doklady 32 : 499-502. In Russian, with English summary.
Toots, H. 1951. Uber einige cryptostome bryozoen aus dem Mitteldevon des Rheinischen
Scheifergebirges. N. Jb. f. Geol. u. Pal., Abh. Bd. 93 : 233-246. In German.
Trizna, V. 1939. New species of the Upper Paleozoic Fenestellidae and Acanthocladiidae
from the Bashkirian Urals. Geol. Oil Inst. Tvans. (sey. A.) 115 : 102-144. In Russian,
with English summary.
Urricu, E.O. 1886. Descriptions of new Silurian and Devonian fossils. Conty. Am. Paleont.
Cincinnati 1 : 3-35.
1890. Paleozoic Bryozoa. Bull. geol. Surv. Illinois 8, Vol. 1 : 283—688; Vol. 2: pls
29-78.
Vine, G. R. 1879. On Palaeocoryne and its development on Fenestella. Sci. Gossip, London
15 : 225-229 and 247-249.
Waters, A.W. 1896. On Mediterranean and New Zealand Reteporae and fenestrate bryozoa.
J. Linn. Soc. (Zool.), London 25 : 255—271.
WHIDBORNE, G. F. 1895. A monograph of the Devonian fauna of the South of England.
Palaeontogr. Soc., Vol. 2 : i-xi, 1-219, 24 pls, London.
Wuittincton, H. B. & Evitt, W. R. 1953. Silicified Middle Ordovician trilobites. Mem.
geol. Soc. Am., Washington 59 : 1-137, 33 pls.
Youne, J. & Younc, J. 1874. On Palaeocoryne and other polyzoal appendages. Q. Jl. geol.
Soc. Lond., London 30 : 684-689.
R. TAVENER-SM1TH, Ph.D.
Dept. of Geology
UNIVERSITY OF NATAL,
KING GEORGE V AVENUE,
DURBAN, SOUTH AFRICA.
PLADE 1
Fenestella frutex M’Coy
Fic. 1. Obverse of large zoarial fragment, with an encrusting stenoporid colony.
PD.5001.* 2:9.
Fics 2, 3, 5,6. Obverse of small zoarial fragments to illustrate common variations in mesh-
work appearance. PD.5002 to PD.5005, x6-0; 5:0; 6:7; 5-6, respectively.
Fic. 4. Stout, barbed spines from the obverse surface to the meshwork. PD.5006. 4:5.
Fics 7, 9. Detail of the obverse surfaces of morphological variants within this species.
PD.5005, 18-0; and PD.5002, x2I-o.
Fic. 8. Reverse side of a zoarial fragment showing longitudinal ‘striae’ and minute nodes.
PD.5007. x5°8.
* The serial numbers of specimens refer to the Bryozoa catalogue numbers of the collections of the
British Museum (Natural History), where the material is stored.
I
PLATE
Bull. By. Mus. nat. Hist. (Geol.) 23, 7
, co ee te
ie ee
_P
a
-
> =, te Fa
EAI
Fenestella ivanovi Shulga-Nesterenko
Fics 1, 3, 5, 7. Obverse of zoarial fragments to show variations in meshwork pattern.
PD.5008, 6-0; and PD.4682 to PD.4684, «5:0; 5:3; 5:0, respectively.
Fic. 2. Proximal part of a zoarium with basal holdfast and lateral supports; the last
developed from marginal branches and dissepiments. PD.4685. 5:2.
Fic. 4. Zoarial fragment with elongate spinose structures which are ‘infertile’ branch
continuations. PD.4686. 95:5.
Fic. 6. Delicate spines with minute barbs growing from the obverse sides of branches.
PD.4687. 5:0.
Fic. 8. Reverse surface of zoarial meshwork. PD.4684. 17:0.
Fic. 9. Detail of obverse side showing the distinctive shape of fenestrules. PD.4685.
< 16:0.
PLATE
Bull. By. Mus. nat. Hist. (Geol.) 23, 7
PLATE 3
Fenestella cf. multispinosa Ulrich
Fic. 1. Reverse of zoarial meshwork showing longitudinal ‘striae’ and nodes on the mid-line
of branches. PD.4688. 5:7.
Fic. 2. Another view of the reverse surface. PD.4689. 5-0.
Fics 3, 7. Obverse of zoarial fragments with slightly different meshwork characteristics.
PD.4689, x 5:0; and PD.4690 7:1, respectively.
Fic. 4. Obverse side showing normal branches passing distally into sterile lateral processes.
JID) AGeie, SK FD
Fics 5, 8. Obverse of meshwork showing localized development of thick, striated, secondary
skeleton. PD.4692, 8-0; and PD.4693, 7-0.
Fic. 6. Reverse of zoarial fragment showing branches passing distally into stout, barbed,
lateral spines. PD.4694. 5°4.
Fic. 9. Detail of obverse surface. PD.4680. x 18-3.
PLATE 3
23,7
Bull. By. Mus. nat. Hist. (Geol.)
PLATE 4
Fenestella modesta Ulrich
Fics 1, 2, 4, 6. Obverse of zoarial fragments illustrating variation in meshwork pattern.
PD.4696, «6-6; PD.4698, «5:0; PD.4695, «6:8; and PD.4697, 6-4, respectively.
Fic. 3. Detail of obverse surface. x 25-0.
Fic. 5. Reverse of zoarial fragment. PD.4695. 6:8.
Fenestella hemispherica M’Coy
Fic. 7. Obverse of zoarial fragment. PD.4699. x 13°5.
Fic. 8. Reverse surface showing uniserial nodes along mid-lines of branches. PD.4704.
46:3.
Fic. 9. Large fragment showing typical meshwork pattern. PD.4701. 4:7.
PLATE 5
Fenestella hemispherica M’Coy
Fics 1, 2. Obverse surface of zoarial fragments. PD.4700, x15:0; and PD.4702, x6-6.
Fic. 3. Reverse side showing nodes. PD.4699. «58.
Fic. 4. Obverse of meshwork showing characteristic mode of branch division. PD.4704.
« 13°8.
Fenestella parallela Hall
Fic. 5. Reverse of zoarial fragment showing striated pattern. PD.4710. 7:3.
Fics 6, 8, 9. Obverse surfaces showing meshwork characteristics. PD.4707 to PD.4709
X70; X7:0; 9:7, respectively.
Fic. 7. Detail of obverse side. PD.4708. 14°5.
PLATE 5
/
Bull. Br. Mus. nat. Hist. (Geol.) 23,
* ;
=
La)
nd
—_
—~ @
.
BA
1D
PAIN AILIS, (6)
Fenestella rudis Ulrich multinodosa subsp. nov.
Fic. 1. Detail of obverse of a zoarial fragment. 14:2.
Fic. 2. Obverse of meshwork with localized presence of thick, secondary skeleton which has
coated branch surfaces and reduced or sealed zooecial apertures. PI).4714 (paratype). 8-8.
Fic. 3. Reverse side of zoarial fragment. PD.4714 (paratype). 8-2.
Fics 4, 6, 7. Obverse of fragments showing slight differences of meshwork pattern.PD.4713
(paratype), 5:0; PD.4716 (paratype), «7-4; and PD.4712 (holotype), 5-0.
Fic. 5. Obverse of partly silicified fragment showing hemi-hexagonal zooecial chambers.
PD.4715 (paratype). 7-9.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 7 PLN ALLE, (6)
PALAIS, 7
Fenestella plebeia M Coy
Fic. 1. Obverse of zoarial fragment showing meshwork pattern. PD.4719. 3-6.
Fics 2, 3. Detail of obverse surface. Fig. 3, from proximal part of colony shows thick
secondary skeletal envelope. Minute punctae along branch medial lines are carinal node bases.
PD.4719, X 11-4; and PD.4717, x 11-3, respectively.
Fic. 4. Obverse of zoarial fragment from proximal part of colony. PD.4717. x 4:5.
Fic. 5. Obverse of partly silicified fragment showing triangular zooecial chambers.
PD.4718. 6-0.
Fic. 6. Zoarial fragment with inflated peristomes projecting into fenestrules. PD.4721.
x 6-4.
Fic. 7. Reverse surface of zoarial fragment. PD.4717. 4:5.
Fenestella ci. arthritica Phillips
Fic. 8. Detail of obverse surface. PD.4722. 15:0.
Fic. 9. Large zoarial fragment giving an impression of the colonial growth form.
PIDFA72 8. ez:
PLATE
23,7
ul. Hist. (Geol.)
Bull. Br. Mus. ne
PAA aS:
Fenestella ci. arthritica Phillips
Fics 1, 3, 4. Obverse of zoarial fragments showing characteristics of meshwork pattern.
Stout carinal nodes are visible along proximal part of middle branch in Fig. 3. PD.4727,
x 3:9; PD.4726, x5:0; and PD.4724, 5:1. respectively.
Fic. 2. Reverse surface of meshwork. PD.4727. 3:9.
Fenestella praemagna Shulga-Nesterenko
Fic. 5. Proximal part of a colony with basal holdfast and lateral supporting struts.
PD.4732. 5-0.
Fic. 6. Strong carinal nodes, which divide at their distal ends, originate along the obverse
mid-lines of branches. PD.4735. 5:1.
Fic. 7. Proximal part of a colony attached to a Pennivetepova branch. PD.4733. X47.
Fic. 8. Obverse of colony showing basal holdfast and supporting processes. PD.4734.
x 10-4.
Fic. 9. Detail of obverse. PD.4731. 14-0.
Fic. 10. Reverse side of zoarial fragment. PD.4730. xO6:1.
Fic. 11. Obvyerse of proximal part of an old colony, with thick secondary skeletal cover.
PID AG PR). >< 7/50)
Bull. Br. Mus. nat. Hist. (Geol.) 23, 7 PLATE 8
PLATE 9
Fenestella fanata Whidborne carrickensis subsp. nov.
Fic. 1. Obverse of meshwork showing characteristic pattern at branch division. PD.4736
(holotype). 10-1.
Fic.2. Zoarial fragment with stout lateral spine developed as a branch continuation.
PD.4737 (paratype). 35.
Fics 3, 5, 7, 8. Obverse of zoarial fragments, showing general meshwork characteristics.
Specimen shown in Fig. 3 (x3:9) is not preserved. The others are numbered: PD.4738
(paratype), x 5:0; PD.4739 (paratype), x 3-8; and PD.4736, x 3:5, respectively.
Fic. 4. Zoarial fragment with ovicells visible in the distal part. PD.4740 (paratype). 3:5.
Fic. 6. Reverse side of a fragment, showing evidence of damage and subsequent repair
during the life of the colony. PD.4741. 3:3.
Fic. 9. Detail of obverse surface. PD.4742 (paratype). 9:9.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 7 PLATE 9
PLATE io
Fenestella cf. spinacristata Moore
Fic. 1. Large zoarial fragment showing overall meshwork pattern. PD.4743. 3:8.
Fic. 2. Reverse surface of meshwork. PD.4746. 5-0.
Fic. 3. Detail of obverse side. PD.4743. 49:5.
Fic. 4. Proximal part of a colony with holdfast attached to a Pennivetepova branch.
PD.4747. X61.
Fic. 5. Obverse of zoarial fragment. PD.4744. x7'1.
Fic. 6. Fragment bearing evidence of structural damage and repair during the life of the
colony. PD.4750. 4:5.
Fenestella cf. funicula Ulrich
Fic. 7. Reverse side of fragment from proximal part of a colony, showing a stout spine
developed from obverse mid-line of a branch. PD.4758. 5°5.
Fic. 8. Obverse of a zoarial fragment. PD.4751. 95:0.
Fic. 9. Reverse side of above. PD.4751. 5:0.
Fic. 10. Fragment from proximal part of an old colony showing thick secondary skeletal
encrustation. PD.4759. 2:9.
PLATE 11
Fenestella cf. funicula Ulrich
Fic. 1. Obverse of zoarial fragment showing ovicells in the distal part. PD.4759. 4:3.
Fics 2, 3. Obverse surfaces showing general meshwork characteristics. PD.4753, 6:8;
and PD.4755, X7:0.
Fic. 4. Obverse surface of meshwork with strong carinal nodes. PD.4756. x 4:0.
Fenestella filistriata Ulrich
Fics 5,7, 9. Obverse of zoarial fragments showing meshwork pattern.
PD.4763, 5:3; PD.4761 x3:8 and PD.4762, x5.0, respectively.
Fic. 6. Reverse surface. PD.4763. 5:5.
Fic. 8. Reverse of zoarial fragment showing longitudinal ‘striae’ and hemi-hexagonal bases
of zooecial chambers. PD.4764. 7-0.
Fic. 10. Detail of obverse.. PD.4761. 12:2.
IIL INARI
Fenestella cf. filistriata Ulrich
Fic. 1. General view of obverse side of a zoarial fragment. PD.4761. 5-2.
Fenestella subspeciosa Shulga-Nesterenko
Fic. 2. Reverse surface. PD.4766. 5:7.
Fics 3, 4, 5. Obverse of zoarial fragments showing general meshwork characteristics.
PD.4765, <5:0; PD.4769, x 4:0; and PD.4767, 5-6.
Fic. 6. Obverse at higher magnification, showing prominent cowl-like peristomes indenting
fenestrule margins. PD.4766. 5-2.
Fic. 7. Reverse of zoarial fragment showing a stout spine which resulted from continued
growth from an abnormally directed dissepiment. This spine later re-united with the mesh-
work, as shown. 7:2.
Fic. 8. Obverse of above fragment showing a local concentration of secondary skeletal
tissue where the aberrant spine (coming from below) rejoined the meshwork. 7-2.
PLATE 13
Fenestella pseudovirgosa Nikiforova
Fic. 1. Detail of obverse. PD.4770. 15:0.
Fic. 2. Reverse side of meshwork. PD.4771. 2:9.
Fic. 3. Zoarial fragment showing general meshwork characteristics. PD.4770. 5:0.
Fic. 4.. The stout spine developed from a branch end in the distal part of this specimen
EID 72, KB),
grew back and re-united with the reverse side of the meshwork.
Fic. 5. This strong spine resulted from the continued growth of an abnormal dissepiment.
PD.4773. X3°7-
Fics 6, 9. Zoarial fragments from proximal parts of colonies, showing thick deposits of
secondary skeletal substance. PD.4774, «5:6; and PD.4775, 5:0.
Fic. 7. Obverse of fragment, showing a strong laterally directed spine. PD.4776. 4:1.
Fic. 8. Reverse surface showing longitudinal ‘striae’, variably developed nodes, and the
triangular base of a zooecial chamber. PD.4777. 4:3.
PA AGE a4:
Fenestella cf. albida Hall
Fics 1, 3. Obverse of zoarial fragments showing general meshwork characteristics.
PD.4783, x 6-7; and PD.4780 x 5:0, respectively.
Fic. 2. Obverse surface showing chain-like pattern due to projection of inflated peristomes
into fenestrules. PD.4781. 6-4.
Fic. 4. Detail of obverse. PD.4780. x 15-0.
Fic. 5. Proximal part of a colony with basal attachment to a Penniveteporva branch.
PD.4779. X3°4-
Fic. 6. Reverse side of meshwork. 5:8.
Fenestella oblongata Koenig
Fic. 7. Obverse of a zoarial fragment. PD.4790. 7-0.
Fic. 8. Reverse surface showing longitudinal ‘striae’, small nodes and the triangular base of a
zooecial chamber. PD.4791. 7°5.
PLATE 15
Fenestella oblongata Koenig
Fic. 1. Detail of obverse of zoarial fragment. PD.4786. 14:8.
Fics 2, 3, 4. Obverse views of fragments showing general aspect of meshwork.
PD.4787, 5:0; PD.4789, x 4:2; and PD.4786, x 6-3, respectively.
Fenestella cf. delicatula Ulrich
Fics 5, 6. Obverse surfaces of zoarial fragments illustrating variation in meshwork charac-
teristics. PD.4797, x6:3; and PD.4794, x7-0.
Fic. 7. Zoarial fragment with strong spines from the reverse side of a branch. The spines
bear minute barbs. PD.4799. 5:0.
Fic. 8. Detail of obverse side. PD.4798. x 16-0.
15
PLS NAE Led
/
(Geol.) 23,
Mus. nat. Hist.
Bull, Br.
PLATE 16
Fenestella cf. delicatula Ulrich
Fic. 1. Reverse side of meshwork. PD.4798. 7-5.
Fic. 2. Obverse of zoarial fragment. PD.4795. 4:8.
Fic. 3. Fragment with strong lateral spine developed asa branch continuation. PD.4798.
K 7:5.
Fenestella polyporata (Phillips)
Fic. 4. Reverse of meshwork showing longitudinal ‘striae’. PD.4802. 7-2.
Fics 5, 7, 8. Obverse surfaces of zoarial fragments showing general features of meshwork.
PD.4805, x 7:0; PD.4802, «3:7; and PD.4804, 5-0, respectively.
Fic. 6. Detail of obverse. PD.4804. <14°9.
Bull. Br. Mus. nat. Hist. (Geol.) 23, 7 PLATE 16
PA ASE, 7
Fenestella polyporata (Phillips)
Fic. 1. Obverse of zoarial fragment.
PAD SOR, <7
Fenestella irregularis Nekhoroshev
FIGS 2, 5.
Obverse of zoarial fragments, showing meshwork features.
and PD.4811, x 3:9, respectively.
PDYASesx< 7-1
Fic. 3. Detail of obverse. PD.4810.
x 13:0.
Fic. 4. Reverse side of zoarial fragment. PD.4809. 7:5.
Fic. 6. Obverse of incompletely silicified fragment showing casts of zooecial chambers
PD.4813. x85.
Fic. 7.
Oblique view of obverse showing strong carinal nodes.
PD.4812. x 4°5.
Levifenestella undecimalis (Shulga-Nesterenko)
Fic. 8. Reverse surface with longitudinal ‘striae’ and grooves excavated by ramifying
Condvanema (ctenostome) stolons. PD.4815. 5:0.
Fic. 9. Detail of obverse. PD.4816. x 16-0.
ATE
IPL
Mus. nat. Hist. (Geol.) 23, 7
Bull. Br.
PATE aS
Levifenestella undecimalis (Shulga-Nesterenko)
PD.4817, x6:6; PD.4814, 4-0; and
PD A816 2 << 7-3:
FIGS 1, 2, 4. Obverse surfaces of zoarial fragments.
PD.4820, x 6-2, respectively.
Reverse of meshwork showing closely spaced longitudinal ‘striae’.
Obverse of large zoarial fragment which gives an idea of the colonial growth form.
Fia. 3.
FIG. 5.
PD.4818. x 2:3.
Minilya plummerae (Moore)
Fic. 6. Small fragment with a stout, barbed spine developed from the reverse surface.
RD A8275 Os:
Obverse of zoarial fragments showing general aspect of meshwork. PD.4825,
Fics 7, 8.
x 6:0; and PD.4824, x7-5, respectively.
PLATE x73
23,7
(Geol.)
Bull. Bry. Mus. nat. Hist.
Pe ACs. 1G)
Minilya plummerae (Moore)
Fic. 1. Detail of obverse of meshwork. Positions of carinal nodes are indicated on a part
of one branch. PD.4823. 22-0.
Figs 2, 4. Obverse of zoarial fragments, showing general meshwork characteristics.
PD.4823, 6-6; and PD.4822, 5-0, respectively.
Fic. 3. Reverse surface. PD.4822. 5:0.
Minilya binodata (Condra)
Fics 5, 7. Obverse of zoarial fragments. PD.4829, «7-3; and PD.4848, 8-5.
Fic. 6. Reverse surface. PD.4828. 5:8.
PEALE, 19
uw. Hist. (Geol.)
Bull. Br. Mus. ne
>
&
ie
6 = = > . 3
* cs 2.
4 > -o: 6.5. J ¥,' a
ei ‘ } sy i
‘
Finny: Hh en CS s é a ¢
5 er S 2
) t re x ~~
b ~ :
A , < A aS s
- ee ad $,' : >
2%
:
a
weeeie
PEALE 20
Minilya binodata (Condra)
Fic. 1. Detail of obverse, with the positions of some carinal nodes indicated. PD.4828.
x 16°5.
Minilya oculata (M’Coy)
Fic. 2. A fragment from the proximal part of an old colony, with thick secondary skeletal
investment. PD.4833. 6:2.
Fic. 3. Detail of obverse showing the biserial arrangement of carinal nodes. PD.4838.
If)
Fics 4, 6, 8. Obverse of zoarial fragments, showing meshwork pattern and growth habit.
PD.4834, «5:0; PD.4837, «6-4; and PD.4835. x 6-9, respectively.
Fic. 5. Reverse side of zoarial fragment. PD.4839. 6:5.
Fic. 7. Proximal part of a colony with holdfast attached to a Rhabdomeson fragment.
PD.4832. x 3:9.
PLATE
AB) 7)
t. Hist. (Geol.)
sy. Mus. na
Bull. I
PACE 2a
Ptiloporella varicosa (M’Coy)
Fics 1, 2, 3, 6, 7. Obverse of zoarial fragments showing prominent development of primary
and secondary branches. PD.4840 to PD.4844 inclusive, x 4:0; X4:-7; X5:0; X2:6; X2°7,
respectively.
Fic. 4. This specimen, with a midrib and symmetrically placed lateral branches, resembles
Ptylopora M’Coy in its growth pattern. PD.4845. 4-6.
Fic. 5. Small fragment with strong spinose structures which originate on the obverse mid-
line. PD.4846. 4:3.
Fic. 8. Reverse Surface, showing the relationship between primary and secondary branches.
IDFA SAV 30:
Fic. 9. Detail of 6bverse. PD.4842. 135.
PAA TE
23,7
(Geol.)
uw. Hist.
Bull. Br. Mus. ne
PLATE 22
Hemitrypa hibernica M’Coy
Fic. 1. Proximal part of a colony attached to a Rhombopora fragment. Ends of supporting
spines are visible. PD.4848. 3:2.
Fic. 2. Reverse surface of zoarial fragment showing a secondarily thickened ‘dissepimental
ALC ED PASAG sass:
Fic. 3. Proximal part of a colony with enclosing superstructure and stout supporting
spines. PD.4850. 5:0.
Fic. 4. Obverse side with superstructure removed. PD.4851. 5-0.
Fic. 5. Obverse with superstructure partly removed to show abnormal branching pattern,
possibly a pathological condition. PD.4852. 5:0.
Fic. 6. Lateral view showing superstructure supported by carinal pillars. Underside of
meshwork is encrusted by a Fistulipova colony. PD.4853. 6-5.
Fic. 7. Reverse surface with distal parts of a long, branched supporting spine which re-
unites with the colonial meshwork. PD.4854. 3:2.
Fic. 8. Obverse of a large fragment, with young Fistulipora colony encrusting the super-
structure. PD.4855. 3:0.
Fic. 9. Obverse of meshwork showing ovicellular concavities. PD.4856. 4-4.
Fic. 10. Detail of obverse. PD.4857. x 18-0.
PLATE
2B
(Geol.)
nat. Hist.
Bull. By. Mus.
~?
a@@e@egee e+e @
eee @
‘¢ ,
ENC PNICIS 23}
Polypora dendroides M’Coy
Fics 1,5. Obverse of zoarial fragments from the proximal parts of colonies. Thick secondary
skeletal tissue is particularly evident at the bases of broken supporting spines, x 5-4; PD.4865,
and PD.4866, x 2-6.
Fics 3, 4. Reverse of meshwork. Fig. 4 shows the proximal parts of supporting spines.
PD.4867, 5-0; and PD.4868 x 2-6.
Fics 2, 6. Obverse surfaces showing general characteristics of meshwork. PD.4867, x 5-0;
and PD.4869, x 4:3.
Fic. 7. Detail of obverse side. PD.4867. 13-0.
Bull, Br. Mus. nat. Hist. (Geol.) 23, 7 PLATE 23
/
PLATE 24
Polypora verrucosa M’Coy
Fic. 1. Detail of obverse. PD.4870. x12°8.
Fics 2, 3,5, 7. Obverse of zoarial fragments showing general features of meshwork and branch
surface. PD.4872, x 3-4; PD.4873, x3:8; PD.4870, x6-6; and PD.4874, x6:2, respectively.
Fic. 4. Fragment from proximal part of a colony, showing prominent peristomes with
terminal apertures, some of which are sealed by the thick secondary skeletal investment.
PD.4871. 6-6.
Fic. 6. Reverse side showing incipient beekitization resulting from the silicification of thick
secondary skeletal accretions. PD.4877. 6-0.
>
Mus. nat. Hist. (Geol.)
Bull. Br.
PLATE 25
Ptylopora pluma M’Coy parva subsp. noy.
Fic. 1. Detail of obverse showing midrib and lateral branches. PD.4880 (paratype).
< 18-6.
Fic. 2. Reverse surface with prominent ‘striae’. PD.4880 (paratype). 1.056:
Fics 3, 5, 8, 10. Obverse of zoarial fragments showing general meshwork characteristics.
PD.4878, «5:0; PD.4879, x 6:0; PD.4881 and PD.4882, 4:1; 4-1 (all are paratypes).
Fic. 4. Proximal part of a colony with thick outer secondary investment. The stalk-like
holdfast has subsidiary attachment structures at its lower end. 5-0.
Fic 6. Midrib of a colony with vestiges of secondary branches. PD.4883 (holotype). x 2-7.
Fics 7, 9. Reverse sides of zoarial fragments. Fig. 9 shows a short lateral spine developed
from a branch end. PD.4884 and PD.4882 (paratype) respectively. Both x 4-1.
PLATE
7
23,
(Geol.)
Bull. By. Mus. nat. Hist.
=F 2 © © & @& o>
ik ia a wm \S F
PLATE 26
Septopora hibernica sp. nov.
Fics 1, 7, 8. Obverse of zoarial fragments. In many places branch surfaces are obscured
and zooecial apertures sealed by thick secondary skeletal deposits. PD.4885 (holotype),
«2:2; PD.4886 (paratype), x 6-1; and PD.4887 (paratype), x 3-6.
Fic. 2. Reverse surface of meshwork. The difference between primary and secondary
branches is very marked. PD.4885 (holotype). x 2:2.
Fic. 3. Obverse with thick investment of secondary skeleton. The ‘striae’ traverse both
branches and dissepiments when traced away from the bases of large spines. PD.4889
(paratype). 4:5.
Fic. 4. Fragment with Ptylopora-like meshwork of midrib and lateral branches.
PD.4888 (paratype). 1-9.
Fic. 5. Zoarial fragments with irregular growth habit. PD.4890 (paratype) and PD.4891.
X4°5-
Fic. 6. Reverse surface showing thick deposits of secondary skeleton. PD.4889 (para-
type). 6-4.
/
Mus. nat. Hist. (Geol.) 23,
Bull. Br.
A LIST OF SUPPLEMENTS
TO THE GEOLOGICAL SERIES
OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya.
Pp. 213; 30 Plates; 2 Text-figures. 1965. 6.
. Et-NaGcGar, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper
Cretaceous—Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley,
Egypt, U.A.R. Pp. 291; 23 Plates; 18 Text-figures. 1966. {1to.
. Davey, R. J., DowntE, C., SARGEANT, W. A. S. & WitiiaMs, G. L. Studies on
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248; 28 Plates; 64 Text-
figures. 1966. £7.
. APPENDIX. Davey, R. J., DowntE, C., SARGEANT, W. A. S. & WILLIAMS, G. L.
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24.
1969. 8op.
. Ertrott, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of the
Middle East. Pp. 111; 24 Plates; 17 Text-figures. 1968. 5.124.
. Ruopes, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni-
ferous) Conodont faunas, and their value in local and continental correlation.
Pp. 315; 31 Plates; 92 Text-figures. I969. II.
. Cuitps, A. Upper Jurassic Rhynchonellid Brachiopods from Northwestern
Europe. Pp. 119; 12 Plates; 40 Text-figures. 1969. £4.75.
. Goopy, P. C. The relationships of certain Upper Cretaceous Teleosts with
special reference to the Myctophorids. Pp. 255; 102 Text-figures. 1969. £6.50.
. OwEN, H. G. Middle Albian Stratigraphy in the Paris Basin. Pp. 164;
3 Plates; 52 Text-figures. 1971. £6.
. Sippigur, Q. A. Early Tertiary Ostracoda of the family Trachyleberididae
from West Pakistan. Pp. 98; 42 Plates; 7 Text-figures. 1971. {8.
Ber Printed in England by Staples Printers Limited at their Kettering, Northants establishment
SAS ere
rer
C9 t Chee
.
ANE ce
ene ee
Rai ecae ata
ty at #4,
y So
ante
sagas
pateeaticgeas
Heit es
ata t
Re roe tie ; Ear ahr acer ary
aa fwiets eat gtg lt ea ae:
sledge tits
nine Seles K Roath: ais x
; 4, ate atid eel)
as
ah
ateeat
fe
~
4
edhe Viton s
timed ae)
phase!
PRISM
tate rite®
eee
++
+
+
+
*
we
ne
eye
‘4
fe i
ane
esety;
shes
x
nn
te