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Full text of "The vertebrate fauna of the Selma Formation of Alabama"

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THE VERTEBRATE FAUNA OF THE 
SELMA FORMATION OF ALABAMA 

PART VII 
THE MOSASAURS 

DALE A. RUSSELL 



PART VIII 
THE FISHES 

SHELTON P. APPLEGATE 



s 




FIELDIANA: GEOLOGY MEMOIRS 

VOLUME 3, NUMBERS 7 AND 8 

Published by 

FIELD MUSEUM OF NATURAL HISTORY 

FEBRUARY 12, 1970 



Geo*. 



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THE VERTEBRATE FAUNA OF THE 
SELMA FORMATION OF ALABAMA 

PART VIII. THE FISHES 



THE VERTEBRATE FAUNA OF THE 
SELMA FORMATION OF ALABAMA 

PART VIII 
THE FISHES 

SHELTON P. APPLECxATE 

Associate Curator of Vertebrate Paleontology 
Los Angeles County Museum of Natural History 



FIELDIANA: GEOLOGY MEMOIRS 

VOLUME 3, NUMBER 8 

Published by 

FIELD MUSEUM OF NATURAL HISTORY 

FEBRUARY 12, 1970 



Library of Congress Catalog Card Number: 53-2305 

PRINTED IN THE UNITED STATES OK AMERICA 
BY FIELD MUSEUM PRESS 



CONTENTS 

PAGE 

Introduction 389 

Systematic Descriptions 389 

Holocephali 389 

Edaphodontidae 389 

Edaphodon barberi, new species 390 

Edaphodon mirificus Leidy 392 

Edaphodon sp 393 

Selachii 393 

Ptychodontidae 393 

Ptychodus polygurus Agassiz 393 

Ptychodus mortoni Mantell 393 

Anacoracidae 393 

Squalicorax falcatus (Agassiz) 393 

Squalicorax pristodontus (Agassiz) 395 

Pseudocorax affinis (Agassiz) 395 

Odontaspididae 395 

Scapanorhynchus rhaphiodon (Agassiz) 395 

Scapanorhynchus rapax (Quaas) 396 

Lamnidae 396 

Lamna appendiculata Agassiz 396 

Isurus mantelli (Agassiz) 397 

Undetermined shark denticles 398 

Actinopterygii 398 

Chondrostei 398 

Acipenseridae 398 

Propenserinae, new subfamily 398 

Propenser, new genus 398 

Propenser hewletti, new species 399 

Holostei 401 

Pycnodontidae 401 

Hadrodus priscus Leidy 401 

Incertae sedis 401 

Pachycormidae 403 

Protosphyraena Leidy 404 

Protosphyraena nitida ? (Cope) 404 

Protosphyraena sp. 1 404 

Protosphyraena sp. 2 404 

Protosphyraena sp. 3 404 

Teleostei 404 

Elopoidei 405 

Elopoidea 405 

Elopidae 405 

Palelops, new genus 405 

Palelops eutawensis, new species 406 

Pachyrhizodontoidea, new superfamily 406 

Pachyrhizodus Dixon 406 

Pachyrhizodus minimus Stewart 408 

Pachyrhizodus caninus Cope 410 

Pachyrhizodus kingi Cope 411 

385 



386 CONTENTS 

PAGE 

Albuloidea 411 

Albulidae 412 

Albula dunklei, new species 412 

Albula sp 413 

Plethodoidea, new superfamily 413 

Bananogmiidae, new family 413 

Bananogmius crieleyi, new species 414 

Bananogmius cf. zitteli (Loomis) 416 

Bananogmius cf. polymicrodus (Stewart) 416 

Bananogmius sp 416 

Moorevillia, new genus 416 

Moorevillia hardi, new species 416 

Clupeoidei 416 

Chirocentroidea, new superfamily 416 

Ichthyodectidae 418 

Ichthyodectes cf. ctenodon Cope 418 

Xiphactinus audax Leidy 418 

Saurodontidae 419 

Saurodon leanus Hays 419 

Saurodon ? sp. 1 420 

Saurodon 1 sp. 2 420 

Saurocephalus cf. lanciformis Harlan 420 

Clupeoidea 420 

Clupeidae indet 420 

Dussumieriinae indet 420 

Myctophoidei 420 

Enchodontidae 420 

Cimolichthys nepaholica (Cope) 420 

Enchodus petrosus Cope 421 

Enchodus cf. saevus Hay 421 

Dercetidae 421 

Stratodus apicalis Cope 421 

Myctophidae 421 

Myctophidae indet 421 

Beryciformes 423 

Trachichthyidae 423 

Hoplopteryx ? sp 423 

Trachichthyidae ? indet 424 

Teleost incertae sedis 1 424 

Teleost incertae sedis 2 424 

Ecology of the Mooreville Chalk 424 

The evidence from the overall geological picture 424 

Ecological evidence from the sediments 425 

Ecological evidence from the plants 425 

Ecological evidence from the invertebrates 426 

Ecological evidence from the fishes 426 

Summary 431 

References 432 



LIST OF ILLUSTRATIONS 

PAGE 

Edaphodon barberi, n. sp., holotype, medial, lateral, and dorsal views of mandible 391 

175. Edaphodon mirificus Leidy, dorsal view of left and right mandibles; Edaphobon sp., medial view of right 

mandible 392 

176. Denticles of sharks, much enlarged : Squalicorax falcatus and undetermined sharks 394 

177. Tooth of Pseudocorax affinis 395 

178. Scapanorhynchus rhaphidon, three anterior teeth; Scapanorhynchus rapax, anterior tooth; Lamna ap- 

pendiculata, three lateral teeth and one posterior tooth ; Isurus mantelli, two anterior teeth and one 

lateral tooth ; Squalicorax falactus, two lateral teeth, and Pseudocorax affinis, lateral tooth .... 396 

179. Tooth of Ptychodus polygurus, vertebral cross-section of Squalicorax falcatus, and vertebral cross-section 

of Lamma appendiculata 397 

180. Reconstruction of skull of Propenser hewletti, dorsal view 398 

181. Propenser hewletti, n. sp., holotype, specimen in collection of Alabama Geological Survey, dorsal view 

of posterior portion of skull 399 

182. Propenser hewletti, n. sp., holotype, rostral plate and body scute 400 

183. Propenser hewletti, n. sp., holotype, hyomandibular and hyoid elements 401 

184. Propenser hewletti, n. sp., holotype, fin rays, shoulder girdle, and dorsal scutes 402 

185. Propenser hewletti, n. sp., holotype, palatoquadrate and part of maxillary, lower jaw, parts of vertebrae 

and neural arches 403 

186. Hadrodus priscus, medial, dorsal, and lateral views of right mandible 404 

187. Premaxillary of possible pycnodontid 404 

188. Paleolps eutawensis, n. sp., scales 405 

189. Pachyrhizodus minimus, nearly complete fish, head and pectoral fins 406 

190. Vertebral cross-sections of Pachyrhizodus: P. minimus, P. caninus, and P. kingi 408 

191. Scales of Pachyrhizodus: P. caninus and P. minimus 409 

192. Reconstruction of Pachyrhizodus minimus 410 

193. Pachyrhizodus caninus, pectoral girdle, vomers, and portion of jaw with attached quadrate 41 1 

194. Albula dunklei, n. sp., holotype, posterior portion of ceratohyal, entopterygoid, portion of skull, and 

other parts 412 

195. Tooth-bearing plate of Albula sp 413 

196. Bananogmius crieleyi, n. sp., holotype, parasphenoid and vomer, right premaxillary with anteriorend 

up, basibranchial, abdominal vertebra, ectopterygoid, and right quadrate with anterior part up . . 414 

197. Moorevillia hardi, n. sp., holotype, left palatine, ventral view, right palatine, dorsal view, portion of 

? three branchial arches, two abdominal vertebrae, and other parts 415 

198. Stratodus apicalis, fragment of operculum, maxillary, palatine, internal view of left lower jaw, fragm ent 

of right lower jaw, and premaxillary; Bananogmius sp., parasphenoid; Bananogmius cf. zitteli, rostrum 417 

199. Jaw of ? Saurodon sp., medial view; Pachyrhizodus caninus, pectoral fin spines 418 

200. Xiphactinus audax, vertebral cross-section; Albula dunklei, vertebral cross-sections; Bananogmius crei- 

leyi, vertebral cross-section; Saurodon leanus, vertebral cross-section 418 

201. Scales: dussumieriine ?, Albula dunklei, Bananogmius cf. polymicrodus 419 

202. Scales: teleost indet, Hoploptery± ? 422 

203. Myctophid scales, four different kinds described in text 423 

204. Operculum referred to family Trachichthyidae ? 424 

387 



THE FISHES 



INTRODUCTION 

In many respects our knowledge of American Cre- 
taceous fossil fishes is still in its infancy. With the ex- 
ception of David's 1946 paper, no work of a faunal 
nature has been done on Upper Cretaceous fishes since 
Hay (1903) and Stewart (1900). Yet unstudied col- 
lections exist in many of our museums and future work 
is bound to modify, if not completely alter, some of 
our present concepts. 

It was first suggested to me by Dr. David H. Dunkle 
that the fossil fishes of the Mooreville Chalk of the 
Selma Formation might form a subject for investiga- 
tion. Field Museum already had a collection of Moore- 
ville fishes collected by Mr. Barber, Dr. William D. 
Turnbull, Mr. R. H. Hard, and Dr. and Mrs. R. Zan- 
gerl. This fine collection has formed the nucleus of the 
present study. 

Through a grant by the Bock Fund of the National 
Academy of Science and with the aid of the University 
of Chicago it was possible for me to visit the Moore- 
ville localities where I collected additional material, 
and made observations concerning the ecology of the 
Chalk. Mr. Bruce Crieley of Chicago accompanied 
me as a field assistant. The Alabama Geological Sur- 
vey, through Dr. R. Jones and Miss Winnie McGlam- 
mery, has not only given advice but has lent specimens. 
My wife, Anne Chase Applegate, has typed and as- 
sisted in the editing of the first draft. Dr. Rainer 
Zangerl and Dr. Robert H. Denison have given a great 
deal of their time editing and discussing this paper. 

The criticism of Dr. Everett C. Olson, Dr. Ralph 
G. Johnson, and Dr. Lore R. David has been valuable. 
Dr. Olson helped in securing the grant for field work. 
Dr. Lore David has confirmed many of the scale identi- 
fications and has let me have the use of her excellent 
collection of Recent teleost scales as well as her scale 
notebooks which include photos of both Recent and 
fossil teleosts. David Techter of Field Museum has 
checked the Museum numbers and has been a great 
help in numerous other ways. Miss Anita Daugherty 
has been most helpful in re-editing this paper. Mrs. 
Myrna L. Patrick has typed the final copy. It was 
first submitted in 1961 as partial fulfillment of a doc- 
toral degree in Paleozoology at the University of Chi- 
cago. 

It should be added that the Selma Formation in 
Alabama is divisible into three mappable units, from 
top to bottom the Demopolis Chalk, the Areola Lime- 



stone, and the Mooreville Chalk. 1 Since the greatest 
majority of fossil fishes have come from the lowest 
member, the conclusions in this paper involve mainly 
this unit. 

The specimens are in the collection of Field Mu- 
seum, unless otherwise specified. 

SYSTEMATIC DESCRIPTIONS 

Class HOLOCEPHALI 

Order CHIMAERIFORMES 

Suborder CHIMAEROIDEI 

Edaphodontidae Owen 

Diagnosis. — Chimaeroidei possessing paired vomer- 
ine, palatine, and mandibular dental elements. The 
beak-like mandibular element has from one to five oval 
tritoral areas per mandible, except for Elasmodus, which 
has two large tritors and numerous smaller tritoral 
areas. The palatine element has from two to four 
tritors with the exception of Ganodus, which has one 
large tritor and numerous small ones. The known vo- 
merine elements possess from six to eight tritors each. 

Discussion. — The greater proportional width and 
depth of the posterior portions of the mandibular den- 
tal elements separate the Edaphodontidae readily from 
the Lower Jurassic Squalorajidae, which have long, thin, 
and naiTow mandibular dental elements. There is no 
likelihood of confusing the Edaphodontidae with the 
known Myriacanthidae, which have mandibular dental 
elements that are shortened in a posterior direction 
but are greatly expanded laterally. Patterson (1965) 
says that in the Myriacanthidae tritors are present 
only on the upper two anterior pairs of dental elements, 
the vomers. The Callorhynchidae, of the Recent fami- 
lies, though lacking recognizable vomerine tritors, lies 
closest to the fossil Edaphodontidae in having only one 
or two tritors per mandibular element. The variable 
Chimaeridae have short, wide, and sharp mandibles 
suitable for chopping food; the tritors are smaller and 
are of a greater number than in the Edaphodontidae. 
The Rhinochimaeridae, both Recent and fossil, have 
the greatest number of small tritors of any of the exist- 
ing families and are perhaps the farthest removed from 

1 According to Keroher and others (1966), the Mooreville 
Chalk has been raised to formational rank and the old Selma 
Formation is now considered to be a group. The Areola Lime- 
stone is included in the Mooreville. However, in keeping with the 
present series I have retained the old usage in the title. 



389 



390 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



the Edaphodontidae. As can be surmised, the genera 
of the Edaphodontidae form a natural group differing 
markedly from the Recent and fossil Chimaeridae, with 
which they have been placed by most recent workers. 
The genera included are: Edaphodon, Isotaenia, Lepto- 
mylus, Ganodus, Ischyodus, and Elasmodns, all of which 
have been discussed by Woodward (1891). The genus 
Psaliodus probably also belongs in this family since 
Woodward believes that it is closely related to Elas- 
modns. 

The Edaphodontidae possess, as far as known, two 
pairs of upper dental elements: posteriorly, two elon- 
gate flattened palatine plates, and, in advance of these, 
two small vomerine elements, one of which was figured 
for Edaphodon by Hussakof (1912, p. 206, fig. 4). The 
lower mandibular dental elements are paired and are 
deep, wide, and generally robust; the over-all shape is 
reminiscent of a hawk-like beak. All the dental ele- 
ments possess one or more tritoral areas, composed of 
tubular dentine (Patterson, 1965), which are raised 
above the surrounding parts of the plate in worn teeth. 
These areas are associated with a crushing or grinding 
habit. 

Leptomylus has only one tritor per mandibular ele- 
ment; Edaphodon, four or five tritors; Ganodus and 
Ischyodus, four tritors; and Elasmodns, at least two 
large tritors and numerous small ones. The mandibular 
dental element is not known in Isotaenia. The palatine 
element in Ischyodus has four tritoral areas; in Edapho- 
don and Elasmodns, three; in Isotaenia, two. In Ganodus 
there is a palatine element reminiscent of the vomerine 
of Edaphodon, with one large tritor which is very elon- 
gate and a row of small tritors that form a U-shape. The 
palatines of Leptomylus are unknown. The vomerine 
plates of Edaphodon are narrow, with from six to eight 
small tritors. The vomerine plates of Ischyodus are 
rectangular, with six tritors in two rows. The vomerine 
teeth have yet to be found in the other genera of the 
Edaphodontidae. 

The Edaphodontidae as defined above range from 
the Jurassic to the Pliocene, probably reaching their 
greatest expansion in number and variety in the Creta- 
ceous, particularly in the Upper Cretaceous. Judging 
from the dentition, they form a homogeneous ancestral 
complex from which the modern chimaeroid families 
were derived. The Chimaeridae in the restricted sense, 
with the separation of the Edaphodontidae, consist of 
Chimaera and Hydrolagus. The former has a Creta- 
ceous to Recent record, the latter is known only from 
the Recent. However, no one has compared the denti- 
tion of these two genera. The Chimaeridae may be 
devived from the Edaphodontidae through Elasmodns. 
The Rhinochimaeridae, consisting of the Recent Rhino- 
chimaera, Harriotta, and Keoharriota, and the fossil Amy- 
lodon and Elasmodectes, appear to branch from the eda- 
phodontid stock. 

The Recent Southern Hemisphere Callorhynchidae 
appear to lie, as stated, closest to Edaphodon. They 
are of much smaller size and inhabit shallow water, 



being known to enter bays (Graham, 1956). Callor- 
hynchus is recorded from the Cretaceous of New Zea- 
land. The trend from the Edaphodontidae to the mod- 
ern families shows an over-all reduction in the size of 
the teeth, probably coinciding with a general reduction 
in body size. The crushing surfaces on the jaws are 
replaced by narrow chopping edges accompanied in 
some cases by the fragmentation of the tritoral areas. 

Edaphodon barberi, new species. Figure 174 

Diagnosis. — The lateral surface of the mandible is 
flattened. The apical tritor is the longest and is elon- 
gate-oval in shape. The antero-intermediate tritor is 
well in advance of and somewhat smaller than the pos- 
tero-intermediate tritor. Above and behind the postero- 
intermediate tritor lies the postero-dorsal tritor. which 
is about equal to the antero-intermediate tritor in area 
and shape. 

Type. — PF 290, a nearly complete left mandible 
from Donald's Farm, Dallas County, Alabama, middle 
Mooreville Chalk. 

Description and Discussion. — The dorsal border of the 
mandible in outline bears anteriorly an apical tritor 
for about one-third its length; in the middle one-third 
it rises gently; in the last one-third the dorsal border 
rises more steeply and then flattens out. The ventral 
border sweeps backward in a smooth arc. The postero- 
intermediate tritor is just under the posterior dorsal 
border. The relationship of these tritors is shown in 
Figure 174. A transverse section of the mandible would 
be approximately triangular. The symphyseal facet 
is indistinct. 

MEASUREMENTS— (in mm.). 

Greatest length of mandible 69 

Greatest thickness of mandible 19 

Length of apical tritor 21 

Width of apical tritor 4.5 

Width of antero-intermediate tritor 3.5 

Width of postero-intermediate tritor 5.5 

Width of postero-dorsal tritor 3.0 

Edaphodon barberi shows superficial similarities to 
Edaphodon stenobryus in shape, but the position of the 
tritors in E. stenobryus is completely different. The 
great length of the apical tritor plus the fact that there 
are separate intermediate tritors places E. barberi out- 
side the variants of Edaphodon mirificus as described 
by Fowler (1911) and Hussakof (1912). 

The back of the type mandible is incomplete, and 
there has been some distortion due to compression along 
with some fracturing. The waterwom appearance is 
most likely due to recent exposure, born out by the 
presence of encrusting lichens on three sides of the 
mandible. 

The species is named for Mr. CM. Barber, who 
was one of the earliest collectors of the Mooreville ver- 
tebrate fauna. 





10 mm 




Fig. 174. Edaphodon barberi, n. sp., PF 290, holotype, mandible; A, medial view; B, lateral view; C, dorsal view. 



391 



392 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 











B 



Fig. 175. A, Edaphodon mirificus Leidy, dorsal view of left and right mandibles (P 27536); B, Edaphodon sp., medial view of right 
mandible (PF 3501). 



Edaphodon mirificus Leidy. Figure 175A 

Referred specimens. — P27536, two mandibles from 
Bank's farm, Greene County, Alabama, lower Moore- 
ville or upper Eutaw. P27529, one left palatine; 
P27537, one right mandible; PF 209, one left palatine; 
all from Moore's farm, Dallas County, Alabama, middle 
Mooreville Chalk. 



Discussion. — Fowler (1911) reports this species from 
the Greensand at Barnsboro and Hornerstown, New 
Jersey, and adds that the Hornerstown Formation is 
probably Cretaceous. Loeblich and Tappan (1957) 
place the Hornerstown in the Paleocene. If they are 
correct and the Edaphodon material is not reworked, 
then this species bridges the Cretaceous-Tertiary boun- 






APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



393 



dary. The genus Edaphodon undoubtedly does bridge 
this gap, but it appears that New Jersey marine ver- 
tebrate Cretaceous or Tertiary records, particularly 
from the Greensand, are subject to question because of 
the possibility of reworking of the material into Ter- 
tiary beds and the apparent lack of precise stratigraphic 
information. The Hornerstown Formation of New Jer- 
sey and the Aquia Formation of Virginia are both 
greensands and are supposed to be Paleocene, but the 
Hornerstown fish fauna has a strong Upper Cretaceous 
aspect while the Aquia shows no such affinities. 1 



(1941). Leidy (1868) lists three Alabama specimens, 
one from Unionville, one from Perry County, and one 
from Greene County. With all these records it would 
not be surprising to find this shark in the Mooreville 
Chalk. Outside of Alabama, P. mortoni is fairly com- 
mon in the Niobrara Chalk of Kansas, and has been 
reported from the Eutaw Formation of Mississippi by 
Stephenson and Monroe (1940). The wide geographi- 
cal range of this species is shown by its occurrence in 
the British Chalk, the Upper Cretaceous of Italy, Mexi- 
co, Mississippi, Alabama, and the Niobrara of Kansas. 



Edaphodon sp. Figure 175B 

Referred specimen. — PF 3501, a right mandible, from 
2 miles W. of West Greene, Greene County, Alabama, 
middle Mooreville Chalk. 

Discussion. — This large mandibular dental element 
has a claw-like appearance and is probably distinct, 
although it may be an extreme variant of Edaphodon 
mirificus. There are two tritoral areas present, a long 
anterior apical area and a very small dorsal tritoral 
area. The general shape, small size, and placement of 
the tritoral areas separate Edaphodon sp. from Eda- 
phodon barberi. The specimen is extremely waterworn 
and much of the back end is missing. 

Class SELACHII 

Order HETERODONTIFORMES 

Suborder HYBODONTOIDEI 

Ptychodontidae Hay 

Discussion. — Casier (1953) has shown that the Pty- 
chodontidae are specialized derivatives of the hybodont 
sharks and not rays at all. Patterson (1965) suggests 
that Hylaeobatis arose from a homodont species of the 
hybodontid genus Lonchidion, and that Hylaeobatis, a 
member of the family Ptychodontidae, gave rise to the 
genus Ptychodus. 

Ptychodus polygyrus Agassiz. Figure 179A 

Referred specimen.— PF 127, one tooth; Bank's Bluff, 
Greene County, Alabama, lowermost Mooreville or up- 
permost Eutaw. 

Discussion. — Cope (1878) reported this species from 
the Rotten Limestone, the old name for the Selma Group 
in Alabama. The Niobrara tooth figured by Williston 
(1900) and assigned to this species actually belongs to 
Ptychodus decurrens. P. polygyrus, the Mooreville spe- 
cies, is known also from the Upper Cretaceous of Eng- 
land, Belgium, and Russia. 

Ptychodus mortoni Mantell. 

Discussion. — This species has not been found in the 
Mooreville member of the Selma Chalk during the pres- 
ent collecting. There is, however, a specimen in the 
Walker Museum collection at Field Museum, U.C. 
14368, from Prairie Bluff, Alabama. The Prairie Bluff 
Chalk lies above the Ripley Formation, which in turn 
lies above the Demopolis Chalk, according to Monroe 



Order GALEIFORMES 
Anacoracidae Gliickman 

Discussion. — The two genera Squalicorax and Pseu- 
docorax have been referred tentatively to the Ceto- 
rhinidae by Woodward (1902-1912) and E. I. White 
(1937). Gliickman (1964) has referred Anacorax, which 
is a synonym of Squalicorax, to a separate family, Ana- 
coracidae. Undescribed material at the Los Angeles 
County Museum suggests that his family reference is 
correct. Therefore, Squalicorax, and with it the closely 
related Pseudocorax, should form the basis of a new 
family. In spite of the fact that Squalicorax has pri- 
ority over Anacorax, the family name Anacoracidae 
must be retained according to Article 40 of the Inter- 
national Code of Zoological Nomenclature. Best indi- 
cations are that these sharks are primitive orectolobid 
derivatives; therefore, I have placed them before the 
Odontaspididae, which are thought to be advanced orec- 
tolobid derivatives. 

Squalicorax falcatus (Agassiz). Figures 176 A-F; 
178 L and N; 179B 

Referred specimens. — PF 3524, four teeth; Ostrea 
layer, Choctaw Bluff, Greene County, Alabama, lower- 
most Mooreville or uppermost Eutaw. PF 3523, four 
teeth; Bank's Bluff, Greene County, Alabama, lower- 
most Mooreville or uppermost Eutaw. PF 124, 29 
vertebrae; PF 3538, four vertebrae; one mile S. of West 
Greene, Greene County, Alabama, middle Mooreville. 
PF 126, 48 vertebrae and dermal denticles; one mile 
NW. of West Greene, Greene County, Alabama, middle 
Mooreville. PF 3537, three vertebrae; 2 miles N., one 
mile W. of West Greene, Greene County, Alabama, 
middle Mooreville. PF 3534, two teeth; P 27496, four 
vertebrae; PF 3533, four vertebrae; P 27448, four ver- 
tebrae; P 27409, 34 vertebrae; P 27446, five vertebrae; 
PF 3528, four vertebrae; PF 3529, six vertebrae; P 
27475, 13 vertebrae and one slide with cross section; 
PF 3530, one vertebra and one tooth; all from Moore's 
farm, Dallas County, Alabama, middle Mooreville. PF 
3536, seven vertebrae, Harrell's Station, Dallas County, 
Alabama, middle Mooreville. PF 3527, one tooth; near 
road, Hewlett's farm, Greene County, Alabama, upper 
Mooreville. PF 3525, one tooth, 8.9 miles E. of Russell 



1 Based on an undescribed Aquia fish fauna collected by me 
and now in Field Museum. 



394 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 









B 










H 







Fig. 176. Denticles of sharks, much enlarged; A-F, Squalicorax falcatus (A-C, PF 126); G-N, undetermined sharks, PF 3521. 



County Line, Route 26, Russell County, Alabama, Bluff- 
town Formation. 

Discussion. — Squalicorax falcatus is now known in 
the Mooreville by its teeth, vertebrae, and denticles. 
The limits of tooth variation are not known because 
of the lack of associated sets. This is one of the most 
common Upper Cretaceous sharks. The vertebral cross- 



section shown in Figure 179 B resembles those of Ceto- 
rhinus in showing concentric rings of calcification plus 
several radial supports. This sort of structure is found 
also in Squatina. While some sets of vertebrae do not 
have associated teeth, there is little doubt that all be- 
long to the same species. In some specimens, partic- 
ularly P 27409, denticles were noted adhering closely 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



395 



to the vertebrae; these evidently settled onto the ver- 
tebrae as the fish decayed. The denticles of another 
specimen, PF 126, display notable variation (fig. 176 A- 
F). Figure 176 A represents the palmate type of den- 
ticle, having five or six backwardly projecting "fingers" ; 
Figure 176 B and C represents the other extreme, a 




I m m 



Fig. 177. Tooth of Pseudocorax affinis, PF 3522. 

smooth bulb-like denticle with only one posterior pro- 
jection. Figure 176 D, E, and F shows the most com- 
mon type of denticle, with three keels on the crown. 
S. falcatus is reported from southern India, Madagas- 
car, northern Europe, England, New Jersey, Kansas, 
and California. 

Squalicorax pristodontus (Agassiz) 

Referred specimen. — PF 660, one tooth; roadbed be- 
tween Thomaston and Safford, Dallas County, Ala- 
bama, Demopolis Chalk. 

Discussion. — Although this species is not known to 
occur in the Mooreville proper, the fact that it does oc- 
cur in the Selma Group gives reason for its inclusion in 
the present paper. The large size and the broad low 
crown of the teeth serve to distinguish them from those 
of S. falcatus. It is unknown in the Niobrara Chalk. 
At present nothing is known of its denticles or verte- 
brae. S. pristodontus is reported from the Upper Cre- 
taceous of India, western Africa, northern Europe, the 
northern and southern borders of the Mediterranean 
Basin, England, and New Jersey. 

Pseudocorax affinis (Agassiz). Figures 177, 178 M 

Referred specimens. — PF 3522, two teeth; Locality 
12A, consolidated layer, Moore's farm, Dallas County, 
Alabama, middle Mooreville. PF 3558, one tooth; Lo- 



cality 1, 6.2 miles W. of Aliceville, Eutaw County, 
Alabama, lower Mooreville. 

Discussion. — The more slender and elevated crown, 
and the prominent notch between the crown and the 
blade, in combination with the absence or weakness of 
serrations, serve to separate Pseudocorax affinis from 
members of the genus Squalicorax. Woodward (1911) 
states that another distinctive character is the slight 
median cleft for a nutritive foramen on the inner face of 
the root. 

There are teeth of this species in an ornamental 
tray on display at the Museum of Fort Hays State Col- 
lege; these constitute the only Niobrara record known 
to me. Elsewhere, the species is known from the Upper 
Cretaceous of Africa, northern Europe, and England. 
The Eocene Alabama record of Hay (1929) is incorrect 
according to White (1956). 

Odontaspididae Miiller and Henle 
Discussion. — Signeux (1949) has amply demonstrated 
the very close affinity between Scapanorhynckus, Mit- 
sukurina, and Odontaspis. In the same paper she has 
shown the distinctions that exist between Scapano- 
rhynckus and Mitsukurina; the existence of such generic 
differences has been questioned by past workers. Iso- 
lated fossil teeth of Scapanorhynckus are very difficult 
to distinguish from similar teeth of Odontaspis, and 
the only good character known at present is the strong 
narrow vertical ridges or striae on the back of the crown. 
The type of the genus Scapanorhynckus is S. lewisi 
from the Upper Cretaceous of Sahel-Alma, Mt. Leb- 
anon, Syria, and is based on well preserved material 
showing body shape, teeth, denticles, and vertebrae. 
Elsewhere this genus is known only from teeth. It 
should be noted that the old genus Carcharias is no 
longer considered valid according to the ruling of the 
International Commission on Zoological Nomenclature, 
no. 723, 1965; the name is superseded by Odontaspis. 

Scapanorhynchus rhaphiodon (Agassiz). Figure 178 
A-C. 

Referred specimens. — PF 3504, three teeth; Choctaw 
Bluff, Greene County, Alabama, lower Mooreville or 
Eutaw. PF 3505, three teeth; Bank's Bluff, Greene 
County, Alabama, lower Mooreville. PF 3502, one 
tooth; Hale's farm, 2 miles N. of West Greene Post 
Office, Greene County, Alabama, middle Mooreville. 
PF 3506, 14 teeth; PF 3509, one tooth; Moore's farm, 
Dallas County, Alabama, middle Mooreville. PF 3511, 
two teeth; Harrell's Station, Dallas County, Alabama, 
middle Mooreville. PF 3503, two teeth ; Hewlett's farm 
near the road, Greene County, Alabama, upper Moore- 
ville. PF 3508, two teeth, 5.1 miles E. of Russell 
County Line, Route 26, Russell County, Alabama, Bluff- 
town Formation. PF 3507, two teeth; PF 3519, one 
tooth; both from 8.9 miles E. of Russell County Line, 
Route 26, Russell County, Alabama, Blufftown For- 
mation. 

Discussion. — The Mooreville specimens agree with 
those teeth figured by Agassiz (1843) under the specific 



396 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



name rhaphiodon. Woodward (1889) erected the genus 
Scapanorhynchus and placed in it Agassiz' species, Lam- 
na rhaphiodon. Williston (1900) put the Niobrara teeth 
of Scapanorhynchus also in this species, with Roemer's 
(1852) Lamna texanus a synonym. Arambourg (1952) 
has considered Lamna texanus a synonym of the very 
different S. rapax. 



the length of the crown, differentiate the teeth of this 
species from those of S. rhaphiodon and agree with 
teeth figured by Quaas (1902) and Arambourg (1952). 
This form may prove to be more characteristic of the 
Eutaw than of the Mooreville but it does occur at the 
Mooreville-Eutaw contact. 5. rapax has previously been 
reported only from Africa, where it occurs in the Seno- 




81 



M 



N 



~M 



20 mm 

Fig. 178. A-C, Scapanorhynchus rhaphiodon, three anterior teeth, PF 3506; D, Scapanorhynchus rapax, anterior tooth, PF 3510; E-H, 
Lamna appendicular, three lateral teeth and one posterior tooth, PF 3513; I-K, Isurtis mantelli, two anterior teeth and one lateral tooth, 
PF 3512; L and N, Squalicorax falcalus, two lateral teeth, PF 3523; M, Pseudocorax affinis, lateral tooth, PF 3522. 



S. rhaphiodon has been reported from the Upper 
Cretaceous of New Zealand, Japan, Australia, the East 
Indies (Timor), India, Africa, both sides of the Medi- 
terranean Basin, Northern Europe, England, the Cari- 
ibbean Islands, and in North America from the New 
Jersey Greensands, the Niobrara of Kansas, and the 
Austin Chalk of Texas. 

Scapanorhynchus rapax (Quaas). Figure 178 D. 

Referred specimens. — PF 3510, two teeth; Ostrea 
layer, Choctaw Bluff, Greene County, Alabama, lower- 
most Mooreville or uppermost Eutaw. 

Discussion. — The large size, wide bases, and poorly 
developed striae, which extend for less than one half 



nian of Libya and the Maestrichtian phosphate beds 
of northern Africa. 

Lamnidae Muller and Henle 
Lamna appendiculata Agassiz. Figures 178 E-H, 
179 C. 

Referred specimens. — PF 3517, one tooth; Ostrea 
layer, Choctaw Bluff, Greene County, Alabama, lower- 
most Mooreville or uppermost Eutaw. PF 3520, two 
teeth; one mile S. of West Greene, Greene County, 
Alabama, middle Mooreville. PF 3535, two teeth; PF 
3532, one tooth; PF 3514, 12 teeth; PF 3515, 50 verte- 
brae and 13 teeth from one individual; PF 3521, one 
vertebra; P 27499, 12 vertebrae; P 27500, two verte- 
brae, one tooth, one vertebral thin section; all from 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



397 






B 



10 mm 

Fig. 179. A, tooth of Ptychodtts polygurus, PF 127; B, vertebral cross-section of Squalicorax falcatus (P 27475); C, vertebral cross 
section of Lamna appendiculata (P 27500). 



Moore's farm, Dallas County, Alabama, middle Moore- 
ville. PF 3516, 15 teeth; Harrell's Station, Dallas 
County, Alabama, middle Mooreville. PF 3518, one 
vertebra; north set of gullies, Hewlett's farm, Green 
County, Alabama, upper Mooreville. PF 3513, five 
teeth; near the road, Hewlett's farm, Greene County, 
Alabama, upper Mooreville. 

Discussion. — Lamna appendiculata is widespread in 
the Mooreville. A direct association of teeth and ver- 
tebrae is known in PF 3515, which includes the re- 
mains of just one individual. No dermal denticles were 
found in association with these remains. A vertebra, 
as seen in cross-section (fig. 179 C), compares favorably 
with the living Isurus oxyrhinchus. 



L. appendiculata is known from the Cretaceous of Ja- 
pan, Australia, New Zealand, the East Indies (Timor), 
Madagascar, both sides of the Mediterranean Basin, 
western Africa, British Columbia, and in the United 
States in Alabama, Kansas, and New Jersey. 

Isurus mantelli (Agassiz). Figure 178 I-K. 

Referred specimens. — PF 3512, eight teeth; Ostrea 
layer, Choctaw Bluff, Greene County, Alabama, lower- 
most Mooreville or uppermost Eutaw. 

Discussion. — The classic distinction between Isurus 
and Lamna has been and still is the absence of lateral 
denticles in the former. Isurus is also characterized 
by having a less rectangular root. In the literature 



398 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



this species has been described only by its teeth; there 
is, however, a large, almost articulated, specimen in the 
Museum of Natural History at the University of Kan- 
sas (no. 199). Associated with the skull of this speci- 
men are smooth denticles simliar to those that we find 
in the snout of the Recent species. In the collections 
at Fort Hays, Kansas, there is now another fine speci- 
men of this species as yet undescribed. 

Isurus mantelli is known from the Upper Creta- 
ceous of the East Indies (Timor), Madagascar, both 
sides of the Mediterranean Basin, northern Europe, 
England, and the Caribbean region. Hay (1929) re- 
ported this species from the Cretaceous of New Jersey. 
Hay's Eocene and Miocene records probably represent 
reworked teeth. The species is also known as stated 
above from the Kansas Chalk. 

Undetermined shark denticles. Figure 176 G-N. 

Referred specimens. — PF 3531, numerous denticles; 
Locality 12A, consolidated layer, Moore's farm, Dallas 
County, Alabama, middle Mooreville. 

Discussion. — At Locality 12A we found a number 
of shark denticles which cannot be placed taxonomi- 
cally with any precision; however, by referring to the 
illustrations given by Bigelow and Schroeder (1948), 
and by examining available Recent specimens, it is 
at least possible to suggest generic affinities. Figure 
176 G shows a denticle which is similar to those of the 
Recent smooth dogfish, Mustelus canis. Figure 176 H 
shows resemblances to several groups; it could be 1am- 
noid. Figure 176 I and J shows three-pronged denticles 
suggestive of those of the family Triakidae. Denticles, 
Figure 176 K and L, are close to those of Mustelus. 
Figure 176 N and M (which is the lateral view of N) 
agrees with those of Rhincodon except for the lack of 
lateral points. Although the best that one can hope 
for from these denticles is an approximate placement, 
they do suggest a shark fauna more varied than that 
which is indicated by teeth and vertebrae. This in 
itself is of some interest. 

Subclass ACTINOPTERYGII 

Infraclass CHONDROSTEI 
Order ACIPENSERIFORMES 

Acipenseridae Bonaparte 

The Acipenseriformes have a fossil record going as 
far back as the Jurassic, with questionably related forms 
to the Pennsylvanian (Romer, 1966). The family Aci- 
penseridae differs from the related family Chondros- 
teidae in having a dorsal series of unpaired splinter- 
like plates on the upper rostral surface, and lateral 
scutes on the body. The sturgeons differ from the 
Polyodontidae (paddlefishes) in lacking teeth in the 
adult as well as in the absence of vacuities in the skull 
so characteristic of the latter. 

Propenserinae, new subfamily 

Diagnosis. — The frontals are expanded laterally and 
border the orbit. The rostrals are numerous and small 




Fig. 180. Reconstruction of skull of Propenser hewletti, dorsal 
view. DS, dermosphenotic; ES, extrascapular; FR, frontal; PA, 
parietal; PO, postorbital; PT, pterotic; SO, supraoccipital. 

in size. The parietals are reduced and separated by a 
very large supraoccipital. The endocranium is ossified, 
with a pronounced fusion of its elements, as is the 
shoulder girdle. The neural arches are fused and each 
arch bears two neural spines. Other characteristics 
are as in the genus. 

Propenser, new genus 

Diagnosis. — As for subfamily. 

Type species. — Propenser hewletti, new species (fig. 
180). 

Description. — The external surfaces of the plates 
are ornamented with equally raised hemispherical tu- 
bercles approximately 5 mm. in diameter. These may 
be fused into lines in a weblike pattern. Ridges with 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



399 



or without the above tubercles are present in various 
plates. The triangular frontals (fig. 180, FR) extend for- 
ward and are rounded on the anterior border; laterally, 
they form the upper border of the orbit. Anterior to 
the frontals are 20 or more rostrals which vary in size 
and shape. (The rostrals were not found in place.) The 
most common rostral shape is elongate lanceolate (fig. 
182 A). The parietals (fig. 180, PA) are small and are 
separated from each other by a huge supraoccipital (fig. 
180, SO), which is deflected downward on each side; 
it is arrow-shaped, with the point toward the rear of 
the skull. Lateral to the parietals lie large pterotics 
(fig. 180, PT) . Posterior to the pterotics are the extra- 
scapulars (fig. 180, ES) , each of which bears a postero- 
lateral forked process which is believed to serve for the 
attachment of the post-temporals. The dermosphe- 
notic (fig. 180, DS) lies just posterior to the postorbital 
(fig. 180, PO). The hyomandibular (fig. 183, A) is ossi- 
fied, as are the other known hyoid elements (fig. 183, B 
through G). The lower jaw (fig. 185, C) is long, thin, 
quite delicate, and toothless. The endocranium is be- 
lieved to have been partly ossified, and a number of 
parts of it are preserved. The palatoquadrate (fig. 185, 
A and B) is well developed. The articular end of the 
quadrate is produced into a definite peg. The shoulder 




girdle is massive (fig. 184, C and D) with both dermal 
and endoskeletal parts ossified and fused. The first 
pectoral fin spine (fig. 184, B) is large, and ornamented 
dorsally with rows of tubercles which merge into small 
ridges. The second spine (fig. 184, A) is of similar size 
but poorly ornamented. The body scutes (fig. 182, B), 
which are thought to be lateral, are elongate rectangular 
and slightly curved to fit the curvature of the body. 
The dorsal scutes (fig. 184, E), of which four are present, 
have the typical tubercles but still are quite sturgeon- 
like. 

Propenser hewletti, new species. Figures 180-185. 

Type. — Specimen in Alabama Geological Survey 
collection, including much of a skull, scutes, shoulder 
girdle, fin spines, and vertebrae; gullies near road, Hew- 
lett's farm, Greene County, Alabama, upper Moore- 
ville. 

Referred specimens. — PF 288, one plate (probably 
from the type specimen) ; Hewlett's farm, Greene County, 
Alabama. A second specimen in Alabama Geological 
Survey collection, part of a skull; near West Greene, 
Greene County, Alabama, middle Mooreville. 

Diagnosis. — Same as for genus. 
I 



20mm 




20 m m 



Fig. 181. Propenser hewletti, n. sp., A, holotype, specimen in collection of Alabama Geological Survey, dorsal view of posterior portion 
of skull; B, second specimen in Alabama Geological Survey collection. 



400 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



Discussion. — The dorsal scutes (fig. 184, E) compare 
favorably, except for their ornamentation, with those 
of other fossil and Recent Acipenseridae. The maxil- 
lary in the type is fused to the palate, which in turn is 
fused to the quadrate (fig. 185, A and B). The rostral 



day analogue in the vertebrae of very large sailfishes, 
which have their abdominal vertebrae coalesced; the 
exact function of this is unknown, but it may serve 
as a spring and an aid in swimming. 

The skull, exclusive of the rostrals, is simpler than 






mm 



20 mm 




20 mm 



Fig. 182. Propenser hewletti, n. sp., holotype; A, rostral plate; B, body scute. 



plates (fig. 182, A) interlock anteriorly and posteriorly 
with a peg and open socket arrangement; laterally, 
they fit together by overlapping and underlapping each 
other, a condition not dissimilar to that found in Aci- 
penser. Propenser differs from Acipenser in the type 
of external ornamentation of the plates (fig. 181), the 
larger size of the frontals (fig. 180), the fact that later- 
ally the frontals form the upper edges of the orbits, the 
smaller size of the parietals, the elongation of the supra- 
occipital, and the more complete ossification of the 
hyomandibular (fig. 183, A), shoulder girdle (fig. 184, 
C-D), neural arches (fig. 185, F), and endocranium. 

The extreme amount of ossification shown partic- 
ularly in the type specimen could be in part due to 
the large size and perhaps old age of this individual, 
but probably represents a primitive condition. The 
fusion of the neural arches (fig. 185, F) finds a present 



in any of the Recent Acipenseridae and agrees more 
closely with that found in the paleoniscoids. The evo- 
lutionary trend in the Acipenseridae has been the frag- 
mentation of the skull bones and the reduction of ossifi- 
cation of many of the elements. Fusion of the maxil- 
lary to the palate and reduction in size of the lower 
jaw have also occurred. In these respects Propenser 
appears to be much more primitive than any living 
genus in this family. 

Each segment of the neural arch of the holostean 
Saurichthys has separate basidorsal and interdorsal ele- 
ments with spines of about equal height (Holmgren 
and Stensio, 1936). Propenser shows the same condi- 
tion except that the bases of these two elements are 
fused into a single unit. In the Acipenseridae the com- 
parable neural arch has been simplified to a single neu- 
ral spine. 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



401 







20mm 





Fig. 183. Propenser heuietti, n. sp., holotype; A, hyomandibular; B-G, hyoid elements. 



It is a pleasure to name this species for Mr. T. G. 
Hewlett, the station master at Boligee, who not only 
carefully collected the type specimen on his farm but 
has shown an active interest over a number of years 
in the fossils which lie close to his home. The second 
and smaller skull was collected by Dr. L. Renger. 

Infraclass HOLOSTEI 
Order PYCNODONTIFORMES 

Pycnodontidae Cope 

Discussion.— The Pycnodontidae are deep-bodied 
angelfish-like holosteans which have strong jaws with 
crushing teeth. Their time of greatest abundance seems 
to have been the Jurassic, but a few persisted into the 
Eocene. 



Hadrodus priscus Leidy. Figure 1£6. 

Referred specimen. — Alabama Geological Survey's 
specimen, right lower jaw; Hewlett's farm, Greene 
County, Alabama, upper Mooreville. 

Discussion. — Leidy (1857, 1873) described a pre- 
maxillary from near Columbia, Mississippi, and Gregory 
(1950) described the premaxillary and splenials of a 
form from the Niobrara Chalk which he called Hadrodus 
marshi. The Mooreville lower jaw. which is complete, 
shows what Gregory called the splenial element, and 
except for the fact that there are only two tooth rows, 
it is very close to Gregory's H. marshi. No sutures 
are discernible on this jaw. 

Incertae sedis. Figure 187. 

Referred specimen .— P 27515, one premaxillary; 
Moore's farm, Dallas County, Alabama, middle Moore- 
ville. 




20mm 







i i 

20 mm 







i i 

20mm 




Fig. 184. Propenser hewlelti, n. sp., holotype; A-B, fin rays; C-D, shoulder girdle; E, dorsal scutes. 



402 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



403 




.tSJjt*' 






$0 




20 mm 




Fig. 185. Propenser hewletli, n. sp., holotype; A-B, palatoquadrate and part of maxillary; C, lower jaw; D-E, parts of vertebrae 
F., neural arches. 



Discussion. — It is most likely that this single pre- 
maxillary belongs to the Pycnodontidae. The presence 
of large pits on the surface appears to place this bone 
closest to Hadrodus. The antero-posterior alignment 
of the five teeth is singular. Medially there is an area 
for the symphyseal attachment with the other pre- 



maxillary. The teeth are broken, with none of them 
bearing the tips of the crown. 

Order AMIIFORMES 

Suborder AMIOIDEI 

Pachycormidae Woodward 



404 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 






20 mm 



Fig. 186. Hadrodus priscus, right mandible; A, medial view; 
B, dorsal view; C, lateral view. 

Protosphyraena Leidy 

Discussion. — Protosphyraena is an Upper Cretaceous 
pachycormid holostean with an elongate and partly 
fused pectoral fin. In the palate are two large anteriorly 
directed fang-like teeth. The snout is produced into 
a spike-like bony rostrum. All indications are that 
these fishes occupied a habitat similar to that of the 
modern billfishes, and must have used the large teeth 
and bill to strike and kill their prey. From the ma- 
terial available there would appear to be several spe- 
cies in the Mooreville Formation, which are here kept 
separate. More material, however, could show that 
there is only one highly variable species. 

Protosphyraena nitida? (Cope) 

Referred specimen. — PF 3547, snout with one tooth 
and part of the base of the skull; Moore's farm, Dallas 
County, Alabama, middle Mooreville. 

Discussion. — A single fang-like upper tooth is pres- 
ent and well preserved; another tooth is in the process 
of being replaced. The skull compares very favorably 



with that of the Niobrara species, P. nitida. A re- 
vision would no doubt reduce the six species from the 
Niobrara, but this is contingent on obtaining better 
materials. 

Protosphyraena sp. (1) 

Referred specimens. — PF 3551, one hypural bone, 
Harrell's Station, Dallas County, Alabama, middle 
Mooreville. PF 3552, one hypural bone; Marion Junc- 
tion, Dallas County, Alabama, middle Mooreville. 

Discussion. — The single hypural bone of Protosphy- 
raena has been figured previously by Woodward (1912), 
and there is no doubt that the Mooreville hypurals 
belong to this genus. 

Protosphyraena sp. (2) 

Referred specimen. — PF 3545, part of a fin; one mile 
N. of store, Hale's farm, Greene County, Alabama, 
middle Mooreville. 

Protosphyraena sp. (3) 

Referred specimens. — P 27363, parts of fin; Craw- 
ford's farm, W. of Alabama 13, Hale County, Alabama, 
middle Mooreville. P 27364, parts of fin, Township 
28, Hale County, W. of Alabama 13, Alabama, middle 
Mooreville. P 27365, fin fragment; Moore's farm, Dal- 
las County, Alabama, middle Mooreville. PF 121, one 
tooth, parts of two jaws; Bank's Bluff, Greene County, 
Alabama, lowermost Mooreville or uppermost Eutaw. 

Discussion. — None of the Mooreville fin fragments 
is as large as in the Niobrara specimens. There are 
two different types of fin rays in the Mooreville, PF 
3545 being smaller than the rest and more like Niobrara 
material in ornamentation. The bone is dense and 
heavy in some of the specimens. 

Infraclass TELEOSTEI 

Discussion. — The concept of the Isospondyli as a 
homogeneous monophyletic group has never rested on 
a firm morphological or paleontological basis. The 
polyphyletic origin of the group has been suggested 
by many, including most recently Greenwood et al. 







10mm 
Fig. 187. Premaxlllary of possible pycnodontid, P 27515 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



405 



(1966). The Isospondyli as recognized, for example, 
by Romer (1945) are more a stage of teleostean devel- 
opment than a distinct phyletic entity. When viewed 
by a student of Cretaceous fishes, the Recent Isospon- 
dyli represent only the surviving members of some of 
the great Jurassic and Lower Cretaceous radiations 
that have achieved varying degrees of success. The 
term Isospondyli, except when used with the above 
reservations, obscures rather than clarifies the relation- 
ships of its various subgroups. 

The suborder Clupeoidea as used by Romer (1945) 
includes both the families Clupeidae and Elopidae, yet 
the latter are two fundamentally distinct lines each of 
which certainly warrants a rank above the family level. 
A traceable phyletic line is that from the Leptolepididae 
to the Elopidae, a complex that has remained distinct 
from the better known leptolepid-chirocentrid-clupeid 
line. If we raise the Leptolepididae to a suborder under 
the Isospondyli (the Leptolepidoidei), the Elopidae and 
related families to a similar suborder (the Elopoidei), 
and the Clupeidae and related families to another sub- 
order (Clupeioidei), realizing that the last two are prob- 
ably derived from the Leptolepidoidei, we have a 
classification based on phylogenetic evidence that clar- 
ifies as it sorts out distinct lines from an ancestral 
group. In the present paper we will deal with the 
Mooreville Elopoidei and the Clupeoidei in turn. The 
Elopoidei are here divided into a number of super- 
families, the Elopoidea, Pachyrhizodontoidea, Albuloi- 
dea, and Plethodoidea ; the Chanoidea and the Osteo- 
glossoidea are thought also to belong to this group. 
The limits of the Elopoidei are so broad that a final 
all-inclusive definition is not at the present possible; 
however, the following definition will serve for the pres- 
ent paper. 

Suborder ELOPOIDEI 

Diagnosis. — These are elongate fusiform fishes. The 
post-temporal fossa is present except in one or two cases 
where it has been lost or reduced to a groove. The 
lateral temporal and subtemporal fossae are generally 
present; no preepiotic nor auditory fossa has been dem- 
onstrated. There is an intercalar-prootic bridge. The 
supraoccipital is small and low, never forming a large 
crest. The circumorbitals extend to the preoperculum. 
The vertebrae are not pierced by the notocord in adults 
and are without transverse processes. The scales are 
longer than deep, and have distinct circuli; distinct 
apical regions are present. There are no ventral ridge 
scales. The ventral fins are post-abdominal. Inter- 
muscular bones are present. 

Superfamily ELOPOIDEA 

Diagnosis. — The parietals meet in the midline; a 
gular plate is present; the maxillaries enter into the 
gape of the mouth; the jaws are toothed, with small 
teeth; the mesethmoid is small; the f rentals are long, 
tapering anteriorly; the nasals are small, not meeting 
and not normally attached to the frontals; the orbito- 




I mm 




I mm 



Fig. 188. Palelops eutairensis, n. sp., scales; A, type, PF 3559; 
B, PF 3560. 



sphenoids and basisphenoids are reduced. There is a 
single dorsal fin. There are two families, Elopidae and 
Megalopidae. 

Discussion. — The scales of this superfamily are very 
uniform and are divided into four quadrants of orna- 
mentation. The apical regions have vermiculate ridges; 
the basal regions have radii which cross the circuli ; the 
dorsal and ventral quadrants have concentric circuli. 

Elopidae Bonaparte 

Discussion. — The characteristics of this family have 
been discussed by Berg (1947) and Dunkle (1940). 
The genus Notelops should be placed in this family 
along with Holcolepis, and probably with Elopoides. 
Elops is known from the Lower Eocene to the Recent. 
The scales of the Elopidae show more than five basal 
radii, whereas the Megalopidae have commonly less 
than five. 

Palelops, new genus 

Type species. — Palelops eutawensis, new species. 
Diagnosis. — Same as for species. 



406 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



Palelops eutawensis, new species. Figure 188. 

Type.— PF 3559, one scale; ', mile E. of Walter 
Dance's farm, 6.2 miles W. of Aliceville, Eutaw County, 
Alabama, lower Mooreville. 

Referred specimens. — PF 3612, two scales; l 4 mile 
E. of Walter Dance's farm, 6.2 miles W. of Aliceville, 
Eutaw County, Alabama, lower Mooreville. PF 3563, 
one scale; Choctaw Bluff, Greene County, Alabama, 
lower Mooreville. PF 3560, four scales; Montgomery's 
farm, 2 miles N. of West Greene, Greene County, Ala- 
bama, middle Mooreville. PF 3561, one scale; Hew- 
lett's farm, gullies near the road, Greene County, Ala- 
bama, upper Mooreville. 

Diagnosis. — The basal regions of the scales have 19 
to 21 radii which tend to be parallel; the circuli in the 
basal regions are distinct ; the dorsal and ventral quad- 
rants have straight circuli. The scale centers have 
coarse raised granules which cover the nuclei of the 
circuli; the area covered by these granulations varies 
in different scales. The apical regions show circuli and 
granulations. These scales differ from those of Elops 
in the greater number of radii, the nonvermiculate 
character of the apical ornamentation, and the basal 
extension of the granulations over the nucleus. 

Discussion. — Dr. Lore David has in one of her scale 
notebooks a photograph of an identical scale from the 
Cretaceous of California; however, locality data are 
lacking, although this scale is believed to be from the 
Panoche Hills, near Fresno, California. Palelops eutaw- 
ensis represents the nearest approach to the genus 
Elops of any scale known in the Mooreville fauna. 
Mr. Marion C. Bonner of Leoti, Kansas, has found an 
example of a scale from the Niobrara Chalk which is 
identical to this species except that it is larger (PF 
3336). Another scale fragment of this species is to be 
found in an undescribed Cretaceous fauna from South 
Dakota in the Field Museum Collection. 

Superfamily PACHYRHIZODONTOIDEA, 

new superfamily 

Diagnosis. — The parietals do not meet in the mid- 
line; the gular plate is absent; the maxillary enters into 
the gape of the mouth; the teeth are large, recurved, 
and conical. The mesethmoid is large and the frontals 
are rectangular in outline. There is a large prominent 
scale bone above the operculum. An enlarged scale is 
present in front of each of the paired fins. There is 
only one recognized family, the Pachyrhizodontidae. 

Discussion. — The Pachyrhizodontoidea represent 
one of the important Cretaceous radiations of the Elo- 
poidei. Some generic relationships are as yet too un- 
certain to assign all the genera to families within this 
superfamily. Among the genera are Pachyrhizodus, 
Thrissopater, Elopopsis, and Rhacolepis. In most the 
parietals are completely separated by the supraoccipital. 
The teeth are well developed, and there is a huge scale 
plate above the operculum. 



Pachyrhizodus Dixon 

Description. — An interfrontal fossa is present in the 
posterior part of the skull roof; it is rectangular in 
shape, with its posterior border formed by the supra- 
occipital. On each side of the supraoccipital lie two 
small parietals. The epiotics form the postero-lateral 
angles of the skull. Anterior to the epiotics are large 
pterotics. There is a portion of a large posttemporal 
in one specimen but its shape is not known. No post- 
frontal has been found. The two pairs of postorbitals 
are large and more or less rectangular. The upper 
postorbital meets the scale bone; the lower one abuts 
against the front edge of the preoperculum. The sub- 
orbital is naiTow, and a prefrontal is present and large. 
The sclerotic bones are evidently two in number. No 
nasal element is known. The premaxillary is consider- 
ably smaller than the maxillary and has two rows of 
teeth. The teeth of both the upper and lower jaw are 
in sockets much as in mosasaurs. The mandible (fig. 
193 D) is large and nearly rectangular in shape but 
narrows anteriorly. A small angular is present; pos- 
terior to the articular surface is a rounded flange. The 
quadrate (fig. 193 D) is triangular except for rounding 
in the posterior apex. The hyomandibular has a large 
single head and a prominent opercular process; ven- 
trally it is very narrow and similar to that of Ziphac- 
tinus (Stewart, 1900). The operculum is covered dor- 
sally by the large scale bone. Externally it has fine 
striae which radiate out from its point of articulation. 
The preoperculum is low and wide, with grooves that 
radiate posteriorly from the juncture of the anterior 
and dorsal arms. The suboperculum has a rounded 
posterior edge; its anterior and dorsal edges are straight 
and form a right angle. The interoperculum is poorly 
known. The branchiostegals are over 20 in number. 
The posterior portion of the skull has on each side a 
large posttemporal fossa. An opisthotic bridge is pres- 
ent. There is no auditoiy foramen such as one finds 
in the Clupeidae. The parasphenoid covers the base 
of the skull and has two small forks posteriorly; its 
anterior extremity is shovel-like. Anterior to the para- 
sphenoid are two rounded thick bones (fig. 193 C), 
thought to be vomers, each of which bears a single 
tooth-like structure. All the palatal elements and gill 
supports bear, where they are exposed in the mouth, 
a continuous cover of small teeth which are curved and 
resemble the jaw teeth but lack distinct sockets. The 
epihyal is roughly semicircular. The ceratohyal (fig. 
193 B) is approximately rectangular but widens at its 
posterior end. The upper part of the shoulder girdle is 
unknown. The lower part (fig. 193 A) has a prominent 
mesocoracoid arch. 

Each pectoral fin has one very large first ray (fig. 
199 B). The ends of the other rays are fringed (fig. 189 
B), branching several times distally. The pelvic rays are 
curved and are of nearly uniform size; each ray is 
divided into joints distally. The anal fin resembles 
the pelvics in these features. The tail has been figured 
by Hay (1903) ; its whole surface is covered with rec- 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



407 




20mm 




Fig. 189. Pachyrhizodus minimus, PF 1697; A, nearly complete fish; B, head and pectoral fins. 



tangular lepidotrichia, has prominent fulcra along the 
edge, and has internally two hypurals. The dorsal fin 
rays have cup-like bases, from which arise a pair of 
lateral projections. There are 50 to 60 vertebrae. 

The genus Pachyrhizodus is also characterized by 
its distinctive scale type (fig. 191) with a triangular 
apical region. In this are dendritic ridges which branch 
toward the apical edge. The lateral and basal areas 



contain strong circuli. 

Discussion. — Some of the Niobrara species of Pachy- 
rhizodus have been based on characters that could fall 
within the range of individual variation. An examina- 
tion of the Niobrara material at the University of Kan- 
sas and at the American Museum of Natural History 
revealed what is believed to be three valid species. 
These are also found in the Mooreville. 



408 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 





10 mm 





10 mm 



10 mm 



Fig. 190. Vertebral cross-sections of Pachyrhizodus; A, P. minimus; B-C, P. caninus; D, P. kingi. 



Pachyrhizodus minimus Stewart. Figures 189, 190 
A, 191 B-C, 192. 

Referred specimens. — P 27489, fragments of dermal 
bone, premaxillary, vertebrae, and lower jaw; Moore's 
farm, Dallas County, Alabama, middle Mooreville. PF 
1697, a nearly complete articulated fish; 5 miles SSW. 
of Clinton, Greene County, Alabama, middle or upper 
Mooreville. 

Discussion. — The two Mooreville specimens, partic- 
ularly the second, are assigned to P. minimus, which 
was described by Stewart (1900) from a lower jaw. 
Other specimens in the University of Kansas Museum, 
which were described as belonging to various families 
and genera by Jordan (1925), are referred to this spe- 



cies; they are Kansanus martini, and Eurychir lindleyi. 
Pachyrhizodus shear eri and lOricardinus tortus may be- 
long to this species but are so poorly preserved as to 
be indeterminate. 

PF 1697 (fig. 189) is the best Mooreville specimen ; 
it is about three feet long and has a total vertebral 
count of 54. There is a very large scale bone; the oper- 
culum shows only faint radiation lines; the preoper- 
culum is large in comparison with that of Elops and 
bears a series of basal striations. The premaxillary is 
only about one-fifth as long as the maxillary. A large 
arrowhead-shaped ethmoid element is conspicuous. The 
pectoral fin has at least 17 rays. There is some evidence 
of a sclerotic ring. The teeth are very small and slen- 
der, one of the better characters for distinguishing this 




I mm 



mmjf^fe - Hl 



V 



\ 








,W\ 



Fig. 191. Scales of Pachyrhizodus; A, P. cam'niis; B-C, P. minimus (C, PF 16971. 



I mm 



409 



410 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 




Fig. 192. Reconstruction of Pachyrhizodus minimus. 



species. The vertebrae show externally very fine longi- 
tudinal striae. 

In cross-section (fig. 190, A), a vertebra shows in 
its center a small core of lamellar bone. About half- 
way from the center is a bony ring. The outer surface 
of the vertebra has a small ring of lamellar bone which 
is indented above and below. 

The apical region of the scales (fig. 191, B and C) 
has externally a triangular area with 17 rays converging 
at the center of the scale. The rest of the scale is 
covered with coarse circuli. 

Pachyrhizodus caninus Cope. Figures 190 B-C, 191 
A, 193, 199 B. 
Referred specimens. — P 27410, lower jaw fragments; 
Banks' farm, Greene County, Alabama, lower Moore- 
ville. P 27416, fin spine, premaxillary, and lower jaw; 
gully near the Choctaw Road, Greene County, Ala- 
bama, lower Mooreville. PF 3554, 19 vertebrae and 
tail; 2 miles N. of West Greene, Greene County, Ala- 
bama, middle Mooreville. PF 1696, jaw and most of 
a skull, pectoral fin, and vertebrae (almost complete 
fish) ; 2 miles W. of West Greene, Greene County, Ala- 
bama, middle Mooreville. PF 128, three vertebrae and 
part of a jaw; 1 6 10 miles N. and ) ■> mile West of West 
Greene. Greene County, Alabama, middle Mooreville. 
PF 137, four vertebrae; PF 138, vertebrae and a fin 
spine; P 27423, one vertebra; P 27451, jaw fragment; 
P 27502, shoulder girdle and fin rays; P 27513, jaw; 
P 27516, jaw fragment; P 27518, quadrate, lower jaw, 
fragments, and fin spines; P 27519, fragments and lower 
jaw; P 27520, two maxillaries; P 27523, part of lower 
jaw; all from Moore's farm, Dallas County, Alabama, 
middle Mooreville. P 27323, almost complete fish; 
gulley near Harrell's Station, Dallas County, Alabama, 
middle Mooreville. PF 134, skull fragments; PF 135, 



ceratohyal; both from Donald's farm, Dallas County, 
Alabama, middle Mooreville. PF 292, jaw fragment, 
Section 33, Township 16 N., Range 16 E., Montgomery 
County, Alabama, middle Mooreville. P 27504, lower 
jaw and three vertebrae; Township 11, W. of High- 
way 13, Hale County, Alabama, middle Mooreville. 
PF 3542, skull fragments, eight vertebrae, and frag- 
ments of fin spine; Hewlett's farm, near road, Locality 
8, Greene County, Alabama, upper Mooreville. P 27532, 
fin spine; Hewlett's farm, Greene County, Alabama, 
upper Mooreville. PF 442, jaw fragments; 2 miles W., 
1 mile N. of West Greene, Greene County, Alabama, 
upper Mooreville. 

Discussion. — Pachyrhizodus caninus is not only the 
largest species of this genus but one of the largest of 
the Mooreville fishes. There can be no question that 
the Mooreville specimens belong to the same species 
as the Niobrara P. caninus. As yet no articulated 
specimens are known from the Mooreville though in- 
dividuals with a great number of associated scales and 
bones do occur. The scales (fig. 191, A) agree with those 
of P. minimus except for the finer and more numerous 
circuli in P. caninus. The apical radii are greater in 
number in P. caninus than in P. minimus and not 
as well defined. The granular ornamentation is more 
prominent in P. caninus, and the scales are relatively 
larger. The cranial bones vary in proportions; this of 
course may be in part due to distortion by post-de- 
positional forces. Among the more stable and easily 
recognizable elements are the large ceratohyals (fig. 
193, B) and the quadrate (fig. 193, D), which has an at- 
tenuated anterior process. The vertebrae are charac- 
terized externally by their smooth sides. In cross-sec- 
tion (fig. 190, B and C) they have a nearly solid bony 
center bordered by an inner ring and a superficial one 
that sends highly irregular processes inward, forming 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



411 




Fig. 193. Pachyrhizodus caninus; A, pectoral girdle, 
attached quadrate. 



20mm 



P 27501; B-D, PF 1696; B, ceratohyals and C, vomers; D, portion of jaw with 



an elaborate pattern similar to fine lacework. The 
overall vertebral cross-section is basically similar, how- 
ever, to that of P. minimus and P. kingi except for 
being more complex in pattern. The jaws of this spe- 
cies (fig. 193, D) have relatively big teeth. 

P. latimentum and P. leptopsis are synonyms of P. 
caninus. 

Pachyrhizodus kingi Cope. Figure 190 D. 

Referred specimens. — P 27411, jaw fragments and 
skull fragments; PF1612, vertebrae and jaw fragments; 
PF 3546, vertebrae and skull fragments; all from Moore's 
farm, Dallas County, Alabama, middle Mooreville. 

Discussion. — Pachyrhizodus kingi is intermediate in 
size between P. caninus and P. minimus. It also occurs 



in the Niobrara. There is a possibility that P. kingi is 
only the juvenile form of P. caninus, but there is a 
decided size gap between these two forms. A vertebral 
cross-section of P. kingi (fig. 190, D) shows differences 
in structure from P. caninus. P. leptognathus and P. 
velox are synonyms of P. kingi. 

Superfamily ALBULOIDEA Hay 

Diagnosis— -The parietals meet in the midline. A 
gular plate is present. The maxillary is excluded from 
the gape of the mouth; the premaxillary is free; the 
jaws, the paired palatal plates, and the gill arches are 
covered with fine conical teeth; the parasphenoid and 
basibranchial bear large button-shaped teeth. Well os- 



412 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



sified orbitosphenoids and basisphenoids are present. 
The frontals are elongate, triangular, and taper anteri- 
orly. The nasals are small and not attached to the 
frontals. The mesethmoid is small. The lateral tem- 
poral fossa is roofed; there is no angular. Very char- 
acteristic are the scales, which are rounded posteriorly 
and bear distinct radial plications anteriorly. The sur- 
face of these plications is ornamented with granules 
arranged in a dendritic pattern or in rows that are 
aligned in a radial direction. The dorsal and ventral 
surfaces of the scale bear fine distinct circuli which are 
closely spaced. Anteriorly the circuli are broken into 
vermieulate ridges as in Elops. 

There are two Recent families that belong in this 
superfamily, the Albulidae and the Pterothrissidae. The 
scales of these two families are almost identical. 

Albulidae Gunther 

Diagnosis. — The dorsal fin is short; the vomer and 
palatines are toothed; and the interorbital septum is 
bony. There are two modern genera, Albula and Dix- 
otiia. 



Albula dunklei, new species. Figures 194, 200 B-C, 
201 B. 

Type. — P 27494, part of "body," scales in place, 
fins, part of a head, and vertebrae; Moore's farm, Dallas 
County, Alabama, middle Mooreville. 

Referred specimens. — PF 3580, scales; Locality 1, 
6.2 miles W. of Aliceville, E. of Walter Dance's farm, 
Eutaw County, Alabama, lower Mooreville. PF 3564, 
scales and vertebrae; Locality 4, Hale's farm, 2 miles 
NE. of West Greene, Greene County, Alabama, middle 
Mooreville. 

Diagnosis. — The bones of the skull are proportion- 
ately much thicker than in the modern species of Albula. 
The pelvic fin has 11 rays. The symplectic is relatively 
larger and the quadrate is more triangular in outline 
than in other species of Albula. The lower jaw bears a 
tooth patch similar to that of Recent species. The 
scales (fig. 201 B) show four basal lobes; the basal regions 
of scales have granules which are arranged in antero- 
posterior rows as opposed to the dendritic pattern in 
Albula vulpes. The apical region of Albula dunklei is 
granulated. 




Fig. 194. Albula dunklei, n. sp., holotype, P 27494; A, posterior portion of ceratohyal; B, entopterygoid; C, portion of skull; D, left 
quadrate with toothed metapterygoid; E, right quadrate with symplectic; F, articulated body scales andjeft pectoral fin; G, left pelvic fin 
lacking first two rays; H, posterior view of left pelvic fin; I, ventral view of right pelvic fin; J, ventral view of right pectoral fin; K, two 
fused branchial elements; L, base of skull with parasphenoid; M, three caudal vertebrae; N, eight abdominal vertebrae. 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



413 



Discussion. — The type (fig. 194), consists of: A, the 
posterior portion of a ceratohyal that in relative size 
compares favorably with the Recent Albula vulpes; B, 
the endopterygoid, which has veiy small, rounded, crush- 
ing teeth; C, probably the palatine element; D, the 
elongate left quadrate in external view; E, the right 
quadrate and symplectic; F, the right pelvic fin sur- 
rounded by body scales in place; G, H, and J, various 
parts of the pelvic fins with at least ten fused rays; J, 
the right pectoral fin; K, two fused branchial elements; 
L, the posterior base of the skull, covered by two wings 
of the parasphenoid; M, three caudal vertebrae with 
processes; N, eight abdominal vertebrae. 

The vertebrae of Albula dunklei show, externally, 
longitudinal striae. The known vertebrae are deeper 
than long. The caudal vertebrae are also deeper than 
wide. In cross-section the thoracic (fig. 200, B) or ab- 
dominal vertebrae show eight groups of radiating bony 
supports, one dorsally with two bands of bone and one 
dorso-laterally on each side with two bands; between 
these groups of bony supports are areas filled with 
spongy bone that meet concave upper borders. The 
lateral groups have three bands; below these there is on 
each side a ventrolateral group with two bands, and 
ventrally there is a wide median group with eight bands 
of bony supports, two pairs of bands on each side of the 
vertebra. Between this median ventral group and the 
ventro-lateral groups are also areas of spongy bone. The 
caudal vertebrae (fig. 200, C) have eight radial bands 
corresponding in position to the eight groups of bands 
of anterior vertebrae. There is a neural and a haemal 
arch fused to the top and bottom of each caudal ver- 
tebra. No areas of spongy bone have been noted in 
the caudal vertebrae. Almost identical vertebrae occur 
in the Recent Albula vulpes. 

This species is named for Dr. David Dunkle, who 
has been instrumental in the identification of the pres- 
ent albulid. 

The Albulidae have a spotty Upper Cretaceous rec- 
ord, due in part to the fact that many of the specimens 
in collections have not been described, while other spe- 
cies are known only from their scales, such as David's 
( 1946) Kleinpellia. Cockerell (1933) described, from a 
scale, a species called Albula antiqua from the Creta- 
ceous of Florida. The University of Kansas Museum 
has a specimen (F 985) from the Niobrara labeled Lep- 
tichthys which is an unquestionable albulid. The U. S. 
National Museum has a similar undescribed albulid, 
also from the Niobrara. Field Museum has a specimen, 
PF 685, from Sahel-Alma, Mt. Lebanon, which is a 
member of the genus Albula. 

Albula sp. Figure 195. 

Referred specimen. — P 27392, part of palate with 
teeth; Moore's farm, Dallas County, Alabama, middle 
Mooreville. 

Discussion. — There is at least one bit of evidence 
for the existence of a second species of Albula in the 
Mooreville. This consists of part of a palate with the 






;. 




10 mm 

Fig. 195. Tooth-bearing plate of Albula sp. (P27392). 

characteristic button-like teeth. The fragment indi- 
cates a fish larger than the Recent Albula vulpes, and 
the teeth are at least twice as large as those in Albula 
dunklei. 

Superfamily PLETHODOIDEA, new superfamily 

Diagnosis. — The parietals meet in the midline; there 
is no gular plate. The maxillaries enter into the gape of 
the mouth. The premaxillaries may be fused to the 
ethmoid. The ethmoid varies in relative size and is 
often huge. The jaws are toothed, and the parasphe- 
noid and basibranchials bear crushing teeth. The fron- 
tals are rectangular; nasals are united anteriorly to the 
frontals. The orbitosphenoids are large. The pectoral 
fins are high on the body. The dorsal fin covers most 
of the back. Scales {Bananogmius) have circuli on basal, 
dorsal, and ventral surfaces; the apical region has granu- 
lations arranged in postero-anterior rows as are the 
radii; the scales do not have a true radial or dendritic 
pattern as in Pachyrhizodus. The vertebrae have ex- 
ternal striations and dorsally two concavities; in cross- 
section they have numerous radial supports separated 
by spongy bone. 

Two families, Bananogmiidae and Plethodidae, are 
included in this superfamily. 

Bananogmiidae, new family 

Diagnosis. —No mucus canals are known on the fron- 
tals. The mandible is deepest at its center. The basi- 
branchial bears only one crushing toothed plate. The 
preoperculum is L-shaped, with the vertical limb nar- 
row and the anterior limb expanded. 

Discussion. — The following three genera are referred 
here: Bananogmius, Paranogmius, and Moorevillia. The 
genus Bananogmius as known at present comprises two 



414 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 




•n&2K 



■ n 









*& 






D E 






20 mm 

Fig. 196. Bananogmius crieleyi, n. sp., holotype, PF 3608. A, parasphenoid and vomer; B, right premaxillary, with anterior end up; 
C, basibranchial; D, abdominal vertebra; E, ectopterygoid; F, right quadrate, with anterior part up; G, ectopterygoid. 



groups. The first includes B. aratus and B. zitteli, both 
with very wide parasphenoid plates. The parasphenoid 
dental plate is oval in B. zitteli and rectangular in B. 
aratus. The parasphenoid dental plate of B. intermedins 
is unknown, but its basibranchials are closer to those 
of the above two species than to those of the second 
group. The premaxillary in B. zitteli is fused to the 
ethmoid. 

The second group of species consists of B. evolutus, 
B. favirostris, B. altus, B. polymicrodus, B. crieleyi, and 
B. sp. Loomis. Here the parasphenoid dental plate is 
elongate and club-shaped. The premaxillaries of B. 
evolutus (Loomis, 1900, pi. XXVI, fig. 6), B. favirostris, 
and B. polymicrodus are all very similar. The jaws of 
B. evolutus and B. polymicrodus are long and low. The 
parasphenoid of B. evolutus is unknown. 



Bananogmius crieleyi, new species. Figures 196, 
200 D. 

Type. — PF 3608, premaxillary, basihyal, parasphe- 
noid and vomer, two ectopterygoids, quadrate?, frag- 
ments of skull, and five vertebrae; Moore's farm, Dallas 
County, Alabama; middle Mooreville. 

Diagnosis. — The parasphenoid (fig. 196 A) is elon- 
gate, with a club-shaped dental patch; fused to it is the 
vomer with an oval tooth patch. The basibranchial 
(fig. 196 C) is very thick, five-sided, and with a deep 
indentation posteriorly. The premaxillary is free and 
bears several rows of small teeth; its external surface 
(fig. 196 B) is ornamented with pits similar to the tooth 
sockets in the basibranchial. Two large elongate tri- 
angular pterygoids (fig. 196 E and G) are present; the 
quadrate (fig. 196 F) is rounded at the top. The verte- 




20 mm 



Fig. 197. Moorerillia hardi, n. sp., holotype, PF 3567. A, left palatine, ventral view; B, right palatine, dorsal view; C, portion of ? 
three branchial arches; D, two abdominal vertebrae; E, part of gill arch or skull; F, portion of skull with two foramina; G, posterior portion 
of left lower jaw; H, upper part of first left pectoral fin ray; I, first right pectoral fin ray; J, posterior portion of parasphenoid; K, medial 
view of left premaxillary; L, lateral view of right premaxillary; M, left pterygoid; N, anterior end of left maxillary; O, anterior end of 
right maxillary; P, dorsal view of left lower jaw; Q, medial view of right lower jaw; R, top of skull; S, portion of base of skull; T, quadrate 
with symplectic; U, tooth patch from right pterygoid. 



415 



416 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



brae (fig. 196 D) have linear pits externally, which are 
deep and irregular; a cross-section of one is illustrated 
on figure 200 D, and shows radiating spongy bone. 

Discussion. — The narrow parasphenoid places this 
form in the second group of the genus Bananogmius. 
The elongated parasphenoid tooth plate and the oval 
tooth plate on the vomer are similar to those of B. 
favirostris. The shape of the vomerine tooth plate is 
similar to that of B. altus. The basibranchial is remi- 
niscent of that of Plethodus pentagon from the British 
Chalk and of B. evolutus. The combination of these 
characters is, however, unique. This species is named 
for Bruce Crieley of Chicago, Illinois, who served as 
my field assistant in Alabama. 

Bananogmius cf. zitteli Loomis. Figure 198 H. 

Referred specimen. — PF 3609, snout; Moore's farm, 
Dallas County, Alabama, middle Mooreville. 

Discussion. — This large pitted snout is too fragmen- 
tary for positive identification; however, it resembles 
closely the rostrum of the fish described by Loomis 
(1900, pp. 229-234, PI. XXI). 

Bananogmius cf. polymicrodus (Stewart). 
Figure 201 C. 
Referred specimen. — PF 3566, tail with scales and 
vertebrae, part of skull; Hale's farm, 2 miles NE. of 
West Greene, Greene County, Alabama, middle Moore- 
ville. 

Discussion. — The scales and vertebrae agree with 
those of B. polymicrodus. The scales bear circuli in 
their basal, dorsal, and ventral regions. The apical 
region has vermiculate granules which lie between smooth 
bands running antero-posteriorly rather than radiating 
from the nucleus. The antero-posterior alignment of 
the bands distinguishes these scales from those of Pachy- 
rhizodus. 

Bananogmius sp. Figure 198 G. 

Referred specimen. — PF 131, parasphenoid and den- 
tal plate; 1 x /i miles W. and % of a mile N. of West 
Greene, Greene County, Alabama, middle Mooreville. 

Discussion. — There is a very close resemblance be- 
tween this parasphenoid plate and that figured by 
Loomis (1900) as Bananogmius sp. As no other material 
is at hand, this fish must still remain specifically in- 
determinate. 

Moorevillia, new genus 

Type species. — Moorevillia hardi, new species. 

Diagnosis. — The premaxillary is free from the eth- 
moid, is small, and lacks prominent external ornamen- 
tation. There is only one row of conical teeth on the 
premaxillary. The lower jaw is long and narrow, with 
an elongate patch of small, recurved, enamel-tipped 
conical teeth. The maxillary is also a long thin element 
bearing a similar patch of teeth. The premaxillary 
fits into the abruptly up-turned anterior end of the 
maxillary. The palatines are small oval plates bearing 
patches of conical teeth, which stand over cup-shaped 
pits in the bone. The pterygoids are very elongate, 



and have a claw shaped posterior end with a large 
tooth patch. The posterior portion of the parasphenoid 
lacks wings. The frontals taper markedly anteriorly. 
The quadrate is high. The first pectoral fin ray is 
robust. The vertebrae are about three times higher 
than long, and show radiating bands of spongy bone 
in cross-section. 

Moorevillia hardi, new species. Figure 197. 

Type. — PF 3567, skull fragments, fin rays, verte- 
brae, and jaws; Moore's farm, Dallas County, Alabama, 
middle Mooreville. 

Diagnosis. — Same as that of the genus. 

Discussion. — This genus agrees with Bananogmius in 
the elongate patch of teeth on the lower jaw and the 
single row of teeth on the premaxillary. No crushing 
tooth pad like that of Bananogmius has been found. 
The premaxillary is decidedly smaller and lacks the 
ornamentation characteristic of Bananogmius. As far 
as can be told in the present specimen the parietals are 
separated by the supraoccipitals. It appears best to 
consider this a member of the Bananogmiidae, which 
it resembles in such features as vertebrae with fine 
external striae, and teeth arranged in characteristic 
patches. American Museum of Natural History speci- 
men No. 8319 from the Niobrara formation probably 
belongs to this genus and species. 

This species is named for Allen M. and Robert H. 
Hard, of Tuscaloosa, Alabama, who collected some of 
the fish material described in this paper. 

Suborder CLUPEOIDEI 

Diagnosis. — No posttemporal fossa is present, the 
parietals are separated from each other by a large crested 
supraoccipital. The orbitosphenoids and basisphenoid 
may be well developed or absent; there are one or two 
supramaxillaries; there is no gular plate. There is no 
adipose fin. Postcleithra are present; there are en- 
larged scales behind the paired fins. The body scales 
are generally wider than long, with circuli usually ob- 
scure. There are at least two superfamilies, the Chiro- 
centroidea and the Clupeoidea. 

Superfamily CHIROCENTROIDEA, 
new superfamily 

Diagnosis. — The supraoccipital crest is very large. 
The pectoral radials are in two rows. The premaxil- 
laries usually bear a few larger teeth than on the maxil- 
laries. Each palatine bone has a hammer-like articular 
process. The parietals are small. The anal fin is oppo- 
site the dorsal. The vertebrae have deep grooves on 
their sides, with two pits above and below for insertion 
of neural and haemal arches. A cross-section of a ver- 
tebra shows spongy bone without radial or concentric 
structures. 

There are three families: Ichthyodectidae, Sauro- 
dontidae, and Chirocentridae. 




^ 








? fXSl 



^efc* 





G.-. % -V. *-T 




20 mm 



Fig. 198. A-F, Siratodiis apicalis; A, fragment of operculum, PF 289; B, maxillary, PF 289; C, palatine, PF 132; D, internal view of 
left lower jaw, PF 289; E, fragment of right lower jaw, PF 289; F, premaxillary; G, Bananogmuis sp., parashpenoid, PF 131 ; H, Hananog- 
mius cf. zitteli, rostrum, PF 3609. 



417 



418 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



Ichthyodectidae Crook 

Diagnosis. —There is no predentary element. The 
scales possess large tubercles and fine circuli; radii are 
present. The teeth are large and rounded in cross- 
section. No nutrient foramina occur below the internal 
alveolar border. The teeth are implanted in sockets. 

Ichthyodectes cf. ctenodon Cope 

Referred specimen. — PF 3576, lower jaw fragment; 
Hale's farm, west gully, Greene County, Alabama, mid- 
dle Mooreville. 

Discussion. — This fragment of a lower jaw agrees 
with that figured by Stewart (1900, PL XLIX, Fig. 5) 
as Ichthyodectes ctenodon. The round cross-section of 
the teeth separates Ichthyodectes from the Saurodon- 
tidae, and the lack of huge anterior teeth serve to differ- 
entiate this form from Xiphactinus audax. 

Xiphactinus audax Leidy. Figure 200 A. 

Referred specimens. — PF 3543, vertebrae, fragmen- 
tary skull; 2 miles N. of West Greene, Hale's farm, 
Locality 4, Greene County, Alabama, middle Moore- 
ville. PF 120, vertebrae; PF 125, lower jaw fragments; 
PF 129, lower jaw fragments; all from 1 1 2 miles W. 




I Omm 




20 mm 



Fig. 199. A, jaw of Saurodon 1 sp., medial view, PF 3585; 
B, Pachyrhizodus caninus, pectoral fin spines, PF 1696. 





5 mm 



10mm 



Fig. 200. A, Xiphactinus audax, vertebral cross-section; B-C, 
Albula dunklei, vertebral cross-sections; D, Bananogmius creileyi, 
vertebral cross-section, PF 3608; E, Saurodon leanus, vertebral 
cross-section. 



and % of a mile N. of West Greene, Greene County, 
Alabama. P 27503, skull fragments and one vertebra; 
P 27525, jaw fragments, skull fragments, and vertebrae ; 
P 27528, skull fragments, vertebrae, and fin spines; 
PF 3568, skull fragments; all from Moore's farm, Dallas 
County, Alabama, middle Mooreville. P 27524, palato- 
quadrate, ' ■> mile S. of Harrell's Station, Dallas County, 
Alabama, middle Mooreville. P 27498, five vertebrae; 
Crawford's farm, Hale County, Alabama; middle Moore- 
ville. P 27531, skull fragments and 11 vertebrae; P 
27534, skull fragments and vertebrae; PF 3541, verte- 
brae and part of a skull; all from Hewlett's farm, Greene 
County, Alabama, upper Mooreville. 

Discussion. — Cope's skull specimen of Portheus mol- 
ossus would make a far superior type to the second 
pectoral fin spine on which Leidy (1870) based Xiph- 
actinus audax, but Leidy's name has priority. There 
is still the possibility that the European generic names 
discussed by Woodward (1901) may be applicable. There 
is little doubt that the Mooreville Xiphactinus is the 
same as the large Niobrara species, X. audax; the other 
described Niobrara species are of questionable validity. 
A cross-section of a vertebra is figured in Figure 200 A. 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



419 




2 mm 




I m m 



2 mm 



Fig. 201. Scales; A, dussumieriine ?, PF 3594; B, Albula dunklei, P 27494; C, Bananogmius cf. polymicrodus, PF 3566. 



Saurodontidae Stewart 

Diagnosis. — A predentary is present. The teeth are 
compressed and are implated in sockets. 

This family has two closely related genera, Saurodon 
and Saurocephalus that have been synonymized by Hay 
(1903), but since no intermediate forms exist among the 
Mooreville specimens, it is thought best to keep the 
genera separate. 

Saurodon leanus Hays. Figure 200 E. 

Referred specimens. — PF 3611, lower jaw and skull 
fragments; 2 miles W., 1 mile N. of West Greene, 
Greene County, Alabama, middle Mooreville. PF 122, 
right lower jaw; 1.6 miles N. of West Greene, Greene 
County, Alabama, middle Mooreville. PF 3540. lower 



jaw and skull fragments; 1 mile W. of West Greene, 
Greene County, Alabama, middle Mooreville. PF 3544, 
jaw fragment; Locality 4, Hale's farm, 2 miles N. of 
West Greene, Greene County, Alabama, middle Moore- 
ville. PF 130, part of lower jaw; 2 miles N. of West 
Greene, Greene County, Alabama, middle Mooreville. 
PF 3548, right lower jaw; P 27413, left lower jaw and 
left premaxillary; P 27414, maxillary; P 27415, pail of 
lower jaw; P 27430, lower jaw and vertebra; P 27435, 
jaws and skull fragments; P 27483, better part of a skull 
and four vertebrae; P 27508, lower jaw and skull frag- 
ment; P 27530, part of skull and two vertebrae; all 
from Moore's farm, Dallas County, Alabama, middle 
Mooreville. P 27506, part of skull and lower jaw; 
P 27507, part of lower jaw; both from Township 11, 
W. of Highway 13, Hale County, Alabama, middle 



420 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



Mooreville. PF 123, skull fragments; l - 2 mile N. of 
Mt. Hebron, near West Greene Road, Greene County, 
Alabama, upper? Mooreville. 

Discussion. — The jaw of Saurodon shows deep notches 
below the alveolar border. The existence of more than 
one species is an open question. The present form is 
referred to S. leanus, the first described species, typi- 
cally from the Upper Cretaceous of New Jersey. A 
sectioned vertebra displaying spongy bone is shown in 
Figure 200 E. 

Saurodon? sp. 1. Figure 199 A. 

Referred specimen. — PF 3585, the better part of a 
small lower jaw; Moore's farm, Dallas County, Ala- 
bama, middle Mooreville. 

Discussion. — This small lower jaw is similar to that 
of Saurodon in jaw shape, but, perhaps because of its 
small size, lacks any of the deep notches of S. leanus. 
This jaw may represent a smaller species or a juvenile 
of S. leanus. 

Saurodon? sp. 2 

Referred specimen. — PF 3550, jaw fragment; Moore's 
farm, Dallas County, Alabama, middle Mooreville. 

Discussion. — This jaw fragment, probably a maxil- 
lary, is similar to the one above, except that the teeth 
are much smaller. 

Saurocephalus cf. lanciformis Harlan 

Referred specimens. — PF 443, upper jaw; W. of West 
Greene, Greene County, Alabama, middle Mooreville. 
PF 3557, portion of lower jaw; Township 11, W. of 
Highway 13, Hale County, Alabama, middle Moore- 
ville. P 27407, jaws; P 27412, lower jaw; P 27505, jaw 
and skull fragments; P 27509, jaws and skull fragments; 
P 27511, skull fragments; all from Moore's farm, Dallas 
County, Alabama, middle Mooreville. 

Discussion. — Saurocephalus is similar to Saurodon ex- 
cept in having foramina instead of deep notches below 
the alveolar border internally. The oldest name is 
used, as the other species are of doubtful validity. 

Superfamily CLUPEOIDEA 

Diagnosis. — The supraoccipital crest is reduced 
though still present; the postcleithra are attached to 
the outer side of the cleithra; the pectoral radiae are 
in one row. The teeth are small or absent. The pre- 
maxillae are "L" shaped. The dorsal fins are anterior 
to the anal fin. Temporal foramina and preepiotic 
fossae are present. The palatines are without hammer- 
like articulations. 

There is one family in the Mooreville; the Clupeidae. 
Clupeidae, indet. 

Referred specimen. — PF 3599, left preoperculum; 
Choctaw Bluff, Greene County, Alabama, lower Moore- 
ville. 



Discussion. — This small preoperculum with three 
very strongly marked sensory canals is of the type 
found commonly only among the Clupeidae. 

Dussumieriinae, indet. Figure 201 A. 

Referred specimen. — PF 3594, two scales; Montgom- 
ery's farm, 2 miles N. of West Greene, Greene County, 
Alabama, middle Mooreville. 

Discussion. — The anterior areas of these scales are 
missing, but the strong posterior radii as well as the 
grooves at right angles to them are typical of scales 
of the modern Dussumieriinae, a subfamily of the Clu- 
peidae. 

Order INIOMI 
Suborder MYCTOPHOIDEI 

Diagnosis. — The maxilla is only weakly if at all 
dentigerous, and is usually excluded from the gape of 
the mouth. The inner teeth in the mouth are generally 
larger than the outer. No mesocoracoid arch is known. 
An adipose fin is commonly present; the ventral fins 
are on the posterior part of the abdomen. 

The following Mooreville families are referred to 
this order: Enchodontidae, Dercetidae, and Myctophi- 
dae. I do not follow Romer's (1966) use of the super- 
order Protacanthopterygii. 

Enchodontidae Loomis 

Diagnosis- -The parietals are separated by a supra- 
occipital. The premaxillaries are large, and the maxil- 
laries, although small, are present in the gape of the 
mouth; each premaxillary has a posterior spine. The 
palatines bear one or two large fangs; each lower jaw 
has two tooth rows, with the outer row much smaller 
than the inner one; the teeth are fused to the jawbones. 
Scutes and scales are present on the body, and the 
surfaces of the cranial plates and scutes are highly 
ornamented. The nasals are small. 

Two Mooreville genera, Enchodus and Cimolichthys, 
are included in this family. 

Cimolichthys nepaholica (Cope) 

Referred specimens. — PF 441, two basal parts of fin 
rays; 2 miles W. of West Greene, Greene County, Ala- 
bama, middle Mooreville. PF 3565, quadrate, basal 
portion of fin ray, and skull fragments; 1 mile W. and 
1 mile N. of West Greene, Greene County, Alabama, 
middle Mooreville. P 27526, two basal parts of fin 
rays, Crawford's farm, Hale County, Alabama, middle 
Mooreville. 

Discussion. — The quadrate of PF 3565 is virtually 
identical with the specialized quadrate of C. nepaholica 
from the Niobrara. A fin ray associated with the quad- 
rate is thought to be either the first pectoral or pelvic 
ray. There is little doubt that Cope's (1872) Empo 
is really Cimolichthys. A number of species have been 
named, but there appears to be only one valid Niobrara 
species, C. nepaholica, as shown by Hay (1903). 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



421 



Enchodus petrosus Cope 

Referred specimens. — PF 3573, palatines, one lower 
jaw, and three teeth; Hale's farm, 2 miles N. of West 
Greene Post Office, Greene County, Alabama, middle 
Mooreville. PF 3574, two palatines; PF 3575, one 
palatine and one lower jaw fragment; Montgomery's 
farm, 2 miles N. of West Greene, Greene County, Ala- 
bama, middle Mooreville. PF 3583, one palatine; 2 
miles W. of West Greene, Greene County, Alabama, 
middle Mooreville. PF 3584, two lower jaw fragments; 
1 mile N., 1 mile W. of West Greene, Greene County, 
Alabama, middle Mooreville. PF 3587, three teeth, 
two jaw fragments, and one palatine; 1 mile N., 2 miles 
W. of West Greene, Greene County, Alabama, middle 
Mooreville. PF 3586, one palatine, two jaw fragments, 
and four teeth, 1 mile S. of West Greene, Greene County, 
Alabama, middle or upper Mooreville. P 27521, one 
tooth; Township 11, W. of Alabama Highway 13, Hale 
County, Alabama, middle Mooreville. PF 136, one 
palatine, jaw fragments, skull fragments, and verte- 
brae; PF 3569, teeth; PF 3570, six teeth; PF 3571, 
two teeth and one palatine; PF 3572, three jaw frag- 
ments; PF 3590, two palatines; PF 3591, one palatine; 
PF 3592, one palatine; PF 3593, two palatines, three 
jaw fragments, and one skull fragment; P 27417, jaw 
fragments and one palatine; all from Moore's farm, 
Dallas County, Alabama, middle Mooreville. PF 
3582, five jaw fragments, one palatine, and one skull 
fragment; Marion Junction, Dallas County, Alabama, 
middle Mooreville. PF 3555, one palatine; PF 3556; 
one palatine, PF 3577, one palatine and one lower jaw; 
PF 3578, one palatine, five jaw fragments, and one 
tooth; PF 3579, three palatines and two teeth; PF 3589, 
two palatines; all from Hewlett's farm, gullies near the 
county road, Greene County, Alabama, upper Moore- 
ville. PF 3581, one tooth; 8.9 miles from the county 
line, Route 26, Russell County, Alabama, Blufftown. 

Discussion. — Enchodus petrosus is one of the com- 
mon species in the Mooreville fauna. Almost any Moore- 
ville locality will produce some evidence, usually a pal- 
atine, of this species. The collected specimens show a 
good deal of variation. 

Enchodus cf. saevus Hay 

Referred specimen. — PF 3588, two lower jaws; 2 
miles W. and 1 mile N. of West Greene, Greene County, 
Alabama, middle Mooreville. 

Discussion. — The more ornate condition of these 
jaws is one difference from E. petrosus, and a similarity 
to the form described by Hay (1903) as Enchodus saevus. 
The relationship of E. petrosus to E. saevus is not at 
the present clear. Hay (1903) and others have used 
the cross-sectional shapes of teeth to distinguish these 
species, but this is not a satisfactory character due to 
individual variation and types of preservation. Hay's 
species was named from Niobrara material. 

Dercetidae Cope 
Diagnosis. — The head and trunk are elongate. The 
parietal bones are of moderate size, meeting in the mid- 



line. The gape of the mouth is wide; the premaxillaries 
form the greater part of the upper edge of the mouth. 
The vertebrae are in the form of constricted cylinders 
pierced by the notochord. Scales are small or wanting. 
A set of elongate scutes is often present. 

One genus occurs in the Mooreville: Stratodus. 

Stratodus apicalis Cope. Figure 198 A-F. 

Referred specimens. — PF 132, one palatine; 1 mile 
NW. of West Greene, Greene County, Alabama, middle 
Mooreville. PF 3610, one palatine; 2 miles W. and 1 
mile N. of West Greene, Greene County, Alabama, 
middle Mooreville. PF 3600, one palatine; P 27424, 
a caudal support; P 27487, a palatine and palatine 
fragments; P 27512, one palatine; all from Moore's 
farm, Dallas County, Alabama, middle Mooreville. PF 
3596, one palatine; Hewlett's farm, north set of gullies, 
Greene County, Alabama, upper Mooreville. PF 289, 
parts of both lower jaws, premaxillary?, opercular frag- 
ment, and skull fragments; Donald's farm, Dallas 
County, Alabama, middle Mooreville. 

Discussion. — There is little doubt that Stratodus oxy- 
pogon Cope is a synonym of S. apicalis; both are based 
on fragmentary material. The lower jaw (fig. 198 D E) 
has a slanting posterior edge with a small cup-shaped 
articular surface for the quadrate. The premaxillary 
(fig. 198 F) has a characteristic patch of teeth. The 
elongate palatines (fig. 198 C) seem the most common 
objects in collections. 

Myctophidae Jordan and Evermann 
Diagnosis. — The upper jaws are bordered by pre- 
maxillaries only; the dentition is usually feeble. The 
ventral fins may be abdominal or under the pectoral 
fins. The parietals are usually separated by a reduced 
supraoccipital. Scales are variable but with distinct 
circuli; their nucleus is apical or central; apical cteni 
are present or absent; an apical field may be present; 
cteni are usually in one row and marginal. 

Myctophid scales are very similar to those of bery- 
coids, and their recognition can be very difficult. The 
myctophids usually possess a single row of cteni or 
or none, whereas the berycoids have usually more than 
one row. The myctophids lack the raised apical field 
which is characteristic of many of the berycoids. 

Myctophidae, indet. Figure 203. 
Referred specimens. - -PF 3598, numerous scales; 
Choctaw Bluff, Greene County, Alabama, lower Moore- 
ville. PF 3601, one scale; PF 3602, lower jaw; both 
fi'om east slope above flood plain of Pintlalla Creek 
on Burksville Road, Montgomery County, Alabama, 
lower Mooreville. PF 3595, one scale; Montgomery's 
farm, 2 miles N. of West Greene, Greene County, Ala- 
bama, middle Mooreville. PF 3597, numerous scales; 
Hewlett's farm, gullies near road, Greene County, Ala- 
bama, upper Mooreville. 

Discussion. — The great number of myctophid scales 
with vertebrae and head plates, the latter almost im- 



422 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 




Fig. 202. Scales; A-B, teleost indet.; A, PF 3607; B, PF 3605; CD, Hoplopteryx ? (C, PF 3603). 



possible to prepare, show differences that leave little 
doubt as to the presence of more than one species in 
the Mooreville; however, since any patch of scales will 
show more than one type of scale, it seems that the 
best course is to describe the predominant scale types. 
Type A (fig. 203 A) is a triangular scale with rounded 



outer angles. The flattened posterior edge of the scale 
bears a number of spines (cteni) in a single row. The 
basal (anterior) portion is the apex of the triangle. 
The nucleus is at or near the base of the scale. The 
circuli are close together but do not normally make 
complete circles anteriorly and become almost straight 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



423 




Fig. 203. Myctophid scales; four different kinds described in text. 



posteriorly. A variant of Type A has a sub-basal nu- 
cleus with the first few circuli complete, and has the 
circuli widely spaced. 

Type B (fig. 203 D) has the nucleus centrally located 
and the circuli placed very far apart. The scale is 
subrectangular with a convex protruding lip in the cen- 
ter of the basal edge. Posteriorly there is one row of 
cteni, ten or more in number. A variant of this type 
shows finely spaced circuli and a posterior nucleus at 
the edges as in Type D. 

Type C (fig. 203 C) is a rounded scale except for a 
convex protruding basal lip as in Type B. The nuclear 
area is very large; circuli are few and widely spaced; 
no cteni are present. A variant of this type shows 
closely spaced circuli very reminiscent of the condition 
met with in the Salmonidae. 

Type D (fig. 203 B) is wider than long, with a 
rounded posterior edge; the basal edge is pectinated. 
The nucleus is posterior, and radii are variable in num- 
ber. The circuli are closely spaced; cteni may or may 
not be present in this type. 

All these scales are very close to what is found in 



the Recent Myctophidae and compare very well with 
the scales in the genus Myctophum. David (1946) has 
described somewhat similar scales from California and 
has assigned them to the genus Sardinioides, placed 
in the Chlorophthalmidae. 

Order BERYCIFORMES 

Diagnosis. — The premaxillaries are protractile and 
the dentition is feeble. There are seven to eight bran- 
chiostegals. The vertebral count is from 24 to 30. The 
scales have a raised apical area and several rows of cteni. 

Trachichthyidae Bleeker 

Hoplopteryx? sp. Figure 202 CD. 

Referred specimens. — PF 3603, scales in coprolite 
with Palelops; Hewlett's farm, north set of gullies, 
Greene County, Alabama, upper Mooreville. PF 3606. 
one scale; Montgomery's farm, 2 miles N. of West 
Greene, Greene County, Alabama, middle Mooreville. 

Discussion. — The Mooreville berycoid scales with 
their raised posterior area and numerous cteni agree 
with the scales of Hoplopteryx superbus figured by Wood- 



424 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



ward (1902, PI. VII, Figs. 3 and 4) from the British 
Chalk. Woodward's scales lack cteni, yet in the Recent 
as well as the fossil Berycidae the cteni easily fall off. 
The basal plications on these scales suggest those found 
in Elops. The fact that scales of this form were found 
intact in a coprolite points to the importance of copro- 
lites in preserving faunal elements that might other- 
wise be lost. Recently Patterson (1965) has demon- 
strated that Hoplopteryx belongs in the family Trachi- 
chthyidae and not the Berycidae. 




Fig. 204. Operculum referred to family Trachichthyidae ?, 
PF 3604. 

Trachichtyidae? indet. Figure 204. 

Referred specimen. — PF 3604, an operculum; 6.2 
miles W. of Aliceville, E. of Walter Dance's farm, Eu- 
taw County, Alabama, lower Mooreville. 

Discussion. — This record is based on an elongate 
opercular element with a distinctively ornamented sur- 
face consisting of ridges topped in places by rounded 
protuberances. The ridges radiate from the upper an- 
terior edge just above the socket for the opercular pro- 
cess of the hyomandibular. The whole operculum is 
higher than long, and the complete element would be 
about 40 mm. high. 

Teleost incertae sedis (1). Figure 202 B. 

Referred specimen. — PF 3605, one scale; Montgom- 
ery's farm, 2 miles N. of West Greene, Greene County, 
Alabama, middle Mooreville. 

Discussion. — This small scale has a basal nucleus 
and four radii which end probably in spine-like points. 
The circuli go in a straight path from one radius to the 
next, except at one radius in the anterior portion of 
the scale where they form an acute angle and approach 
the nucleus. In many respects this scale agrees with 
that of Caulolepis longidens, a Recent berycoid, yet till 
more evidence is available it is best to list this very 
unusual scale as incertae sedis. 



Teleost incertae sedis (2) . Figure 202 A. 

Referred specimen. — PF 3607, one scale; east slope 
above flood plain of Pintlalla Creek on Burksville Road, 
Montgomery County, Alabama, lower Mooreville. 

Discussion. — This indeterminate but highly distinc- 
tive scale is closest to those of the myctophids. The 
nucleus is possibly posterior. The surface is covered 
by distinctive granules. There is no evidence of cteni. 

ECOLOGY OF THE MOOREVILLE CHALK 

The Evidence From the Overall 

Geological Picture 

The geological setting of the Mooreville Formation 
of the Selma group has been discussed in some detail 
by Monroe (1941) and by Zangerl (1948), who gives a 
stratigraphic column. The Upper Cretaceous sediments 
of Alabama in their area of outcrop lie on an eroded 
surface of rocks ranging in age from the Pennsylvanian 
to the Pre-Cambrian. The Tuscaloosa Formation rests 
upon this basement complex and is the basal Upper 
Cretaceous Formation in most of the area in question. 
According to Monroe (1941), the Tuscaloosa is com- 
posed of irregularly bedded clays, sands, and gravels; 
these sediments are said by him to have been laid 
down on stream deltas and in part on bay shores and 
marshes. Berry (1919) has described a number of ter- 
restrial plant fossils from this formation based upon 
leaf remains. Fossil wood is reported to be common. 
Mollusks are rare. Marine facies of this formation are 
reported down dip in oil company drill holes, and some 
drill cores at the Alabama Geological Survey contain 
scales of marine fishes. 

Overlying the Tuscaloosa is the Eutaw Formation, 
which consists of fine gravels at the base that grade 
into highly crossbedded glauconitic sands (Monroe, 
1941). At the top of the Eutaw, the Eutaw-Mooreville 
contact is not sharply defined. There is a great re- 
duction of glauconitic sands at the base of what is gen- 
erally considered to be Mooreville, and in a few feet 
above these beds chalk dominates. The glauconitic 
sands continue, though in sharply decreasing amount, 
into the Mooreville chalk; minute amounts of glau- 
conite are present throughout the Mooreville and even 
the overlying Areola member. 

If we consider this sequence of beds from Tuscaloosa 
through the Mooreville as a unit, there is an overall 
decrease in sediment particle size from the bottom to 
the top, perhaps connected with a diminishing of cur- 
rent phenomena; this is accompanied by a shift from 
heterogeneous sediments to relatively homogeneous sed- 
iments, which could be indicative of an increase in 
depth. The situation suggests a slowly submerging 
coast line: Tuscaloosa-like sediments are followed by 
shallow water Eutaw-like sediments which in turn are 
followed by deeper water Mooreville-like sediments. It 
seems likely that at any one time all these three types 
were being deposited, and that the Eutaw sediments 
were seaward of the Tuscaloosa, as the Mooreville were 
seaward of the Eutaw. 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



425 



The Areola Limestone member is a very thin layer 
(five feet at the type locality) which overlies the chalk; 
the contact is not sharply defined. The upper part of 
the chalk has numerous limy lenses which become 
more abundant as one nears the Areola. The Areola 
Limestone might represent the deepest-water sediment 
of all the formations that have been mentioned. 

The Mooreville is strikingly homogeneous along its 
area of outcrop. The lower layers look identical in 
composition even when seen at localities a hundred 
miles or more from each other. The upper layers, al- 
though not examined over such great distances, give 
the same impression. These remarks apply not only to 
the sediments but also to the fauna. One must assume 
fairly deep water to explain this great conformity over 
such wide areas. 

We have at present no information concerning con- 
temporaneous shoreward deposits directly north of the 
Mooreville outcrop area. Toward the northwest, in 
Tennessee, the Mooreville intertongues with the Coffee 
Sands, which resemble the Eutaw in fauna as well as in 
sediments. To the east the Mooreville is replaced by 
the Blufftown, a formation containing a more clay-like 
sediment which has abundant Ostrea and Exogyra reefs, 
probably an indication of shallower water. A more 
diverse molluscan fauna is present although poorly pre- 
served. Down dip and subsurface in the direction of 
Florida and southern Alabama the Mooreville is re- 
placed by a series of limestones and glauconites which 
may indicate either shallower depths or deep water 
glauconites. 

If the present strike of the sediments is close to the 
original shore line, the Mooreville was deposited at the 
end of a peninsula which was formed by pre-existing 
Appalachian structure; the bulk of deposition of clastic 
material lies to the northwest and northeast of the 
Mooreville. In the Mooreville outcrop area there must 
have been a lack of major stream entry, resulting in 
minimal clastic sedimentation. 

Ecological Evidence From the Sediments 

The remarkable homogeneity of the Mooreville sedi- 
ments lends itself to an attempted environmental re- 
construction. Any environmental analysis should con- 
sider inorganic as well as organic evidence. This anal- 
ysis is here confined to the Mooreville area of outcrop 
in Alabama. Since this is obviously a marine sediment, 
other environments are not considered. 

Mooreville Chalk sediments are very similar to other 
Upper Cretaceous chalks. In total bulk, clay materials, 
in the clay size range, make up the greater portion of 
the chalk. Calcareous particles ranging from less than 
one micron to two or three millimeters form the next 
highest percentage. A small percentage of quartz grains 
is present in all the samples. Some chert, similar if 
not identical to that of the Alabama Mississippian for- 
mations, occurs in the larger size ranges. Glauconite 
pellets and mica can be found in any sample of the 
Mooreville. There is a suite of heavy minerals all of 



less than two millimeters in their greatest diameter. 
Secondary minerals include gypsum crystals, calcite 
crystals, lenses of limestone, and nodules of chalcopy- 
rite. The latter range from less than one-fourth of an 
inch to nine inches in their greatest diameter, and show 
their secondary nature in replacement of organic por- 
tions of the chalk. 

The calcareous portion of the chalk has a high or- 
ganic content consisting of bones, plates, and tests of 
marine organisms. Coccoliths and rhabdoliths, which 
are the calcareous disks and platelets of Chrysophyta, 
are an important constituent. These algal platelets 
have a size range of from one micron to about thirty 
microns. Every handful of chalk normally contains a 
number of Inoceramus shell prisms, ostracod tests, Fora- 
minifera shells, small Ostrea, and bone fragments. Phos- 
phate nodules in the form of worm coprolites occur in 
vast numbers. If one puts such a sample of Mooreville 
chalk in water it will be altered to a very fine ooze. It 
is highly probable that the ocean bottom was made up 
of such an ooze throughout most of the deposition of 
the chalk. The oozey nature of the sediments would 
account for the restricted epifauna. A lack of oxygen 
characteristic of present oozes would account in part 
for the scarcity of an infauna. The small size of inor- 
ganic particles suggests a great distance from areas of 
active erosion as well as from major drainage systems. 
The small clastic portion of the sediment might have 
been swept in by turbidity currents. 

Only at Moore's farm (Locality 12A) is there any 
good evidence of current action. At this locality there 
is an exposed layer of shell and bone conglomerate less 
than an acre in extent and less than one foot thick. 
In this layer, presumably due to removal of the finer 
particles, there has been a concentration of larger or- 
ganic remains such as Inoceramus prisms, ostracods, 
teleost scales, vertebrae, and teeth, shark denticles and 
teeth, small Ostrea, small gastropods, and coiled cepha- 
lopods. The abundance of vertebrate material makes 
this layer almost a bone bed. In the Fort Hays, Kansas 
State College collection there are samples of a similar 
consolidated layer from the Niobrara. 

If the Mooreville Formation had been deposited in 
shallow water, i.e., between one and 300 feet in depth, 
one would expect a sediment notably disturbed by wave 
and tidal action. The absence of current phenomena 
(except at Locality 12A) and the great homogeneity of 
the Mooreville Chalk are not compatible with a shallow 
water interpretation. 

Ecological Evidence From the Plants 

Plant remains as a whole are rare in the Mooreville 
except for the flood of microscopic "armor" plates, coc- 
coliths and rhabdoliths, from single celled algae be- 
longing to the phylum Chrysophyta, family Coccolitho- 
phoridae. Members of this group of flagellates abound 
today in temperate and tropic open ocean surface wa- 
ters. The small size of the plates and the small size of 
the organisms that bear them has until recently re- 



426 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



tarded the study of this group from a biological as well 
as a geological viewpoint. The coccoliths are common 
in numerous Jurassic, Cretaceous, Tertiary, and Re- 
cent sediments. The organisms themselves are so num- 
erous that they may well serve as the primary synthe- 
sizers, at the base of the open sea food chain. 

At Choctaw Bluff and Hewlett's farm obscure chal- 
copyritized impressions of what evidently were marine 
algae are present but not common. The fragmentary 
nature of these remains suggests that they may have 
floated into the area of deposition. Fossil wood is 
found at a number of Mooreville localities but it is 
rather rare; the fact that it is bored by marine mollusks 
indicates that it had floated for some length of time. 
No leaves have been found in the Mooreville though 
the Field Museum collection contains a few specimens 
tentatively identified as willow galls, which could prob- 
ably have floated with ease. None of the above evi- 
dence necessarily suggest immediate proximity to land 
nor particularly shallow water. 

Ecological Evidence From the Invertebrates 

The common invertebrates that can be found in 
any sample of Mooreville Chalk are the calcareous and 
arenaceous Foraminifera, worm pellets, Inoceramus 
prisms, and Ostrea spat. At any locality one can expect 
Ostrea congesta and usually another species of Ostrea, 
the large shells of Inoceramus, and the plates or spines 
of echinoderms. At a number of localities there were 
worm tubes belonging to Hamulus and shells of the 
pelecypod Durania. Encrusting Bryozoa (Zangerl, 
1948, PI. 2) were present at four localities. 

This fauna seems to be composed of pelagic, neritic, 
epifaunal, and infaunal elements. The pelagic forms 
are Teredo in driftwood, a number of the Foraminifera, 
the small ammonites, gastropods, and ostracods. The 
genus Pecten may be a neritic element at least part of 
the time, but the small size is suggestive of the deep 
water pectens that occur off our coasts today. The 
epifaunal elements such as Exogyra, Gryphaea, Para- 
nomia, Inoceramus, and the radiolite Durania, some 
Foraminifera, and probably some of the ostracods, are 
those adapted to soft bottom conditions. Hamulus oc- 
curs as encrusting species as well as unattached species. 
The encrusting forms in addition to Hamulus include 
Ostrea congesta, Bryozoa, the boring sponge Cliona, and 
the barnacle Scapellum. Worm burrows and pellets 
represent the infaunal elements. 

Ostrea congesta occurs commonly on Inoceramus 
shells, rarely on vertebrate remains. In both the Nio- 
brara and the Mooreville Formations the large species 
of Inoceramus have Ostrea congesta on the outer sides 
of both valves, even when the two valves are found in 
articulation. This could occur if the huge thin-shelled 
Inoceramus had a habit of keeping its outer edge up 
and hinge down in the ooze. 

The large conical Durania may have existed par- 
tially buried in or on top of the bottom ooze. Some 
individuals of Durania have other Durania shells at- 
tached to them, showing a tendency toward bioherm 



formation but not, however, toward the formation of 
true reefs. 

It should be noted that although reef-forming genera 
such as Ostrea and Exogyra are found, no reefs occur 
in the Mooreville. The typical near-shore Upper Cre- 
taceous molluscan fauna, such as found at Coon Creek, 
Tennessee, is lacking or represented by few individuals. 
Burrowing mollusks are not present; this may be due 
to the suffocating ooze conditions or to the depth of 
water, perhaps both. 

Invertebrates were taken from 18 Mooreville lo- 
calities. At every locality, Foraminifera, worm pellets, 
ostracods, echinoderm remains, Ostrea congesta, Ostrea 
sp. spat, and large Inoceramus shells were collected. 
Durania austinia was taken from 10 localities. Exogyra 
ponderosa, Ostrea sp., and encrusting Bryozoa were 
found at four localities. Ostrea cretacea, a small shelled 
Inoceramus, the wood boring Teredo, and the barnacle 
Scapellum were collected from three localities. A soli- 
tary coral, the worm Serpula sp., worm burrows, the 
brachiopod Lingula sp., Ostrea plumosa, 0. bleckensis, 0. 
mesentarica, Exogyra uptoiensis, Gryphaea vesicularis, 
G. vomer, and Pecten sp. were taken at two localities. 
Each of the following is known from a single locality: 
Cliona sp., the boring sponge, Hamulus major, H. onyx, 
Nucula sp., Ostrea lava, 0. falcata, 0. spatulata, Pecten. 
cf. simplex, Paranomia scabra, small undetermined gas- 
tropods, the nautiloid Eutrephoceras dekayi, small am- 
monites, a single fragment of a large ammonite, and 
several crab claws. 

An infauna characteristic of modern shallow muddy 
bottoms is generally absent in chalk deposits, though 
other contemporaneous Upper Cretaceous clays show 
many of the pelecypod and gastropod genera that still 
live today in muddy bottom sediments. The absence 
of such an infauna in the chalks may be due to the depth 
at the time of deposition. In depths of less than 200 
feet one would expect to find some evidence of reef 
formation, either by Exogyra, Ostrea, radiolites, corals 
or algae. Though the oozy nature of the sediment could 
be an inhibiting factor, the shells that do occur, or 
even the vertebrate remains, could have acted as a 
temporary platform for reef formation. If the depth 
were too great, however, the lack of light and food 
would preclude this possibility. The vast amount of 
planktonic material in the Mooreville must have been 
formed in an open sea environment, possibly beyond 
the depositional area, and may have been carried into 
the Mooreville area of deposition by a Gulf Stream-like 
current moving at the surface. In my opinion, the 
evidence of the sediments, plants, and invertebrates 
indicates a depth of more than 600 feet or 100 fathoms, 
if it is to be compared with Recent conditions, for it is 
beyond the 100 fathom line that Recent deep-water 
conditions begin to appear. 

The Ecological Evidence From the Fishes 
Conditions of Preservation 

The Mooreville fish remains, with the exception of 
those from the Moore farm Locality 12A, consist of 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



427 



isolated and generally largely disarticulated fish evi- 
dently deposited at random in the sediment. Only 
one nearly complete articulated specimen has been col- 
lected. The remains generally consist of scattered 
patches of scales, vertebrae, and skull elements, all of 
which show some degree of articulation. Sharks may 
have teeth, denticles, and vertebrae associated. In- 
dividual scales or teeth often occur alone. The scatter- 
ing of remains is most likely due to (1) predation in 
life, leaving only uneaten parts of fishes to fall to the 
bottom, (2) decay of dead fishes while still floating, or 
(3) disturbance by scavengers before burial. 

In the following sections each of the groups of fishes 
in the Mooreville is discussed as to its ecology as indi- 
cated by the modern and Tertiary counterparts. 

Chimaeroids 

The Edaphodontidae are extinct. The known depth 
distribution of the six modern genera of the order Chi- 
maeriformes is given below (in fathoms) : 



Chimaera 

Hydrolagus 

Callorhynchus 

Rhinochimaera 

Harriotta 

Neoharriotta 



1 to 600 

1 to 1290 

1 to 100 

200 

375 to 1422 

300 plus 



The first three genera have a wide depth range, 
though they have been found most commonly in shallow 
water well up on the continental shelf. The last three 
genera are specialized deep water forms. As has been 
stated earlier, the Edaphodontidae are particularly close 
to the Recent genus Callorhynchus. Graham (1956) 
has discussed the natural history of this form in New 
Zealand. There are five recognized species, which are 
circumpolar and confined to the Southern Hemisphere. 
During the warm portion of the year, the New Zealand 
species moves into the bays and even ascends rivers; 
during the cold season it moves out into deeper water. 
Graham (1956) states that this species needs highly 
oxygenated water. Callorhynchus feeds on mollusks, 
fish, arthropods, crabs, shrimp, and jellyfish (Graham, 
1939). One may conclude from the size of the dental 
plates in Edaphodon that the Mooreville forms reached 
gigantic proportions as compared with the Recent Cal- 
lorhynchus. 

Large chimaeroids today are characteristic of deep 
water, while shallow or coastal water forms seldom ex- 
ceed two feet, as in Callorhynchus. In the Recent fauna 
four and five foot long deep-water specimens of Hy- 
drolagus and Chimaera are known. The Mooreville 
edaphodontids were probably much larger and may 
have reached a length of over eight feet, a size favoring 
a deep water habit. 

Sharks 

The Selachii in the Mooreville are represented by 
isolated individuals and are thinly distributed through- 
out the formation. (The one known exception to this 
is Locality 12A. where a consolidated layer shows, among 



other things, a great many mixed shark teeth, verte- 
brae, and denticles.) The fact that the denticles adhere 
to the vertebrae, a phenomenon common in the Nio- 
brara specimens at the University of Kansas as well as 
in the Mooreville specimens, speaks for fairly rapid 
burial by sinking into the bottom ooze, then decay in 
situ, with a lack of strong bottom currents. 

The Ptychodontidae are a predominantly Upper 
Cretaceous group of sharks, known only by isolated 
teeth, tooth sets, and doubtful vertebrae. The wide 
geographic range of Ptychodus speaks for a broad oceanic 
distribution. While nothing is known of the feeding 
habits of this group, the battery of row upon row of 
large teeth (over 600 in P. mortoni according to Wood- 
ward, 1902-1912) would have made this shark an ex- 
cellent invertebrate feeder, perhaps living on Crustacea 
and pelagic mollusks. 

The Anacoracidae have been thought to show a 
reduction in dentition leading toward the Cetorhinidae. 
Most recent evidence supports the theory that this 
family may represent a direct ancestor of the primitive 
orectolobids and is not related to Cetorhinus at all. 
Its teeth were well adapted for fish eating, and the well 
calcified vertebral column suggests that it was a power- 
ful swimmer. The wide distribution indicates that this 
may have been a pelagic shark. 

The Odontaspididae are represented in the Moore- 
ville by the genus Scapanorhynchus. The large size of 
the teeth indicates a shark of about ten feet in length. 
Modern odontaspids occur in both coastal and deep 
water; at least one is known to be truly pelagic. The 
wide distribution of Scapanorhynchus is in keeping with 
a pelagic habit. 

Probably the most common shark tooth in the Moore- 
ville is that of Lamna appendiculata. The Recent spe- 
cies of Lamna are confined to temperate and boreal 
regions (Bigelow and Schroeder, 1948). Off southern 
California this species is taken in deep water, but occurs 
near the surface in the northern Pacific, suggesting that 
in warmer waters it lives at greater depth. This genus 
today is considered to be a truly pelagic shark. 

Isurus is known today by two species (Garrick, 
1966). Both are open ocean sharks and reach lengths 
of about 12 feet (Bigelow and Schroeder, 1948; Apple- 
gate, 1966). The Cretaceous specimens from the Nio- 
brara Chalk had a length of at least 22 feet. The 
modern Isurus feeds on fish ; the large Cretaceous forms 
could have fed also on marine reptiles. 

None of the sharks occurring in the Mooreville be- 
long to groups that could not be pelagic. So far there 
have been found none of the small sharks that normally 
occur in shallow water such as Squalus and the orecto- 
lobids, which do occur in other Cretaceous sediments. 
Of note is the lack of rays of any sort, a group which 
is known to exist at a much earlier period, but again 
is more characteristic of shallow water except for the 
family Rajidae and several small-toothed forms which 
either had not evolved by Cretaceous times or must 
have left few remains. 



428 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



Sturgeons and Holosteans 

The sturgeon, Propenser, undoubtedly represents a 
marine form. Sea sturgeons today are wide-ranging 
and are taken in the open sea. They have been taken 
in Scandinavian waters at a depth of 25 fathoms, and 
there is no reason to believe they would not be found 
in deeper water. 

The pycnodont genus Hadrodus is known from both 
the Niobrara and the Mooreville, as well as from the 
Upper Cretaceous of Mississippi. This huge fish had 
crushing jaws and perhaps fed on invertebrates. If it 
was similar to the other pycnodonts it might have been 
deep-bodied and could have had a habitat similar to 
that of the present-day deep-bodied moonfish, Lampris, 
which is found in the open ocean. 

The pachycormid Protosphyraena, with its exceed- 
ingly long bill and partly fused pectoral fin, can be 
compared adaptively with the billfishes or istiophorids. 
The large fang-like teeth indicate a carnivorous diet. 
The long pectoral fin would perhaps stabilize the fish 
for the thrashing used in food-getting, as well as serving 
as a weapon of offense or defense. The caudal hypurals 
are coalesced into a single element, a feature occurring 
in the living istiophorids and associated with powerful 
swimming. Protosphyraena is known from the English 
chalk, New Jersey, Kansas, and California. The wide 
distribution and the modern analogs are at least con- 
sistent with the interpretation that this was an oceanic 
or pelagic species. 

The sturgeons and holosteans are to be considered 
as survivors of more archaic groups. Protosphyraena 
seems to be the most specialized. Only the sturgeon 
might be considered indicative of a shallow water en- 
vironment, yet this may be due to an incomplete knowl- 
edge of the living species, which probably at times go 
very deep or swim over deep areas. Recent tagging 
studies show that the modern sturgeon can move great 
distances. 

Teleosts 

The Elopidae are represented today by the single 
genus Elops, which has a worldwide tropic and south 
temperate distribution. Elops is common in the open 
sea along the coast of the southeastern United States. 
It also frequently enters brackish water, being taken 
from the mouths of rivers and from bays. Gill (1907) 
records the presence of shrimp in Elops stomachs, and 
Darnell (1958) reports a predominance of small fish 
and penaeid shrimps. The Recent species appears to 
be wide-ranging. This appears to be true also of the 
fossil Cretaceous genera assigned to this family. 

The related family Albulidae is represented today 
by the genera Albula and Dixonia. According to Hilde- 
brand (1963), the extent to which Albula enters deep 
offshore water is unknown. Bonefish do occur in the 
shallows, feeding in the mud. The geographic distri- 
bution of the modern species is similar to that of Elops; 
it is found in all warm seas. Frizell (1965) believes 
that the Eocene species was abundant in deep water. 



Pachyrhizodontidae are extinct, which forces one to 
resort to structural adaptations to elucidate their prob- 
able habits. The chalk species show a proportionally 
large sickle-shaped tail. The pectoral fins are large and 
powerful with a large stout first pectoral spine. These 
characters are adaptations of a powerful swimmer. The 
large incurved teeth indicate a carnivorous diet. The 
Scombridae would appear to be the modern ecological 
counterpart of this family. The Pachyrhizodontidae, 
like the similar Scombridae, must have been open sea 
pelagic fishes with nearly worldwide distribution. 

The Bananogmiidae are another extinct group. In 
fin structure and shape they are similar to the Pachy- 
rhizodontidae; however, the unique crushing dentition 
is closest to that of the Albulidae. In the European 
Cretaceous this family is absent and may have been 
ecologically replaced by the closely related and similar 
family, the Plethodidae. There is notable uncertainty 
as to the morphology and habits of these groups of 
fishes. The crushing dentition and good swimming abil- 
ity could be correlated with feeding on shrimp and 
other open sea arthropods as well as on nautiloids and 
ammonites. 

The fossil Ichthyodectidae are very close to the 
modern Chirocentridae. The latter are large powerful- 
swimming predaceous fishes that have a wide ecological 
range; they are found in both coastal and open ocean 
waters. The Ichthyodectidae were even larger fish than 
the Pachyrhizodontidae with Xiphactinus being per- 
haps the largest bony fish that ever lived. 

The Saurodontidae resemble the Ichthyodectidae in 
general structure but possess a predentary which may 
have served in a manner similar to the elongate rostrum 
of Protosphyraena. The thin blade-like teeth indicate 
a predaceous habit. 

One of the interesting aspects of Cretaceous chalk 
faunas is the almost complete absence of the Clupeidae. 
So far, only one preoperculum in the Mooreville can 
be referred to this family. Clupeids are also absent in 
the British chalk and the Niobrara. They have, how- 
ever, a good Cretaceous record and they dominate the 
Upper Cretaceous of California and Syria. Their ab- 
sence in the chalks is one of the most striking examples 
of the great differences in modern as well as fossil fish 
faunas in different ecological situations. 

The Enchodontidae are not only a common Creta- 
ceous family but are probably the second most common 
group of the Mooreville fishes. The predatory encho- 
dontids 1 seem to be closely related to the present day 
Omosudidae and Alepisauridae, which are deep water 
Iniomi. Enchodus is known from Cretaceous sediments 
of undoubted shallow water. Cimolichthys is a large 
powerful fish which could certainly be an oceanic form. 

The Dercetidae are represented in the Mooreville 
by Stratodus apicalis which is a fairly small predaceous 
fish; the great number of little sharp teeth on the pala- 
tines and jaws would appear to be very effective for 
holding its prey. This species is reported from phos- 

1 Specimens preserved with prey within their bodies are known. 



APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



429 



Depth Range in Fathoms (F) of Genera and Families of Mooreville Fishes 



GENUS 

Edaphodon 
Ptychodus 

Scapanorhynchus 

Lamna 

Isurus 

Squalicorax 

Pseudocorax 

Propenser 

Hadrodus 

Protosphyraena 

Palehps 
Paehyrhizodus 

Albula 
Bananogmius 

Moorerillia 

Ichthyodeetes 

Xiphactinus 

Saurodon 

Saurocephalus 

Enchodus 

Cimoliehthys 

Stratodus 

Myctophidae 

Trachichthyidae 

TOTAL— 



0-50 F 
SHALLOW 



50-100 F 
MODERATE 



14 



x 
x 

X 
X 

22 



100+ F 
DEEP 



? x x 

T x x 

XXX 
XXX 
X X ? 

? X X 

? X X 

X X 

X X 

X X 

X X 



X 
X 
X 
X 
X 

21 



MODERN ANALOGS DISCUSSED IN TEXT 

Large deep sea chimaeroids, several genera. 

None; modern heterodonts occur in shallow to deep 
water. 

None; modern Odontaspis comprises shallow, pelagic, 
and deep water species. 

Pelagic over moderate depths and in deep water. 

Pelagic over moderate depths and in deep water. 

None; probably pelagic. 

None; probably pelagic. 

Coastal ; could occur over deep water. 

None; probably pelagic over deep water. 

None; like modern swordfishes, mostly pelagic over 
moderate and deep water. 

Modern Elops mostly in shallow to moderate depths. 

None; like scombroids, in general over moderate to 
deep water. 

Modern Albula in shallow to moderate depths. 

None; like scombroids in general, over moderate to 
deep water. 

None; like scombroids in general, over moderate to 
deep water. 

None; modern Chirocenlrus shallow water and pelagic 
species. 

None; modern Chirocenlrus shallow water and pelagic 
species. 

None; modern Chirocenlrus shallow water and pelagic 
species. 

None; modern Chirocenlrus shallow water and pelagic 
species. 

None; modern related genera deep water. 

None; modern related genera deep water. 

None; however, could perhaps be in deep water. 

Modern genera in fairly deep water. 

Modern genera in deep water. 



phate beds in North Africa, which may indicate a shal- 
low water habit but again the wide distribution might 
also indicate a pelagic existence. 

The Myctophidae or lantern fishes represent the 
most common Mooreville fish group. Small patches 
of myctophid scales occur at most of the localities in 
vast quantities. The comparatively fragile vertebrae 
and head plates are also abundant. The modern mem- 
bers of this family are deep water plankton feeders. 
They move up at night and down during the day in 
deep oceanic waters, probably following the periodic 
movements of zooplankton; some species commonly 
come to the surface at night, where they are taken in 
great numbers. 

Goode and Bean (1895) place the family Trachich- 
thyidae among the deep water beryeoids, with Traeh- 
ichthys being taken from depths of from 200 to 400 
fathoms. Smith (1961) likewise places this family with 



the deep sea beryeoids, and says that it ranges from 50 
to 500 fathoms in depth. 

If one is willing to assume that at least most of the 
common genera of Mooreville fishes have been pre- 
served and are represented in the present collection, 
then we may make some inferences concerning the pos- 
sible ecological relationships among these forms. Rank- 
ing the families or genera in order of abundance of 
individuals we have as follows: abundant, the Mycto- 
phidae, Enchodus. Squalicorax, Scapanorhynchus, Lamna. 
Paehyrhizodus. Xiphactinus; common. Saurodon, Sauro- 
cephalus. Bananogmius; rare. Stratodus, Protosphyraena. 
Edaphodon. Palehps. Albula. Cimoliehthys. Isurus; very 
rare. Pseudocorax, Ichthyodectes. and Clupeidae. The 
last two are known by only one or two individuals. 

Feeding habits may be inferred from the stomach 
contents of the fossils (none among the Mooreville spec- 
mens), the food habits of their modern analogs, and the 



430 



FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 



shape, size, and number of teeth. The Mooreville gen- 
era with crushing teeth are Edaphodon, Ptychodus, Had- 
rodus, and Albula. Any or all of these could be con- 
sidered mollusk feeders, as is the Recent Albula. Those 
forms having pointed teeth suitable for biting and sev- 
ering are Scapanorhyrichus, Isurus, Lamna, Squalicorax, 
Pseudocorax, Protosphyraena, Pachyrhizodus, Xiphac- 
tinus, Ichthyodectes, Saurodon, Saurocephalus, Cimolich- 
thys, and Enchodus. Those fishes having numerous 
pointed teeth capable of holding the prey and assisting 
in the swallowing process are Moorevillia, Stratodus, 
and Hoplopteryx. The Myctophidae have small teeth 
suitable for feeding on plankton. The sturgeon, Pro- 
penser, lacked teeth altogether; Acipenser, its modem 
counterpart, is a detritus feeder. 

Ranking these fish by size, those with a general size 
range in the adults of from six to fifteen feet are Pro- 
penser, Xiphactinus, Pachyrhizodus caninus, Isurus, 
Scapanorhyrichus, Squalicorax, Ichthyodectes, and Lamna. 
Fish with a range of around three to six feet are Pachy- 
rhizodus kingi, at least some species of Bananogmius, 
Cimolichthys, and (at the lowermost limit of the range) 
Pachyrhizodus minimus. Fishes ranging from one to 
three feet are Stratodus and Enchodus. Less than one 
foot are Hoplopteryx, the Clupeidae, and the Mycto- 
phidae. 

In the Mooreville, as stated, the myctophids occur 
as patches of scales with vertebrae and disassociated 
plates. These patches may represent stomach ejects 
from other fish, particularly sharks. Myctophid scales 
are also very common in the Mooreville coprolites. The 
great abundance of this family makes it a likely can- 
didate to form the base of the Mooreville fish food 
chain. The Clupeidae and the berycoids play a minor 
role. All four groups may be considered as the basal 
grade, which may be called Grade 4; they fed mostly 
on invertebrates and could have been the prey of Grade 
3, including Cimolichthys, Enchodus, and Stratodus. 
Grade 2 consists of Ichthyodectes, Pachyrhizodus kingi, 
Squalicorax, Scapanorhynchus, Bananogmius, and Pachy- 
rhizodus minimus. The top of this pyramid, Grade 1, 
would be the largest carnivores, Pachyrhizodus caninus, 
Xiphactinus, and the sharks, Isurus and Lamna. 

Below this fish food chain there is a more fundamen- 
tal chain with the base consisting of phytoplankton. 
The phytoplankton is represented in the sediment by 
Chrysophyta, which might have been fed on by a di- 
verse zooplankton consisting in part of Foraminifera, 
ostracods, and small gastropods. There may have been 
a vast number of forms at this level in the food chain 
which either did not fossilize or are yet to be collected. 
This zooplankton was fed upon by the myctophids 
which in turn were fed upon by the other three grades of 
fishes plus other higher vertebrates such as turtles, mosa- 
saurs, and plesiosaurs. Aside from this chain there 
were some fish with crushing dentition which evidently 
fed directly on the molluscan fauna or the larger zoo- 
plankton. 



In conclusion, the Mooreville fish fauna is a mixed 
one consisting of undoubted pelagic open sea forms 
along with some possible near shore and perhaps shallow 
water fishes, but showing a dominance of deep water 
fishes. The probable currents maintained the open 
water surface feeders, and the close proximity to shore 
(within a few miles) or near shore environments con- 
tributed the near shore fishes in this assemblage. The 
intermixture of these two faunas with an emphasis on 
the open sea fishes seems characteristic not only of the 
Mooreville but of Cretaceous chalks in general. 

At the present time many Cretaceous chalk faunas 
are still poorly known but all indications point to the 
fact that these chalk faunas throughout the world have 
assemblages closer to each other than to nearby for- 
mations of equivalent age but different sediment types. 

In general, chalks show a fauna consisting of a large 
number of huge predaceous teleosts such as Xiphac- 
tinus, Ichthyodectes, and Pachyrhizodus. Sharks are pres- 
ent but only moderately abundant. The Clupeidae are 
rare or absent. Myctophidae are common. 

In number of similar species the Mooreville Forma- 
tion's fish fauna is closest to that of the Niobrara Chalk, 
which has been studied mainly by Williston (1900), 
Stewart (1900), Loomis (1900), and Hay (1903). In 
number of similar genera the Mooreville's fish fauna is 
closest to that of the British Chalk, the best known 
Upper Cretaceous fauna due to the long history of col- 
lecting culminating in the excellent monographic treat- 
ment of Woodward (1902-1912). 

With the exception of the Niobrara, Upper Creta- 
ceous chalks either intertongue with or are underlain 
by greensands. The common occurrence of Ostrea and 
Exogyra reefs in the greensands suggests shallower 
water. The greensand fish faunas show, to use the New 
Jersey fauna as an example, abundant sharks, rays, and 
Chimaeridae. Teleosts are present but not well known ; 
the large chalk species seem to be absent. Of the 41 
species of New Jersey fish, seven are similar to Moore- 
ville species and four of the seven are wide-ranging 
sharks. 

The littoral zone in the Upper Cretaceous is repre- 
sented by the North African phosphate beds and by a 
poorly known South Dakota fish fauna mentioned by 
Zangerl and Sloan (1960). These beds differ from the 
Mooreville in the great concentration of vertebrate ma- 
terial in the form of disassociated teeth, vertebrae, head 
plates, and, in South Dakota, of scales, to form a bone- 
bed conglomerate. The phosphate beds are known to 
cover extensive areas, and in contrast to the chalks have 
a good pre- and post-Cretaceous history. There is no 
similar Recent deposit, but the Pliocene and Pleistocene 
phosphate beds of Florida and the "bone beds" of the 
Miocene Calvert in Virginia and Temblor of California 
are the same general type of deposit and evidently were 
formed in very shallow water in or near the intertidal 
zone. The Upper Cretaceous North African phosphate 
beds contain five species which are found in the Moore- 






APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 



431 



ville, four sharks and one teleost, Stratodus apicalis. 
The North African phosphate fauna is predominantly 
a shark fauna, with sharks comprising 75 per cent of all 
vertebrate remains (Arambourg, 1952). The sharks in 
the Mooreville are well under 25 per cent. The pre- 
dominance of shark teeth is also true of other phosphate 
beds mentioned above. The North African phosphate 
beds lack ptychodontids, although other rays are abun- 
dant, as are sawfish; the Mooreville, however, contains 
only ptychodontids and no evidence of rays or sawfish. 

SUMMARY 

The Mooreville formation contains a rich fossil fish 
fauna consisting of at least 17 families, 28 genera, and 
42 species. Future field work will undoubtedly add to 
this assemblage. Our present knowledge of the ecology 



of chalks leaves much to be desired. The evidently 
oozy bottom has resulted in a restricted bottom fauna, 
a fact that complicates any interpretation as to depth. 
One is however faced with two possible models which 
might represent the condition found in the Mooreville: 
a shallow water ooze around 3 to 30 feet deep, or a 
deep water ooze over 100 fathoms. Certainly as far 
as the present evidence is considered, the Mooreville 
Chalk seems to fit the second model and was probably 
deposited in water over 100 fathoms or 600 feet deep. 
At the same time, the terrestrial remains which have 
been found in the chalk, a bird and several dinosaurs, 
suggest a nearby shore. This seems probable, as the 
Mooreville beds lie at the southern end of the Appala- 
chian belt which probably formed a headland with few 
if any large streams at the southern end during Creta- 
ceous times. 



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