The Vertebrate Fauna of the Selma Formation of Alabama

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THE VERTEBRATE FAUNA OF THE SELMA FORMATION OF ALABAMA

PART VII THE MOSASAURS DALE A. RUSSELL

PART VIII THE FISHES

SHELTON P. APPLEGATE

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- lateral pendiculata, three teeth and one posterior tooth ; Isurus mantelli, two anterior teeth and one tooth lateral lateral ; Squalicorax falactus, two 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 stone, and the Mooreville Chalk. 1 Since the greatest majority of fossil fishes have come from the lowest In many respects our knowledge of American Cre- member, the conclusions in this paper involve taceous fossil fishes is still in its infancy. With the ex- mainly this unit. ception of David's 1946 paper, no work of a faunal The are in nature has been done on Upper Cretaceous fishes since specimens the collection of Field Mu- seum, unless otherwise Hay (1903) and Stewart (1900). Yet unstudied col- specified. lections exist in many of our museums and future work is bound to if not of modify, completely alter, some SYSTEMATIC DESCRIPTIONS our present concepts. Class It was first suggested to me by Dr. David H. Dunkle HOLOCEPHALI that the fossil fishes of the Mooreville Chalk of the Order CHIMAERIFORMES Selma Formation might form a subject for investiga- Suborder CHIMAEROIDEI tion. Field Museum already had a collection of Moore- Edaphodontidae Owen ville fishes collected by Mr. Barber, Dr. William D. Turnbull, Mr. R. H. Hard, and Dr. and Mrs. R. Zan- Diagnosis.—Chimaeroidei possessing paired vomer- gerl. This fine collection has formed the nucleus of the ine, palatine, and mandibular dental elements. The present study. beak-like mandibular element has from one to five oval tritoral areas Through a grant by the Bock Fund of the National per mandible, except for Elasmodus, which has two tritors and numerous smaller Academy of Science and with the aid of the University large tritoral areas. The element has from two to of Chicago it was possible for me to visit the Moore- palatine four tritors with the of ville localities where I collected additional material, exception Ganodus, which has one tritor and numerous small ones. The vo- and made observations concerning the ecology of the large known merine elements from six to Chalk. Mr. Bruce Crieley of Chicago accompanied possess eight tritors each. me as a field assistant. The Alabama Geological Sur- Discussion.—The greater proportional width and vey, through Dr. R. Jones and Miss Winnie McGlam- depth of the posterior portions of the mandibular den- mery, has not only given advice but has lent specimens. tal elements separate the Edaphodontidae readily from My wife, Anne Chase Applegate, has typed and as- the Lower Jurassic Squalorajidae, which have long, thin, sisted in the editing of the first draft. Dr. Rainer and naiTow mandibular dental elements. There is no Zangerl and Dr. Robert H. Denison have given a great likelihood of confusing the Edaphodontidae with the deal of their time editing and discussing this paper. known Myriacanthidae, which have mandibular dental elements that are shortened in a The criticism of Dr. Everett C. Olson, Dr. Ralph posterior direction G. Johnson, and Dr. Lore R. David has been valuable. but are greatly expanded laterally. Patterson (1965) that in the tritors are Dr. Olson helped in securing the grant for field work. says Myriacanthidae present on the two of Dr. Lore David has confirmed many of the scale identi- only upper anterior pairs dental elements, fications and has let me have the use of her excellent the vomers. The Callorhynchidae, of the Recent fami- collection of Recent teleost scales as well as her scale lies, though lacking recognizable vomerine tritors, lies closest to the fossil in one notebooks which include photos of both Recent and Edaphodontidae having only fossil teleosts. David Techter of Field Museum has or two tritors per mandibular element. The variable Chimaeridae have and mandibles checked the Museum numbers and has been a great short, wide, sharp suitable for the tritors are smaller and help in numerous other ways. Miss Anita Daugherty chopping food; are of a number than in the has been most helpful in re-editing this paper. Mrs. greater Edaphodontidae. The both Recent and have Myrna L. Patrick has typed the final copy. It was Rhinochimaeridae, fossil, the number of small tritors of of the exist- first submitted in 1961 as partial fulfillment of a doc- greatest any families and are the farthest removed from toral degree in Paleozoology at the University of Chi- ing perhaps cago. 1 According to Keroher and others (1966), the Mooreville It should be added that the Selma Formation in Chalk has been raised to formational rank and the old Selma Formation is now considered to be a The Areola Lime- Alabama is divisible into three mappable units, from group. stone is included in the Mooreville. However, in keeping with the top to bottom the Demopolis Chalk, the Areola Lime- 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 being known to enter bays (Graham, 1956). Callor- of the Edaphodontidae form a natural group differing hynchus is recorded from the Cretaceous of New Zea- markedly from the Recent and fossil Chimaeridae, with land. The trend from the Edaphodontidae to the mod- which they have been placed by most recent workers. ern families shows an over-all reduction in the size of The genera included are: Edaphodon, Isotaenia, Lepto- the teeth, probably coinciding with a general reduction mylus, Ganodus, Ischyodus, and Elasmodns, all of which in body size. The crushing surfaces on the jaws are have been discussed by Woodward (1891). The genus replaced by narrow chopping edges accompanied in Psaliodus probably also belongs in this family since some cases by the fragmentation of the tritoral areas. Woodward believes that it is closely related to Elas- modns. Edaphodon barberi, new species. Figure 174 The as far as two Edaphodontidae possess, known, Diagnosis.—The lateral surface of the mandible is of dental elements: two elon- pairs upper posteriorly, flattened. The apical tritor is the longest and is elon- flattened and, in advance of gate palatine plates, these, gate-oval in shape. The antero-intermediate tritor is two small vomerine elements, one of which was figured well in advance of and somewhat smaller than the pos- for Hussakof The Edaphodon by (1912, p. 206, fig. 4). tero-intermediate tritor. Above and behind the postero- lower mandibular dental elements are and are paired intermediate tritor lies the postero-dorsal tritor. which and the over-all is deep, wide, generally robust; shape is about equal to the antero-intermediate tritor in area reminiscent of a hawk-like beak. All the dental ele- and shape. ments possess one or more tritoral areas, composed of Type.—PF 290, a nearly complete left mandible tubular dentine (Patterson, 1965), which are raised from Donald's Farm, Dallas County, Alabama, middle above the surrounding parts of the plate in worn teeth. Mooreville Chalk. These areas are associated with a crushing or grinding habit. Description and Discussion.—The dorsal border of the mandible in outline bears an tritor Leptomylus has only one tritor per mandibular ele- anteriorly apical for about one-third its length; in the middle one-third ment; Edaphodon, four or five tritors; Ganodus and it rises gently; in the last one-third the dorsal border Ischyodus, four tritors; and Elasmodns, at least two rises more steeply and then flattens out. The ventral large tritors and numerous small ones. The mandibular border sweeps backward in a smooth arc. The postero- dental element is not known in Isotaenia. The palatine intermediate tritor is under the dorsal element in Ischyodus has four tritoral areas; in Edapho- just posterior border. The relationship of these tritors is shown in don and Elasmodns, three; in Isotaenia, two. In Ganodus Figure 174. A transverse section of the mandible would there is a palatine element reminiscent of the vomerine be approximately triangular. The symphyseal facet of Edaphodon, with one large tritor which is very elon- is indistinct. gate and a row of small tritors that form a U-shape. The palatines of Leptomylus are unknown. The vomerine MEASUREMENTS— mm.). plates of Edaphodon are narrow, with from six to eight (in small tritors. The vomerine of are plates Ischyodus Greatest length of mandible 69 rectangular, with six tritors in two rows. The vomerine Greatest thickness of mandible 19 teeth have yet to be found in the other genera of the Length of apical tritor 21 Edaphodontidae. Width of apical tritor 4.5 Width of antero-intermediate tritor 3.5 The Edaphodontidae as defined above range from Width of postero-intermediate tritor 5.5 the Jurassic to the Pliocene, probably reaching their Width of postero-dorsal tritor 3.0 greatest expansion in number and variety in the Creta- ceous, particularly in the Upper Cretaceous. Judging barberi shows from the dentition, they form a homogeneous ancestral Edaphodon superficial similarities to complex from which the modern chimaeroid families Edaphodon stenobryus in shape, but the position of the tritors in E. is were derived. The Chimaeridae in the restricted sense, stenobryus completely different. The of the tritor the with the separation of the Edaphodontidae, consist of great length apical plus fact that there are Chimaera and Hydrolagus. The former has a Creta- separate intermediate tritors places E. barberi outside the variants ceous to Recent record, the latter is known only from of Edaphodon mirificus as described the Recent. However, no one has compared the denti- by Fowler (1911) and Hussakof (1912). tion of these two genera. The Chimaeridae may be The back of the type mandible is incomplete, and devived from the Edaphodontidae through Elasmodns. there has been some distortion due to compression along The Rhinochimaeridae, consisting of the Recent Rhino- with some fracturing. The waterwom appearance is chimaera, Harriotta, and Keoharriota, and the fossil Amy- most likely due to recent exposure, born out by the lodon and Elasmodectes, appear to branch from the eda- presence of encrusting lichens on three sides of the phodontid stock. mandible.

The Recent Southern Hemisphere Callorhynchidae The species is named for Mr. CM. Barber, who appear to lie, as stated, closest to Edaphodon. They was one of the earliest collectors of the Mooreville ver- are of much smaller size and inhabit shallow water, tebrate fauna. 10 mm

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

391 392 FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3

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 Discussion.—Fowler (1911) reports this species from Referred specimens.—P27536, two mandibles from the Greensand at Barnsboro and Hornerstown, New Bank's farm, Greene County, Alabama, lower Moore- Jersey, and adds that the Hornerstown Formation is ville or upper Eutaw. P27529, one left palatine; probably Cretaceous. Loeblich and Tappan (1957) P27537, one right mandible; PF 209, one left palatine; place the Hornerstown in the Paleocene. If they are all from Moore's farm, Dallas County, Alabama, middle correct and the Edaphodon material is not reworked, Mooreville Chalk. then this species bridges the Cretaceous-Tertiary boun- APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 393 dary. The genus Edaphodon undoubtedly does bridge (1941). Leidy (1868) lists three Alabama specimens, this gap, but it appears that New Jersey marine ver- one from Unionville, one from Perry County, and one tebrate Cretaceous or Tertiary records, particularly from Greene County. With all these records it would from the Greensand, are subject to question because of not be surprising to find this shark in the Mooreville the possibility of reworking of the material into Ter- Chalk. Outside of Alabama, P. mortoni is fairly com- tiary beds and the apparent lack of precise stratigraphic mon in the Niobrara Chalk of Kansas, and has been information. The Hornerstown Formation of New Jer- reported from the Eutaw Formation of Mississippi by sey and the Aquia Formation of Virginia are both Stephenson and Monroe (1940). The wide geographi- greensands and are supposed to be Paleocene, but the cal range of this species is shown by its occurrence in Hornerstown fish fauna has a strong Upper Cretaceous the British Chalk, the Upper Cretaceous of Italy, Mexi- aspect while the Aquia shows no such affinities. 1 co, Mississippi, Alabama, and the Niobrara of Kansas.

Edaphodon sp. Figure 175B Order GALEIFORMES —PF a mandible, from Referred specimen. 3501, right Anacoracidae Gliickman 2 miles W. of West Greene, Greene County, Alabama, — middle Mooreville Chalk. Discussion. The two genera Squalicorax and Pseu- docorax have been referred tentatively to the Ceto- Discussion.—This large mandibular dental element rhinidae by Woodward (1902-1912) and E. I. White has a claw-like appearance and is probably distinct, (1937). Gliickman (1964) has referred Anacorax, which although it may be an extreme variant of Edaphodon is a synonym of Squalicorax, to a separate family, Ana- mirificus. There are two tritoral areas present, a long coracidae. Undescribed material at the Los Angeles anterior apical area and a very small dorsal tritoral County Museum suggests that his family reference is area. The general shape, small size, and placement of correct. Therefore, Squalicorax, and with it the closely the tritoral areas separate Edaphodon sp. from Eda- related Pseudocorax, should form the basis of a new phodon barberi. The specimen is extremely waterworn family. In spite of the fact that Squalicorax has pri- and much of the back end is missing. ority over Anacorax, the family name Anacoracidae must be retained to Article 40 of the Inter- Class SELACHII according national Code of Zoological Nomenclature. Best indi- Order HETERODONTIFORMES cations are that these sharks are primitive orectolobid Suborder HYBODONTOIDEI derivatives; therefore, I have placed them before the which are to be advanced orec- Ptychodontidae Hay Odontaspididae, thought tolobid derivatives. Discussion.—Casier (1953) has shown that the Pty- chodontidae are specialized derivatives of the hybodont Squalicorax falcatus (Agassiz). Figures 176 A-F; sharks and not at all. Patterson (1965) suggests rays 178 L and N; 179B that Hylaeobatis arose from a homodont species of the Referred specimens.—PF 3524, four teeth; Ostrea hybodontid genus Lonchidion, and that Hylaeobatis, a Choctaw Bluff, Greene County, Alabama, lower- member of the family Ptychodontidae, gave rise to the layer, most Mooreville or uppermost Eutaw. PF 3523, four genus Ptychodus. teeth; Bank's Bluff, Greene County, Alabama, lowermost Mooreville or uppermost Eutaw. PF 124, 29 Ptychodus polygyrus Agassiz. Figure 179A vertebrae; PF 3538, four vertebrae; one mile S. of West Referred specimen.—PF 127, one tooth; Bank's Bluff, Greene, Greene County, Alabama, middle Mooreville. Greene County, Alabama, lowermost Mooreville or up- PF 126, 48 vertebrae and dermal denticles; one mile permost Eutaw. NW. of West Greene, Greene County, Alabama, middle Discussion.— this from Cope (1878) reported species Mooreville. PF 3537, three vertebrae; 2 miles N., one the old name for the Selma the Rotten Limestone, Group mile W. of West Greene, Greene County, Alabama, in Alabama. Niobrara tooth Williston The figured by middle Mooreville. PF 3534, two teeth; P 27496, four and to this to (1900) assigned species actually belongs vertebrae; PF 3533, four vertebrae; P 27448, four ver- decurrens. P. the Mooreville Ptychodus polygyrus, spe- tebrae; P 27409, 34 vertebrae; P 27446, five vertebrae; is also from the Cretaceous of cies, known Upper Eng- PF 3528, four vertebrae; PF 3529, six vertebrae; P and Russia. land, Belgium, 27475, 13 vertebrae and one slide with cross section; PF 3530, one vertebra and one tooth; all from Moore's mortoni Mantell. Ptychodus farm, Dallas County, Alabama, middle Mooreville. PF Discussion.—This species has not been found in the 3536, seven vertebrae, Harrell's Station, Dallas County, Mooreville member of the Selma Chalk during the pres- Alabama, middle Mooreville. PF 3527, one tooth; near ent collecting. There is, however, a specimen in the road, Hewlett's farm, Greene County, Alabama, upper Walker Museum collection at Field Museum, U.C. Mooreville. PF 3525, one tooth, 8.9 miles E. of Russell 14368, from Prairie Bluff, Alabama. The Prairie Bluff which in turn lies above the 1 Chalk Ripley Formation, Based on an undescribed Aquia fish fauna collected by me lies above the Demopolis Chalk, according to Monroe and now in Field Museum. 394 FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3

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- section shown in Figure 179 B resembles those of Ceto- town Formation. rhinus in showing concentric rings of calcification plus Discussion.—Squalicorax falcatus is now known in several radial supports. This sort of structure is found the Mooreville by its teeth, vertebrae, and denticles. also in Squatina. While some sets of vertebrae do not The limits of tooth variation are not known because have associated teeth, there is little doubt that all be- of the lack of associated sets. This is one of the most long to the same species. In some specimens, partic- common Upper Cretaceous sharks. The vertebral cross- ularly P 27409, denticles were noted adhering closely APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 395

to the vertebrae; these evidently settled onto the ver- cality 1, 6.2 miles W. of Aliceville, Eutaw County, tebrae as the fish decayed. The denticles of another Alabama, lower Mooreville. specimen, PF 126, display notable variation (fig. 176 A- Discussion.—The more slender and elevated crown, F). Figure 176 A represents the palmate type of den- and the prominent notch between the crown and the five or six ticle, having backwardly projecting "fingers" ; blade, in combination with the absence or weakness of 176 B and C the other a Figure represents extreme, 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. I m m 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 Fig. 177. Tooth of Pseudocorax affinis, PF 3522. showing body shape, teeth, denticles, and vertebrae. smooth bulb-like denticle with only one posterior pro- Elsewhere this genus is known only from teeth. It jection. Figure 176 D, E, and F shows the most com- should be noted that the old genus Carcharias is no mon type of denticle, with three keels on the crown. longer considered valid according to the ruling of the S. falcatus is reported from southern India, Madagas- International Commission on Zoological Nomenclature, car, northern Europe, England, New Jersey, Kansas, no. 723, 1965; the name is superseded by Odontaspis. and California. Scapanorhynchus rhaphiodon (Agassiz). Figure 178 Squalicorax pristodontus (Agassiz) A-C. — Referred specimen.—PF 660, one tooth; roadbed be- Referred specimens. PF 3504, three teeth; Choctaw tween Thomaston and Safford, Dallas County, Ala- Bluff, Greene County, Alabama, lower Mooreville or bama, Demopolis Chalk. Eutaw. PF 3505, three teeth; Bank's Bluff, Greene Alabama, lower Mooreville. PF 3502, one Discussion.—Although this species is not known to County, Hale's farm, 2 miles N. of West Greene Post occur in the Mooreville proper, the fact that it does oc- tooth; Greene Alabama, middle Mooreville. cur in the Selma Group gives reason for its inclusion in Office, County, PF 3506, 14 teeth; PF 3509, one tooth; Moore's farm, the present paper. The large size and the broad low Dallas Alabama, middle Mooreville. PF 3511, crown of the teeth serve to distinguish them from those County, two teeth; Harrell's Station, Dallas County, Alabama, of S. falcatus. It is unknown in the Niobrara Chalk.

middle Mooreville. PF 3503, two teeth ; Hewlett's farm At present nothing is known of its denticles or verte- near the road, Greene County, Alabama, upper Moore- brae. S. pristodontus is reported from the Upper Cre- ville. PF 3508, two teeth, 5.1 miles E. of Russell taceous of India, western Africa, northern Europe, the Route Russell Bluff- northern and southern borders of the Mediterranean County Line, 26, County, Alabama, town Formation. PF 3507, two teeth; PF 3519, one Basin, England, and New Jersey. tooth; both from 8.9 miles E. of Russell County Line, Pseudocorax affinis (Agassiz). Figures 177, 178 M Route 26, Russell County, Alabama, Blufftown For- Referred specimens.—PF 3522, two teeth; Locality mation. 12A, consolidated layer, Moore's farm, Dallas County, Discussion.—The Mooreville specimens agree with Alabama, middle Mooreville. PF 3558, one tooth; Lo- those teeth figured by Agassiz (1843) under the specific 396 FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3 name rhaphiodon. Woodward (1889) erected the genus the length of the crown, differentiate the teeth of this Scapanorhynchus and placed in it Agassiz' species, Lam- species from those of S. rhaphiodon and agree with na rhaphiodon. Williston (1900) put the Niobrara teeth teeth figured by Quaas (1902) and Arambourg (1952). of Scapanorhynchus also in this species, with Roemer's This form may prove to be more characteristic of the (1852) Lamna texanus a synonym. Arambourg (1952) Eutaw than of the Mooreville but it does occur at the has considered Lamna texanus a synonym of the very Mooreville-Eutaw contact. 5. rapax has previously been different S. rapax. reported only from Africa, where it occurs in the Seno-

~M 81 M N

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 nian of Libya and the Maestrichtian phosphate beds Cretaceous of New Zealand, Japan, Australia, the East of northern Africa. Indies (Timor), India, Africa, both sides of the Medi- Lamnidae Muller and Henle terranean Basin, Northern Europe, England, the Cari- Lamna appendiculata Agassiz. Figures 178 E-H, ibbean Islands, and in North America from the New 179 C. Jersey Greensands, the Niobrara of Kansas, and the Referred specimens. —PF 3517, one tooth; Ostrea Austin of Texas. Chalk layer, Choctaw Bluff, Greene County, Alabama, lowermost Mooreville or uppermost Eutaw. PF 3520, two Scapanorhynchus rapax (Quaas). Figure 178 D. — teeth; one mile S. of West Greene, Greene County, Referred specimens. PF 3510, two teeth; Ostrea Alabama, middle Mooreville. PF 3535, two teeth; PF layer, Choctaw Bluff, Greene County, Alabama, lower- 3532, one tooth; PF 3514, 12 teeth; PF 3515, 50 verte- most Mooreville or uppermost Eutaw. brae and 13 teeth from one individual; PF 3521, one Discussion.—The large size, wide bases, and poorly vertebra; P 27499, 12 vertebrae; P 27500, two verte- developed striae, which extend for less than one half brae, one tooth, one vertebral thin section; all from APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 397

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- L. appendiculata is known from the Cretaceous of Ja- ville. PF 3516, 15 teeth; Harrell's Station, Dallas pan, Australia, New Zealand, the East Indies (Timor), County, Alabama, middle Mooreville. PF 3518, one Madagascar, both sides of the Mediterranean Basin, vertebra; north set of gullies, Hewlett's farm, Green western Africa, British Columbia, and in the United County, Alabama, upper Mooreville. PF 3513, five States in Alabama, Kansas, and New Jersey. teeth; near the road, Hewlett's farm, Greene County, Isurus mantelli 178 I-K. Alabama, upper Mooreville. (Agassiz).— Figure Discussion.—Lamna appendiculata is widespread in Referred specimens. PF 3512, eight teeth; Ostrea the Mooreville. A direct association of teeth and ver- layer, Choctaw Bluff, Greene County, Alabama, lower- tebrae is known in PF 3515, which includes the re- most Mooreville or uppermost Eutaw. mains of just one individual. No dermal denticles were Discussion.—The classic distinction between Isurus found in association with these remains. A vertebra, and Lamna has been and still is the absence of lateral as seen in cross-section (fig. 179 C), compares favorably denticles in the former. Isurus is also characterized with the living Isurus oxyrhinchus. 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 specimen 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 specimen 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) reported 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 Fig. 180. Reconstruction of skull of Propenser hewletti, dorsal Order ACIPENSERIFORMES view. DS, dermosphenotic; ES, extrascapular; FR, frontal; PA, parietal; PO, postorbital; PT, pterotic; SO, supraoccipital. Acipenseridae Bonaparte in size. The parietals are reduced and separated by a The Acipenseriformes have a fossil record going as very large supraoccipital. The endocranium is ossified, far back as the Jurassic, with questionably related forms with a pronounced fusion of its elements, as is the to the Pennsylvanian (Romer, 1966). The family Aci- shoulder girdle. The neural arches are fused and each penseridae differs from the related family Chondros- arch bears two neural spines. Other characteristics teidae in having a dorsal series of unpaired splinter- are as in the genus. like plates on the upper rostral surface, and lateral scutes on the body. The sturgeons differ from the Propenser, new genus —As for Polyodontidae (paddlefishes) in lacking teeth in the Diagnosis. — subfamily. adult as well as in the absence of vacuities in the skull Type species. Propenser hewletti, new species (fig. so characteristic of the latter. 180). Description.—The external surfaces of the plates Propenserinae, new subfamily — are ornamented with equally raised hemispherical tu- Diagnosis. The frontals are expanded laterally and bercles approximately 5 mm. in diameter. These may border the orbit. The rostrals are numerous and small 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 girdle is massive (fig. 184, C and D) with both dermal plates. The triangular frontals (fig. 180, FR) extend for- and endoskeletal parts ossified and fused. The first ward and are rounded on the anterior border; laterally, pectoral fin spine (fig. 184, B) is large, and ornamented they form the upper border of the orbit. Anterior to dorsally with rows of tubercles which merge into small the frontals are 20 or more rostrals which vary in size ridges. The second spine (fig. 184, A) is of similar size and shape. (The rostrals were not found in place.) The but poorly ornamented. The body scutes (fig. 182, B), most common rostral shape is elongate lanceolate (fig. which are thought to be lateral, are elongate rectangular 182 A). The parietals (fig. 180, PA) are small and are and slightly curved to fit the curvature of the body. separated from each other by a huge supraoccipital (fig. The dorsal scutes (fig. 184, E), of which four are present, 180, SO), which is deflected downward on each side; have the typical tubercles but still are quite sturgeon- it is arrow-shaped, with the point toward the rear of like. the skull. Lateral to the parietals lie large pterotics

(fig. 180, PT) . Posterior to the pterotics are the extra- Propenser hewletti, new species. Figures 180-185. of a (fig. 180, ES) , each which bears postero- scapulars Type.—Specimen in Alabama Geological Survey lateral forked process which is believed to serve for the collection, including much of a skull, scutes, shoulder attachment of the post-temporals. The dermosphe- girdle, fin spines, and vertebrae; gullies near road, Hew- notic lies to the (fig. 180, DS) just posterior postorbital lett's farm, Greene County, Alabama, upper Moore- The is ossi- (fig. 180, PO). hyomandibular (fig. 183, A) ville. fied, as are the other known hyoid elements (fig. 183, B Referred specimens.—PF 288, one plate (probably through G). The lower jaw (fig. 185, C) is long, thin, from the Hewlett's type specimen) ; farm, Greene County, quite delicate, and toothless. The endocranium is be- Alabama. A second specimen in Alabama Geological lieved to have been partly ossified, and a number of Survey collection, part of a skull; near West Greene, parts of it are preserved. The palatoquadrate (fig. 185, Greene County, Alabama, middle Mooreville. A and B) is well developed. The articular end of the quadrate is produced into a definite peg. The shoulder Diagnosis.—Same as for genus.

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 day analogue in the vertebrae of very large sailfishes, favorably, except for their ornamentation, with those which have their abdominal vertebrae coalesced; the of other fossil and Recent Acipenseridae. The maxil- exact function of this is unknown, but it may serve lary in the type is fused to the palate, which in turn is as a spring and an aid in swimming. fused to the quadrate (fig. 185, A and B). The rostral The skull, exclusive of the rostrals, is simpler than

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 in any of the Recent Acipenseridae and agrees more with a peg and open socket arrangement; laterally, closely with that found in the paleoniscoids. The evo- they fit together by overlapping and underlapping each lutionary trend in the Acipenseridae has been the frag- other, a condition not dissimilar to that found in Aci- mentation of the skull bones and the reduction of ossifi- penser. Propenser differs from Acipenser in the type cation of many of the elements. Fusion of the maxil- of external ornamentation of the plates (fig. 181), the lary to the palate and reduction in size of the lower larger size of the frontals (fig. 180), the fact that later- jaw have also occurred. In these respects Propenser ally the frontals form the upper edges of the orbits, the appears to be much more primitive than any living smaller size of the parietals, the elongation of the supra- genus in this family. occipital, and the more complete ossification of the Each segment of the neural arch of the holostean hyomandibular (fig. 183, A), shoulder girdle (fig. 184, Saurichthys has separate basidorsal and interdorsal ele- C-D), neural arches (fig. 185, F), and endocranium. ments with spines of about equal height (Holmgren The extreme amount of ossification shown partic- and Stensio, 1936). Propenser shows the same condi- ularly in the type specimen could be in part due to tion except that the bases of these two elements are the large size and perhaps old age of this individual, fused into a single unit. In the Acipenseridae the com- but probably represents a primitive condition. The parable neural arch has been simplified to a single neu- fusion of the neural arches (fig. 185, F) finds a present 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. Hadrodus priscus Leidy. Figure 1£6. Hewlett, the station master at Boligee, who not only Referred specimen.—Alabama Geological Survey's carefully collected the type specimen on his farm but specimen, right lower jaw; Hewlett's farm, Greene has shown an active interest over a number of years County, Alabama, upper Mooreville. in the fossils which lie close to his home. The second Discussion.— Leidy (1857, 1873) described a pre- and smaller skull was collected by Dr. L. Renger. maxillary from near Columbia, Mississippi, and Gregory (1950) described the premaxillary and splenials of a Infraclass HOLOSTEI form from the Niobrara Chalk which he called Hadrodus marshi. The Mooreville lower jaw. which is Order PYCNODONTIFORMES complete, shows what Gregory called the splenial element, and for the fact that there are two Pycnodontidae Cope except only tooth rows, it is very close to Gregory's H. marshi. No sutures Discussion.— The Pycnodontidae are deep-bodied are discernible on this jaw. angelfish-like holosteans which have strong jaws with crushing teeth. Their time of greatest abundance seems Incertae sedis. Figure 187. to have been the Jurassic, but a few persisted into the Referred specimen .— P 27515, one premaxillary; Eocene. 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 premaxillary. The teeth are broken, with none of them maxillary belongs to the Pycnodontidae. The presence bearing the tips of the crown. of on the surface to this bone large pits appears place Order AMIIFORMES closest to Hadrodus. The antero-posterior alignment Suborder AMIOIDEI of the five teeth is singular. Medially there is an area for the symphyseal attachment with the other pre- Pachycormidae Woodward 404 FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3

with that of the Niobrara species, P. nitida. A revision 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 20 mm heavy in some of the specimens.

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

Discussion.—The concept of the Isospondyli as a Protosphyraena Leidy homogeneous monophyletic group has never rested on Discussion.— is an Cretaceous Protosphyraena Upper a firm morphological or paleontological basis. The holostean with an elongate and partly pachycormid polyphyletic origin of the group has been suggested fused pectoral fin. In the are two large palate anteriorly by many, including most recently Greenwood et al. 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 material available there would appear to be several species 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 tooth is single fang-like upper pres- 10mm ent and well preserved; another tooth is in the process of being replaced. The skull compares very favorably 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 relationships 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 line is that from the phyletic Leptolepididae I mm 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 suborder (Clupeioidei), realizing that the last two are probably derived from the Leptolepidoidei, we have a classification based on phylogenetic evidence that clarifies 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 superfamilies, 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;

I mm however, the following definition will serve for the present paper. Fig. 188. Palelops eutairensis, n. sp., scales; A, type, PF 3559; B, PF 3560. Suborder ELOPOIDEI

Diagnosis.—These are elongate fusiform fishes. The sphenoids and basisphenoids are reduced. There is a post-temporal fossa is present except in one or two cases single dorsal fin. There are two families, Elopidae and where it has been lost or reduced to a groove. The Megalopidae. lateral and fossae are temporal subtemporal generally Discussion.—The scales of this superfamily are very no nor present; preepiotic auditory fossa has been dem- uniform and are divided into four quadrants of orna- onstrated. There is an The intercalar-prootic bridge. mentation. The apical regions have vermiculate ridges; supraoccipital is small and low, never forming a large the cross the circuli the basal regions have radii which ; crest. circumorbitals extend to the The preoperculum. dorsal and ventral quadrants have concentric circuli. The vertebrae are not pierced by the notocord in adults and are without transverse processes. The scales are Elopidae Bonaparte than longer deep, and have distinct circuli; distinct Discussion.—The characteristics of this family have are There are no ventral apical regions present. ridge been discussed by Berg (1947) and Dunkle (1940). scales. The ventral fins are Inter- post-abdominal. The genus Notelops should be placed in this family muscular bones are present. along with Holcolepis, and probably with Elopoides. Elops is known from the Lower Eocene to the Recent. Superfamily ELOPOIDEA The scales of the Elopidae show more than five basal whereas the have less Diagnosis.—The parietals meet in the midline; a radii, Megalopidae commonly than five. gular plate is present; the maxillaries enter into the gape of the mouth; the jaws are toothed, with small Palelops, new genus teeth; the mesethmoid is small; the frentals are long, — new tapering anteriorly; the nasals are small, not meeting Type species. Palelops eutawensis, species. and not normally attached to the frontals; the orbito- Diagnosis.—Same as for species. 406 FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3

Palelops eutawensis, new species. Figure 188. Pachyrhizodus Dixon PF one mile E. Type.— 3559, scale; ', of Walter Description.—An interfrontal fossa is present in the Dance's 6.2 miles W. of farm, Aliceville, Eutaw County, posterior part of the skull roof; it is rectangular in lower Mooreville. Alabama, shape, with its posterior border formed by the supra- Referred specimens.—PF 3612, two scales; l4 mile occipital. On each side of the supraoccipital lie two E. of Walter Dance's farm, 6.2 miles W. of Aliceville, small parietals. The epiotics form the postero-lateral Eutaw County, Alabama, lower Mooreville. PF 3563, angles of the skull. Anterior to the epiotics are large one scale; Choctaw Bluff, Greene County, Alabama, pterotics. There is a portion of a large posttemporal lower Mooreville. PF 3560, four scales; Montgomery's in one specimen but its shape is not known. No post- farm, 2 miles N. of West Greene, Greene County, Ala- frontal has been found. The two pairs of postorbitals bama, middle Mooreville. PF 3561, one scale; Hew- are large and more or less rectangular. The upper lett's farm, gullies near the road, Greene County, Ala- postorbital meets the scale bone; the lower one abuts bama, upper Mooreville. against the front edge of the preoperculum. The sub- orbital is and a is Diagnosis.—The basal regions of the scales have 19 naiTow, prefrontal present and large. The sclerotic bones are two in number. to 21 radii which tend to be parallel; the circuli in the evidently No basal are distinct the nasal element is known. The premaxillary is consider- regions ; dorsal and ventral quad- smaller than the rants have straight circuli. The scale centers have ably maxillary and has two rows of teeth. The teeth of both the coarse raised granules which cover the nuclei of the upper and lower jaw are in sockets much as in mosasaurs. circuli; the area covered by these granulations varies The mandible (fig. 193 is and in in different scales. The apical regions show circuli and D) large nearly rectangular shape but narrows A small is granulations. These scales differ from those of Elops anteriorly. angular present; posterior to the articular surface is a rounded in the greater number of radii, the nonvermiculate flange. The 193 is character of the apical ornamentation, and the basal quadrate (fig. D) triangular except for rounding in the The extension of the granulations over the nucleus. posterior apex. hyomandibular has a large single head and a ven- Discussion.—Dr. Lore David has in one of her scale prominent opercular process; trally it is very narrow and similar to that of notebooks a photograph of an identical scale from the Ziphac- tinus (Stewart, 1900). The operculum is covered dor- Cretaceous of California; however, locality data are sally by the large scale bone. Externally it has fine lacking, although this scale is believed to be from the striae which radiate out from its point of articulation. Panoche Hills, near Fresno, California. Palelops eutaw- The preoperculum is low and wide, with that ensis represents the nearest approach to the genus grooves radiate posteriorly from the juncture of the anterior Elops of any scale known in the Mooreville fauna. and dorsal arms. The suboperculum has a rounded Mr. Marion C. Bonner of Leoti, Kansas, has found an posterior edge; its anterior and dorsal edges are straight example of a scale from the Niobrara Chalk which is and form a right angle. The interoperculum is poorly identical to this species except that it is larger (PF known. The branchiostegals are over 20 in number. 3336). Another scale fragment of this species is to be The posterior of the skull has on each side a found in an undescribed Cretaceous fauna from South portion large fossa. An is Dakota in the Field Museum Collection. posttemporal opisthotic bridge present. There is no auditoiy foramen such as one finds in the Clupeidae. The parasphenoid covers the base Superfamily PACHYRHIZODONTOIDEA, of the skull and has two small forks posteriorly; its anterior extremity is shovel-like. Anterior to the new superfamily parasphenoid are two rounded thick bones (fig. 193 C), thought to be vomers, each of which bears a single Diagnosis.—The parietals do not meet in the mid- tooth-like structure. All the palatal elements and gill line; the gular plate is absent; the maxillary enters into supports bear, where they are exposed in the mouth, the gape of the mouth; the teeth are large, recurved, a continuous cover of small teeth which are curved and and conical. The mesethmoid is large and the frontals resemble the jaw teeth but lack distinct sockets. The are rectangular in outline. There is a large prominent epihyal is roughly semicircular. The ceratohyal (fig. scale bone above the operculum. An enlarged scale is 193 B) is approximately rectangular but widens at its present in front of each of the paired fins. There is posterior end. The upper part of the shoulder girdle is only one recognized family, the Pachyrhizodontidae. unknown. The lower part (fig. 193 A) has a prominent Discussion. —The Pachyrhizodontoidea represent mesocoracoid arch. one of the important Cretaceous radiations of the Elo- Each fin has one first poidei. Some generic relationships are as yet too un- pectoral very large ray (fig. certain to assign all the genera to families within this 199 B). The ends of the other rays are fringed (fig. 189 superfamily. Among the genera are Pachyrhizodus, B), branching several times distally. The pelvic rays are Thrissopater, Elopopsis, and Rhacolepis. In most the curved and are of nearly uniform size; each ray is parietals are completely separated by the supraoccipital. divided into joints distally. The anal fin resembles The teeth are well developed, and there is a huge scale the pelvics in these features. The tail has been figured above the its is plate operculum. by Hay (1903) ; whole surface 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 contain strong circuli. and has two fin edge, internally hypurals. The dorsal Discussion.—Some of the Niobrara species of Pachy- rays have cup-like bases, from which arise a pair of rhizodus have been based on characters that could fall lateral projections. There are 50 to 60 vertebrae. within the range of individual variation. An examina- The genus Pachyrhizodus is also characterized by tion of the Niobrara material at the University of Kan- its distinctive scale type (fig. 191) with a triangular sas and at the American Museum of Natural History apical region. In this are dendritic ridges which branch revealed what is believed to be three valid species. toward the apical edge. The lateral and basal areas 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 cies; they are Kansanus martini, and Eurychir lindleyi. A, 191 B-C, 192. Pachyrhizodus sheareri and lOricardinus tortus may belong to this species but are so poorly preserved as to Referred specimens.—P 27489, fragments of dermal be indeterminate. bone, premaxillary, vertebrae, and lower jaw; Moore's PF 1697 is the best Mooreville farm, Dallas County, Alabama, middle Mooreville. PF (fig. 189) specimen ; it is about three feet and has a total vertebral 1697, a nearly complete articulated fish; 5 miles SSW. long count of 54. There is a scale the of Clinton, Greene County, Alabama, middle or upper very large bone; operculum shows faint radiation the Mooreville. only lines; preoperculum is large in comparison with that of Elops and Discussion.—The two Mooreville specimens, partic- bears a series of basal striations. The premaxillary is the are to P. which ularly second, assigned minimus, only about one-fifth as long as the maxillary. A large was described by Stewart (1900) from a lower jaw. arrowhead-shaped ethmoid element is conspicuous. The Other specimens in the University of Kansas Museum, pectoral fin has at least 17 rays. There is some evidence which were described as belonging to various families of a sclerotic ring. The teeth are very small and slen- and genera by Jordan (1925), are referred to this spe- der, one of the better characters for distinguishing this - V I mm Hl mmjf^fe \

,W\ I mm

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

409 410 FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3

Fig. 192. Reconstruction of Pachyrhizodus minimus. species. The vertebrae show externally very fine longi- ceratohyal; both from Donald's farm, Dallas County, tudinal striae. Alabama, middle Mooreville. PF 292, jaw fragment,

In cross-section (fig. 190, A), a vertebra shows in Section 33, Township 16 N., Range 16 E., Montgomery its center a small core of lamellar bone. About half- County, Alabama, middle Mooreville. P 27504, lower way from the center is a bony ring. The outer surface jaw and three vertebrae; Township 11, W. of High- of the vertebra has a small ring of lamellar bone which way 13, Hale County, Alabama, middle Mooreville. is indented above and below. PF 3542, skull fragments, eight vertebrae, and fragments of fin Hewlett's near The apical region of the scales (fig. 191, B and C) spine; farm, road, Locality has externally a triangular area with 17 rays converging 8, Greene County, Alabama, upper Mooreville. P 27532, at the center of the scale. The rest of the scale is fin spine; Hewlett's farm, Greene County, Alabama, covered with coarse circuli. upper Mooreville. PF 442, jaw fragments; 2 miles W., 1 mile N. of West Greene, Greene County, Alabama, Pachyrhizodus caninus Cope. Figures 190 B-C, 191 upper Mooreville. A, 193, 199 B. Discussion.—Pachyrhizodus caninus is not only the Referred specimens.—P 27410, lower jaw fragments; largest species of this genus but one of the largest of Banks' farm, Greene County, Alabama, lower Moore- the Mooreville fishes. There can be no question that ville. P 27416, fin spine, premaxillary, and lower jaw; the Mooreville specimens belong to the same species gully near the Choctaw Road, Greene County, Ala- as the Niobrara P. caninus. As yet no articulated bama, lower Mooreville. PF 3554, 19 vertebrae and specimens are known from the Mooreville though in- tail; 2 miles N. of West Greene, Greene County, Ala- dividuals with a great number of associated scales and bama, middle Mooreville. PF 1696, jaw and most of bones do occur. The scales (fig. 191, A) agree with those a skull, pectoral fin, and vertebrae (almost complete of P. minimus except for the finer and more numerous of in P. radii in fish) ; 2 miles W. West Greene, Greene County, Ala- circuli caninus. The apical are greater bama, middle Mooreville. PF 128, three vertebrae and number in P. caninus than in P. minimus and not 6 > part of a jaw; 1 10 miles N. and ) mile West of West as well defined. The granular ornamentation is more Greene. Greene County, Alabama, middle Mooreville. prominent in P. caninus, and the scales are relatively PF 137, four vertebrae; PF 138, vertebrae and a fin larger. The cranial bones vary in proportions; this of spine; P 27423, one vertebra; P 27451, jaw fragment; course may be in part due to distortion by post-de- P 27502, shoulder girdle and fin rays; P 27513, jaw; positional forces. Among the more stable and easily

P 27516, jaw fragment; P 27518, quadrate, lower jaw, recognizable elements are the large ceratohyals (fig. fragments, and fin spines; P 27519, fragments and lower 193, B) and the quadrate (fig. 193, D), which has an at- jaw; P 27520, two maxillaries; P 27523, part of lower tenuated anterior process. The vertebrae are charac- jaw; all from Moore's farm, Dallas County, Alabama, terized externally by their smooth sides. In cross-sec- middle Mooreville. P 27323, almost complete fish; tion (fig. 190, B and C) they have a nearly solid bony gulley near Harrell's Station, Dallas County, Alabama, center bordered by an inner ring and a superficial one middle Mooreville. PF 134, skull fragments; PF 135, that sends highly irregular processes inward, forming APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 411

20mm

Fig. 193. Pachyrhizodus caninus; A, pectoral girdle, P 27501; B-D, PF 1696; B, ceratohyals and C, vomers; D, portion of jaw with attached quadrate. an elaborate pattern similar to fine lacework. The in the Niobrara. There is a possibility that P. kingi is overall vertebral cross-section is basically similar, how- only the juvenile form of P. caninus, but there is a ever, to that of P. minimus and P. kingi except for decided size gap between these two forms. A vertebral being more complex in pattern. The jaws of this spe- cross-section of P. kingi (fig. 190, D) shows differences cies (fig. 193, D) have relatively big teeth. in structure from P. caninus. P. leptognathus and P. P. latimentum and P. leptopsis are synonyms of P. velox are synonyms of P. kingi. caninus. Superfamily ALBULOIDEA Hay Pachyrhizodus kingi Cope. Figure 190 D. Referred specimens.—P 27411, jaw fragments and Diagnosis— -The parietals meet in the midline. A skull fragments; PF1612, vertebrae and jaw fragments; gular plate is present. The maxillary is excluded from PF 3546, vertebrae and skull fragments; all from Moore's the gape of the mouth; the premaxillary is free; the farm, Dallas County, Alabama, middle Mooreville. jaws, the paired palatal plates, and the gill arches are Discussion.—Pachyrhizodus kingi is intermediate in covered with fine conical teeth; the parasphenoid and size between P. caninus and P. minimus. It also occurs basibranchial bear large button-shaped teeth. Well os- 412 FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3

sified orbitosphenoids and basisphenoids are present. Albula dunklei, new species. Figures 194, 200 B-C, The frontals are elongate, triangular, and taper anteri- 201 B. The nasals are small and not attached to the orly. Type.—P 27494, part of "body," scales in place, frontals. The mesethmoid is small. The lateral tem- fins, part of a head, and vertebrae; Moore's farm, Dallas poral fossa is roofed; there is no angular. Very char- County, Alabama, middle Mooreville. acteristic are the scales, which are rounded posteriorly Referred specimens.—PF 3580, scales; Locality 1, and bear distinct radial plications anteriorly. The sur- 6.2 miles W. of Aliceville, E. of Walter Dance's farm, face of these is ornamented with plications granules Eutaw County, Alabama, lower Mooreville. PF 3564, arranged in a dendritic pattern or in rows that are scales and vertebrae; Locality 4, Hale's farm, 2 miles in a radial direction. The dorsal and ventral aligned NE. of West Greene, Greene County, Alabama, middle surfaces of the scale bear fine distinct circuli which are Mooreville. closely spaced. Anteriorly the circuli are broken into Diagnosis.—The bones of the skull are proportion- vermieulate ridges as in Elops. ately much thicker than in the modern species of Albula. There are two Recent families that belong in this The pelvic fin has 11 rays. The symplectic is relatively superfamily, the Albulidae and the Pterothrissidae. The larger and the quadrate is more triangular in outline scales of these two families are almost identical. than in other species of Albula. The lower jaw bears a Albulidae Gunther tooth patch similar to that of Recent species. The scales (fig. 201 B) show four basal lobes; the basal regions Diagnosis.—The dorsal fin is short; the vomer and of scales have granules which are arranged in antero- palatines are toothed; and the interorbital septum is posterior rows as opposed to the dendritic pattern in bony. There are two modern genera, Albula and Dix- Albula vulpes. The apical region of Albula dunklei is otiia. 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, crushing 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 abdominal 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 10 mm lateral groups have three bands; below these there is on each side a ventrolateral group with two bands, and Fig. 195. Tooth-bearing plate of Albula sp. (P27392). ventrally there is a wide median group with eight bands button-like teeth. The indi- of bony supports, two pairs of bands on each side of the characteristic fragment vertebra. Between this median ventral group and the cates a fish larger than the Recent Albula vulpes, and twice as as those in Albula ventro-lateral groups are also areas of spongy bone. The the teeth are at least large dunklei. caudal vertebrae (fig. 200, C) have eight radial bands in to the groups of bands corresponding position eight Superfamily PLETHODOIDEA, new superfamily of anterior vertebrae. There is a neural and a haemal —The meet in the there arch fused to the top and bottom of each caudal ver- Diagnosis. parietals midline; tebra. No areas of spongy bone have been noted in is no gular plate. The maxillaries enter into the gape of the caudal vertebrae. Almost identical vertebrae occur the mouth. The premaxillaries may be fused to the ethmoid. The ethmoid varies in relative size and is in the Recent Albula vulpes. often huge. The jaws are toothed, and the parasphe- This species is named for Dr. David Dunkle, who noid and basibranchials bear crushing teeth. The fron- has been instrumental in the identification of the pres- tals are nasals are united to the ent albulid. rectangular; anteriorly frontals. The orbitosphenoids are large. The pectoral The Albulidae have a spotty Upper Cretaceous rec- fins are high on the body. The dorsal fin covers most ord, due in part to the fact that many of the specimens of the back. Scales {Bananogmius) have circuli on basal, in collections have not been described, while other spe- dorsal, and ventral surfaces; the apical region has granu- cies are known only from their scales, such as David's lations arranged in postero-anterior rows as are the ( 1946) Kleinpellia. Cockerell (1933) described, from a radii; the scales do not have a true radial or dendritic scale, a species called Albula antiqua from the Creta- pattern as in Pachyrhizodus. The vertebrae have ex- ceous of Florida. The University of Kansas Museum ternal striations and dorsally two concavities; in cross- has a specimen (F 985) from the Niobrara labeled Lep- section they have numerous radial supports separated tichthys which is an unquestionable albulid. The U. S. by spongy bone. National Museum has a similar undescribed albulid, Two families, Bananogmiidae and Plethodidae, are also from the Niobrara. Field Museum has a specimen, included in this superfamily. PF 685, from Sahel-Alma, Mt. Lebanon, which is a member of the genus Albula. Bananogmiidae, new family

Diagnosis. —No mucus canals are known on the fron- Albula sp. Figure 195. — tals. The mandible is deepest at its center. The basi- Referred specimen. P 27392, part of palate with branchial bears only one crushing toothed plate. The teeth; Moore's farm, Dallas County, Alabama, middle preoperculum is L-shaped, with the vertical limb nar- Mooreville. row and the anterior limb expanded. Discussion.—There is at least one bit of evidence Discussion. —The following three genera are referred for the existence of a second species of Albula in the here: Bananogmius, Paranogmius, and Moorevillia. The Mooreville. This consists of part of a palate with 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 Bananogmius crieleyi, new species. Figures 196, with very wide parasphenoid plates. The parasphenoid 200 D. dental plate is oval in B. zitteli and rectangular in B. Type.—PF 3608, premaxillary, basihyal, parasphe- aratus. The parasphenoid dental plate of B. intermedins noid and vomer, two ectopterygoids, quadrate?, frag- is unknown, but its basibranchials are closer to those ments of skull, and five vertebrae; Moore's farm, Dallas of the above two species than to those of the second County, Alabama; middle Mooreville. The in B. zitteli is fused to the group. premaxillary Diagnosis.—The parasphenoid (fig. 196 A) is elon- ethmoid. gate, with a club-shaped dental patch; fused to it is the The second group of species consists of B. evolutus, vomer with an oval tooth patch. The basibranchial B. favirostris, B. altus, B. polymicrodus, B. crieleyi, and (fig. 196 C) is very thick, five-sided, and with a deep B. sp. Loomis. Here the parasphenoid dental plate is indentation posteriorly. The premaxillary is free and elongate and club-shaped. The premaxillaries of B. bears several rows of small teeth; its external surface

evolutus (Loomis, 1900, pi. XXVI, fig. 6), B. favirostris, (fig. 196 B) is ornamented with pits similar to the tooth and B. polymicrodus are all very similar. The jaws of sockets in the basibranchial. Two large elongate tri- B. evolutus and B. polymicrodus are long and low. The angular pterygoids (fig. 196 E and G) are present; the

parasphenoid of B. evolutus is unknown. 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 and have a claw shaped posterior end with a large deep and irregular; a cross-section of one is illustrated tooth patch. The posterior portion of the parasphenoid on figure 200 D, and shows radiating spongy bone. lacks wings. The frontals taper markedly anteriorly. Discussion. —The narrow parasphenoid places this The quadrate is high. The first pectoral fin ray is form in the second group of the genus Bananogmius. robust. The vertebrae are about three times higher The elongated parasphenoid tooth plate and the oval than long, and show radiating bands of spongy bone tooth plate on the vomer are similar to those of B. in cross-section. favirostris. The shape of the vomerine tooth plate is 197. similar to that of B. altus. The basibranchial is remi- Moorevillia hardi, new species. Figure niscent of that of Plethodus pentagon from the British Type.—PF 3567, skull fragments, fin rays, verte- Chalk and of B. evolutus. The combination of these brae, and jaws; Moore's farm, Dallas County, Alabama, characters is, however, unique. This species is named middle Mooreville. of who served as for Bruce Crieley Chicago, Illinois, Diagnosis.—Same as that of the genus. my field assistant in Alabama. Discussion.—This genus agrees with Bananogmius in Bananogmius cf. zitteli Loomis. Figure 198 H. the elongate patch of teeth on the lower jaw and the Referred specimen. —PF 3609, snout; Moore's farm, single row of teeth on the premaxillary. No crushing Dallas County, Alabama, middle Mooreville. tooth pad like that of Bananogmius has been found. The is smaller lacks Discussion.—This large pitted snout is too fragmen- premaxillary decidedly and the ornamentation characteristic of As far tary for positive identification; however, it resembles Bananogmius. as can be told in the the are closely the rostrum of the fish described by Loomis present specimen parietals the It best to (1900, pp. 229-234, PI. XXI). separated by supraoccipitals. appears consider this a member of the Bananogmiidae, which cf. Bananogmius polymicrodus (Stewart). it resembles in such features as vertebrae with fine 201 C. Figure external striae, and teeth arranged in characteristic —PF 3566, tail with scales and Referred specimen. patches. American Museum of Natural History speci- of Hale's 2 miles NE. of vertebrae, part skull; farm, men No. 8319 from the Niobrara formation probably West Greene middle Moore- Greene, County, Alabama, belongs to this genus and species. ville. This species is named for Allen M. and Robert H. Discussion. —The scales and vertebrae agree with Hard, of Tuscaloosa, Alabama, who collected some of those of B. polymicrodus. The scales bear circuli in the fish material described in this paper. their basal, dorsal, and ventral regions. The apical has vermiculate granules which lie between smooth region Suborder CLUPEOIDEI bands running antero-posteriorly rather than radiating from the nucleus. The antero-posterior alignment of Diagnosis.—No posttemporal fossa is present, the the bands distinguishes these scales from those of Pachy- parietals are separated from each other by a large crested rhizodus. supraoccipital. The orbitosphenoids and basisphenoid may be well developed or absent; there are one or two Bananogmius sp. Figure 198 G. supramaxillaries; there is no gular plate. There is no Referred specimen.—PF 131, parasphenoid and den- adipose fin. Postcleithra are present; there are en- tal plate; 1 /ix miles W. and % of a mile N. of West larged scales behind the paired fins. The body scales Greene, Greene County, Alabama, middle Mooreville. are generally wider than long, with circuli usually ob- Discussion.—There is a very close resemblance be- scure. There are at least two superfamilies, the Chiro- tween this parasphenoid plate and that figured by centroidea and the Clupeoidea. Loomis (1900) as Bananogmius sp. As no other material is at hand, this fish must still remain specifically indeterminate. Superfamily CHIROCENTROIDEA, new superfamily Moorevillia, new genus — Type species. —Moorevillia hardi, new species. Diagnosis. The supraoccipital crest is very large. radials are in rows. Diagnosis.—The premaxillary is free from the eth- The pectoral two The premaxillaries bear a few teeth than on moid, is small, and lacks prominent external ornamen- usually larger the maxillaries. bone has tation. There is only one row of conical teeth on the Each palatine a hammer-like articular The are small. anal fin is premaxillary. The lower jaw is long and narrow, with process. parietals The oppo- site the dorsal. vertebrae on an elongate patch of small, recurved, enamel-tipped The have deep grooves conical teeth. The maxillary is also a long thin element their sides, with two pits above and below for insertion of neural and haemal arches. cross-section of ver- bearing a similar patch of teeth. The premaxillary A a fits into the abruptly up-turned anterior end of the tebra shows spongy bone without radial or concentric structures. maxillary. The palatines are small oval plates bearing patches of conical teeth, which stand over cup-shaped There are three families: Ichthyodectidae, Sauro- pits in the bone. The pterygoids are very elongate, 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 PF of lower 131 jaw, 289; E, fragment right jaw, PF 289; F, premaxillary; G, Bananogmuis sp., parashpenoid, PF ; 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, middle 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 differentiate this form from Xiphactinus audax.

Xiphactinus audax Leidy. Figure 200 A. Referred specimens.—PF 3543, vertebrae, fragmentary 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;

1 PF 129, lower jaw fragments; all from 1 2 miles W.

10mm 5 mm

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.

I Omm and % of a mile N. of West Greene, Greene County, Alabama. P 27503, skull fragments and one vertebra; skull P 27525, jaw fragments, 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, palatoquadrate, ' > 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, vertebrae 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 20 mm is little doubt that the Mooreville Xiphactinus is the same as the large Niobrara species, X. audax; the other described Niobrara are of Fig. 199. A, jaw of Saurodon 1 sp., medial view, PF 3585; species questionable validity. B, Pachyrhizodus caninus, pectoral fin spines, PF 1696. A cross-section of a vertebra is figured in Figure 200 A. APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 419

2 mm

2 mm I m m

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

Saurodontidae Stewart jaw and skull fragments; 1 mile W. of West Greene, Greene County, Alabama, middle Mooreville. PF 3544, Diagnosis.—A predentary is present. The teeth are 4, Hale's farm, 2 miles N. of compressed and are implated in sockets. jaw fragment; Locality West Greene, Greene County, Alabama, middle Moore- This family has two closely related genera, Saurodon ville. PF 130, part of lower jaw; 2 miles N. of West and that have been Saurocephalus synonymized by Hay Greene, Greene County, Alabama, middle Mooreville. (1903), but since no intermediate forms exist the among PF 3548, right lower jaw; P 27413, left lower jaw and Mooreville specimens, it is best to the thought keep left premaxillary; P 27414, maxillary; P 27415, pail of genera separate. lower jaw; P 27430, lower jaw and vertebra; P 27435, jaws and skull fragments; P 27483, better part of a skull Saurodon leanus Figure 200 E. Hays. and four vertebrae; P 27508, lower jaw and skull frag- Referred specimens.—PF 3611, lower jaw and skull ment; P 27530, part of skull and two vertebrae; all fragments; 2 miles W., 1 mile N. of West Greene, from Moore's farm, Dallas County, Alabama, middle Greene County, Alabama, middle Mooreville. PF 122, Mooreville. P 27506, part of skull and lower jaw; right lower jaw; 1.6 miles N. of West Greene, Greene P 27507, part of lower jaw; both from Township 11, County, Alabama, middle Mooreville. PF 3540. lower W. of Highway 13, Hale County, Alabama, middle 420 FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3

l Mooreville. PF 123, skull fragments; 2- mile N. of Discussion.—This small preoperculum with three Mt. Hebron, near West Greene Road, Greene County, very strongly marked sensory canals is of the type Alabama, upper? Mooreville. found commonly only among the Clupeidae. Discussion. —The jaw of Saurodon shows deep notches Dussumieriinae, indet. Figure 201 A. below the alveolar border. The existence of more than — one species is an open question. The present form is Referred specimen. PF 3594, two scales; Montgom- referred to S. leanus, the first described species, typi- ery's farm, 2 miles N. of West Greene, Greene County, cally from the Upper Cretaceous of New Jersey. A Alabama, middle Mooreville. sectioned vertebra displaying spongy bone is shown in Discussion.—The anterior areas of these scales are Figure 200 E. missing, but the strong posterior radii as well as the grooves at right angles to them are typical of scales Saurodon? sp. 1. Figure 199 A. of the modern Dussumieriinae, a subfamily of the Clu- Referred specimen.—PF 3585, the better part of a peidae. small lower jaw; Moore's farm, Dallas County, Ala- bama, middle Mooreville. Order INIOMI Suborder Discussion.—This small lower jaw is similar to that MYCTOPHOIDEI of Saurodon in jaw shape, but, perhaps because of its — Diagnosis. The maxilla is only weakly if at all small lacks of the notches of S. leanus. size, any deep dentigerous, and is usually excluded from the gape of This jaw represent a smaller species or a juvenile may the mouth. The inner teeth in the mouth are generally of S. leanus. larger than the outer. No mesocoracoid arch is known. An adipose fin is commonly present; the ventral fins Saurodon? 2 sp. — are on the posterior part of the abdomen. Referred specimen. PF 3550, jaw fragment; Moore's The following Mooreville families are referred to farm, Dallas County, Alabama, middle Mooreville. this order: Enchodontidae, Dercetidae, and Myctophi- dae. I do not follow Romer's Discussion. —This jaw fragment, probably a maxil- (1966) use of the super- order lary, is similar to the one above, except that the teeth Protacanthopterygii. are much smaller. Enchodontidae Loomis

Saurocephalus cf. lanciformis Harlan Diagnosis- -The parietals are separated by a supraoccipital. The premaxillaries are large, and the maxil- Referred specimens. —PF 443, upper jaw; W. of West laries, although small, are in the of the Greene, Greene County, Alabama, middle Mooreville. present gape mouth; each premaxillary has a posterior spine. The PF 3557, portion of lower jaw; Township 11, W. of palatines bear one or two large fangs; each lower jaw Highway 13, Hale County, Alabama, middle Moore- has two tooth rows, with the outer row much smaller ville. P 27407, jaws; P 27412, lower jaw; P 27505, jaw than the inner one; the teeth are fused to the and skull fragments; P 27509, jaws and skull fragments; jawbones. Scutes and scales are present on the and the P 27511, skull fragments; all from Moore's farm, Dallas body, surfaces of the cranial and scutes are County, Alabama, middle Mooreville. plates highly ornamented. The nasals are small. Discussion. —Saurocephalus is similar to Saurodon ex- Two Mooreville genera, Enchodus and Cimolichthys, cept in having foramina instead of deep notches below are included in this family. the alveolar border internally. The oldest name is used, as the other are of doubtful species validity. Cimolichthys nepaholica (Cope) Referred specimens.—PF 441, two basal parts of fin Superfamily CLUPEOIDEA rays; 2 miles W. of West Greene, Greene County, Ala- Diagnosis. —The supraoccipital crest is reduced bama, middle Mooreville. PF 3565, quadrate, basal though still present; the postcleithra are attached to portion of fin ray, and skull fragments; 1 mile W. and the outer side of the cleithra; the pectoral radiae are 1 mile N. of West Greene, Greene County, Alabama, in one row. The teeth are small or absent. The pre- middle Mooreville. P 27526, two basal parts of fin maxillae are "L" shaped. The dorsal fins are anterior rays, Crawford's farm, Hale County, Alabama, middle to the anal fin. Temporal foramina and preepiotic Mooreville. fossae are are without present. The palatines hammer- Discussion.—The quadrate of PF 3565 is virtually like articulations. identical with the specialized quadrate of C. nepaholica There is one family in the Mooreville; the Clupeidae. from the Niobrara. A fin ray associated with the quadrate is thought to be either the first pectoral or pelvic Clupeidae, indet. ray. There is little doubt that Cope's (1872) Empo Referred specimen.—PF 3599, left preoperculum; is really Cimolichthys. A number of species have been Choctaw Bluff, Greene County, Alabama, lower Moore- named, but there appears to be only one valid Niobrara ville. species, C. nepaholica, as shown by Hay (1903). APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 421

Enchodus petrosus Cope line. The gape of the mouth is wide; the premaxillaries form Referred specimens.—PF 3573, palatines, one lower the greater part of the upper edge of the mouth. jaw, and three teeth; Hale's farm, 2 miles N. of West The vertebrae are in the form of constricted cylinders Greene Post Office, Greene County, Alabama, middle pierced by the notochord. Scales are small or wanting. Mooreville. PF 3574, two palatines; PF 3575, one A set of elongate scutes is often present. palatine and one lower jaw fragment; Montgomery's One genus occurs in the Mooreville: Stratodus. farm, 2 miles N. of West Greene, Greene County, Ala- middle Mooreville. one 2 bama, PF 3583, palatine; Stratodus apicalis Cope. Figure 198 A-F. miles W. of West Greene, Greene County, Alabama, Referred specimens.—PF 132, one palatine; 1 mile middle Mooreville. PF 3584, two lower jaw fragments; NW. of West Greene, Greene County, Alabama, middle 1 mile N., 1 mile W. of West Greene, Greene County, Mooreville. PF 3610, one palatine; 2 miles W. and 1 Alabama, middle Mooreville. PF 3587, three teeth, mile N. of West Greene, Greene County, Alabama, two jaw fragments, and one palatine; 1 mile N., 2 miles middle Mooreville. PF 3600, one palatine; P 27424, W. of West Greene, Greene County, Alabama, middle a caudal support; P 27487, a palatine and palatine Mooreville. PF 3586, one palatine, two jaw fragments, fragments; P 27512, one palatine; all from Moore's and four teeth, 1 mile S. of West Greene, Greene County, farm, Dallas County, Alabama, middle Mooreville. PF Alabama, middle or upper Mooreville. P 27521, one 3596, one palatine; Hewlett's farm, north set of gullies, tooth; Township 11, W. of Alabama Highway 13, Hale Greene County, Alabama, upper Mooreville. PF 289, County, Alabama, middle Mooreville. PF 136, one parts of both lower jaws, premaxillary?, opercular frag- skull and verte- palatine, jaw fragments, fragments, ment, and skull fragments; Donald's farm, Dallas PF 3569, PF six PF 3571, brae; teeth; 3570, teeth; County, Alabama, middle Mooreville. two teeth and one palatine; PF 3572, three jaw frag- Discussion.—There is little doubt that Stratodus ments; PF 3590, two palatines; PF 3591, one palatine; oxy- is a of S. both are based PF 3592, one palatine; PF 3593, two palatines, three pogon Cope synonym apicalis; on material. The lower 198 D E) jaw fragments, and one skull fragment; P 27417, jaw fragmentary jaw (fig. has a slanting posterior edge with a small cup-shaped fragments and one palatine; all from Moore's farm, articular surface for the The Dallas County, Alabama, middle Mooreville. PF quadrate. premaxillary (fig. 198 F) has a characteristic patch of teeth. The 3582, five jaw fragments, one palatine, and one skull 198 seem the most common fragment; Marion Junction, Dallas County, Alabama, elongate palatines (fig. C) objects in collections. middle Mooreville. PF 3555, one palatine; PF 3556; one palatine, PF 3577, one palatine and one lower jaw; Myctophidae Jordan and Evermann PF 3578, one five and one palatine, jaw fragments, Diagnosis.—The upper jaws are bordered by pre- PF three and two PF tooth; 3579, palatines teeth; 3589, maxillaries only; the dentition is usually feeble. The two all from Hewlett's near the palatines; farm, gullies ventral fins may be abdominal or under the pectoral road, Greene Moore- county County, Alabama, upper fins. The parietals are usually separated by a reduced ville. PF one 8.9 miles from the 3581, tooth; county supraoccipital. Scales are variable but with distinct line, Route 26, Russell County, Alabama, Blufftown. — circuli; their nucleus is apical or central; apical cteni Discussion. Enchodus petrosus is one of the com- are present or absent; an apical field may be present; mon species in the Mooreville fauna. Almost any Moore- cteni are usually in one row and marginal. ville locality will produce some evidence, usually a pal- Myctophid scales are very similar to those of bery- atine, of this species. The collected specimens show a coids, and their recognition can be very difficult. The good deal of variation. myctophids usually possess a single row of cteni or or whereas the have more than Enchodus cf. saevus Hay none, berycoids usually one row. The myctophids lack the raised apical field Referred specimen.—PF 3588, two lower jaws; 2 which is characteristic of many of the berycoids. miles W. and 1 mile N. of West Greene, Greene County, Alabama, middle Mooreville. Myctophidae, indet. Figure 203. Discussion.—The more ornate condition of these Referred specimens. - -PF 3598, numerous scales; jaws is one difference from E. petrosus, and a similarity Choctaw Bluff, Greene County, Alabama, lower Moore- to the form described by Hay (1903) as Enchodus saevus. ville. PF 3601, one scale; PF 3602, lower jaw; both The relationship of E. petrosus to E. saevus is not at fi'om east slope above flood plain of Pintlalla Creek the present clear. Hay (1903) and others have used on Burksville Road, Montgomery County, Alabama, the cross-sectional shapes of teeth to distinguish these lower Mooreville. PF 3595, one scale; Montgomery's species, but this is not a satisfactory character due to farm, 2 miles N. of West Greene, Greene County, Ala- individual variation and types of preservation. Hay's bama, middle Mooreville. PF 3597, numerous scales; species was named from Niobrara material. Hewlett's farm, gullies near road, Greene County, Ala- Dercetidae Cope bama, upper Mooreville. Diagnosis.—The head and trunk are elongate. The Discussion.—The great number of myctophid scales im- parietal bones are of moderate size, meeting in the mid- with vertebrae and head plates, the latter almost 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 outer angles. The flattened posterior edge of the scale doubt as to the presence of more than one species in bears a number of spines (cteni) in a single row. The the Mooreville; however, since any patch of scales will basal (anterior) portion is the apex of the triangle. show more than one type of scale, it seems that the The nucleus is at or near the base of the scale. The best course is to describe the predominant scale types. circuli are close together but do not normally make Type A (fig. 203 A) is a triangular scale with rounded 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- the Recent Myctophidae and compare very well with cleus with the first few circuli complete, and has the the scales in the genus Myctophum. David (1946) has circuli widely spaced. described somewhat similar scales from California and Type B (fig. 203 D) has the nucleus centrally located has assigned them to the genus Sardinioides, placed and the circuli placed very far apart. The scale is in the Chlorophthalmidae. subrectangular with a convex protruding lip in the center of the basal edge. Posteriorly there is one row of Order BERYCIFORMES cteni, ten or more in number. A variant of this type Diagnosis.—The premaxillaries are protractile and shows finely spaced circuli and a posterior nucleus at the dentition is feeble. There are seven to eight bran- the edges as in D. Type chiostegals. The vertebral count is from 24 to 30. The C (fig. 203 C) is a rounded scale for a Type except scales have a raised apical area and several rows of cteni. convex protruding basal lip as in Type B. The nuclear Trachichthyidae Bleeker area is very large; circuli are few and widely spaced; no cteni are present. A variant of this type shows Hoplopteryx? sp. Figure 202 CD. closely spaced circuli very reminiscent of the condition Referred specimens.—PF 3603, scales in coprolite met with in the Salmonidae. with Palelops; Hewlett's farm, north set of gullies,

Type D (fig. 203 B) is wider than long, with a Greene County, Alabama, upper Mooreville. PF 3606. rounded posterior edge; the basal edge is pectinated. one scale; Montgomery's farm, 2 miles N. of West The nucleus is posterior, and radii are variable in num- Greene, Greene County, Alabama, middle Mooreville. ber. The circuli are closely spaced; cteni may or may Discussion.—The Mooreville berycoid scales with not be present in this type. their raised posterior area and numerous cteni agree All these scales are very close to what is found in 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 Teleost incertae sedis (2) . Figure 202 A. lack in the Chalk. Woodward's scales cteni, yet Recent Referred specimen.—PF 3607, one scale; east slope fossil the fall off. as well as the Berycidae cteni easily above flood plain of Pintlalla Creek on Burksville Road, The basal plications on these scales suggest those found Montgomery County, Alabama, lower Mooreville. in The fact that scales of this form were found Elops. Discussion.—This indeterminate but highly distinc- intact in a to the importance of copro- coprolite points tive scale is closest to those of the myctophids. The lites in faunal elements that might other- preserving nucleus is possibly posterior. The surface is covered wise be lost. Patterson has demon- Recently (1965) by distinctive granules. There is no evidence of cteni. strated that Hoplopteryx belongs in the family Trachi- chthyidae and not the Berycidae. 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 composed 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 terrestrial 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. Fig. 204. Operculum referred to family Trachichthyidae ?, PF 3604. Overlying the Tuscaloosa is the Eutaw Formation, consists of fine at the base Trachichtyidae? indet. Figure 204. which gravels that grade into highly crossbedded glauconitic sands (Monroe, Referred specimen.—PF 3604, an operculum; 6.2 1941). At the top of the Eutaw, the Eutaw-Mooreville miles W. of Aliceville, E. of Walter Dance's farm, Eu- contact is not defined. There is a re- taw County, Alabama, lower Mooreville. sharply great duction of glauconitic sands at the base of what is gen- Discussion.—This record is based on an elongate erally considered to be Mooreville, and in a few feet opercular element with a distinctively ornamented sur- above these beds chalk dominates. The glauconitic face consisting of ridges topped in places by rounded sands continue, though in sharply decreasing amount, protuberances. The ridges radiate from the upper an- into the Mooreville chalk; minute amounts of glau- terior edge just above the socket for the opercular pro- conite are present throughout the Mooreville and even cess of the hyomandibular. The whole operculum is the overlying Areola member. higher than long, and the complete element would be If we consider this sequence of beds from Tuscaloosa about 40 mm. high. through the Mooreville as a unit, there is an overall Teleost incertae sedis (1). Figure 202 B. decrease in sediment particle size from the bottom to — one Referred specimen. PF 3605, scale; Montgom- the top, perhaps connected with a diminishing of cur- 2 ery's farm, miles N. of West Greene, Greene County, rent phenomena; this is accompanied by a shift from Mooreville. Alabama, middle heterogeneous sediments to relatively homogeneous sed- Discussion.—This small scale has a basal nucleus iments, which could be indicative of an increase in and four radii which end probably in spine-like points. depth. The situation suggests a slowly submerging The circuli go in a straight path from one radius to the coast line: Tuscaloosa-like sediments are followed by next, except at one radius in the anterior portion of shallow water Eutaw-like sediments which in turn are the scale where they form an acute angle and approach followed by deeper water Mooreville-like sediments. It the nucleus. In many respects this scale agrees with seems likely that at any one time all these three types that of Caulolepis longidens, a Recent berycoid, yet till were being deposited, and that the Eutaw sediments more evidence is available it is best to list this very were seaward of the Tuscaloosa, as the Mooreville were unusual scale as incertae sedis. seaward of the Eutaw. APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 425

The Areola Limestone member is a very thin layer less than two millimeters in their greatest diameter. (five feet at the type locality) which overlies the chalk; Secondary minerals include gypsum crystals, calcite the contact is not sharply defined. The upper part of crystals, lenses of limestone, and nodules of chalcopy- the chalk has numerous limy lenses which become rite. The latter range from less than one-fourth of an more abundant as one nears the Areola. The Areola inch to nine inches in their greatest diameter, and show Limestone might represent the deepest-water sediment their secondary nature in replacement of organic por- of all the formations that have been mentioned. tions of the chalk.

The Mooreville is strikingly homogeneous along its The calcareous portion of the chalk has a high or- area of outcrop. The lower layers look identical in ganic content consisting of bones, plates, and tests of composition even when seen at localities a hundred marine organisms. Coccoliths and rhabdoliths, which miles or more from each other. The upper layers, al- are the calcareous disks and platelets of Chrysophyta, though not examined over such great distances, give are an important constituent. These algal platelets the same impression. These remarks apply not only to have a size range of from one micron to about thirty the sediments but also to the fauna. One must assume microns. Every handful of chalk normally contains a fairly deep water to explain this great conformity over number of Inoceramus shell prisms, ostracod tests, Fora- such wide areas. minifera shells, small Ostrea, and bone fragments. Phos- We have at present no information concerning con- phate nodules in the form of worm coprolites occur in vast numbers. If one such a of Mooreville temporaneous shoreward deposits directly north of the puts sample Mooreville outcrop area. Toward the northwest, in chalk in water it will be altered to a very fine ooze. It Tennessee, the Mooreville intertongues with the Coffee is highly probable that the ocean bottom was made up Sands, which resemble the Eutaw in fauna as well as in of such an ooze throughout most of the deposition of the chalk. nature of the sediments would sediments. To the east the Mooreville is replaced by The oozey account for the restricted lack of the Blufftown, a formation containing a more clay-like epifauna. A oxygen characteristic of oozes would account in sediment which has abundant Ostrea and Exogyra reefs, present part probably an indication of shallower water. A more for the scarcity of an infauna. The small size of inor- a distance from areas of diverse molluscan fauna is present although poorly pre- ganic particles suggests great active erosion as well as served. Down dip and subsurface in the direction of from major drainage systems. Florida and southern Alabama the Mooreville is re- The small clastic portion of the sediment might have been in currents. placed by a series of limestones and glauconites which swept by turbidity may indicate either shallower depths or deep water Only at Moore's farm (Locality 12A) is there any glauconites. good evidence of current action. At this locality there If the present strike of the sediments is close to the is an exposed layer of shell and bone conglomerate less than an acre in extent and less than one foot thick. original shore line, the Mooreville was deposited at the In this due to removal of the finer end of a peninsula which was formed by pre-existing layer, presumably Appalachian structure; the bulk of deposition of clastic particles, there has been a concentration of larger or- material lies to the northwest and northeast of the ganic remains such as Inoceramus prisms, ostracods, Mooreville. In the Mooreville outcrop area there must teleost scales, vertebrae, and teeth, shark denticles and small small and coiled have been a lack of major stream entry, resulting in teeth, Ostrea, gastropods, cepha- minimal clastic sedimentation. lopods. The abundance of vertebrate material makes this layer almost a bone bed. In the Fort Hays, Kansas Ecological Evidence From the Sediments State College collection there are samples of a similar consolidated layer from the Niobrara. The remarkable homogeneity of the Mooreville sedi- If the Mooreville Formation had been in ments lends itself to an attempted environmental re- deposited shallow between one and 300 feet in construction. Any environmental analysis should con- water, i.e., depth, one would a sediment disturbed wave sider inorganic as well as organic evidence. This anal- expect notably by and tidal action. The absence of current ysis is here confined to the Mooreville area of outcrop phenomena at and the of in Alabama. Since this is obviously a marine sediment, (except Locality 12A) great homogeneity other environments are not considered. the Mooreville Chalk are not compatible with a shallow water interpretation. Mooreville Chalk sediments are very similar to other Cretaceous chalks. In total Upper bulk, clay materials, Ecological Evidence From the Plants in the clay size range, make up the greater portion of the chalk. Calcareous particles ranging from less than Plant remains as a whole are rare in the Mooreville one micron to two or three millimeters form the next except for the flood of microscopic "armor" plates, coc- highest percentage. A small percentage of quartz grains coliths and rhabdoliths, from single celled algae be- is present in all the samples. Some chert, similar if longing to the phylum Chrysophyta, family Coccolitho- not identical to that of the Alabama Mississippian for- phoridae. Members of this group of flagellates abound mations, occurs in the larger size ranges. Glauconite today in temperate and tropic open ocean surface wa- size of pellets and mica can be found in any sample of the ters. The small size of the plates and the small Mooreville. There is a suite of heavy minerals all 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 formation but not, however, toward the formation of as a geological viewpoint. The coccoliths are common true reefs. in numerous Jurassic, Cretaceous, and Re- Tertiary, It should be noted that although reef-forming genera cent sediments. The themselves are so num- organisms such as Ostrea and Exogyra are found, no reefs occur erous that well serve as the they may primary synthe- in the Mooreville. The typical near-shore Upper Cre- sizers, at the base of the sea food chain. open taceous molluscan fauna, such as found at Coon Creek, At Bluff Hewlett's Choctaw and farm obscure chal- Tennessee, is lacking or represented by few individuals. of what were copyritized impressions evidently marine Burrowing mollusks are not present; this may be due are but not common. algae present The fragmentary to the suffocating ooze conditions or to the depth of nature of these remains suggests that they may have water, perhaps both. floated into the area of Fossil wood is deposition. Invertebrates were taken from 18 Mooreville lo- found at a number of Mooreville localities but it is calities. At every locality, Foraminifera, worm pellets, rather rare; the fact that it is bored by marine mollusks ostracods, echinoderm remains, Ostrea congesta, Ostrea indicates that it had floated for some length of time. sp. spat, and large Inoceramus shells were collected. No leaves have been found in the Mooreville though Durania austinia was taken from 10 localities. Exogyra the Field Museum collection contains a few specimens ponderosa, Ostrea sp., and encrusting Bryozoa were tentatively identified as willow which could galls, prob- found at four localities. Ostrea cretacea, a small shelled ably have floated with ease. None of the above evi- Inoceramus, the wood boring Teredo, and the barnacle dence immediate to land necessarily suggest proximity Scapellum were collected from three localities. A soli- nor particularly shallow water. tary coral, the worm Serpula sp., worm burrows, the Ecological Evidence From the Invertebrates brachiopod Lingula sp., Ostrea plumosa, 0. bleckensis, 0.

The common invertebrates that can be found in mesentarica, Exogyra uptoiensis, Gryphaea vesicularis, G. vomer, and Pecten were taken at two localities. any sample of Mooreville Chalk are the calcareous and sp. Each of the following is known from a single locality: arenaceous Foraminifera, worm pellets, Inoceramus Cliona sp., the boring sponge, Hamulus major, H. onyx, prisms, and Ostrea spat. At any locality one can expect Nucula sp., Ostrea lava, 0. falcata, 0. spatulata, Pecten. Ostrea congesta and usually another species of Ostrea, cf. simplex, Paranomia scabra, small undetermined gas- the large shells of Inoceramus, and the plates or spines the nautiloid small am- of echinoderms. At a number of localities there were tropods, Eutrephoceras dekayi, monites, a single fragment of a large ammonite, and worm tubes belonging to Hamulus and shells of the several crab claws. pelecypod Durania. Encrusting Bryozoa (Zangerl, An infauna characteristic of modern shallow 1948, PI. 2) were present at four localities. muddy bottoms is generally absent in chalk This fauna seems to be composed of pelagic, neritic, deposits, though other Cretaceous show epifaunal, and infaunal elements. The pelagic forms contemporaneous Upper clays of the and that still are Teredo in driftwood, a number of the Foraminifera, many pelecypod gastropod genera live in bottom sediments. The absence the small ammonites, gastropods, and ostracods. The today muddy of such an infauna in the chalks be due to the genus Pecten may be a neritic element at least part of may depth at the time of In of less than 200 the time, but the small size is suggestive of the deep deposition. depths feet one would to find some evidence of reef water pectens that occur off our coasts today. The expect formation, either Ostrea, radiolites, corals epifaunal elements such as Exogyra, Gryphaea, Para- by Exogyra, or the nature of the sediment could nomia, Inoceramus, and the radiolite Durania, some algae. Though oozy be an factor, the shells that do occur, or Foraminifera, and probably some of the ostracods, are inhibiting even the vertebrate could have acted as a those adapted to soft bottom conditions. Hamulus oc- remains, temporary platform for reef formation. If the curs as encrusting species as well as unattached species. depth were too the lack of and food The encrusting forms in addition to Hamulus include great, however, light would preclude this The vast amount of Ostrea congesta, Bryozoa, the boring sponge Cliona, and possibility. planktonic material in the Mooreville must have been the barnacle Scapellum. Worm burrows and pellets formed in an sea represent the infaunal elements. open environment, possibly beyond the depositional area, and may have been carried into Ostrea congesta occurs commonly on Inoceramus the Mooreville area of deposition by a Gulf Stream-like shells, rarely on vertebrate remains. In both the Nio- current moving at the surface. In my opinion, the brara and the Mooreville Formations the large species evidence of the sediments, plants, and invertebrates of Inoceramus have Ostrea congesta on the outer sides indicates a depth of more than 600 feet or 100 fathoms, of both valves, even when the two valves are found in if it is to be compared with Recent conditions, for it is articulation. This could occur if the huge thin-shelled beyond the 100 fathom line that Recent deep-water Inoceramus had a habit of keeping its outer edge up conditions begin to appear. and hinge down in the ooze. The Ecological Evidence From the Fishes The large conical Durania may have existed par- tially buried in or on top of the bottom ooze. Some Conditions of Preservation individuals of Durania have other Durania shells at- The Mooreville fish remains, with the exception of tached to them, showing a tendency toward bioherm those from the Moore farm Locality 12A, consist of APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 427 isolated and generally largely disarticulated fish evi- other things, a great many mixed shark teeth, verte- dently deposited at random in the sediment. Only brae, and denticles.) The fact that the denticles adhere one nearly complete articulated specimen has been col- to the vertebrae, a phenomenon common in the Nio- lected. The remains generally consist of scattered brara specimens at the University of Kansas as well as patches of scales, vertebrae, and skull elements, all of in the Mooreville specimens, speaks for fairly rapid which show some degree of articulation. Sharks may burial by sinking into the bottom ooze, then decay in have teeth, denticles, and vertebrae associated. In- situ, with a lack of strong bottom currents. dividual scales or teeth often occur alone. The scatter- The Ptychodontidae are a predominantly Upper ing of remains is most likely due to (1) predation in Cretaceous group of sharks, known only by isolated fall to life, leaving only uneaten parts of fishes to the teeth, tooth sets, and doubtful vertebrae. The wide bottom, (2) decay of dead fishes while still floating, or geographic range of Ptychodus speaks for a broad oceanic (3) disturbance by scavengers before burial. distribution. While nothing is known of the feeding In the following sections each of the groups of fishes habits of this group, the battery of row upon row of in the Mooreville is discussed as to its ecology as indi- large teeth (over 600 in P. mortoni according to Wood- cated by the modern and Tertiary counterparts. ward, 1902-1912) would have made this shark an excellent invertebrate feeder, perhaps living on Crustacea Chimaeroids and pelagic mollusks. The to The Edaphodontidae are extinct. The known depth Anacoracidae have been thought show a distribution of the six modern genera of the order Chi- reduction in dentition leading toward the Cetorhinidae. Most recent evidence the that this maeriformes is given below (in fathoms) : supports theory family may represent a direct ancestor of the primitive Chimaera 1 to 600 orectolobids and is not related to Cetorhinus at all. Hydrolagus 1 to 1290 Its teeth were well adapted for fish eating, and the well Callorhynchus 1 to 100 Rhinochimaera 200 calcified vertebral column suggests that it was a power- Harriotta 375 to 1422 ful swimmer. The wide distribution indicates that this Neoharriotta 300 plus may have been a pelagic shark. The are in the Moore- The first three genera have a wide depth range, Odontaspididae represented ville the The size of though they have been found most commonly in shallow by genus Scapanorhynchus. large the teeth indicates a shark of about ten feet in water well up on the continental shelf. The last three length. Modern odontaspids occur in both coastal and deep genera are specialized deep water forms. As has been water; at least one is known to be truly pelagic. The stated earlier, the Edaphodontidae are particularly close wide distribution of is in keeping with to the Recent genus Callorhynchus. Graham (1956) Scapanorhynchus a habit. has discussed the natural history of this form in New pelagic Zealand. There are five recognized species, which are Probably the most common shark tooth in the Moore- circumpolar and confined to the Southern Hemisphere. ville is that of Lamna appendiculata. The Recent spe- During the warm portion of the year, the New Zealand cies of Lamna are confined to temperate and boreal Off southern species moves into the bays and even ascends rivers; regions (Bigelow and Schroeder, 1948). during the cold season it moves out into deeper water. California this species is taken in deep water, but occurs Graham (1956) states that this species needs highly near the surface in the northern Pacific, suggesting that oxygenated water. Callorhynchus feeds on mollusks, in warmer waters it lives at greater depth. This genus is considered to be a shark. fish, arthropods, crabs, shrimp, and jellyfish (Graham, today truly pelagic 1939). One may conclude from the size of the dental Isurus is known today by two species (Garrick, plates in Edaphodon that the Mooreville forms reached 1966). Both are open ocean sharks and reach lengths gigantic proportions as compared with the Recent Cal- of about 12 feet (Bigelow and Schroeder, 1948; Apple- lorhynchus. gate, 1966). The Cretaceous specimens from the Nio- Large chimaeroids today are characteristic of deep brara Chalk had a length of at least 22 feet. The Isurus feeds on fish the Cretaceous forms water, while shallow or coastal water forms seldom ex- modern ; large could fed also on marine ceed two feet, as in Callorhynchus. In the Recent fauna have reptiles. four and five foot long deep-water specimens of Hy- None of the sharks occurring in the Mooreville be- drolagus and Chimaera are known. The Mooreville long to groups that could not be pelagic. So far there edaphodontids were probably much larger and may have been found none of the small sharks that normally have reached a length of over eight feet, a size favoring occur in shallow water such as Squalus and the orecto- a deep water habit. lobids, which do occur in other Cretaceous sediments. Of note is the lack of rays of any sort, a group which Sharks is known to exist at a much earlier period, but again The Selachii in the Mooreville are represented by is more characteristic of shallow water except for the isolated individuals and are thinly distributed through- family Rajidae and several small-toothed forms which out the formation. (The one known exception to this either had not evolved by Cretaceous times or must is Locality 12A. where a consolidated layer shows, among have left few remains. 428 FIELDIANA: GEOLOGY MEMOIRS, VOLUME 3

Sturgeons and Holosteans Pachyrhizodontidae are extinct, which forces one to The sturgeon, Propenser, undoubtedly represents a resort to structural adaptations to elucidate their prob- marine form. Sea sturgeons today are wide-ranging able habits. The chalk species show a proportionally and are taken in the open sea. They have been taken large sickle-shaped tail. The pectoral fins are large and in Scandinavian waters at a depth of 25 fathoms, and powerful with a large stout first pectoral spine. These there is no reason to believe they would not be found characters are adaptations of a powerful swimmer. The in deeper water. large incurved teeth indicate a carnivorous diet. The Scombridae would to be the modern The pycnodont genus Hadrodus is known from both appear ecological of this the Niobrara and the Mooreville, as well as from the counterpart family. The Pachyrhizodontidae, like the similar must have been Upper Cretaceous of Mississippi. This huge fish had Scombridae, open sea fishes with worldwide distribution. crushing jaws and perhaps fed on invertebrates. If it pelagic nearly was similar to the other pycnodonts it might have been The Bananogmiidae are another extinct group. In deep-bodied and could have had a habitat similar to fin structure and shape they are similar to the Pachy- that of the present-day deep-bodied moonfish, Lampris, rhizodontidae; however, the unique crushing dentition which is found in the open ocean. is closest to that of the Albulidae. In the European Cretaceous this family is absent and have been The pachycormid Protosphyraena, with its exceed- may ecologically replaced by the closely related and similar ingly long bill and partly fused pectoral fin, can be family, the Plethodidae. There is notable compared adaptively with the billfishes or istiophorids. uncertainty as to the and habits of these of The large fang-like teeth indicate a carnivorous diet. morphology groups fishes. The crushing dentition and abil- The long pectoral fin would perhaps stabilize the fish good swimming ity could be correlated with feeding on and for the thrashing used in food-getting, as well as serving shrimp other open sea arthropods as well as on nautiloids and as a weapon of offense or defense. The caudal hypurals ammonites. are coalesced into a single element, a feature occurring The fossil are close to the in the living istiophorids and associated with powerful Ichthyodectidae very modern Chirocentridae. The latter are swimming. Protosphyraena is known from the English large powerful- fishes that have a wide chalk, New Jersey, Kansas, and California. The wide swimming predaceous ecological are found in both coastal and distribution and the modern analogs are at least con- range; they open ocean waters. The were even fish sistent with the interpretation that this was an oceanic Ichthyodectidae larger than the with or pelagic species. Pachyrhizodontidae Xiphactinus being perhaps the largest bony fish that ever lived. The sturgeons and holosteans are to be considered The Saurodontidae resemble the in as survivors of more archaic groups. Protosphyraena Ichthyodectidae general structure but possess a predentary which seems to be the most specialized. Only the sturgeon may have served in a manner similar to the rostrum might be considered indicative of a shallow water en- elongate of Protosphyraena. The thin blade-like teeth indicate vironment, yet this may be due to an incomplete knowl- a predaceous habit. edge of the living species, which probably at times go One of the of Cretaceous chalk very deep or swim over deep areas. Recent tagging interesting aspects faunas is the almost absence of the studies show that the modern sturgeon can move great complete Clupeidae. distances. So far, only one preoperculum in the Mooreville can be referred to this family. Clupeids are also absent in Teleosts the British chalk and the Niobrara. They have, however, a good Cretaceous record and they dominate the The Elopidae are represented today by the single Upper Cretaceous of California and Syria. Their ab- genus Elops, which has a worldwide tropic and south sence in the chalks is one of the most striking examples temperate distribution. Elops is common in the open of the great differences in modern as well as fossil fish sea along the coast of the southeastern United States. faunas in different ecological situations. It also frequently enters brackish water, being taken The Enchodontidae are not only a common Creta- from the mouths of rivers and from bays. Gill (1907) ceous family but are probably the second most common records the presence of shrimp in Elops stomachs, and group of the Mooreville fishes. The predatory encho- Darnell (1958) reports a predominance of small fish dontids 1 seem to be closely related to the present day and penaeid shrimps. The Recent species appears to Omosudidae and Alepisauridae, which are deep water be wide-ranging. This appears to be true also of the Iniomi. Enchodus is known from Cretaceous sediments fossil Cretaceous genera assigned to this family. of undoubted shallow water. Cimolichthys is a large The related family Albulidae is represented today powerful fish which could certainly be an oceanic form. by the genera Albula and Dixonia. According to Hilde- The Dercetidae are represented in the Mooreville brand (1963), the extent to which Albula enters deep by Stratodus apicalis which is a fairly small predaceous offshore water is unknown. Bonefish do occur in the fish; the great number of little sharp teeth on the pala- shallows, feeding in the mud. The geographic distri- tines and jaws would appear to be very effective for bution of the modern species is similar to that of Elops; holding its prey. This species is reported from phos- it is found in all warm seas. Frizell (1965) believes that the Eocene species was abundant in deep water. 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

0-50 F 50-100 F 100+ F GENUS SHALLOW MODERATE DEEP MODERN ANALOGS DISCUSSED IN TEXT

Edaphodon Large deep sea chimaeroids, several genera.

Ptychodus None; modern heterodonts occur in shallow to deep water.

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

Lamna ? x x Pelagic over moderate depths and in deep water.

Isurus T x x Pelagic over moderate depths and in deep water. Squalicorax XXX None; probably pelagic. Pseudocorax XXX None; probably pelagic. X ? Propenser X Coastal ; could occur over deep water.

Hadrodus ? X X None; probably pelagic over deep water.

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

Palehps X X Modern Elops mostly in shallow to moderate depths.

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

Albula X X Modern Albula in shallow to moderate depths.

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

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

Ichthyodeetes None; modern Chirocenlrus shallow water and pelagic species.

Xiphactinus None; modern Chirocenlrus shallow water and pelagic species.

Saurodon None; modern Chirocenlrus shallow water and pelagic species.

Saurocephalus None; modern Chirocenlrus shallow water and pelagic species.

Enchodus x X None; modern related genera deep water.

Cimoliehthys x X None; modern related genera deep water. Stratodus X X None; however, could perhaps be in deep water.

Myctophidae X X Modern genera in fairly deep water.

Trachichthyidae X Modern genera in deep water. TOTAL— 14 22 21

phate beds in North Africa, which may indicate a shal- the deep sea beryeoids, and says that it ranges from 50 low water habit but again the wide distribution might to 500 fathoms in depth. also indicate a existence. pelagic If one is willing to assume that at least most of the The Myctophidae or lantern fishes represent the common genera of Mooreville fishes have been pre- most common Mooreville fish group. Small patches served and are represented in the present collection, of myctophid scales occur at most of the localities in then we may make some inferences concerning the pos- vast quantities. The comparatively fragile vertebrae sible ecological relationships among these forms. Rank- and head plates are also abundant. The modern mem- ing the families or genera in order of abundance of bers of this family are deep water plankton feeders. individuals we have as follows: abundant, the Mycto- They move up at night and down during the day in phidae, Enchodus. Squalicorax, Scapanorhynchus, Lamna. deep oceanic waters, probably following the periodic Paehyrhizodus. Xiphactinus; common. Saurodon, Sauro- movements of zooplankton; some species commonly cephalus. Bananogmius; rare. Stratodus, Protosphyraena. come to the surface at night, where they are taken in Edaphodon. Palehps. Albula. Cimoliehthys. Isurus; very great numbers. rare. Pseudocorax, Ichthyodectes. and Clupeidae. The last two are known one or two individuals. Goode and Bean (1895) place the family Trachich- by only thyidae among the deep water beryeoids, with Traeh- Feeding habits may be inferred from the stomach ichthys being taken from depths of from 200 to 400 contents of the fossils (none among the Mooreville spec- fathoms. Smith (1961) likewise places this family with 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- In conclusion, the Mooreville fish fauna is a mixed era with crushing teeth are Edaphodon, Ptychodus, Had- one consisting of undoubted pelagic open sea forms rodus, and Albula. Any or all of these could be con- along with some possible near shore and perhaps shallow sidered mollusk feeders, as is the Recent Albula. Those water fishes, but showing a dominance of deep water forms having pointed teeth suitable for biting and sev- fishes. The probable currents maintained the open ering are Scapanorhyrichus, Isurus, Lamna, Squalicorax, water surface feeders, and the close proximity to shore Pseudocorax, Protosphyraena, Pachyrhizodus, Xiphac- (within a few miles) or near shore environments con- tinus, Ichthyodectes, Saurodon, Saurocephalus, Cimolich- tributed the near shore fishes in this assemblage. The thys, and Enchodus. Those fishes having numerous intermixture of these two faunas with an emphasis on pointed teeth capable of holding the prey and assisting the open sea fishes seems characteristic not only of the in the swallowing process are Moorevillia, Stratodus, Mooreville but of Cretaceous chalks in general. have and Hoplopteryx. The Myctophidae small teeth At the present time many Cretaceous chalk faunas suitable for on The Pro- feeding plankton. sturgeon, are still poorly known but all indications point to the penser, lacked teeth altogether; Acipenser, its modem fact that these chalk faunas throughout the world have counterpart, is a detritus feeder. assemblages closer to each other than to nearby for- Ranking these fish by size, those with a general size mations of equivalent age but different sediment types. in the adults of from six to fifteen feet are Pro- range In general, chalks show a fauna consisting of a large penser, Xiphactinus, Pachyrhizodus caninus, Isurus, number of huge predaceous teleosts such as Xiphac- and Lamna. Scapanorhyrichus, Squalicorax, Ichthyodectes, tinus, Ichthyodectes, and Pachyrhizodus. Sharks are pres- Fish with a range of around three to six feet are Pachy- ent but only moderately abundant. The Clupeidae are rhizodus kingi, at least some species of Bananogmius, rare or absent. Myctophidae are common. and the lowermost limit of the Cimolichthys, (at range) In number of similar species the Mooreville Forma- Fishes one to Pachyrhizodus minimus. ranging from tion's fish fauna is closest to that of the Niobrara Chalk, feet Enchodus. Less than one three are Stratodus and which has been studied mainly by Williston (1900), foot are Hoplopteryx, the Clupeidae, and the Mycto- Stewart (1900), Loomis (1900), and Hay (1903). In phidae. number of similar genera the Mooreville's fish fauna is In the Mooreville, as stated, the myctophids occur closest to that of the British Chalk, the best known as patches of scales with vertebrae and disassociated Upper Cretaceous fauna due to the long history of col- plates. These patches may represent stomach ejects lecting culminating in the excellent monographic treat- of from other fish, particularly sharks. Myctophid scales ment Woodward (1902-1912). are also very common in the Mooreville coprolites. The With the exception of the Niobrara, Upper Creta- great abundance of this family makes it a likely can- ceous chalks either intertongue with or are underlain didate to form the base of the Mooreville fish food by greensands. The common occurrence of Ostrea and chain. The Clupeidae and the berycoids play a minor Exogyra reefs in the greensands suggests shallower role. All four groups may be considered as the basal water. The greensand fish faunas show, to use the New grade, which may be called Grade 4; they fed mostly Jersey fauna as an example, abundant sharks, rays, and on invertebrates could have been the of are well and prey Grade Chimaeridae. Teleosts present but not known ; 3, including Cimolichthys, Enchodus, and Stratodus. the large chalk species seem to be absent. Of the 41 Grade 2 consists of Ichthyodectes, Pachyrhizodus kingi, species of New Jersey fish, seven are similar to Moore- Squalicorax, Scapanorhynchus, Bananogmius, and Pachy- ville species and four of the seven are wide-ranging rhizodus minimus. The top of this pyramid, Grade 1, sharks. be the would largest carnivores, Pachyrhizodus caninus, The littoral zone in the Upper Cretaceous is repre- Xiphactinus, and the sharks, Isurus and Lamna. sented by the North African phosphate beds and by a Below this fish food chain there is a more fundamen- poorly known South Dakota fish fauna mentioned by tal chain with the base consisting of phytoplankton. Zangerl and Sloan (1960). These beds differ from the The phytoplankton is represented in the sediment by Mooreville in the great concentration of vertebrate ma- Chrysophyta, which might have been fed on by a di- terial in the form of disassociated teeth, vertebrae, head verse zooplankton consisting in part of Foraminifera, plates, and, in South Dakota, of scales, to form a bone- ostracods, and small gastropods. There may have been bed conglomerate. The phosphate beds are known to a vast number of forms at this level in the food chain cover extensive areas, and in contrast to the chalks have which either did not fossilize or are yet to be collected. a good pre- and post-Cretaceous history. There is no This zooplankton was fed upon by the myctophids similar Recent deposit, but the Pliocene and Pleistocene which in turn were fed upon by the other three grades of phosphate beds of Florida and the "bone beds" of the fishes plus other higher vertebrates such as turtles, mosa- Miocene Calvert in Virginia and Temblor of California saurs, and plesiosaurs. Aside from this chain there are the same general type of deposit and evidently were were some fish with crushing dentition which evidently formed in very shallow water in or near the intertidal fed directly on the molluscan fauna or the larger zoo- zone. The Upper Cretaceous North African phosphate plankton. 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. of chalks leaves much to be desired. The evidently The North African phosphate fauna is predominantly oozy bottom has resulted in a restricted bottom fauna, a shark fauna, with sharks comprising 75 per cent of all a fact that complicates any interpretation as to depth. vertebrate remains (Arambourg, 1952). The sharks in One is however faced with two possible models which the Mooreville are well under 25 per cent. The pre- might represent the condition found in the Mooreville: dominance of shark teeth is also true of other phosphate a shallow water ooze around 3 to 30 feet deep, or a beds mentioned above. The North African phosphate deep water ooze over 100 fathoms. Certainly as far beds lack ptychodontids, although other rays are abun- as the present evidence is considered, the Mooreville dant, as are sawfish; the Mooreville, however, contains Chalk seems to fit the second model and was probably only ptychodontids and no evidence of rays or sawfish. deposited in water over 100 fathoms or 600 feet deep. At the same time, the terrestrial remains which have SUMMARY been found in the chalk, a bird and several dinosaurs, suggest a nearby shore. This seems probable, as the The Mooreville formation contains a rich fossil fish Mooreville beds lie at the southern end of the Appala- fauna consisting of at least 17 families, 28 genera, and chian belt which probably formed a headland with few 42 species. Future field work will undoubtedly add to if any large streams at the southern end during Creta- this assemblage. Our present knowledge of the ecology ceous times. REFERENCES

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432 APPLEGATE: VERTEBRATE FAUNA OF SELMA FORMATION 433

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