LIBRARY OF THE
UNIVERSITY OF ILLINOIS
AT URBANA-CHAMPAIGN
%
550
v. !->>
GEOLOGY
The person charging this material is re-
sponsible for its return to the library from
which it was withdrawn on or before the
Latest Date stamped below.
Theft, mutilation, and underlining of books are reasons
for dlstiplinary attion and may result In dismissal from
the University.
To renew call Telephone Center, 333-8400
UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMF-AIGN
L161— O-1096
! )l5<
3
Q
i^-?
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*.
OQy
Lib,
Mf^
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; l4 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 12 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-
dontids1 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.
REFERENCES
Agassiz, L.
1833-1843. Recherches sur les poissons fossiles. 3 (Placoides),
390 + 34 pp. Atlas, 83 pis., 1833-1845. Neuchatel.
Applegate, S. P.
1966. A possible record-sized bonito shark, Isurus oxyrinchus
Rafinesque, from southern California. Calif. Fish Game, 52
(3), pp. 204-207.
Arambourg, C.
1952. Les vertebres fossiles des gisements de phosphates (Maroc
— Algerie — Tunisie). Notes et Mem. Serv. Geol. Maroc.
No. 92, 372 pp., 62 figs. 46 pis., 7 tables (appendix by Ber-
gounioux, pp. 375-396).
Berg, L.
1947. Classification of fishes both recent and fossil. (1940,
original Russian text, Moscow, pp. 87-345). Ann Arbor,
Michigan. Pp. 346-517.
Berry, E. W.
1919. Upper Cretaceous floras of the eastern Gulf region in
Tennessee, Mississippi, Alabama and Georgia. U. S. Geol.
Surv., Prof. Paper 112, pp. 1-177, figs. 1-10, pis. 1-33.
Bigelow, B. and Schroeder, W. C.
1948. Fishes of the western North Atlantic, Part 1, Mem.
Sears Foundation for Marine Research, No. 1, Chap. 3,
Sharks, pp. 59-576, figs. 6-106.
Casier, E.
1953. Origine des Ptychodontes. Mem. Inst. Sci. Nat. Belg.,
Bruxelles (2) 49, pp. 1-51, 2 pis.
COCKERELL, T. D. A.
1933. The antiquity of Albula. Copeia, 1933 (4), pp. 157-244.
Cope, E. D.
1872. On the families of fishes of the Cretaceous formation in
Kansas. Proc. Amer. Philos. Soc, 12, pp. 327-357.
1878. Descriptions of fishes from the Cretaceous and Tertiary
deposits west of the Mississippi River. Bull. U.S. Geol.
Geog. Surv. Terrs., 4, pp. 67-77.
Darnell, R.
1958. Food habits of fishes and larger invertebrates of Lake
Ponchartrain, Louisiana, an estuarine community. Inst. Ma-
rine Sci., 5, pp. 353-416, figs. 1-17.
David, L.
1946. Upper Cretaceous fish remains from the western border of
the San Joaquin Valley, California. Carnegie Inst. Wash.
Publ. 551, pp. 83-112, 11 figs., 3 pis.
DUNKLE, D. H.
1940. The cranial osteology of Notelops brama (Agassiz), an
elopid fish from the Cretaceous of Brazil. Lloydia, 3, pp.
157-190, 9 figs.
Fowler, H. W.
1911. A description of the fossil fish remains of the Cretaceous
Eocene and Miocene formations of New Jersev. Geol. Surv.
New Jersey, Bull. 4, vi + 182 pp., 108 figs.
Frizell, D. L.
1965. Otolith-based genera and lineages of fossil bonefishes
(Clupeiformes, Albulidae). Senckenbergiana, 46a, pp. 85-
110.
Garrick, J. A. F.
1966. Revision of sharks of genus Isurus with description of a
new species (Galeoidea, Lamnidae). Proc. U. S. Nat. Mus.,
118, pp. 663-690, 9 figs., 2 tables, 4 pis.
Gill, T. N.
1907. The Tarpon and lady-fish and their relatives. Smith-
sonian Misc. Coll., 48, Part 3, pp. 31-46, pis. 17-21.
Gluckman, L.S.
1964. Paleogene sharks and their stratigraphic significance.
Acad. Sci. USSR, Div. Earth Sciences, Monographic Col-
lections, Moscow-Leningrad. Pp. 1-230.
Goode, G. B., and Bean, T. H.
1895. Oceanic ichthyology. Smithsonian Inst., U.S. Nat. Mus.,
Spec. Bull., xxxv + 553 pp., plus Atlas with 417 figs.
Graham, D. H.
1939. Food of the fishes of Otago Harbor and adjacent seas.
Trans. Proc. Roy. Soc. New Zealand, 68, pp. 421-436.
1956. A treasury of New Zealand fishes, 2nd ed. Wellington,
New Zealand., 424 pp.
Greenwood, P. H., Rosen, D. E., et al
1966. Phyletic studies of teleostean fishes, with a provisional
classification of living forms. Bull. Amer. Mus. Nat. Hist.,
131, Article 4, pp. 341-455.
Gregory, J. T.
1950. A large pycnodont from the Niobrara Chalk. Postilla,
No. 5, pp. 1-10, 2 figs.
Hay, O. P.
1903. On certain genera and species of North American Creta-
ceous actinopterous fishes. Bull. Amer. Mus. Nat. Hist.,
19, 1-95, pis. 1-5, 72 text figs.
1929. Second bibliography and catalogue of the fossil Verte-
brata of North America. Carnegie Inst. Washington, Publ.
39, 1, vii + 916 pp.
HlLDEBRAND, S. F.
1963. Family Albulidae. In Fishes of the western North At-
lantic. Sears Foundation for Marine Research, Yale Uni-
versity, Part 3, pp. 132-147.
Holmgren, N. and Stensio, E.
1936. Kranium und Visceralskelett der Akranier, Cyclostomen
und Fische. In Bolk, Goppert, Kallius, Lubosch. Handbuch
der vergleich. Anat. Wirbeltiere, 4, pp. 247-296.
Hussakof, L.
1912. The Cretaceous chimaeroids of North America. Bull.
Amer. Mus. Nat. Hist., 31, pp. 195-227, 21 figs., pis. 14-20.
International Commission on Zoological Nomenclature
1965. Bull. Zool. Nomencl., No. 723, pp. 32-36
Jordan, D. S.
1925. A collection of fossil fishes in the University of Kansas
from the Niobrara Formation of the Cretaceous. Kansas
Univ., Sci. Bull., 15, No. 2, Dec, pp. 219-234, pis. XIII to
XXIII.
Keroher, G. C, and others
1966. Lexicon of geologic names of the United States for 1936-
1960. U. S. Geol. Surv. Bull. 1200, Part 2, G-O: pp. 2581-
2582.
Leidy, J.
1857. Notices of some remains of extinct fishes. Proc. Acad.
Nat. Sci. Phila., 1857, pp. 167-168.
1868. Notice of American species of Ptychodus. Proc. Acad.
Nat. Sci., Phila., 1868, pp. 205-208.
1870. [Remarks on ichthyodorulites from Kansas and Ten-
nessee and on mammalian remains from Illinois.] Proc. Acad.
Nat. Sci. Phila., 1870, pp. 12-13.
432
APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION
433
1873. Contributions to the extinct vertebrate fauna of the
Western Territories. Rept. U. S. Geol. Surv. Terr., 1,
Part 1, pp. 14-358, pis. 1-37.
Loeblich, A. R. Jr. and Tappan, H.
1957. Correlation of the Gulf and Atlantic Coastal Plain Pale-
ocene and Lower Eocene formations by means of planktonic
Foraminifera. Jour. Paleontol., 31, pp. 1109-1137, 5 text figs.
Loomis, F.
1900. Die Anatomie und die Verwandtschaft der Ganoid —
und Knochen-Fische aus der Kreide Formation von Kansas.
Palaeontographica, 46, Lief 5-6, pp. 213-283, pis. 19-27,
13 text figs.
Monroe, W.
1941. Notes on deposits of Selma and Ripley age in Alabama.
Geol. Surv. Ala. Bull., 48, 150 pp., 15 figs., 2 maps.
Patterson, C.
1965. The phylogeny of the chimaeroids. Phil. Trans. Royal
Soc. London, Series B, Biol. Sci., 249, No. 757, pp. 101-219.
Quaas, A.
1902. Beitrag zur Kenntniss der Fauna der obersten Kreide-
bildungen in der libyschen Wiiste (overwegischichten und
Bliitterthone). Palaeontographica, 30,(2) pp. 153-336, pis.
20-33.
ROEMER, F.
1852. Die Kreidebildungen von Texas und ihre organischen
Einschlusse. Mit einem die Beschreibung von Versteiner-
ungen aus palaozoischen und tertiaren Schichten enthaltenden
Anhange und mit 11 von C. Hohe nach der Natur auf Stein
gezeichneten Tafeln. Bonn. Pp. i-vii, 1-100.
Romer, A. S.
1945. Vertebrate Paleontology, 2nd ed., Univ. of Chicago
Press, vii +687 pp., 377 figs.
1966. Vertebrate Paleontology, 3rd ed. Univ. of Chicago Press.
viii+468 pp., 443 figs., 4 tables.
SlGNEUX, J.
1949. Notes paleoichthyologiques. Bull. Mus. Nat. Hist.,
Paris, 21, pp. 633-638, 3 figs.
Smith, J. L. B.
1961. The Sea Fishes of Southern Africa, 4th ed. Cape Town.
xvi+580 pp., 1232+22+38 figs., Ill pis.
Stephenson, L. and Monroe, W.
1940. The Upper Cretaceous deposits. Mississippi State Geol.
Surv., Bull. 40, 296 pp., 48 figs., 15 pis.
Stewart, A.
1900. Teleosts of the Upper Cretaceous. Univ. Geol. Surv.
Kansas, 6, pp. 257-403, 6 text figs., pis. 33-73.
White, E. G.
1937. Interrelationships of the elasmobranchs with a key to
the order Galea. Bull. Amer. Mus. Nat. Hist., 74, pp. 25-
138, 66 figs., 51 pis.
White, E. I.
1956. The Eocene fishes of Alabama. Bull. Amer. Paleontol.,
36, no. 156, pp. 123-152, figs. 1-86, pi. 11.
Williston, S. W.
1900. Cretaceous fishes. Selachians and Pycnodonts. Univ.
Geol. Surv. Kansas, 6, pp. 237-256, pis. 24-32.
Woodward, A. S.
1889. Catalogue of the fossil fishes in the British Museum.
Part I. Containing the Elasmobranchii. Brit. Mus. Nat.
Hist., pp. i-xlvii +1-474, pis. 1-17, 13 woodcuts.
1891. Catalogue of the fossil fishes in the British Museum.
Part II. Containing the Elasmobranchii (Acanthodii), Holo-
cephali, Ichthyodorulites, Ostracodermi, Dipnoi, and Tele-
ostomi (Crossopterygii), and chondrostean Actinopterygii.
Brit. Mus. Nat. Hist., pp. i-xliv + 1-567, pis. 1-16, 57 text
figs.
1901. Catalogue of the fossil fishes in the British Museum.
Part IV. Containing the actinopterygian Teleostomi of the
suborders Isospondyli (in part), Ostariophysi, Apodes, Per-
cesoces, Hemibranchii, Acanthopterygii, and Anacanthini.
Brit. Mus. Nat. Hist., pp. i-xxxviii+ 1-636, pis. 1-19, 22
text figs.
1902-1912. The fishes of the English chalk. Palaeontogr.
Soc. London, pp. 1-264, pis. 1-54, text figs. 1-79.
Zangerl, R.
1948. The vertebrate fauna of the Selma formation of Alabama.
I. Introduction. II. The pleurodiran turtles. Fieldiana: Geol.
Mem., 3, 56 pp., 16 figs., 4 pis.
Zangerl, R. and Sloan R. E.
1960. A new specimen of Desmatochelys lorn Williston; a prim-
itive cheloniid sea turtle from the Cretaceous of South Da-
kota. Fieldiana: Geol., 14, pp. 7-40, 2 pis.