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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 


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FIELDIANA:  GEOLOGY  MEMOIRS 

VOLUME  3,  NUMBERS  7  AND  8 

Published  by 

FIELD  MUSEUM  OF  NATURAL  HISTORY 

FEBRUARY  12,  1970 


Geo*. 


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Lib, 


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THE  VERTEBRATE  FAUNA  OF  THE 
SELMA  FORMATION  OF  ALABAMA 

PART  VIII.    THE  FISHES 


THE  VERTEBRATE  FAUNA  OF  THE 
SELMA  FORMATION  OF  ALABAMA 

PART  VIII 
THE  FISHES 

SHELTON  P.  APPLECxATE 

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


FIELDIANA:  GEOLOGY  MEMOIRS 

VOLUME  3,  NUMBER  8 

Published  by 

FIELD  MUSEUM  OF  NATURAL  HISTORY 

FEBRUARY  12,  1970 


Library  of  Congress  Catalog  Card  Number:  53-2305 

PRINTED    IN   THE   UNITED   STATES   OK   AMERICA 
BY   FIELD   MUSEUM    PRESS 


CONTENTS 

PAGE 

Introduction 389 

Systematic  Descriptions     389 

Holocephali      389 

Edaphodontidae 389 

Edaphodon  barberi,  new  species 390 

Edaphodon  mirificus  Leidy 392 

Edaphodon  sp 393 

Selachii 393 

Ptychodontidae 393 

Ptychodus  polygurus  Agassiz 393 

Ptychodus  mortoni  Mantell 393 

Anacoracidae 393 

Squalicorax  falcatus  (Agassiz) 393 

Squalicorax  pristodontus  (Agassiz) 395 

Pseudocorax  affinis  (Agassiz) 395 

Odontaspididae 395 

Scapanorhynchus  rhaphiodon  (Agassiz) 395 

Scapanorhynchus  rapax  (Quaas) 396 

Lamnidae 396 

Lamna  appendiculata  Agassiz 396 

Isurus  mantelli  (Agassiz) 397 

Undetermined  shark  denticles 398 

Actinopterygii 398 

Chondrostei 398 

Acipenseridae 398 

Propenserinae,  new  subfamily 398 

Propenser,  new  genus 398 

Propenser  hewletti,  new  species 399 

Holostei 401 

Pycnodontidae 401 

Hadrodus  priscus  Leidy 401 

Incertae  sedis 401 

Pachycormidae      403 

Protosphyraena  Leidy 404 

Protosphyraena  nitida  ?  (Cope) 404 

Protosphyraena  sp.  1 404 

Protosphyraena  sp.  2 404 

Protosphyraena  sp.  3 404 

Teleostei 404 

Elopoidei 405 

Elopoidea 405 

Elopidae 405 

Palelops,  new  genus      405 

Palelops  eutawensis,  new  species 406 

Pachyrhizodontoidea,  new  superfamily 406 

Pachyrhizodus  Dixon 406 

Pachyrhizodus  minimus  Stewart 408 

Pachyrhizodus  caninus  Cope 410 

Pachyrhizodus  kingi  Cope      411 

385 


386  CONTENTS 

PAGE 

Albuloidea 411 

Albulidae 412 

Albula  dunklei,  new  species 412 

Albula  sp 413 

Plethodoidea,  new  superfamily 413 

Bananogmiidae,  new  family 413 

Bananogmius  crieleyi,  new  species 414 

Bananogmius  cf.  zitteli  (Loomis) 416 

Bananogmius  cf.  polymicrodus  (Stewart) 416 

Bananogmius  sp 416 

Moorevillia,  new  genus 416 

Moorevillia  hardi,  new  species 416 

Clupeoidei 416 

Chirocentroidea,  new  superfamily 416 

Ichthyodectidae 418 

Ichthyodectes  cf.  ctenodon  Cope 418 

Xiphactinus  audax  Leidy 418 

Saurodontidae 419 

Saurodon  leanus  Hays 419 

Saurodon  ?  sp.  1      420 

Saurodon  1  sp.  2      420 

Saurocephalus  cf.  lanciformis  Harlan 420 

Clupeoidea 420 

Clupeidae  indet 420 

Dussumieriinae  indet 420 

Myctophoidei 420 

Enchodontidae 420 

Cimolichthys  nepaholica  (Cope) 420 

Enchodus  petrosus  Cope 421 

Enchodus  cf.  saevus  Hay 421 

Dercetidae 421 

Stratodus  apicalis  Cope 421 

Myctophidae      421 

Myctophidae  indet 421 

Beryciformes 423 

Trachichthyidae 423 

Hoplopteryx  ?  sp 423 

Trachichthyidae  ?  indet 424 

Teleost  incertae  sedis  1      424 

Teleost  incertae  sedis  2      424 

Ecology  of  the  Mooreville  Chalk     424 

The  evidence  from  the  overall  geological  picture 424 

Ecological  evidence  from  the  sediments 425 

Ecological  evidence  from  the  plants 425 

Ecological  evidence  from  the  invertebrates 426 

Ecological  evidence  from  the  fishes 426 

Summary 431 

References 432 


LIST  OF  ILLUSTRATIONS 

PAGE 

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

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

mandible      392 

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

177.  Tooth  of  Pseudocorax  affinis 395 

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

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

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

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

of  Lamma  appendiculata 397 

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

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

of  posterior  portion  of  skull 399 

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

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

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

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

and  neural  arches 403 

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

187.  Premaxillary  of  possible  pycnodontid 404 

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

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

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

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

192.  Reconstruction  of  Pachyrhizodus  minimus 410 

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

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

other  parts      412 

195.  Tooth-bearing  plate  of  Albula  sp 413 

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

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

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

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

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

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

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

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

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

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

202.  Scales:  teleost  indet,  Hoploptery±  ? 422 

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

204.  Operculum  referred  to  family  Trachichthyidae  ? 424 

387 


THE   FISHES 


INTRODUCTION 

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

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

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

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

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


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

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

SYSTEMATIC  DESCRIPTIONS 

Class  HOLOCEPHALI 

Order  CHIMAERIFORMES 

Suborder  CHIMAEROIDEI 

Edaphodontidae  Owen 

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

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

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


389 


390 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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

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

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

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

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


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

Edaphodon  barberi,  new  species.    Figure  174 

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

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

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

MEASUREMENTS— (in  mm.). 

Greatest  length  of  mandible 69 

Greatest  thickness  of  mandible 19 

Length  of  apical  tritor 21 

Width  of  apical  tritor 4.5 

Width  of  antero-intermediate  tritor 3.5 

Width  of  postero-intermediate  tritor 5.5 

Width  of  postero-dorsal  tritor 3.0 

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

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

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


10  mm 


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


391 


392 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


B 


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


Edaphodon  mirificus  Leidy.    Figure  175A 

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


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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


393 


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


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


Edaphodon  sp.    Figure  175B 

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

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

Class  SELACHII 

Order  HETERODONTIFORMES 

Suborder  HYBODONTOIDEI 

Ptychodontidae  Hay 

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

Ptychodus  polygyrus  Agassiz.  Figure  179A 

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

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

Ptychodus  mortoni  Mantell. 

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


Order  GALEIFORMES 
Anacoracidae  Gliickman 

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

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

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


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


394 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


B 


H 


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


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

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


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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


395 


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


I  m  m 


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

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

Squalicorax  pristodontus  (Agassiz) 

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

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

Pseudocorax  affinis  (Agassiz).    Figures  177,  178  M 

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


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

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

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

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

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

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

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


396 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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


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


81 


M 


N 


~M 


20  mm 

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


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

Scapanorhynchus  rapax  (Quaas).    Figure  178  D. 

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

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


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

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

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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


397 


B 


10  mm 

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


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

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


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

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

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

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


398 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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

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

Undetermined  shark  denticles.    Figure  176  G-N. 

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

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

Subclass  ACTINOPTERYGII 

Infraclass  CHONDROSTEI 
Order  ACIPENSERIFORMES 

Acipenseridae  Bonaparte 

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

Propenserinae,  new  subfamily 

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


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

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

Propenser,  new  genus 

Diagnosis. — As  for  subfamily. 

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

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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


399 


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


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

Propenser  hewletti,  new  species.    Figures  180-185. 

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

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

Diagnosis. — Same  as  for  genus. 
I 


20mm 


20  m  m 


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


400 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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


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

The  skull,  exclusive  of  the  rostrals,  is  simpler  than 


mm 


20    mm 


20    mm 


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


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

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


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

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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


401 


20mm 


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


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

Infraclass  HOLOSTEI 
Order  PYCNODONTIFORMES 

Pycnodontidae  Cope 

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


Hadrodus  priscus  Leidy.    Figure  1£6. 

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

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

Incertae  sedis.    Figure  187. 

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


20mm 


i  i 

20  mm 


i  i 

20mm 


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


402 


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


403 


.tSJjt*' 


$0 


20   mm 


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


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


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

Order  AMIIFORMES 

Suborder  AMIOIDEI 

Pachycormidae  Woodward 


404 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


20  mm 


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

Protosphyraena  Leidy 

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

Protosphyraena  nitida?  (Cope) 

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

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


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

Protosphyraena  sp.  (1) 

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

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

Protosphyraena  sp.  (2) 

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

Protosphyraena  sp.  (3) 

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

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

Infraclass  TELEOSTEI 

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


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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


405 


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

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

Suborder  ELOPOIDEI 

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

Superfamily  ELOPOIDEA 

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


I  mm 


I  mm 


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


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

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

Elopidae  Bonaparte 

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

Palelops,  new  genus 

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


406 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


Palelops  eutawensis,  new  species.    Figure  188. 

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

Referred  specimens. — PF  3612,  two  scales;  l4  mile 
E.  of  Walter  Dance's  farm,  6.2  miles  W.  of  Aliceville, 
Eutaw  County,  Alabama,  lower  Mooreville.  PF  3563, 
one  scale;  Choctaw  Bluff,  Greene  County,  Alabama, 
lower  Mooreville.  PF  3560,  four  scales;  Montgomery's 
farm,  2  miles  N.  of  West  Greene,  Greene  County,  Ala- 
bama, middle  Mooreville.  PF  3561,  one  scale;  Hew- 
lett's farm,  gullies  near  the  road,  Greene  County,  Ala- 
bama, upper  Mooreville. 

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

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

Superfamily  PACHYRHIZODONTOIDEA, 

new  superfamily 

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

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


Pachyrhizodus  Dixon 

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

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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


407 


20mm 


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


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

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


contain  strong  circuli. 

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


408 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


10  mm 


10  mm 


10  mm 


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


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

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

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


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

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


I  mm 


mmjf^fe  -Hl 


V 


\ 


,W\ 


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


I    mm 


409 


410 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


Fig.  192.    Reconstruction  of  Pachyrhizodus  minimus. 


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

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

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

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


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

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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


411 


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


20mm 


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


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

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

Pachyrhizodus  kingi  Cope.   Figure  190  D. 

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

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


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

Superfamily  ALBULOIDEA  Hay 

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


412 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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

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

Albulidae  Gunther 

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


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

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

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

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


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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


413 


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

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

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

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

Albula  sp.    Figure  195. 

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

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


;. 


10  mm 

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

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

Superfamily  PLETHODOIDEA,  new  superfamily 

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

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

Bananogmiidae,  new  family 

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

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


414 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


•n&2K 


■  n 


*& 


D  E 


20  mm 

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


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

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


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

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

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


20  mm 


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


415 


416 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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

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

Bananogmius  cf.  zitteli  Loomis.    Figure  198  H. 

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

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

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

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

Bananogmius  sp.    Figure  198  G. 

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

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

Moorevillia,  new  genus 

Type  species. — Moorevillia  hardi,  new  species. 

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


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

Moorevillia  hardi,  new  species.    Figure  197. 

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

Diagnosis. — Same  as  that  of  the  genus. 

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

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

Suborder  CLUPEOIDEI 

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

Superfamily  CHIROCENTROIDEA, 
new  superfamily 

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

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


^ 


?fXSl 


^efc* 


G.-.%-V.    *-T 


20  mm 


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


417 


418 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


Ichthyodectidae  Crook 

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

Ichthyodectes  cf.  ctenodon  Cope 

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

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

Xiphactinus  audax  Leidy.    Figure  200  A. 

Referred  specimens. — PF  3543,  vertebrae,  fragmen- 
tary skull;  2  miles  N.  of  West  Greene,  Hale's  farm, 
Locality  4,  Greene  County,  Alabama,  middle  Moore- 
ville. PF  120,  vertebrae;  PF  125,  lower  jaw  fragments; 
PF  129,  lower  jaw  fragments;  all  from  1  12  miles  W. 


I  Omm 


20  mm 


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


5  mm 


10mm 


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


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

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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


419 


2  mm 


I  m  m 


2  mm 


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


Saurodontidae  Stewart 

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

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

Saurodon  leanus  Hays.    Figure  200  E. 

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


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


420 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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

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

Saurodon?  sp.  1.    Figure  199  A. 

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

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

Saurodon?  sp.  2 

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

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

Saurocephalus  cf.  lanciformis  Harlan 

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

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

Superfamily  CLUPEOIDEA 

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

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

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


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

Dussumieriinae,  indet.    Figure  201  A. 

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

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

Order  INIOMI 
Suborder  MYCTOPHOIDEI 

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

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

Enchodontidae  Loomis 

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

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

Cimolichthys  nepaholica  (Cope) 

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

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


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421 


Enchodus  petrosus  Cope 

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

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

Enchodus  cf.  saevus  Hay 

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

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

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


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

One  genus  occurs  in  the  Mooreville:  Stratodus. 

Stratodus  apicalis  Cope.    Figure  198  A-F. 

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

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

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

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

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

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


422 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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


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


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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


423 


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


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

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

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

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

All  these  scales  are  very  close  to  what  is  found  in 


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

Order  BERYCIFORMES 

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

Trachichthyidae  Bleeker 

Hoplopteryx?  sp.    Figure  202  CD. 

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

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


424 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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


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

Trachichtyidae?  indet.    Figure  204. 

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

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

Teleost  incertae  sedis  (1).    Figure  202  B. 

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

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


Teleost  incertae  sedis  (2) .    Figure  202  A. 

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

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

ECOLOGY  OF  THE  MOOREVILLE  CHALK 

The  Evidence  From  the  Overall 

Geological  Picture 

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

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

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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


425 


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

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

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

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

Ecological  Evidence  From  the  Sediments 

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

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


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

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

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

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

Ecological  Evidence  From  the  Plants 

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


426 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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

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

Ecological  Evidence  From  the  Invertebrates 

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

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

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

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


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

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

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

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

The  Ecological  Evidence  From  the  Fishes 
Conditions  of  Preservation 

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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


427 


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

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

Chimaeroids 

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


Chimaera 

Hydrolagus 

Callorhynchus 

Rhinochimaera 

Harriotta 

Neoharriotta 


1  to  600 

1  to  1290 

1  to  100 

200 

375  to  1422 

300  plus 


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

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

Sharks 

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


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

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

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

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

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

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

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


428 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


Sturgeons  and  Holosteans 

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

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

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

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

Teleosts 

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

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


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

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

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

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

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

The  Enchodontidae  are  not  only  a  common  Creta- 
ceous family  but  are  probably  the  second  most  common 
group  of  the  Mooreville  fishes.  The  predatory  encho- 
dontids1  seem  to  be  closely  related  to  the  present  day 
Omosudidae  and  Alepisauridae,  which  are  deep  water 
Iniomi.  Enchodus  is  known  from  Cretaceous  sediments 
of  undoubted  shallow  water.  Cimolichthys  is  a  large 
powerful  fish  which  could  certainly  be  an  oceanic  form. 

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

1  Specimens  preserved  with  prey  within  their  bodies  are  known. 


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


429 


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


GENUS 

Edaphodon 
Ptychodus 

Scapanorhynchus 

Lamna 

Isurus 

Squalicorax 

Pseudocorax 

Propenser 

Hadrodus 

Protosphyraena 

Palehps 
Paehyrhizodus 

Albula 
Bananogmius 

Moorerillia 

Ichthyodeetes 

Xiphactinus 

Saurodon 

Saurocephalus 

Enchodus 

Cimoliehthys 

Stratodus 

Myctophidae 

Trachichthyidae 

TOTAL— 


0-50  F 
SHALLOW 


50-100  F 
MODERATE 


14 


x 
x 

X 
X 

22 


100+  F 
DEEP 


?  x  x 

T  x  x 

XXX 
XXX 
X  X  ? 

?  X  X 

?  X  X 

X  X 

X  X 

X  X 

X  X 


X 
X 
X 
X 
X 

21 


MODERN  ANALOGS  DISCUSSED  IN  TEXT 

Large  deep  sea  chimaeroids,  several  genera. 

None;  modern  heterodonts  occur  in  shallow  to  deep 
water. 

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

Pelagic  over  moderate  depths  and  in  deep  water. 

Pelagic  over  moderate  depths  and  in  deep  water. 

None;  probably  pelagic. 

None;  probably  pelagic. 

Coastal ;  could  occur  over  deep  water. 

None;  probably  pelagic  over  deep  water. 

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

Modern  Elops  mostly  in  shallow  to  moderate  depths. 

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

Modern  Albula  in  shallow  to  moderate  depths. 

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

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

None;  modern  Chirocenlrus  shallow  water  and  pelagic 
species. 

None;  modern  Chirocenlrus  shallow  water  and  pelagic 
species. 

None;  modern  Chirocenlrus  shallow  water  and  pelagic 
species. 

None;  modern  Chirocenlrus  shallow  water  and  pelagic 
species. 

None;  modern  related  genera  deep  water. 

None;  modern  related  genera  deep  water. 

None;   however,  could   perhaps  be  in   deep  water. 

Modern  genera  in  fairly  deep  water. 

Modern  genera  in  deep  water. 


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

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

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


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

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

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


430 


FIELDIANA:  GEOLOGY  MEMOIRS,  VOLUME  3 


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

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

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

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


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

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

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

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

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

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


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


431 


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

SUMMARY 

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


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


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Applegate,  S.  P. 

1966.  A  possible  record-sized  bonito  shark,  Isurus  oxyrinchus 
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Arambourg,  C. 

1952.  Les  vertebres  fossiles  des  gisements  de  phosphates  (Maroc 
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1958.  Food  habits  of  fishes  and  larger  invertebrates  of  Lake 
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David,  L. 

1946.  Upper  Cretaceous  fish  remains  from  the  western  border  of 
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DUNKLE,  D.  H. 

1940.  The  cranial  osteology  of  Notelops  brama  (Agassiz),  an 
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Fowler,  H.  W. 

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432 


APPLEGATE:  VERTEBRATE  FAUNA  OF  SELMA  FORMATION 


433 


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