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S, 


BULLETIN  OF 

THE  BRITISH  MUSEUM 

(NATURAL  HISTORY) 


GEOLOGY 

VOL.  XVII 

1968-1969 


TRUSTEES  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

LONDON:  1970 


DATES    OF    PUBLICATION    OF    THE    PARTS 

No.  i.  igth  November       .          .         .          .          .  1968 

No.  2.   igth  November        .....  1968 

No.  3.   1 7th  January           .          .          ,          .          .  1969 

No.  4.   ryth  January           .          .          .          .          .  1969 

No.  5.  aist  January            .....  1969 

No.  6.  7th  March       ......  1969 

No.  7.   I5th  April       ......  1969 

No.  8.  i6th  May        ......  1969 


PRINTED  IN  GREAT  BRITAIN 
BY  ALDEN  &  MOWBRAY  LTD 
AT  THE  ALDEN  PRESS,  OXFORD 


CONTENTS 

GEOLOGY  VOLUME  XVII 

No.  i.  On  the  Cretaceous  age  of  the  so-called  Jurassic  Cheilostomatus  Polyzoa 
(Bryozoa).  A  contribution  to  the  knowledge  of  the  polyzoa-fauna  of 
the  Maastrichtian  in  the  Cotentin  (Manche).  E.  VOIGT  I 

No.  2.     The    caudal    skeleton    in    Mesozoic    Acanthopterygian    fishes.     C. 

PATTERSON  47 

No.  3.     Non-calcareous  microplankton  from  the  Cenomanian  of  England, 

Northern  France  and  North  America.     Part  I  :     R.  J.  DAVEY  103 

No.  4.     A  redescription  of  W.  Carruthers  '  type  Graptolites.     I.  STRACHAN         181 

No.  5.     A  revision  of  the  English  Wealden  Flora,   I  Charales-Ginkgoales. 

J.  WATSON  207 

No.  6.     Two  new  Dicynodonts  from  the  Triassic  Ntawere  Formation,  Zambia. 

C.  B.  Cox  255 

No.  7.     Lower  Cambrian  Archaeocyatha  from  the  Ajax  Mine,  Beltana,  South 

Australia.     F.  DEBRENNE  295 

No.  8.     Some  Bathonian  Ostracoda  of  England  with  a  revision  of  the  Jones 

1884,  and  Jones  &  Sherborn  1888  Collections.     R.  H.  BATE  377 

Index  to  Volume  XVII  439 


ON  THE  CRETACEOUS  AGE  OF 

THE  SO-CALLED  JURASSIC 

CHEILOSTOMATUS  POLYZOA 

(BRYOZOA) 

A  CONTRIBUTION  TO  THE  KNOWLEDGE  OF 

THE  POLYZOA-FAUNA  OF  THE  MAASTRICHTIAN 

IN  THE  COTENTIN  (MANCHE) 


E.  VOIGT 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
GEOLOGY  Vol.  17  No.  i 

LONDON:  1968 


ON  THE  CRETACEOUS  AGE  OF  THE  SO-CALLER  ' 
JURASSIC  CHEILOSTOMATOUS  POLYZOA 

(BRYOZOA) 

A  CONTRIBUTION  TO  THE  KNOWLEDGE  OF  THE 

POLYZOA-FAUNA  OF  THE  MAASTRICHTIAN 

IN  THE  COTENTIN  (MANCHE) 


BY 

EHRHARD  VOIGT 


Pp.  1-45;  8  Plates;  2  Text-figures 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

GEOLOGY  Vol.  17  No.  i 

LONDON:  1968 


THE    BULLETIN    OF    THE    BRITISH    MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they  become 
ready.  Volumes  will  contain  about  three  or  four 
hundred  pages,  and  will  not  necessarily  be  completed 
within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  17,  No.  I  of  the  Geological 
Palaeontological  series.  The  abbreviated  titles  of  the 
Periodicals  cited  follow  those  of  the  World  List  of 
Scientific  Periodicals. 


World  List  abbreviation 
Bull.  Br.  Mus.  nat.  Hist.  (Geol.). 


©  Trustees  of  the  British  Museum  (Natural  History)  1968 


TRUSTEES    OF 
THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 

Issued  19  November  1968  Price  £2 


ON  THE  CRETACEOUS  AGE  OF  THE  SO-CALLED 
JURASSIC  CHEILOSTOMATOUS  POLYZOA 

(BRYOZOA) 

A  CONTRIBUTION  TO  THE  KNOWLEDGE  OF  THE 

POLYZOA-FAUNA  OF  THE  MAASTRICHTIAN 

IN  THE  COTENTIN  (MANCHE) 

By  EHRHARD  VOIGT 

SYNOPSIS 

The  supposed  Jurassic  cheilostomatous  Polyzoa  described  by  J.  W.  Gregory  (1894)  as  Mem- 
branipora  jurassica  and  Onychocella  bathonica  from  the  Bathonian  of  Ranville  (Calvados),  are 
of  Cretaceous  age  and  must  have  come  from  the  Maastrichtian  of  the  Cotentin  (Manche,  France). 
The  matrices  of  the  type  specimens  yielded  thirty-three  further  species  of  Maastrichtian  Polyzoa 
of  which  nine  are  Cyclostomata  and  twenty-four  Cheilostomata.  Three  new  species  Radulopora 
minor  n.  sp.,  Rosseliana  thomasi  n.  sp.  and  Frurionella  fertilis  n.  sp.,  and  the  new  genus  Radulo- 
pora are  described. 

CONTENTS 

I.     INTRODUCTION  AND  ACKNOWLEDGMENTS        .....  3 

II.     CHEILOSTOMATA  OF  CRETACEOUS  AGE  DESCRIBED   FROM   JURASSIC 

BEDS      ..........     4 

III.  POLYZOA  FROM  JURASSIC  BEDS  WHICH  HAVE  BEEN  ERRONEOUSLY 

REFERRED  TO  THE  CHEILOSTOMATA     ......  y 

IV.  ON  THE  ORIGIN  OF  GREGORY'S  "  JURASSIC  "  CHEILOSTOMATA   .  .  IO 

V.     PALAEOZOIC  CHEILOSTOMATA?       .          .          .          .          .          .          .  13 

VI.     THE  ACCOMPANYING  FAUNA  OF  THE  ROCK-MATRIX  OF  GREGORY'S 

"  JURASSIC  "  CHEILOSTOMATA  .          .          .          .          .          .          .  14 

VII.     REFERENCES      ..........  42 

I.     INTRODUCTION   AND    ACKNOWLEDGMENTS 

THE  predominant  orders  of  Polyzoa  (Bryozoa)  of  the  Cretaceous  and  Cainozoic 
periods  are  the  Cyclostomata  and  the  Cheilostomata.  The  Cheilostomata  have  been 
increasing  ever  since  an  explosive  development  in  the  Upper  Cretaceous,  whereas 
the  Cyclostomata  have  decreased  during  the  Tertiary  and  Quaternary  following  a 
flourishing  period  in  the  Cretaceous.  Today  there  exists  only  a  comparatively 
small  relic  of  this  cyclostomatous  fauna  stem  which,  in  Jurassic  times,  was  the 
only  living  group  of  Polyzoa  apart  from  some  rare  species  of  boring  Ctenostomata. 

This  last  statement  contradicts  all  textbooks  of  palaeontology  in  which  the 
Cheilostomata  are  being  erroneously  recorded  as  beginning  at  the  latest  in  the 
Jurassic,  an  opinion  repeated  by  R.  S.  Bassler  (1953). 

It  is  the  purpose  of  this  paper  to  prove  that  the  so-called  Jurassic  Cheilostomata 
described  by  Gregory  (1894)  from  the  Bathonian  of  Ranville  (Calvados),  are  in  fact 

GEOL.   17,  I.  I 


4  UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

Upper  Cretaceous  in  age  and  must  have  their  origin  in  the  Maastrichtian  of  the 
Cotentin  (Manche)  in  Normandy. 

Other  Jurassic  Polyzoa  described  as  Cheilostomata  by  several  authors  do  not 
belong  to  that  order  but  are  true  Cyclostomata.  In  any  case  it  must  be  emphasized 
that,  if  they  are  undoubtedly  Jurassic,  they  are  not  Cheilostomata,  or  if  they  belong 
to  that  order  they  are  not  Jurassic. 

I  wish  to  record  my  warm  appreciation  of  the  late  Dr.  Dighton  Thomas  of  the 
British  Museum  (Natural  History)  to  whom  I  am  very  much  indebted  for  studies 
in  the  Museum  collection,  for  the  loan  of  specimens,  for  discussion,  and  for  his  help 
and  advice.  At  the  time  of  his  death  he  had  begun  tidying  the  English  of  the 
manuscript,  and  this  was  kindly  finished  by  Dr.  J.  M.  Hancock.  I  am  also  grateful 
to  Dr.  A.  B.  Hastings  for  interesting  discussions  about  the  subject.  Grateful  thanks 
are  due  to  Prof.  J.  Lehman,  Dr.  J.  Sornay  and  Dr.  E.  Buge  of  the  Musee  d'Histoire 
Naturelle  (Paris)  for  permission  to  study  and  to  photograph  the  types  of  Bryozoa 
in  the  d'Orbigny-Collection.  I  also  wish  to  thank  Dr.  J.  Roger,  Paris,  for  kindly 
accompanying  me  and  guiding  me  in  the  Maastrichtian  territory  of  the  Cotentin 
and  other  French  classic  localities.  Finally  my  special  thanks  are  due  to  the 
Deutsche  Forschungsgemeinschaft  for  financial  help  to  further  this  research  and 
their  support  for  my  studies  on  Cretaceous  Polyzoa. 


II.    CHEILOSTOMATA    OF   CRETACEOUS    AGE   DESCRIBED    FROM 

JURASSIC   BEDS 

Considering  first  the  ages  of  undoubted  Cheilostomata  which  have  been  described 
as  Jurassic  forms,  we  can  enumerate  the  following  three  species:  Flustra flabelliformis 
Lamouroux  1821,  Onychocella  bathonica  Gregory  1894,  and  Membranipora  jurassica 
Gregory  1894. 


i.     "  Flustra  "  flabelliformis  Lamouroux  1821 
1821     Flustra  flabelliformis  Lamouroux:  113,  pi.  76,  figs.  11-13. 

This  species,  described  in  a  footnote  and  figured  by  Lamouroux,  apparently  from 
Ranville,  is  a  fanlike  fragment  of  a  bilaminar  Onychocella.  The  very  simple  figure 
shows  pyriform  zooecia  with  round  orifices  increasing  in  size  to  the  periphery  of  the 
frond.  An  avicularium,  rounded  distally,  seems  to  be  indicated.  Below  the  broken 
cryptocyst  the  rectangular  or  box-like  pattern  of  the  zooecia  is  visible.  The  original 
diagnosis  of  Lamouroux  is: 

"Flustra  en  forme  d'eventail,  fossile,  epaisse,  a  bords  entiers,  composee  de 
deux  membranes,  une  superieure  mince,  un  peu  translucide,  divisee  en  alveoles 
profonds,  a  bords  irreguliers  avec  un  oscule  rond  dans  le  centre,  qui  communique 
a  une  cellule  en  forme  de  carre  long,  tres  regulier,  avec  des  cloisons  epaisses 
et  solides,  les  transversales  alternant  entre  elles,  les  longitudinales  se  prolongeant 
sans  interruption  de  la  base  aux  extremites;  grandeur,  2  a  3  centimetres; 
epaisseur,  environ  i  millimetre." 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN  5 

Unfortunately  there  is  no  possibility  of  restudying  the  typespecimen  because, 
according  to  Sherborn,  Lamouroux's  collection  has  been  lost.  Nevertheless,  there 
is  no  doubt  that  the  specimen  is  not  from  the  Bathonian  of  Ranville  as  supposed 
by  Gregory,  but  must  be  regarded  as  a  Cretaceous  Onychocella,  likely  to  have  come 
from  one  of  the  Maastrichtian  localities  of  the  Cotentin  (Manche),  as  is  demonstrated 
for  the  two  following  species.  Lamouroux  himself  gives  as  the  origin  only  the 
neighbourhood  of  Caen,  which  perhaps  includes  not  only  Ranville  but  also  the 
Cotentin. 

Gregory  (1896  :  214)  in  his  Catalogue  of  the  Jurassic  Bryozoa  in  the  British 
Museum  suppressed  his  specific  name  bathonica  for  an  Onychocella,  regarded  by  him 
in  1894  as  a  new  species  from  the  Bathonian  of  Ranville,  in  favour  of  Lamouroux's 
species  because  he  believed  that  the  two  forms  were  identical. 

As  will  be  shown,  Gregory's  Onychocella  bathonica  is  the  well-known  Maastrichtian 
Onychocella  piriformis  Goldfuss  1826  first  described  from  the  "  Maastrichter  Tuff- 
kreide  "  in  the  Netherlands.  If  Gregory  was  correct  in  identifying  his  Onychocella 
bathonica  with  Flustra  flabelliformis  Lamouroux,  the  latter  name  has  priority.  On 
the  other  hand  Lamouroux's  figure  is  rather  unsatisfactory,  as  Gregory  himself 
stated,  for  a  conclusive  identification  of  Flustra  flabelliformis  Lamouroux  with 
Onychocella  piriformis  Goldfuss.  There  are  so  many  similar  species  of  Onychocella 
in  the  Upper  Cretaceous  that  it  seems  to  be  quite  impossible  to  find  out  which  species 
was  intended  by  Lamouroux  under  this  name.  Gregory  notes  that  Lamouroux's 
figure  is  so  indefinite  that  Pictet  gave  a  figure  of  a  form,  which  he  referred  to  this 
species,  which  was  really  Diastopora  lamellosa  Michelin.  It  is  probable  that  Gregory, 
in  identifying  his  Onychocella  bathonica  with  Flustra  flabelliformis,  was  influenced 
by  his  supposition  that  they  are  of  Bathonian  age,  and  as  Jurassic  species  of  Onycho- 
cella must  be  very  scarce,  he  believed  that  they  must  be  identical. 


2.     "  Onychocella  bathonica  "  Gregory  1894 

( =  Onychocella  piriformis  Goldfuss  1826) 

(PL  5,  figs.  11-12) 

1826  Eschar  a  piriformis  Goldfuss:  23,  pi.  8,  fig.  10. 

1851  Eschara  piriformis  (Goldfuss)  von  Hagenow:  75,  pi.  9,  fig.  6  and  pi.  n,  fig.  6. 

1894  Onychocella  bathonica  Gregory  :  63,  fig.  2 

1896  Onychocella  flabelliformis  (Lamouroux)  ;  Gregory  :  214,  fig.  22. 

1930  Onychocella  piriformis  (Goldfuss);  Voigt:  454,  pi.  16,  figs.  1-2. 

This  is  the  supposed  Jurassic  cheilostomate  which  was  described  and  figured  by 
Gregory  under  this  name  in  1894  and,  as  stated  above,  later  treated  by  him  as  a 
synonym  of  "  Flustra  "flabelliformis  Lamouroux.  The  first  mention  of  a  "  Jurassic 
Onychocella  "  was  made  by  Gregory  (1893  :  239).  It  is  represented  in  the  British 
Museum  Collection  by  two  specimens,  D.iSi  (type)  and  0.480,  both  from  the 
Tesson-Collection.  As  previously  suggested  by  the  present  author  (Voigt  1930  :  454), 
these  two  specimens  are  not  from  the  Bathonian  "  Calcaire  a  polypiers  "  as  recorded 


6  UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

by  Gregory,  but  both  are  Upper  Cretaceous  in  age.  This  is  proved  firstly  by  a 
Cheilostomatous  Polyzoan,  Stamenocella  marginata  (d'Orbigny)  which  is  visible  in 
the  matrix  of  specimen  D.i8i  (PL  3,  fig.  i)  and  which  was  overlooked  by  Gregory, 
and  secondly  by  a  large  fauna  of  other  Maastrichtian  Polyzoa  which  has  been 
isolated  from  the  adherent  matrix  of  the  block  D.iSi  and  which  is  described  in 
section  VI.  This  study  shows  clearly  that  all  these  supposed  Bathonian  Polyzoa 
are  in  fact  of  Upper  Cretaceous  age  as  already  indicated  for  "  Membraniporajurassica  " 
by  Lang  (1916  :  96,  97  and  1922  :  197-198)  and  Larwood  (1962  :  223). 

Onychocella  bathonica  Gregory  is  the  same  species  as  Onychocella  piriformis  (Gold- 
fuss)  1826  from  Maastricht.  It  is  quite  commonly  found  in  various  localities  of  the 
French  Cotentin  (Manche).  Although  Gregory  has  discussed  Goldfuss's  species 
and  written  that  it  has  a  lower  zooecial  aperture,  while  the  avicularian  aperture  is 
larger  and  the  front  wall  occurs  only  above  and  not  on  both  sides  of  this,  a  comparison 
with  a  specimen  from  Chef  du  Pont  indicates  that  they  are  synonyms  (pi.  5,  fig.  n). 
The  supposed  differences  are  not  significant  and  they  are  not  found  when  material 
from  Maastricht  or  from  the  Cotentin  localities  is  used  for  comparison.  Some  of  the 
opesiae1  in  the  figured  (pi.  5,  fig.  12)  British  Museum  specimen  D.iSi  are  a  little 
smaller  than  those  of  the  figured  specimen  from  Chef  du  Pont  (pi.  5,  fig.  n),  but 
there  are  variations  within  the  same  specimen.  Gregory's  figure  is  deceptive  because 
it  shows  a  small  quadrangular  fragment  with  only  five  whole  zooecia  and  one  avicu- 
larium;  it  does  not  correspond  in  size  to  his  two  originals.  The  Holotype  of  Onycho- 
cella bathonica  Gregory  is  a  large  bent  unilaminar  frond  of  nearly  3  cm.  length  (pi.  5, 
fig.  12).  It  contains  a  dozen  avicularia  and  shows,  on  some  zooecia,  the  very  small 
characteristic  endozooecial  ovicells  which  are  just  visible  as  minute  swellings  at 
the  distal  ends  of  the  zooecia  figured  here  (pi.  5,  fig.  12).  The  other  specimen 
0.480  is  a  large  unilaminar  fragment  of  n  x  8-5  cm.  size  with  four  avicularia. 

In  discussing  the  affinities  of  Onychocella  flabelliformis  (Lamouroux)  Gregory 
maintains  that  its  nearest  ally  may  be  von  Hagenow's  Cellepora  (Discopora)  konincki- 
ana  (1851  :  95,  pi.  n,  figs.  11-12)  from  Maastricht,  a  species  which,  with  its  straight 
rows  of  avicularia  and  small  zooecia,  is  very  differently  shaped.  In  discussing 
Onychocella  (  =  Cellepora}  koninckiana  (1896  :  215)  he  distinguished  the  form 
figured  by  von  Hagenow  in  his  fig.  n  as  a  new  species — Onychocella  hagenowi.  He 
assigned  it  to  a  separate  species  because  he  believed  that  it  has  larger  elliptical 
opesiae  with  the  longer  axis  longitudinal,  an  entire  lower  margin  of  the  opesia  and 
much  larger  avicularia.  This  example  shows  how  dangerous  it  is  to  judge  the 
variability  of  species  on  the  evidence  of  figures  alone.  Von  Hagenow  was  quite 
correct  in  considering  the  two  forms  as  only  one  species  because  they  can  often  be 
observed  in  the  same  zoarium  as  confirmed  by  Voigt  (1930  :  460). 

The  first  known  species  of  Onychocella  are  from  the  Cenomanian,  and  they  are 
small  and  of  a  lower  level  of  evolution.  Judging  from  this  point  of  view  it  would 
be  very  odd  if  the  oldest  species  should  have  the  largest  zooecia  of  the  genus,  more 
than  i  mm.  long,  and  big  avicularia  1-3-1 -4  mm.  in  length  as  these  are  otherwise 
developed  only  at  the  acme  of  the  Onychocellids  in  the  late  Upper  Cretaceous. 

1  I  have  followed  the  advice  of  Dr.  Hastings  in  using  "  opesia  ",  plural  "  opesiae  ",  rather  than 
"  opesium  ",  plural  "  opesia  ". 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN  7 

3.     Castanopora  jur assica  (Gregory)  1894 
(PI.  7,  figs.  4-7) 

1894  M  embranipora  jur  assica  Gregory:  62,  text-fig,  i. 

1896  Membranipora  jurassica  Gregory:  212,  text-fig.  21,  p.  213. 

1916  Rhiniopora  jurassica  (Gregory)  Lang:  96. 

1922  Rhiniopora  scabra  Lang:  196,  pi.  4,  fig.  7,  text-fig.  62. 

1922  Rhiniopora  jurassica  (Gregory)  Lang:  197. 

1962  Castanopora  jurassica  (Gregory)  Larwood:  223,  pi.  17,  figs.  3-5,  text-figs.  108-109. 

Holotype  D .  180,  large  bilaminar  fragment  of  damaged  zoarium  partly  embedded 
in  matrix.  Upper  Maastrichtian,  Cotentin,  Manche,  France  (not  Maastricht, 
Netherlands) . 

Lang  recognized  the  cribrimorph  nature  of  this  species.  It  was  overlooked  by 
Gregory  that  the  cribrimorph  structure  of  the  frontal-shield  was  preserved  quite 
well  in  some  zooecia  of  the  type  specimen  D.iSo  (Tesson  Coll.)  which  was  figured 
by  Larwood  (1962  :  224,  text-fig.  108,  pi.  17,  fig.  5).  Although  Gregory  described 
it  correctly  as  "  erect  foliaceous,  bilaminate  ",  Lang  defined  this  form  as  "  encrusting 
unilaminar  "  as  did  Larwood  (1962).  Dr.  Dighton  Thomas  and  Dr.  A.  Hastings 
were  kind  enough  to  confirm  my  first  observation  made  in  the  Museum  collection 
that  Gregory's  type  specimen  is  bilaminar  and  not  encrusting. 

Gregory,  misled  by  the  erroneous  data  of  Tesson's  label,  recorded  the  species  as 
coming  from  the  Bathonian  of  Calvados.  Lang  (1922  :  197)  recognizing  the  Maas- 
trichtian age,  supposed  that  it  originated  from  the  Dutch  locality  Maastricht  itself, 
and  recorded  the  distribution  of  this  species  as,  "  Senonian,  Maastrichtian,  Maastricht, 
Limburg,  Holland  ",  specimen  0.3313  being  labelled  "  Maastricht,  Old  collection  ". 
Larwood  (1962,  pi.  17,  fig.  3)  followed  Lang  and  gave  a  photograph  of  this  specimen. 
I  am  much  indebted  to  Dr.  Dighton  Thomas  for  lending  me  0.3313,  for  there  is  no 
other  record  of  Castanopora  jurassica  from  Maastricht,  and  the  specimen  may  have 
come  from  the  same  locality  as  D.iSo.  Dr.  Dighton  Thomas  compared  the 
matrices  of  these  two  specimens:  they  are  very  similar  in  colour,  grain  size,  and 
in  size  of  fossil  debris,  and  they  could  have  come  from  the  same  locality  in  the 
Cotentin. 

As  will  be  shown  in  section  IV  of  this  paper,  the  type  locality  cannot  be  Maastricht 
in  Holland,  but  must  be  the  same  as  that  of  the  last  species,  a  locality  in  the  Cotentin. 
The  matrix  of  the  type  specimen  D .  180  is  a  hard,  Polyzoan-bearing,  detrital  lime- 
stone, resembling  very  much  the  "  Craie  a  Baculites  "  or  "  Craie  a  Thecidees  "  of 
the  Cotentin.  Lang  has  already  recognized  some  other  Cheilostomata  in  the  matrix 
of  this  block.  They  are  described  and  figured  here  under  the  name  of  Multicrescis 
laxata  d'Orbigny,  Rosseliana  thomasi  n.  sp.  and  Pliophloea  sp.  These  species  are 
unknown  from  Maastricht  itself,  and  combined  with  the  results  above  on  "  Onycho- 
cella  bathonica" ,  there  can  be  no  doubt  that  the  two  species  both  come  from  the 
Cotentin. 

All  this  is  now  confirmed  by  the  fact  that  I  have  found  four  fragments  of  "  Casta- 
nopora "  jurassica  (Gregory)  in  my  material  from  Chef  du  Pont  (Cotentin,  Manche). 


8  UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

Dr.  Dighton  Thomas  kindly  compared  them  with  the  holotype,  and  he  has  no  doubt 
that  they  represent  the  same  species.  They  agree  in  measurements,  in  number 
of  costae,  and  all  other  characteristics.  All  are  bilaminar  with  the  exception  of  a 
young  zoarium  which  is  unilaminar  and  whose  zooecia  are  only  0-09-1-00  mm. 
long.  In  the  others  the  very  large  zooecia  are  about  1-2-1-4  mm.  long.  There  are 
about  twenty-six  to  thirty-four  costae  with  eight  lateral  costal  fusions  as  stated  by 
Larwood.  PL  7,  fig.  5  shows  an  instructive  view  with  some  zooecia,  one  of  which 
has  preserved  its  undamaged  cribrimorph  front  wall;  the  others  are  broken  and 
show  the  "  membranimorph  "  habit  of  Gregory's  figure.  On  the  reverse  side  of 
the  specimen  (PI.  7,  fig.  4)  are  some  zooecia  with  the  characteristic  ovicells  of  the 
genus  which  were  previously  not  known  in  full  preservation.  They  are  hyper- 
stomial  and  globular,  prominent,  and  overlapping  the  distal  zooecium  as  presumed 
by  Larwood.  They  are  perforated  by  isolated  fine  pores  (invisible  in  my  figure) 
as  in  Castanopora  guascoi  Ubaghs  from  Maastricht. 

It  must  be  emphasized  that  Castanopora  jurassica  (Gregory)  has  never  been 
found  near  Maastricht,  although  I  have  studied  the  Polyzoa  of  the  Maastrichtian 
Tuffkreide  for  40  years.  Near  Maastricht  another  allied  bilaminar  form  is  repre- 
sented, Cnstanopora  guascoi  (Ubaghs)  (1865  :  51,  pi.  2,  fig.  3),  whose  zooecia  in  some 
cases  reach  a  length  of  1-5  mm.  but  which  has  fewer  costae  (fifteen  to  twenty-two). 
Specimens  like  these  were  described  by  von  Hagenow  as  Cellepora  (Dermatopora) 
faujasi  (von  Hagenow  :  1851,  p.  99,  pi.  10,  fig.  19).  It  is  impossible  to  mistake 
Castanopora  guascoi  for  Castanopora  jurassica  (Gregory),  but  it  is  of  interest  that, 
if  the  costae  are  broken  down,  this  species  gives  the  appearance  of  Membranipora 
bipunctata  (Goldfuss  1826  :  26,  pi.  9,  figs.  7a-b),  traces  of  costae  being  no  longer 
visible.  The  Cotentin  fauna  contains  further  allied  bilaminar  and  unilaminar 
species  which  could  be  mistaken  for  Castanopora  jurassica,  but  there  is  little  room 
to  discuss  them.  Rhiniopora  scabra  Lang  1916  from  Riigen  is,  according  to  Larwood, 
a  synonym  of  Castanopora  jurassica  Gregory,  because  it  agrees  in  general  with  the 
number  of  costae,  and  in  having  8  lateral  costal  fusions  and  pelmatidia. 

In  a  recent  work  (Voigt  1968  :  65)  finished  before  the  issue  of  the  present  paper, 
the  genus  Rhiniopora  Lang  1916,  united  with  Castanopora  by  Larwood  (1962),  is 
maintained  at  least  as  a  subgenus  of  Castanopora. 

SPECIMENS : 

0.3313.  Large  fragment  partly  embedded  in  matrix,  recorded  by  Lang  (1922  : 
197)  and  figured  by  Larwood  (1962,  pi.  17,  fig.  3),  labelled  "  Maastricht,  Old  col- 
lection ",  but  certainly  from  a  locality  of  the  Cotentin-Maastrichtian,  Manche, 
France. 

0.49724.  Small  bilaminar  fragment  with  three  ovicelled  zooecia.  Upper 
Maastrichtian,  Chef  du  Pont,  Cotentin,  Manche,  France.  Collection  E.  Voigt. 

0.49725.  Young  unilaminar  zoarium  with  some  damaged  zooecia.  Horizon 
and  locality  as  above.  Collection  E.  Voigt. 

Some  fragments.  Horizon  and  locality  as  above.  In  Collection  E.  Voigt,  Ham- 
burg, Nr.  3585  and  3924. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIX  g 

III.  POLYZOA  FROM  JURASSIC  BEDS  WHICH  HAVE  BEEN 
REFERRED  TO  THE  CHEILOSTOM  AT  A 

It  is  now  evident  that  the  above  mentioned  Cheilostomata  are  not  Jurassic. 
There  are  scattered  records  in  the  literature  of  Cheilostomatous  Polyzoa  whose 
Jurassic  age  is  undoubted.  In  all  such  cases  suspicion  rises  that  they  are  not 
Cheilostomata.  The  fallibility  of  many  of  these  records  depends  on  the  progress 
of  science  and  today  nobody  can  take  seriously  mention  of  Cheilostomatous  genera 
like  Hippothoa,  Eschara,  Cdlaria  or  Cellepora  etc.  from  the  Jurassic  by  early  authors. 
However,  Gregory  (1894  :  61)  in  his  note  on  the  Jurassic  Cheilostomata  considered 
Eschara  ranvilliana  Michelin  from  the  Bathonian  of  Ranville,  and  Cellar ia  smithi 
Phillips  from  the  Cornbrash  of  Scarborough,  as  true  Cheilostomata  and  cited  them 
in  support  of  his  theory  of  the  existence  of  Jurassic  Cheilostomata.  In  his  Catalogue 
of  the  Jurassic  Bryozoa  (1896  :  56  and  127)  Cellaria  smithi  Phillips  is  assigned  to 
Stomatopora  and  Eschara  ranvilliana  Michelin  to  Diastopora  with  no  mention  that 
they  were  regarded  as  Cheilostomes  by  himself  two  years  earlier. 

F.  D.  Longe  (1881)  was  still  convinced  that  Eschara  ranvilliana  Michelin  from  the 
Bathonian  of  Ranville  belonged  to  the  Cheilostomata  when  he  wrote:  "  It  is  perfectly 
clear,  however,  that  some  of  the  Oolitic  Escharoids  themselves  possess  the  charac- 
teristic cell  features  of  the  Cheilostomata  in  a  marked  degree;  and  their  affinity  to 
the  Cheilostomatous  Escharidae  has  been  recognized  by  no  less  authorities  than 
d'Orbigny  and  Michelin  ". 

He  gave  a  detailed  analysis  of  the  supposed  Cheilostomatous  features  of  this 
species  and  related  forms,  and  tried  to  derive  the  Cheilostomata  from  certain  Oolitic 
Diastoporids  whose  zooecia  remind  one,  by  their  oval  or  polygonal  shape,  of  the 
Cheilostomata.  ("  The  decumbent  cells  in  Diastopora  may  be  regarded  as  ancestral 
Cheilostomatous  cells,  and  Diastopora  itself  as  the  parent  stock  from  which  many, 
if  not  all,  of  the  families  of  the  Chalk  and  subsequent  periods,  grouped  as  Cheilo- 
stomata, have  been  derived  ".). 

Walford  (1894)  published  a  note  "  On  Cheilostomatous  Bryozoa  from  the  Middle 
Lias  ".  Under  the  new  generic  name  Cisternifera  he  described  some  species  of  cyclo- 
stomatous  Bryozoa  with  large  heterozooecia — so-called  "  cistern-cells  " — whose 
relationship  with  the  ovicells  of  the  Cyclostomata  was  already  assumed  by  Walford. 
They  were  thought  to  bear,  occasionally,  minute  avicularia  on  the  upper  lip  of  the 
zooecia.  Apart  from  the  question  of  whether  these  structures  are  avicularia  or  not, 
the  different  forms  attributed  to  Cisternifera  are  true  Cyclostomata,  and  Gregory 
(1896)  himself  has  distributed  them  amongst  the  genera  Diastopora  and  Entalophora. 

Cellepora  davaiacensis  Lissajous  1923  from  the  Bathonian  of  the  Macon  district 
(France),  from  whose  generic  name  one  would  expect  it  to  be  a  Cheilostomatous 
Polyzoan,  and  which  Lissajous  has  compared  with  the  Cheilostome  Cellepora  poly- 
thele  Quenstedt,  has  been  recognized  by  David  (1952)  as  belonging  to  the  Calcispongia, 
probably  of  the  genus  Synopelta  Zittel.  The  list  of  literature  on  Jurassic  Polyzoa 
published  by  David  (1960)  contains  many  references  to  records  of  Cheilostomata  in 
other  works. 

From  these  investigations  it  is  now  certain  that  all  supposed  species  of  Jurassic 
Cheilostomata  prove  to  be  mistaken  identifications  or  erroneous  stratigraphic  records. 


10 


UPPER   CRETACEOUS    POLYZOA   FROM   COTENTIN 


IV.    ON    THE   ORIGIN   OF   GREGORY'S    "    JURASSIC   CHEILOSTOM  AT  A    ' 

In  the  Cotentin  (Manche)  Upper  Maastrichtian  a  lithology  similar  to  the  "  Tuff- 
kreide  "  from  Maastricht  itself  has  long  been  known.  The  fades  in  the  Cotentin 
is  a  "  tuffaceous  "  detrital  limestone  with  remains  of  many  Foraminifera,  Polyzoa, 
Echinoderms,  Brachiopods  (Craie  a  Thecidees  ")  or  hard  limestones  (Craie  a  Bacu- 
lites).  It  is  true  that  certain  blocks  of  these  rocks  can  be  easily  mistaken  for  genuine 
"  Maastrichter  Tuffkreide  ",  because  many  of  the  small  fossils  which  make  up  the 
rock  are  common  to  both  strata. 

An  important  difference  is  the  absence  of  any  larger  foraminifera  like  Orbitoides, 
Lepidorbitoides ,  Siderolites  or  Omphalocyclus  etc.  which  are  distinctive  of  the  Upper 
Maastrichtian  in  Holland  and  Belgium.  Hofker  (1959)  in  his  monograph  on  the 
Foraminifera  of  the  Cotentin  Maastrichtian  has  shown  that  these  beds  must  be 
intermediate  in  age  between  horizons  Cr  4  and  Mb  in  the  terminology  of  Uhlenbroeck 
for  the  Upper  Cretaceous  in  South  Limburg.  This  would  indicate  a  stratigraphic 
position  between  the  phosphatic  chalk  of  Ciply  and  the  base  of  the  Tuffaceous  chalk 
of  St.  Symphorien  in  the  Mons  basin,  or  an  equivalent  of  the  higher  beds  of  Folx-les- 
Caves  and  Orp-le-Petit  in  northern  Belgium.  This  could  explain  the  lack  of  larger 
Foraminifera  in  this  region  which  did  not  invade  the  northern  regions  before  the 
higher  horizons  of  the  Maastrichtian.  The  Maastrichtian  age  is  based  upon  the 
occurrence  of  Scaphites  constrictus  J.  Sowerby  in  the  area  of  Valogne.  For  the  other 
fauna  see  Vieillard  &  Dollfus  1875. 

In  1957  I  visited  the  Cotentin  region  in  order  to  study  the  classic  Senonian  localities 
of  d'Orbigny  (1850-54)  who  had  described  from  here,  chiefly  from  Nehou  and  from 
Sainte  Colombe,  some  hundred  species  of  Polyzoa.  Although  there  were  no  extant 
exposures  in  the  immediate  neighbourhood  of  these  villages,  some  larger  quarries 
near  Fresville  and  Port  Filiolet  and  an  excavation  for  the  dairy  of  Chef  du  Pont 
supplied  material  which  furnished  examples  of  a  considerable  part  of  the  Maas- 
trichtian Polyzoa  which  were  described  and  figured  by  d'Orbigny.  (For  the  site 


Valognes 


Nthou 


zter 


Fresville 

*      ^& 


S*  Colombe 


Rauvilte  - 
la  Place  °*j 
Rue-Tourville 

24      6     d     10 


) 

Chef  du  Pont 


Port  Filiolet 


km 


FIG.  i.  Map  of  the  outcrops  (black)  and  the  localities  of  the  Maastrichtian  in  the  Cotentin 
area  (Normandy).  (After  Carte  geologique  detaillee  de  la  France  (i  :  80000)  Feuille  28, 
St.  L6  (2  erne  Ed.)  1926. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN  n 

of  these  localities  see  Text-fig,  i.)  This  material  was  augmented  by  some  samples 
given  by  Dr.  F.  Schmid  (Hannover)  and  Polyzoa  from  Fresville  presented  by  Dr. 
P.  Marie  (Paris).  It  is  interesting  that  many  different  and  new  forms  not  mentioned 
by  d'Orbigny  were  found  at  these  localities,  and  by  contrast,  a  large  number  of 
d'Orbigny's  species  could  not  be  found  again.  Perhaps  there  are  some  horizons  with 
a  different  fauna,  and  the  beds  of  Sainte  Colombe  and  Nehou  may  belong  to  such 
levels,  whose  exposures  are  abandoned  today,  or  the  different  localities  represent 
heterogeneous  ecologic  biotopes. 

The  Polyzoan  fauna  of  this  region  is  the  most  important  one  in  the  northern  area 
and  its  knowledge  is  fundamental  to  our  knowledge  of  the  Maastrichtian  Polyzoa. 
With  the  exception  of  the  inadequate  revision  of  the  Cretaceous  Polyzoa  of  d'Orbigny 
as  a  whole,  which  was  undertaken  by  Pergens  (1889)  and  by  Canu  (1900),  nobody 
has  concerned  himself  with  the  Polyzoan  fauna  of  the  Cotentin  for  no  years.  An 
up-to-date  revision  of  this  fauna  is  therefore  an  urgent  task,  but  it  cannot  be  under- 
taken without  a  re-study  of  d'Orbigny's  types  which  are  preserved  in  the  Musee 
d'Histoire  naturelle  in  Paris. 

The  most  obvious  obstacle  for  a  successful  revision  is  the  fact  that  many  of 
d'Orbigny's  type  specimens  are  missing,  or,  if  specimens  are  present,  it  is  often 
impossible  to  say  with  certainty  whether  they  are  true  "  types  "  or  not.  On  the 
other  hand  many  of  the  drawings  do  not  agree  well  with  the  originals ;  they  may  be 
stylized,  restored  or  improved  and  consequently  it  is  often  rather  difficult  to  give  a 
satisfactory  identification  of  d'Orbigny's  species  (see  Canu  1900  :  335).  His  omission 
to  specify  the  exact  locality  of  the  figured  specimen  in  cases  when  there  is  more  than 
one  such  locality,  increases  the  difficulty  of  identifying  the  types.  There  are  about 
300  photographs  of  d'Orbigny's  polyzoan  types  of  the  Paleontologie  Fran9aise  in 
the  author's  collection,  and  this  stock,  together  with  a  rich  collection  of  French 
Cretaceous  Polyzoa,  are  the  basis  for  the  following  discussion. 

The  proofs  for  a  Cotentin  origin  for  Gregory's  "  Jurassic  Cheilostomata  "  are: 

1.  The  matrix  of  sediment  adhering  to  Gregory's  originals  is  identical  to  that  of 
the  above-listed  Maastrichtian  localities  of  Cotentin.     The  matrix  detached  from 
block   D.i8i   with   Onychocella  piriformis   Goldfuss  has   yielded   two   specimens 
(66.42981-82)  of  the  characteristic  fossil  Thecidea  papillata  V.  Schlottheim  (pi.  i, 
figs.  9-10)  and  it  is  evident  that  it  has  been  collected  from  the  so-called  "  Craie  a 
Thecidees  "  of  the  Cotentin. 

2.  The  objection  that  this  brachiopod  is  also  common  at  Maastricht  and  in  Belgium 
in  a  similar  facies  is  weakened  by  the  fact  that  the  accompanying  fauna  in  this 
block,  consisting  of  thirty-one  species  of  Polyzoa,  contains  some  species  which  are 
confined  to  the  Cotentin  fauna  and  have  never  been  found  in  Holland  and  Belgium, 
although  indeed  both  strata  have  many  species  in  common.     Examples  of  species 
not  known  from  these  countries  are:  Multicrescis  laxata  d'Orbigny,  Membranipora 
unipora   Marsson,   Stamenocella  cf.    marginata    (d'Orbigny),    Castanopora  jurassica 
(Gregory),  Onychocella  bellona  (d'Orbigny),  Onychocella  cf.  cepha  d'Orbigny,  Onycho- 
cellaria  caecilia  sp.,  Semiescharinella  complanata  d'Orbigny,  Lunulites  sp.,  Rosseliana 
thomasi  n.  sp.,  Frurionella  fertilis  n.  sp.  and  others. 


12          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

3.  A  Cotentin  origin  is  further  suggested  by  another  Cretaceous  Polyzoan  in  the 
museum  collection  which  is  embedded  in  a  typical  sample  of  "  Craie  a  Thecidees  ". 
It  is  a  large  specimen  of  a  Reticrisina,  determined  as  the  Jurassic  "  Reticulipora 
dianthus  (Blainville)  "  (6.4569  Old  collection)  and  labelled  "  Bathonian  Fresville  ". 
This  example  gives  a  good  idea  of  how  the  error  could  have  arisen :  Bathonian  rocks 
being  absent  near  Fresville,  Cretaceous  Polyzoa  were  mistaken  for  Bathonian  fossils. 
Gregory's  "  Jurassic  Cheilostomata  "  both  belong  to  the  Tesson-Collection,  which 
according  to  Gregory's  Catalogue  contains  many  Bathonian  Polyzoa  from  the  famous 
locality  of  Ranville  (Calvados). 

The  two  localities  are  about  100  km.  apart,  and  it  is  probable  that  the  labels 
were  confused  or  that  the  Cretaceous  Polyzoan  limestone  was  mistaken  for  the  not 
dissimilar  Coral  and  Polyzoan-limestone  ("  Calcaire  a  polypiers  ")  of  the  Bathonian. 

The  Tesson-Collection  to  which  Gregory's  types  belong  was  acquired  in  1857 
(Gregory  1896  :  35)  and  the  fossils  were  probably  collected  many  years  ago  at  a 
time  when  geological  mapping  of  this  region  was  still  in  its  infancy. 

It  is  not  impossible  that  Gregory's  types  both  came  from  Fresville  like  the  Reti- 
crisina mentioned  above.  The  rock  matrix  of  D.iSi  is  very  like  that  of  the  "  Craie 
a  Thecidees  "  of  Fresville,  but  as  Dr.  Dighton  Thomas  informed  me,  that  from  Chef 
du  Pont  is  even  closer  in  appearance.  But  such  lithological  differences  may  occur 
in  the  same  sequence,  and  are  of  no  great  importance. 

At  least  one  other  possible  explanation  of  this  error  should  be  mentioned.  Between 
the  Maastrichtian  outcrops  of  Sainte  Colombe  and  Rue  Tourville  is  the  village 
Rauville-la-Place  (see  Text-fig,  i).  Possibly  a  label  "  Rauville  "  was  misunderstood 
for  "  Ranville  ",  Calvados. 

It  must  be  reckoned  that  errors  like  these  are  more  common  in  old  collections,  and 
it  may  be  recalled  that  a  considerable  number  of  Reuss'  "  Cenomanian  Polyzoa  of 
Saxony"  came  from  the  Vincentown  Limesand  in  New  Jersey  (U.S.A.),  of  Dano- 
Paleocene  age  (Voigt  1942).  Another  mistake  is  the  supposedly  new  "  Polyphyma 
bulbosa  "  Hamm  (1881)  from  Maastricht  which  is  a  Hauterivian  form  from  northern 
Germany,  and  which  was  described  in  1839  by  Roemer  as  Alveolites  heteropora 
(Voigt  1953  :  57).  Certain  suspicious  anachronisms  of  phylogenetic  level  may  be 
explained  in  this  manner  as  has  been  shown  by  the  present  author  with  some  of 
Reuss'  types. 

4.  I  have  collected  much  material  of  Bathonian  Polyzoa  at  the  classic  locality  of 
Ranville  (Calvados),  but  I  have  never  found  any  trace  of  Cheilostomata.     The 
objection,  that  the  Cheilostomata  may  originate  in  Jurassic  times  cannot  be  con- 
tested; but  we  have  no  remains  of  them  earlier  than  the  Lower  Cretaceous.     It  is 
true  that  in  the  last  decade  several  important  groups  of  fossils  have  been  proved  to 
be  older  than  was  formerly  assumed.     Today  we  know  of  Cambrian  bivalves, 
Carboniferous  belemnites,  Triassic  frogs,  and  these  examples  could  be  augmented. 
It  is  possible  that  Jurassic  Cheilostomata  may  be  found  one  day.     But  in  such  a 
case  we  should  expect  primitive  forms  like  Membrammorphs  as  are  found  in  the 
lower  Cretaceous  and  not  highly  developed  forms  like  Onychocella,  or  highly  special- 
ized forms  like  the  Cribrimorph  Castanopora. 

Lower  Cretaceous  Cheilostomata  are  of  the  greatest  phylogenetic  interest,  and  all 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN          13 

forms  described  to  date  are  rare.     Most  of  them  are  Albian  and  the  oldest  is  recorded 
from  the  Neocomian.     Their  number  is  so  small  that  they  can  be  listed  in  a  few  lines : 

1.  Membranipora  neocomiensis  d'Orbigny  1853, 

Neocomian,  Saint-Sauveur,  Yonne,  France.  The  only  specimen  was  too  bad 
to  be  figured  by  d'Orbigny.  It  must  be  regarded  as  a  nomen  nudum. 

2.  "Membranipora"  constricta  d'Orbigny  1853, 

Aptian,  Les  Croutes  (Yonne). 

Remarks:  I  have  photographed  the  supposed  "  type  "  of  this  species  (pi.  7, 
fig.  8).  It  is  without  doubt  a  "  Membranipora  "  s.L,  but  it  does  not  cor- 
respond with  the  original  figure. 

3.  Rhammatopora  (?)  johnstoniana  Mantell  1844, 

Aptian,  Lower  Greensand,  Shanklin-Sand,  Kent. 

4.  Rhammatopora  gaultina  Vine  1890,  with  his  synonyms  Rhammatopora  vinei 

Lang  and  Rhammatopora  pembrokiae  Lang  (c.f.  Thomas  &  Larwood  1960) 
Albian-Cenomanian,  England. 

5.  Charixa  vennensis  Lang  1915,  Albian,  Dorset. 

6.  Pyripora  texana  Thomas  &  Larwood  1956,  Albian,  Texas  (U.S.A.). 

7.  Wilbertopora  mutabilis  Cheetham  1954,  Albian,  Texas  (U.S.A.). 

Vine  (1890)  mentioned  Membranipora  fragilis  d'Orbigny,  Membranipora  ?  obliqua 
d'Orbigny,  Membranipora  elliptica  v.  Hagenow  and  Hippothoa  simplex  d'Orbigny 
from  the  Red  Chalk  of  Hunstanton.  The  identification  of  these  forms  must  be 
revised,  but  nevertheless  all  the  named  species  are  primitive  encrusting  membrani- 
morphs  of  the  Division  Malacostega,  suborder  Anasca,  and  half  of  them  are  mono- 
or  oligoserial,  and  lack  either  ovicells  or  avicularia  as  we  must  theoretically  presume 
if  our  ideas  about  the  evolution  of  the  early  Cheilostomata  are  correct.  They  are 
followed  in  the  Cenomanian  by  the  first  primitive  Cribrimorphs  and  Coilostega  with 
the  families  of  Onychocellidae  and  Microporidae.  Therefore  it  is  hard  to  under- 
stand why  Canu  &  Bassler  (1920  :  318)  stated  that  following  the  Membraniporae, 
the  Acroporidae  Canu  1913  (=  Porinidae  d'Orbigny  1852)  of  the  suborder  Asco- 
phora  are  the  most  ancient  Cheilostome  fossils.  The  main  evolution  of  this  family 
is  in  the  later  Upper  Cretaceous,  and  the  only  described  species  of  Porina  from  the 
Cenomanian  is  P.  cenomana  Lecointre  (1912)  whose  origin  and  inner  structure  needs 
revising. 

V.    PALAEOZOIC   CHEILOSTOMATA? 

In  this  connection  the  question  of  the  systematic  position  of  the  north  American 
families  Worthenoporidae  Ulrich  1893  (Carboniferous)  and  Palescharidae  Miller  1889 
(Ordovician-Devonian)  cannot  be  neglected.  These  were  established  for  the  single 
genera  Worthenopora  Ulrich  1889  and  Paleschara  Hall  1874  which  resemble  Cheilo- 
stome Polyzoa  in  some  features.  Ulrich  (1890)  stated  that  the  affinities  of  that  genus 
are  nearer  to  the  Membraniporidae  than  any  other  and  that  his  present  views 
would  admit  it  being  placed  in  the  Cheilostomata.  Nickles  &  Bassler  (1900)  regarded 
both  families  without  any  restriction  as  Cheilostomata.  This  classification  was 
followed  by  many  authors,  but  in  recent  times  this  opinion  seems  to  have  been 


I4          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

abandoned.  Bassler  (1953)  has  placed  them  among  the  Cryptostomata,  but  con- 
siders Worthenopora  "  may  belong  among  cheilostomes  ". 

Worthenopora,  indeed,  has  the  outward  aspect  of  a  Cheilostome  with  its  triangular 
or  semielliptical  apertures,  with  posterior  raised  margin  and  spine  bases.  Paleschara 
with  its  simple  short  polygonal  zooecia  is  like  a  Membranipora  which  possesses 
completely  opened  opesiae  without  any  trace  of  a  gymnocyst. 

Dr.  Dighton  Thomas  has  kindly  lent  me  some  specimens  of  both  genera  from  the 
museum  collection.  Although  there  is  no  space  here  to  go  into  details,  further  studies 
of  this  problem  are  intended.  I  have  got  the  impression  that  they  cannot  be  attri- 
buted to  the  Cheilostomata.  Paleschara  must  be  regarded  as  a  very  primitive 
Cryptostome,  and  Worthenopora,  in  contrast,  as  a  specialized  one.  The  latter  has 
reached  a  level  of  evolution  which  reminds  one  of  certain  Cheilostomata.  It  has 
not  been  found  in  beds  younger  than  Mississippian,  and  there  are  no  intermediate 
forms  between  it  and  the  Cretaceous  Cheilostomata.  Consequently  it  seems  to  be 
impossible  to  regard  it  as  an  ancestor  of  the  true  earliest  membranimorph  Cheilo- 
stomata of  the  Lower  Cretaceous. 

Paleschara,  with  its  network  of  rather  simple  polygonal  zooecia,  may  perhaps  be 
primitive  enough  to  give  rise  to  Cheilostomata-like  forms.  But  it  could  be  better 
regarded  as  a  cryptostomatous  form  corresponding  to  a  membranimorph  level  of 
primitive  Cheilostomata. 

We  have  no  palaeontological  evidence  for  a  descent  of  the  Cheilostomata  from  the 
Cryptostomata  or  from  the  Ctenostomata.  As  Borg  (1930  :  54)  and  Cori  (1941) 
regard  the  Ctenostomata  as  emanating  from  primitive  Cheilostomata,  it  must  be 
emphasized  that  the  Ctenostomata  are  the  older  group,  represented  by  fossils  from 
the  Ordovician  onwards  and  that  there  is  no  possibility  of  deriving  the  Ctenostomata 
from  the  Cheilostomata.  Silen  (1942)  regards  the  Cheilostomata  and  the  Cteno- 
stomata as  closely  allied  ("  Cheilo-Ctenostomata  ")  and  believes  that  both  have 
common  ancestors. 

Silen  has  established  an  interesting  theory  about  the  origin  of  the  Cheilostomata 
from  hypothetical  primitive  forms  like  the  recent  Labiostomella  which  he  calls 
Protocheilostomata.  These  have  erect  zoaria  with  frontal  budding  and  other 
primitive  features,  and  it  would  mean  that  the  encrusting  growth  of  many  Cheilo- 
stomata and  the  lateral  budding  as  existing  in  all  other  Cheilostomata,  are  secondary. 
It  should  be  emphasized  that  the  oldest  known  Cheilostomata  from  the  Lower 
Cretaceous  do  not  show  this  primitive  character.  All  are  encrusting,  and  they 
must  already  have  passed  the  evolutionary  level  of  the  "  Protocheilostomata  ", 
which  according  to  Silen  were  feebly  or  not  at  all  calcified.  If  this  is  admitted, 
there  remains  no  possibility  of  regarding  Worthenopora  or  Paleschara  as  early  Cheilo- 
stomata of  the  Palaeozoic. 

VI.    THE   ACCOMPANYING  FAUNA   OF  THE  MATRIX   OF  GREGORY'S 
"   JURASSIC   CHEILOSTOMATA  " 

It  is  satisfactory  that  in  the  matrix  of  Gregory's  "  Jurassic  Cheilostomata  "  are 
enclosed  other  Bryozoa  which  were  overlooked  by  Gregory.  In  block  D.iSo, 
containing  Castanopora  jurassica,  Multicrescis  laxata  d'Orbigny,  Rosseliana  thomasi 
n.  sp.  and  Pliophoea  sp.  could  also  be  observed. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN          15 

I  am  much  indebted  to  Dr.  Dighton  Thomas  for  his  permission  to  remove  about 
2  cm.3  of  the  matrix  of  block  D.iSi  with  Onychocella  bathonica.  This  piece  was 
cracked  under  a  press  and  the  washing  of  the  residue  has  yielded  a  fauna  of  thirty- 
three  species  of  Polyzoa  and  one  Brachiopod.  This  fauna  is  described  in  detail  in 
the  following  part  of  this  paper.  Most  of  the  treated  specimens  are  figured  although 
the  state  of  preservation  is  rather  poor  because  the  hard  rock  is  unfavourable  for 
clean  preparation.  All  specimens  are  somewhat  damaged,  or  rolled  and,  unfortu- 
nately, recrystallized  or  covered  with  minute  calcite  crystals  as  is  often  observed  in 
the  Cotentin  material,  and  it  is  nearly  impossible  to  stain  them  with  colour.  This 
explains  any  mediocre  photos.  In  many  cases  figures  of  comparable  specimens  of 
the  same  species  from  the  Cotentin  are  given  for  comparison.  If  they  do  not  always 
present  exactly  the  same  picture  as  the  specimens  from  the  original  French  localities, 
this  is  because  there  often  exists  a  great  variability  in  size  or  growth-stage,  and 
in  preservation,  and  that  the  identification  is  based  upon  a  vast  quantity  of  material. 
It  is  surprising  that  in  only  2  cm.3  of  rock  from  D .  181  some  small  fragments  of  new 
species  were  found,  although  these  have  long  been  known  to  the  author  from  the 
Maastrichtian  of  the  Cotentin.  The  opportunity  is  taken  here  to  describe  them. 
But  it  should  not  be  forgotten  that  this  small  fauna  from  only  a  few  cm.3  matrix 
must  represent  a  very  small  part  of  the  rich  Polyzoan  fauna  of  the  Cotentin  Maas- 
trichtian, and  that  its  composition  is  purely  accidental. 

The  list  given  in  table  i,  p.  41,  contains  thirty-three  Polyzoan  species  of  which 
twenty-four  are  Cheilostomata.  It  is  not  complete  because  in  some  cases  a  com- 
plete identification  could  not  be  made,  and  for  a  few  forms  no  identification  was 
possible.  Excepting  Pliophloea  sp.  all  forms  have  been  found  by  the  author  in 
the  Maastrichtian  of  the  Cotentin.  Twenty-six  species  were  found  at  Chef  du 
Pont,  eighteen  at  Port  Filiolet  and  nine  at  Fresville.  But  it  should  be  taken  into 
account  that  the  investigated  material  is  too  poor  for  important  deductions.  Many 
species  very  common  at  all  Cotentin  localities  are  not  represented,  and  others  are 
new  for  the  Cotentin.  The  affinities  with  the  classic  locality  of  Maastricht  are 
proved  by  fourteen  species  but  this  figure  may  be  too  large  because  several  forms  of 
both  regions  may  be  identical.  Von  Hagenow's  types  from  Maastricht  having  been 
lost  during  the  second  world  war,  it  is  not  yet  possible  to  decide  this  question  with 
certainty. 

I.     POLYZOA 

A.     CYCLOSTOMATA 

Genus  BERENICEA  Lamouroux  1821 

i.     Berenicea  sp. 

One  small  incomplete  zoarium  which  is  indeterminable,  encrusts  Reteporidea 
lichenoides  Goldfuss. 

SPECIMENS.  A  minute  incomplete  zoarium  encrusting  Reteporidea  lichenoides 
Goldfuss  (=0.49569). 


16          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

Genus  ENTALOPHORA  Lamouroux  1821 
2.     Entalophora  benedeniana  (von  Hagenow) 

1851     Pustulopora  benedeniana  von  Hagenow:  17,  pi.  i,  fig.  6. 

1899     Entalophora  madreporacea  Goldfuss  var.  benedeni  von  Hagenow;  Gregory:  239. 

1964     Entalophora  benedeniana  (von  Hagenow)  Voigt:  422,  pi.  i,  figs.  1-7. 

One  small  fragment  embedded  in  matrix  has  been  referred  to  this  well-known 
species  from  Maastricht  which  is  represented  in  the  author's  collection  from  the 
Maastrichtian  of  Chef  du  Pont  (Manche).  For  morphological  details  and  ovicells 
see  Voigt  (1964). 

SPECIMENS.  0.49560.  One  poorly  preserved  fragment  in  matrix  from  D.iSi. 
Upper  Maastrichtian,  Cotentin,  Manche,  France. 

Genus  IDMIDRONEA  Canu  &  Bassler  1920 

3.  Idmidronea  macitenta  (von  Hagenow) 

1851  Idmonea  macilenta  von  Hagenow:  29,  pi.  2,  fig.  4. 

?i85i  Idmonea  ramosa  d'Orbigny:  736,  pi.  6n,  figs.  11-15. 

1899  Retecava  ramosa  (d'Orbigny)  Gregory:  192,  pro  parte. 

1951  Idmidronea  macilenta  Voigt:  38,  pi.  4,  figs.  14-17. 

One  small  distal  fragment  of  this  abundant  species  of  the  Upper  Maastrichtian 
belongs  to  /.  macilenta.  These  thin  distal  branches  do  not  have  the  numerous 
firmatopores  which  are  well  developed  on  the  reverse  side  of  the  broader  and  older 
stems.  Therefore  they  are  very  similar  to  Idmonea  (Tubigera)  antiqua  Defrance 
(d'Orbigny  1853  :  722,  pi.  613,  figs.  11-15,  figured  under  the  name  of  Stichopora 
regularis  d'Orbigny).  Idmonea  disticha  Goldfuss  (sensu  von  Hagenow  1851  :  30, 
pi.  2,  fig.  8)  may  be  mistaken  for  this  form  also,  but  always  it  is  not  so  flat  as  the 
distal  ends  of  Idmidronea  macilenta.  The  species  cited  by  d'Orbigny  from  Sainte- 
Colombe  under  the  latter  name  may  be  Idmonea  macilenta. 

If  this  species  is  con-specific  with  Idmonea  ramosa  d'Orbigny,  the  latter  name  has 
priority  and  must  be  preferred.  It  is  represented  at  Fresville  (Cotentin,  Manche). 

STRATIGRAPHICAL  RANGE.     Upper  Maastrichtian. 

SPECIMENS.  0.49561.  A  worn  fragment  from  matrix  of  D.iSr.  Upper 
Maastrichtian,  Cotentin,  Manche,  France. 

0.49843.  A  fragment,  upper  Maastrichtian,  Chef  du  Pont,  Cotentin,  Manche, 
France.  E.  Voigt  Collection. 

Genus  HETEROCRISINA  Gabb  &  Horn  1860,  em.  Voigt  1964 

4.  Heterocrisina  communis  (d'Orbigny) 

1853  Idmonea  communis  d'Orbigny:  745,  pi.  750,  figs.  6— 10. 

1887  Idmonea  pseudodisticha  (non  von  Hagenow)  Marsson:  28,  pi.  2,  fig.  8. 

1860  Heterocrisina  abbottii  Gabb  &  Horn:  404,  pi.  69,  figs.  45-47. 

1899  Retecava  abbottii  (Gabb  &  Horn)  Gregory:  205. 

non  1907  Idmonea  abbottii  (Gabb  &  Horn)  Ulrich  &  Bassler:  321,  pi.  22,  figs.  3-4. 

1964  Heterocrisina  communis  (d'Orbigny)  Voigt:  432  pi.  3,  figs.  i-io. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN  17 

Two  small  worn  fragments  belong  to  this  species,  which  is  very  distinctive  because 
of  its  large  frontal  ovicell,  although  it  has  often  been  mistaken  for  other  species. 
For  further  information  about  its  generic  position  and  morphological  details  see 
Voigt  (1964). 

This  form  is  common  at  all  Maastrichtian  localities  in  the  Cotentin  (Fresville, 
Port  Filiolet  and  Chef  du  Pont). 

STRATIGRAPHICAL  RANGE.     Campanian-Maastrichtian. 

SPECIMENS.  D . 49562-63.  From  matrix  of  D .  181.  Upper  Maastrichtian,  Coten- 
tin, Manche,  France. 

Genus  CRISISINA  d'Orbigny  1847 

5.     Crisisina  carinata  (Roemer) 

(PL  i,  figs.  4-5) 

1840     Idmonea  carinata  Roemer:  21,  pi.  5,  fig.  20. 

1964     Crisisina  carinata  (Roemer)  Voigt:  429,  pi.  4,  figs.  1-7.     (See  full  references.) 

Three  worn  fragments  belong  to  this  very  common  and  widespread  species  of  the 
Upper  Cretaceous.  The  synonymy  is  very  confused — see  Voigt  (1964) — where  all 
known  synonymies  are  given  and  the  generic  classification  is  discussed. 

Very  abundant  at  all  Maastrichtian  Cotentin  localities. 

STRATIGRAPHICAL  RANGE.     Cenomanian-Paleocene. 

SPECIMENS.  0.49564-66.  From  matrix  of  D.iSr.  Upper  Maastrichtian,  Coten- 
tin, France. 

0.49567.  A  fragment  for  comparison  with  0.49566.  Upper  Maastrichtian, 
Port  Filiolet,  Cotentin,  Manche,  France,  Voigt  Collection. 

Genus  OSCULIPORA  d'Orbigny  1847 
6.     Osculipora  truncata  (Goldfuss) 

1826  Retepora  truncata  Goldfuss:  28,  pi.  9,  fig.  14. 

1851  Truncatula  truncata  (Goldfuss)  v.  Hagenow  :  35,  pi.  3,  fig.  2. 

1851  Truncatula  tetrasticha  von  Hagenow:  34,  pi.  3,  fig.  3. 

1909  Osculipora  truncata  (Goldfuss)  Gregory:  58. 

1922  Osculipora  truncata  (Goldfuss)  Canu  &  Bassler:  57,  pi.  23,  figs.  1-6. 

One  small  worn  fragment  may  be  referred  to  0.  truncata;  it  is  in  the  condition  of 
Truncatula  tetrasticha  von  Hagenow,  which  represents  highly  worn  branches  of 
0.  truncata  (Goldfuss.) 

Further  investigations  are  necessary  to  check  whether  the  older  citations  of  this 
form  from  the  Cenomanian  by  Reuss  (1872)  are  correct.  In  the  author's  collection 
from  Chef  du  Pont,  Port  Filiolet  and  Fresville  are  numerous  fragments  of  this 
species,  which  was  not  recorded  by  d'Orbigny  from  any  of  his  Cotentin  localities. 

STRATIGRAPHICAL  RANGE.     Cenomanian  (?)  to  Maastrichtian. 

SPECIMENS.  0.49568.  A  very  poorly  preserved  worn  fragment  from  the  matrix 
of  D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

GEOL.   17,   I.  2 


i8          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

Genus  RETEPORIDEA  d'Orbigny  1849 
7.     Reteporidea  lichenoides  (Goldfuss) 

1826     Retepora  lichenoides  Goldfuss:  29,  pi.  9,  figs,  isa-b. 

1851     Idmonea  lichenoides  (Goldfuss)  von  Hagenow:  28,  pi.  2,  fig.  6. 

1899     Retecava  lichenoides  (Goldfuss)  Gregory:  194,  fig.  16,  p.  195. 

One  very  poor  fragment  which  is  encrusted  by  a  small  young  Berenicea,  has  been 
recognized  as  this  species  which  is  very  abundant  in  the  Upper  Maastrichtian.  It 
is  represented  in  the  author's  collection  from  Port  Filiolet  by  some  specimens.  It 
has  not  previously  been  recorded  from  the  Cotentin  Maastrichtian. 

STRATIGRAPHICAL  RANGE.     Maastrichtian. 

SPECIMENS.  0.49569.  A  small  worn  fragment  with  an  encrusting  Berenicea 
from  the  matrix  of  D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

Genus  PETALOPORA  Lonsdale  1850 

8.     Petalopora  sp. 

(PI.  i,  figs.  1-3) 

There  are  a  few  fragments  of  a  badly  preserved  Petalopora  which  are  conspecific 
with  similar  specimens  from  Chef  du  Pont  (pi.  i,  fig.  2) .  Their  identification  involves 
some  difficulties  because  it  is  impossible  to  identify  them  from  published  figures. 
They  are  allied  to  Heteropora  reiiculata  Marsson  (1887  :  26,  pi.  2,  fig.  4)  in  the  size 
of  the  branches  and  diameter  of  the  apertures  (about  0-14-0-15  mm.),  but  the 
mesopores  are  much  less  conspicuous,  and  on  the  figured  fragment  (pi.  i,  fig.  i) 
they  look  a  little  like  longitudinally-oriented  lines  which  are  straight  or  sinuously 
bent  and  interspersed  between  fine  ribs.  On  the  specimen  from  Chef  du  Pont  the 
mesopores  are  larger  and  they  show  a  more  longitudinally-oriented  inconspicuous 
network  (pi.  i,  fig.  3).  I  have  no  doubt  that  these  three  specimens  are  conspecific 
in  spite  of  this  difference  but  I  dare  not  identify  them  with  any  known  species. 

SPECIMENS.  0.49570-72.  Three  small  worn  fragments  from  the  matrix  of 
D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

0.49573.  A  larger  fragment  from  the  matrix  of  D.iSi.  Upper  Maastrichtian, 
Cotentin,  Manche,  France. 

0.49574.  A  larger  fragment.  Upper  Maastrichtian,  Port  Filiolet,  Cotentin, 
Manche,  France.  Voigt  Collection.  One  branched  fragment.  Upper  Maastrich- 
tian, Chef  du  Pont,  Cotentin,  Manche,  France.  Voigt  Collection,  Hamburg,  Nr.  3969. 

Genus  MULTICRESCIS  d'Orbigny  1854 

9.     Multicrescis  laxata  d'Orbigny 

(PL  i,  figs.  6-8) 

1854     Multicrescis  laxata  d'Orbigny:  1077,  pi.  800,  figs.  10-11. 

HOLOTYPE.  Upper  Maastrichtian,  Sainte  Colombe,  Cotentin,  Manche,  France. 
d'Orbigny  Collection,  Paris,  Musee  d'Histoire  Naturelle  Nr.  8416. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN  19 

There  is  a  fragment  of  a  "  heteroporid  "  Polyzoan  with  an  encrusting  colony  of 
Rosseliana  thomasi  n.  sp.  embedded  in  the  matrix  of  Castanopora  jurassica  Gregory 
(D.i8i).  It  cannot  be  distinguished  from  d'Orbigny's  type  specimen  of  Multi- 
crescis  laxata,  whose  apertures  and  the  mesopores  are  exactly  the  same.  Unfortu- 
nately there  are  no  other  specimens  of  this  species  in  d'Orbigny's  collection  which 
would  allow  confirmation  of  the  inner  structures  by  sections.  Specimens  from 
Chef  du  Pont  (Manche),  which  seem  to  be  conspecific  with  d'Orbigny's  species  have 
a  median  lamella  like  Grammascosoecia  Canu  &  Bassler  1922.  Pergens  (1889  :  373) 
and  Canu  &  Bassler  (1922  :  119)  have  included  this  species  in  the  synonymy  of 
Grammascosoecia  dichotoma  (Goldfuss)  from  Maastricht  (see  von  Hagenow  1851  :  47, 
pi.  5,  fig.  15).  This  might  be  correct;  some  specimens  from  Fresville  can  hardly  be 
distinguished  from  the  Maastricht  species.  On  the  other  hand  there  are  some  fifty 
fragments  from  Fresville,  and  none  show  the  characteristic  pattern  of  small  regular 
smooth  quadrangles  which  grow  from  the  calcified  mesopores  (cf.  von  Hagenow's 
fig.  I5i  and  Voigt  1951  pi.  4,  fig.  i)  and  which  can  be  observed  in  the  majority  of 
the  Maastricht  specimens.  Therefore  I  still  hesitate  to  unite  it  with  Gramma- 
scosoecia dichotoma  (Goldfuss). 

D'Orbigny  attributed  his  species  to  his  genus  Multicrescis  which  is  multilamellar. 
But  this  is  not  the  case  in  M.  laxata,  although  d'Orbigny  noted  two  layers  in  his 
type-specimen  from  Sainte-Colombe,  which  is  a  basal  fragment  (pi.  I,  figs.  7-8). 
It  is  well  known  that  the  basal  stems  often  develop  more  than  one  layer  of  zooecia, 
and  therefore  there  is  no  reason  to  place  this  form  in  the  genus  Multicrescis.  Many 
specimens  from  Fresville  have  radiating  rows  of  peristomes  as  in  Multicavea,  which 
can  also  be  observed  in  Grammascosoecia  dichotoma  (Goldfuss).  But  the  ovicell  of 
the  Cotentin  form  has  not  yet  been  discovered.  Therefore  I  prefer  to  leave  this 
form  provisionally  under  the  name  given  by  d'Orbigny. 

It  should  be  noted  that  the  median  lamella  in  Grammascosoecia  dichotoma  (Gold- 
fuss)  is  not  constant.  It  is  lacking  in  many  specimens  from  Maastricht,  and  there 
is  no  reason  to  assume  a  different  species. 

STRATIGRAPHICAL  RANGE.     Upper  Maastrichtian. 

SPECIMENS.  0.49575.  A  worn  fragment  with  encrusting  Rosseliana  thomasi 
n.  sp.  embedded  in  matrix  of  Castanopora  jurassica  (Gregory)  (D.iSo).  Upper 
Maastrichtian,  Cotentin,  Manche,  France.  Labelled  erroneously  by  Gregory  as 
"  Bathonian,  Ranville,  Calvados  ". 


B.     CHEILOSTOMATA 

MEMBRANIPORA  Blainville  1830  (sensu  lato) 

10.     "  Membranipora  "  unipora  (Marsson) 

(PL  2,  figs.  7-8) 

1852     Flustrella  simplex  d'Orbigny:  293,  pi.  699,  figs.  14-16. 

1887     Biflustra  unipora  Marsson:  52. 

1929     Membranipora  genucia  Brydone:  37,  pi.  13,  figs.  10-11. 


20          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

1930     Membranipora  unipora  (Marsson) :  Voigt  420,  pi.  4,  fig.  5. 
1925     Membranipora  unipora  (Marsson):  Levinsen  329,  pi.  2,  fig.  22. 

HOLOTYPE.  Upper  Maastrichtian,  Nehou,  Cotentin,  Manche,  France.  d'Orbigny 
Collection,  Paris.  Musee  d'Histoire  Naturelle  Nr.  8130. 

One  unilaminar  specimen  embedded  in  matrix,  showing  some  zooecia  with  the 
characteristic  avicularium  on  the  gymnocyst  below  the  opesia  and  the  narrow 
helmet-shaped  hyperstomial  ovicell,  agrees  very  well  with  d'Orbigny's  type  from 
Nehou  (pi.  2,  fig.  8)  and  another  well  preserved  specimen  from  the  Maastrichtian  of 
Port  Filiolet  (Manche).  The  zoarial  length  is  from  0-80-1-20  mm.  The  spines  of 
the  well  raised  margin  of  the  opesia,  which  is  0-45-0-50  mm.  long,  number  about 
twenty,  but  they  are  inconspicuous  and  often  hidden  by  recrystallization  of  calcite. 
The  zooecia  and  the  opesiae  of  my  specimen  of  this  species  from  Port  Filiolet  are  a 
little  larger  than  those  of  the  Museum-specimen  but  this  lies  within  the  range  of 
variation  of  this  form.  Membranipora  genucia,  described  by  Brydone  from  the 
upper  Campanian  of  Meudon  near  Paris,  is  a  synonym  as  shown  by  comparison 
with  Cotentin  and  Rugen  specimens  with  those  from  Meudon. 

Marsson  who  translated  this  species  from  Flustrella  to  Biflustra,  has  changed 
the  species  name  simplex  to  unipora  because  there  existed  already  a  recent  Biflustra 
simplex  d'Orbigny  1839.  Otherwise  another  Cretaceous  Membranipora  simplex 
d'Orbigny  exists  too.  It  is  clear  that  the  placing  of  this  species  in  "  Membranipora  " 
is  only  provisional;  a  revision  of  the  Cretaceous  Membraniporae  might  put  it  in 
another  genus. 

STRATIGRAPHICAL  RANGE.     Upper  Campanian-Maastrichtian. 

SPECIMENS.  0.49576.  Fragment  with  ovicelled  zooecia  from  the  matrix  of 
D.iSr.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

Genus  AMPHIBLESTRELLA  Prud'homme  1960 

ii.     Amphiblestrella  elegans  (von  Hagenow) 

(PL  4,  figs.  1-3) 

1851  Siphonella  elegans  von  Hagenow:  84,  pi.  6,  fig.  7. 

1851  Flustrella  baculina  d'Orbigny:  291,  pi.  699,  figs.  4-6. 

1930  Amphiblestrum  elegans  (von  Hagenow)  Voigt:  448,  pi.  13,  figs.  13-16. 

1960  Amphiblestrella  elegans  (von  Hagenow):  Prud'homme:  949. 

1962  Amphiblestrum  elegans  (von  Hagenow)  Berthelsen:  100,  pi.  9,  figs.  1-5. 

There  is  only  one  small  fragment  which  is  1-7  mm.  long  and  shows  eight  rows 
of  zooecia;  normally  there  are  ten  to  sixteen.  The  zoarial  dimensions  are  smaller 
than  those  of  specimens  from  the  type-locality  of  Maastricht,  and  from  the  Cotentin, 
where  this  species  is  very  common.  The  axial  canal  of  the  hollow  zoaria,  clearly 
visible  in  the  thicker  zoaria,  is  much  reduced  in  the  slender  branches,  and  may  disap- 
pear almost  completely  as  shown  in  d'Orbigny's  type  specimen  of  his  Flustrella 
baculina  (pi.  4,  fig.  2)  from  Nehou.  The  zooecia  from  Danian  material  are  mostly 
longer  than  those  from  the  Maastrichtian  (about  0-7  mm.  instead  of  0-6  mm.).  The 
size  of  the  opesiae  is  very  variable  in  this  species.  Canu  (1900)  in  his  revision  of 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN          21 

d'Orbigny  (1851-54)  has  incorrectly  regarded  this  species  as  a  synonym  of  Flustrella 
irregularis  d'Orbigny. 

In  the  Cotentin  localities  this  species  is  represented  from  Fresville  and  Chef  du 
Pont  and  by  d'Orbigny's  type  specimen  of  Flustrella  baculina  from  Nehou. 

STRATI  GRAPHICAL  RANGE.     Maastrichtian-Danian. 

SPECIMENS.  D. 49577.  A  worn  fragment  from  matrix  of  D.iSi.  Upper 
Maastrichtian,  Cotentin,  Manche,  France. 

D. 49578.  A  well  preserved  branched  fragment.  Upper  Maastrichtian,  Chef  du 
Pont,  Cotentin,  Manche,  France.  Voigt  Collection. 

Type-specimen  of  Flustrella  baculina  d'Orbigny.  Upper  Maastrichtian  Nehou, 
Cotentin,  Manche,  France.  In  d'Orbigny  Collection,  Paris,  Musee  d'Histoire 
Naturelle  Nr.  8127. 

Genus  RADULOPORA  nov. 

DERIVATIO  NOMINIS.  Derived  from  the  species-name  of  Biflustra  radula  Marsson 
1887. 

DIAGNOSIS.  Zoarium  bilaminar,  dichotomously  branched  and  probably  radicelled 
at  the  base.  Zooecia  dimorphic,  the  marginal  zooecia  of  the  acute  edges  of  the 
branches  being  larger,  and  having  larger  opesiae  than  the  normal  zooecia.  Crypto- 
cyst  well  developed,  finely  granulated;  opesiae  small  with  straight  proximal  rim 
and  occasionally  developed  lip.  Distal  interzooecial  asymmetrical  vibracula  above 
the  opesiae  with  long  elliptical  opening  and  a  small  thornlike  process  going  out  from 
the  left  or  right  inner  margin.  Ovicells  inconspicuous  exteriorly,  deeply  immersed, 
endozooecial. 

Type  species:  Biflustra  radula  Marsson  1887,  Lower  Maastrichtian  Riigen 
(Germany) . 

REMARKS.  This  new  genus  comprises  three  characteristic  species  in  the  Maastrich- 
tian, which  cannot  be  attributed  to  any  other  genus  hitherto  known.  Biflustra 
radula  Marsson  was  assigned  incorrectly  to  Amphiblestrum  by  Voigt  (1930).  It 
differs  from  Amphiblestrum  not  only  in  its  bilaminar  and  apparently  radicelled 
zoarium  with  dimorphic  zooecia  and  semicircular  opesia  in  which  a  proximal  lip  is 
originally  developed,  but  also  by  its  avicularian-like  vibraculum.  This  has  no 
pivot  for  the  articulation  of  the  mandibula  as  in  true  avicularia,  but  asymmetrical 
curved  thorn-like  processes  on  the  inner  left  or  right  edge  of  the  vibracula  (Text- 
fig.  2)  and  an  ellipsoidal  cavity  in  the  proximal  part  adapted  for  the  motion  of  the 
seta  of  the  vibraculum.  These  structures  are  very  well  shown  in  the  type-species, 
R.  radula  (Marsson)  (pi.  3,  figs.  11-12),  while  they  are  indicated  in  R.  minor  only 
in  some  better  preserved  specimens. 

Owing  to  the  loss  of  the  Marsson  Collection  during  the  last  war,  a  neotype  for 
R.  radula  Marsson  should  be  erected.  The  specimen  of  Biflustra  radula  figured 
here  could  represent  a  good  neotype,  but  it  was  collected  from  a  chalk  block  in 
glacial  drift  and  therefore  a  topotype  from  the  Lower  Maastrichtian  of  Riigen 
would  be  preferred. 

It  is  difficult  to  decide  if  this  genus  should  be  regarded  as  belonging  to  the  Mala- 


22 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 


costega  or  Coilostega.  The  cryptocyst  is  strongly  calcified  and  the  opesia  is  so 
small  that  the  Malacostega  level  has  probably  been  exceeded.  Opesiules  are  absent 
as  in  many  Coilostega.  Nevertheless  Marsson  described  it  as  Biflustra,  and  perhaps 
he  was  right  to  assign  it  to  the  Membranimorphs,  although  there  are  many  inter- 
mediate forms  between  these  two  groups.  This  genus  seems  to  be  rather  isolated 
and  it  is  difficult  to  attribute  it  to  one  particular  family  of  the  Malacostega. 


FIG.  2.     Radulopora  radula  (Marsson).     Some  zooecia  and  vibracularia.      x6o. 

12.     Radulopora  minor  sp.  n. 

(PI.  3,  figs.  6-io) 

Holotype  Upper  Maastrichtian,  St.  Pietersberg  near  Maastricht  (Netherlands). 
0.49844.  Voigt  Collection. 

DERIVATIO  NOMINIS.  The  name  refers  to  the  smaller  zooecial  dimensions  in 
contrast  to  the  type-species  R.  radula  (Marsson)  (pi.  3,  figs.  11-12). 

DIAGNOSIS.  Radulopora  with  slender  dichotomous  branches  0-7-1 -2  mm.  wide, 
consisting  of  about  three  to  seven  alternating  transverse  rows  of  zooecia  about 
0-5-0 -6  mm.  long.  The  edges  of  the  zoarium  are  formed  by  the  larger  marginal 
zooecia:  other  zooecia  not  clearly  distinct  at  their  margins,  elongate  and  pyriform 
with  distal  raised  margin  and  a  well  developed  cryptocyst  deeply  immersed  proxi- 
mally.  Opesiae  rounded  quadrangular  or  high-semicircular  rounded  distally  and 
truncated  proximally,  showing  in  some  specimens  a  well  developed  proximal  lip. 
Distal  vibracula  small,  the  peak  turned  obliquely  downwards  but  symmetrically 
oriented  outwards  from  the  median  line  toward  the  edges  of  the  branches.  Ovicells 
form  inconspicuous  slight  swellings  above  the  opesiae  (pi.  3,  fig.  9). 

DESCRIPTION.  The  bilaminar  fronds  have  narrow  cylindrical  or  prismatic  basal 
rods  which  are  pointed  toward  their  proximal  ends  and  suggest  an  articulated  basal 
attachment  of  the  zoarium.  The  zoarial  and  opesial  dimensions  are  smaller  in  these 
proximal  parts  of  the  zoarium  than  in  the  more  distal  branches.  The  shape  of  the 
opesia  varies  from  nearly  semi-circular  to  high-oval,  trapezoidal  or  oval:  if  it  is 
oval  it  is  narrowed  proximally  but  it  is  never  circular.  The  straight  proximal 
edge  of  the  opesia  is  deeply  immersed  in  old  zooecia  and  may  disappear.  The  distinct 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN          23 

proximal  lip  observed  regularly  in  some  specimens  has  its  origin  from  this  straight 
proximal  edge.  The  opesia  occupies  about  one  quarter  to  one  fifth  of  the  length 
of  the  zooecium.  The  cryptocyst  is  highly  calcified  and  often  appears  in  the  proximal 
part.  The  marginal  zooecia  appears  to  be  larger  than  they  actually  are  because 
they  are  not  narrowed  in  their  proximal  part,  and  the  opesiae  are  always  distinctly 
larger  than  in  the  other  zooecia,  although  there  may  be  transitions  between  the  size 
of  the  opesiae  of  the  normal  and  those  of  the  marginal  zooecia.  The  ovicelled 
zooecia  never  have  vibracula.  The  ovicells  are  very  flat  swellings  above  the  opesiae. 
If  their  roof  is  broken  away  a  large  deep  hollow  is  revealed  showing  that  the  ovicell 
is  deeply  immersed  although  it  must  be  regarded  as  endozooecial. 

A  minute  fragment  of  eight  zooecia  belongs  to  this  species  which  corresponds  very 
well  with  the  abundant  material  of  the  author's  collection  from  the  Cotentin  localities 
and  from  Maastricht. 

MEASUREMENTS. 

Lz  (=  Distance  between  the  opesia) :  0-50-0-67  mm. 

ho  =  0-10-0-12  mm.    ""I 

>  normal  zooecia. 
lo  =  0-075-0-12  mm.  J 

ho  =  0-14-0-17  mm.    ~\  .     , 

,  >  marginal  zooecia. 

lo  =  0-15-0-16  mm.    / 

STRATIGRAPHICAL  RANGE.     Upper  Maastrichtian. 

SPECIMENS.  0.49579.  A  small  worn  fragment.  From  matrix  of  D.iSi. 
Upper  Maastrichtian,  Cotentin,  Manche,  France. 

0.49580-81.  Two  fragments.  Upper  Maastrichtian,  Chef  du  Pont,  Cotentin, 
Manche,  France.  Voigt  Collection. 

0.49582.  Fragment  with  ovicelled  zooecia.  Upper  Maastrichtian,  Cotentin, 
Manche,  France.  Voigt  Collection. 

Genus  HAGENOWINELLA  Canu  1900 
13.     Hagenowinella  cf.  incrassata  (d'Orbigny) 

(PI.  2,  figs.  1-3) 
1853     Flustrellaria  incrassata  d'Orbigny:  527,  pi.  726,  figs.  5-8. 

A  large  unilaminar  fragment  consisting  of  more  than  twenty  zooecia  may  belong 
to  this  species,  although  the  opesiae  are  oval  and  not  so  broad  and  truncated  as  in 
d'Orbigny 's  type-specimen  from  Sainte-Colombe  (Manche).  A  fragment  of  this 
form  collected  by  the  author  near  Chef  du  Pont  (Manche),  shows,  however,  that  the 
shape  of  the  opesiae  can  vary  considerably  and  that  in  certain  parts  of  the  zoarium 
the  opesiae  are  fairly  oval  as  in  0.49583,  in  which  two  zooecia  have  the  cryptocyst 
broken  giving  the  false  appearance  of  avicularia.  The  hyperstomial  ovicells  shown 
on  the  specimen  from  Chef  du  Pont  are  broad  and  low.  The  horse  shoe-like  lamella 
in  the  interior  of  the  zooecia  seems  to  be  hidden  by  a  fine  calcitic  crust,  but  it  is 
indicated  in  pi.  2,  figs.  2-3,  by  two  small  knob-like  processes  at  the  inner  margin  of 
the  opesia. 


24          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

SPECIMENS.  0.49583.  A  small  fragment  of  zoaiium,  embedded  in  matrix  of 
D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

0.49845.  Fragment  of  zoarium.  Upper  Maastrichtian,  Fresville,  Cotentin, 
Manche,  France.  Voigt  Collection.  Fragment  with  ovicelled  zooecia.  Upper 
Maastrichtian,  Chef  du  Pont,  Cotentin,  Manche,  France.  Voigt  Collection,  Ham- 
burg, Nr.  3909. 

STRATIGRAPHICAL  RANGE.     Upper  Maastrichtian. 

Genus  BACTRELLARIA  Marsson  1887 
14.     Bactrellaria  rugica  Marsson 

(PI.  2,  figS.  9-14) 

1887     Bactrellaria  rugica  Marsson  59,  pi.  5,  fig.  18. 

1930     Bactrellaria  rugica  Marsson;  Voigt:  444,  pi.  12,  figs.  25-26. 

The  species  is  represented  by  one  very  poor  fragment  showing  four  zooecia  of  the 
frontal  face  with  worn  avicularia  forming  cavities  between  the  opesiae.  Although 
the  specimen  is  very  small  and  worn,  the  species  is  so  characteristic  that  there  is 
no  doubt  about  the  identification.  It  has  been  found  by  the  author  at  Maastricht 
and  Port  Filiolet  (Manche)  where  it  is  rare.  The  frontal  avicularium  below  the 
opesia  is,  if  well  preserved,  rather  prominent,  elongate  beak-like  and  is  proximally 
oriented  with  a  raised  spatulate  peak.  Marsson's  figure  showing  only  a  round 
pore  is  inadequate,  but  he  makes  mention  of  the  tube-like  avicularia  which  occasion- 
ally occur  on  the  lateral  edges  of  the  zoaria.  The  frontal  avicularium  below  the 
opesia,  is,  if  present  and  well  preserved,  rather  prominent,  tube-like,  elongate  and 
proximally  oriented.  Its  beak  is  dilated  and  spatulate  but  is  mostly  broken  away 
or  damaged.  It  is  rather  well  preserved  on  a  specimen  from  an  Upper  Maastrichtian 
chalk-bearing  erratic  flint-boulder  found  in  the  gravel-pit  of  Wulmstorf  near  Harburg 
(pi.  2,  fig.  13).  If  the  dilated  spatulate  avicularian  beak  is  destroyed  there  remains 
an  oval  ring  or  a  scar  like  the  ones  in  many  specimens  of  the  Cotentin  (pi.  2,  figs.  9 
and  12).  The  small  pit  on  Marsson's  figure,  however,  is  not  the  cicatrix  of  a  destroyed 
avicularium,  but  corresponds  to  the  spot  where  the  avicularium  is  usually  developed. 
Three  pairs  of  oral  spines  are  present  in  well  preserved  specimens.  The  ovicells  are 
hyperstomial.  Many  fragments  of  the  band-like  zoaria  from  the  Cotentin  localities 
have  four  to  six  rows  of  zooecia  instead  of  three;  their  diameter  varies  between 
0-7-2-0  mm.  The  length  of  the  zooecia  is  between  0-6-0-8  mm.,  that  of  the  opesia 
c.  0-35  mm.  In  addition  to  the  band-like  specimens  there  occur,  at  Maastricht, 
prismatic  vincularian  rods  with  five  or  more  rows  of  zooecia  showing  absolutely  the 
same  features  and  size  of  zooecia,  avicularia  and  ovicells.  They  are  very  like 
Pithodella  and  it  is  possible  that  they  represent  another  mode  of  growth  of  the  same 
species.  Eschara  gaimardi  von  Hagenow  (1851  :  82,  pi.  12,  fig.  10)  is  thought  to 
be  an  Eschara-like  stem  development  of  this  genus. 

STRATIGRAPHICAL  RANGE.     Maastrichtian. 

SPECIMENS.     0.49584.     A    small    worn    fragment.     From    matrix    of    D.iSi. 
Upper  Maastrichtian,  Cotentin,  Manche,  France. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN          25 

0.49585.  A  well  preserved  fragment.  Upper  Maastrichtian,  Port  Filiolet, 
Cotentin,  Manche,  France.  E.  Voigt  Collection.  A  worn  fragment.  Upper  Maas- 
trichtian, Port  Filiolet,  Cotentin,  Manche,  France.  Voigt  Collection,  Hamburg 
Nr.  3914. 

A  well  preserved  fragment.  Upper  Maastrichtian  Chalk  from  Flint-boulder  in 
Pleistocene  Drift.  Wulmstorf  near  Harburg-Hamburg,  Germany.  Voigt  Collection 
Nr.  4146. 

Genus  STAMENOCELLA  Canu  &  Bassler  1917 

15.     Stamenocella  marginata  (d'Orbigny) 

(PI.  3,  figs.  1-5) 

1852     Flusirella  marginata  d'Orbigny:  295,  pi.  700,  figs.  7-9. 
1852     Flustrella  convexa  d'Orbigny:  290,  pi.  699,  figs.  1-3. 
1852     Biflustra  tesselata  d'Orbigny:  271,  pi.  694,  figs.  7-9. 

HOLOTYPE.  A  small  fragment  of  Flustrella  marginata  d'Orbigny.  Upper  Maas- 
trichtian, Sainte  Colombe,  Cotentin,  Manche,  France.  In  d'Orbigny  Collection, 
Paris,  Musee  d'Histoire  Naturelle,  Nr.  8134. 

A  worn  fragment  of  this  species  embedded  in  the  matrix  of  Onychocella  bathonica 
Gregory  (D .  181)  agrees  in  all  essential  characteristics  with  the  two  specimens  which 
were  described  by  d'Orbigny  as  Flustrella  marginata  from  Sainte-Colombe  and 
Biflustra  tesselata  from  Nehou,  and  which  are  figured  here  for  comparison.  The 
latter  is  a  worn  specimen  of  Stamenocella  marginata  showing  traces  of  ovicells  and 
represents  the  same  stage  of  preservation  as  figs.  3  and  8  of  pi.  31  of  Canu  &  Bassler 
(1920)  where  the  ovicells  and  the  avicularia  of  Stamenocella  mediaviculifera  and 
Stamenocella  inferaviculifera  cause  two  shallow  cavities  between  the  opesiae. 

PI.  3,  fig.  5,  shows  a  rather  well  preserved  specimen  which  seems  to  be  intermediate 
between  Flustrella  marginata  d'Orbigny  and  Flustrella  convexa  d'Orbigny.  The 
basal  region,  which  tapers  proximally  and  shows  a  radicelled  base,  has  zooecia  cor- 
responding to  those  of  Flustrella  convexa.  They  are  closed  by  a  calcareous  lamella 
and  pierced  by  elliptical  or  round  openings  as  can  also  be  observed  in  other  species 
of  Stamenocella. 

The  variation  in  the  size  of  the  opesiae  is  enormous  in  this  species  as  shown  in 
pi.  3,  fig.  5,  and  the  following  measurements: 

LZ  =  0-50-0-63  mm. 

lz  =  0-17-0-25  mm. 
L0  =  0-17-0-33  mm. 

10  =  0-07-0-15  mm. 

STRATIGRAPHICAL  RANGE.     Maastrichtian. 

SPECIMENS.  0.47322.  A  fragment  embedded  in  the  matrix  of  Onychocella 
bathonica  Gregory  (D.iSi).  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

D .  49586.  A  fragment  from  the  matrix  of  D .  181.  Upper  Maastrichtian,  Cotentin, 
Manche,  France. 


26          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

0.49587.  A  small  worn  fragment.  Locality  and  horizon  as  above.  A  well 
preserved  fragment  showing  the  stage  of  Flustrella  convexa  d'Orbigny  at  the  base  of 
the  zoarium.  Upper  Maastrichtian,  Chef  du  Pont,  Cotentin,  Manche,  France, 
Voigt  Collection,  Hamburg,  Nr.  4128. 

Type  specimen  of  Biflustra  tesselata  d'Orbigny.  Upper  Maastrichtian,  Nehou, 
Cotentin,  Manche,  France.  d'Orbigny  Collection,  Paris,  Musee  d'Histoire  Naturelle. 
Nr.  8053. 

Genus  THYRACELLA  Voigt  1930 

16.     Thyracella  cf .  meudonensis  (d'Orbigny) 

(PI.  2,  figs.  4-6) 

cf.  1851     Biflustra  meudonensis  d'Orbigny:  263,  pi.  692,  figs.  4-6. 

1951     Thyracella  cf.  meudonensis  (d'Orbigny)  Voigt:  59,  pi.  9,  figs.  4-5. 

Three  fragments  of  this  bilaminar  species  are  conspecific  with  a  "  Biflustra  " 
which  is  very  abundant  in  the  Maastrichtian  of  Port  Filiolet  (Manche).  It  has  a 
very  prominent  large  avicularium  as  is  typical  of  Thyracella  (pi.  2,  fig.  4).  I  think 
it  could  be  identified  with  Biflustra  meudonensis  d'Orbigny  recorded  by  him  from 
Ne"hou  and  Meudon.  In  the  catalogue  of  the  d'Orbigny  collection  only  one  fragment 
from  the  Chalk  of  Meudon  is  registered  under  Nr.  8090  although  I  possess  more 
than  fifty  specimens  from  this  locality  which  would  fit  very  well  into  that  species. 
It  is  worn  and  does  not  correspond  with  the  figure;  it  does  not  look  like  a  chalk 
fossil  from  Meudon,  but  seems  to  have  come  from  a  more  littoral  facies  resembling 
the  Maastrichtian  of  the  Cotentin.  The  worn  specimens,  described  and  figured  by 
the  present  author  from  the  Maastrichtian  of  Kunrade  (S-Limburg)  and  Ilten 
(northern  Germany)  give  a  different  impression  from  that  of  the  well  preserved 
material  from  the  Cotentin  localities  in  the  author's  collection.  But  there  are 
all  intermediate  stages.  For  comparison  a  photograph  of  a  characteristic  specimen 
with  a  large  avicularium,  from  Port  Filiolet  (Manche),  is  given  (pi.  2,  fig.  4).  The 
length  of  the  avicularium  is  c.  0-8  mm.,  and  that  of  the  zooecia  c.  0-5-0-6  mm. 
The  well  preserved  zooecia  always  have  a  sharp  and  distinct  margin  in  the  distal 
part  as  shown  in  the  figure  of  d'Orbigny. 

STRATIGRAPHICAL  RANGE.     Maastrichtian  and  (fide  d'Orbigny)  Upper  Campanian. 

SPECIMENS.  D  .49588.  A  damaged  fragment  from  the  matrix  of  D .  181.  Upper 
Maastrichtian,  Cotentin,  Manche,  France. 

0.49846.  A  fragment  embedded  in  matrix  of  D.iSi.  Upper  Maastrichtian, 
Cotentin,  Manche,  France. 

0.49589-90.  Two  fragments  from  the  matrix  of  D.iSi.  Upper  Maastrichtian, 
Cotentin,  Manche,  France. 

A  well  preserved  branched  fragment  with  an  avicularium.  Upper  Maastrichtian, 
Port  Filiolet,  Cotentin,  Manche,  France.  Voigt  Collection,  Hamburg.  Nr.  3919. 

A  worn  fragment  labelled  Meudon  but  likely.  Upper  Maastrichtian,  locality 
uncertain  (?  Nehou,  Cotentin,  Manche).  d'Orbigny  Collection,  Paris,  Musee  d'Histoire 
Naturelle  Nr.  8040. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN          27 

Genus  VINCULARIA  (auct.) 

The  genus  Vincularia  Defrance  is  here  understood  in  the  sense  of  the  older  authors, 
although  this  is  not  correct  according  to  the  rules  of  nomenclature.  Vincularia  is 
now  restricted  to  those  forms  which  Canu  1907  named  Heterocella.  But  Vincularia 
fragilis  Defrance  1829  from  the  Eocene  is  the  type  species  of  the  genus  of  Vincularia 
(cf.  Bassler  1953  :  157)  and  the  name  Vincularia  must  be  reserved  for  this  group. 
Heterocella,  therefore,  as  a  synonym  of  Vincularia,  must  be  dropped. 

The  consequence  of  this  is  that  there  is  no  name  available  for  the  many  Cretaceous 
species  of  "  Vincularia  "  of  d'Orbigny,  Marsson,  Brydone  and  other  authors.  Canu 
(1900)  has  united  most  species  of  Vincularia  under  Smittipora  Jullien  (1881),  which 
Bassler  (1953)  considered  con-generic  with  Diplopholeos ,  Rectonychocella  and  Velu- 
mella  Canu  &  Bassler.  But  the  definition  of  Smittipora  does  not  fit  most  "  Vin- 
cularias  ".  Admitted  that  Vincularia  has  been  established  primarily  as  a  zoarial 
growth-form  for  rod-like  stems,  there  still  exists  a  natural  group  of  forms  for  which 
this  name  has  been  used  and  for  which  another  name  does  not  yet  exist.  On  the 
other  hand  it  seems  to  be  necessary  to  distinguish  the  forms  which  are  radicelled 
or  articulated  at  their  base  from  those  which  are  attached  by  an  encrusting  base. 
This  is  not  the  place  to  give  a  new  classification  of  Vincularia,  and  it  is  necessary 
to  retain  the  old  name  provisionally. 


17.     Vincularia  canalifera  von  Hagenow 
(PI.  5,  figs.  7-10  and  PI.  8,  figs.  9-12) 

1851      Vincularia  canalifera  von  Hagenow:  61,  pi.  6,  fig.  14. 
?i85i      Vincularia  flexuosa  d'Orbigny:  76,  pi.  656,  figs.  16-18. 
1930     Vincularia  canalifera  von  Hagenow;  Voigt,  467,  pi.  17,  fig.  18. 

More  than  thirty  fragments  of  this  species  were  found  in  the  matrix  of  D.iSi. 
They  are  conspecific  with  the  most  common  Vincularia-species  of  Maastricht  and 
Kunrade  (Netherlands)  which  was  described  by  von  Hagenow  as  Vincularia  canali- 
fera. Von  Hagenow's  figure  is  not  quite  typical  because  the  length  of  the  opesia 
is  relatively  large,  about  one  third  or  one  quarter  of  the  length  of  the  zooecium. 
Among  some  hundred  fragments  there  are  only  two  which  correspond  to  von  Hage- 
now's figure,  but  this  species  can  show  great  variation  in  size  and  shape  of  the 
zooecia  and  opesiae.  The  opesia  varies  between  an  oval  (pi.  5,  figs.  7,  9,  10)  or 
more  semicircular  (pi.  5,  fig.  8,  pi.  8,  figs.  9-11)  opening  which  is  truncated  more  or 
less  proximally.  In  well  preserved  specimens  it  has  a  slight  margin,  but  never  a 
proximal  lip.  In  the  proximal  basal  region  the  zooecia  are  0-25  mm.  long,  and  when 
fully  developed  0-5  mm. ;  the  length  of  the  opesiae  is  0-10-0-17  mm.  The  avicularia, 
which  have  not  been  figured  until  now,  reach  the  length  of  the  zooecia  or  exceed  it 
(pi.  5,  fig.  8,  pi.  8,  figs.  10,  n).  They  are  rare,  straight,  and  appear  at  the  beginning 
of  a  new  row  of  zooecia,  or  are  normally  enclosed  in  these.  They  are  broader  than  the 
autozooecia  and  have  a  flatly  rounded  prominent  distal  rim  and  a  small  elliptical 


28          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

opesia  which  is  longitudinally  oriented.  Common  to  the  zooecia  and  the  avicularia 
is  a  furrow-like  deepening  in  the  median  axis  of  the  cryptocyst  from  which  the 
specific  name  is  derived.  Ovicells  have  never  been  observed. 

The  rods  have  a  diameter  of  0-5-0 -7  mm.  and  consist  of  eight  to  fourteen  rows 
of  zooecia.  They  taper  proximally  and  were  articulated  or  radicelled  at  their  base. 
This  is  shown  by  some  of  the  earliest  zooecia  which  seem  to  be  modified  to  lodge 
chitinous  rootlets  (pi.  5,  fig.  7).  Their  opesiae  are  smaller  and  their  upper  half  is 
covered  by  a  bent  calcitic  lamella  as  is  seen  also  in  other  articulated  or  radicelled 
forms. 

This  species  is  also  common  in  the  Maastrichtian  of  the  Cotentin  near  Port  Filiolet, 
Fresville  and  Chef  du  Pont  from  which  region  d'Orbigny  described  some  very  similar 
vincularian  species. 

I  take  it  that  Vincularia  flexuosa  d'Orbigny  is  conspecific  with  this  species,  which 
is  cited  by  d'Orbigny  from  Nehou  in  the  Cotentin  and  from  the  Santonian  of  Ven- 
dome.  I  have  studied  the  type  (Nr.  7752  in  the  d'Orbigny  collection)  and  I  cannot 
find  any  significant  differences.  It  is  figured  on  pi.  8,  fig.  14  for  comparison  with 
specimens  from  Maastricht  (pi.  5,  figs.  7  and  8  and  pi.  8,  figs,  n  and  12).  The  reason 
why  I  hesitate  to  place  this  species  under  d'Orbigny's  name  is,  that  according  to  the 
label  the  type  specimen  is  from  the  Santonian  of  Vendome  (as  stated  also  in  his 
catalogue)  although  I  suspect  that  it  comes  from  Nehou  in  the  Cotentin.  Because 
it  is  not  clear  if  the  holotype  comes  from  the  Cotentin,  I  prefer  the  name  Vincularia 
canalifem  which  is  given  by  von  Hagenow  in  the  same  year  as  d'Orbigny's  name. 

Another  form  from  the  Cotentin  which  is  very  similar  to  Vincularia  canalifera 
is  V.  concinna  (d'Orbigny  1851  :  79,  pi.  657,  figs.  10-12).  It  may  be  another  syno- 
nym. D'Orbigny's  holotype  is  figured  on  pi.  8,  fig.  13;  this  is  the  only  specimen 
of  this  species  in  the  d'Orbigny  Collection  of  Nr.  7756).  This  specimen,  recorded  by 
Canu  (1900  :  420)  as  "insuffisant ",  has  opesiae  also  which  are  about  0.11-0.17  mm. 
long,  but  the  cryptocyst  seems  to  be  less  deepened  than  in  Vincularia  canalifera  von 
Hagenow.  Unfortunately  it  shows  no  avicularia,  knowledge  of  which  is  very  import- 
ant for  the  discrimination  of  many  species  of  Vincularia  which  are  very  similar 
and  cannot  be  identified  from  the  work  of  d'Orbigny.  D'Orbigny  never  figured 
or  mentioned  in  his  species  the  avicularia,  which  are  rather  rare  and  similar  to  the 
autozooecia. 

STRATIGRAPHICAL  RANGE.     Maastrichtian. 

SPECIMENS.  0.49732.  Branched  fragment,  Upper  Maastrichtian,  Geulem  near 
Berg,  Geul- Valley  near  Maastricht,  Netherlands.  Voigt  Collection. 

D. 49733,  0.49735-36,  0.49737-66.  Fragments  from  the  matrix  of  D.iSi, 
Upper  Maastrichtian,  Cotentin,  Manche. 

0.49734.  Fragment,  Upper  Maastrichtian  Chef  du  Pont,  Cotentin,  Manche. 
Voigt  Collection. 

0.49840-41.  Two  fragments,  Upper  Maastrichtian,  St.  Pietersberg  near  Maas- 
tricht, Netherlands.  Voigt  Collection. 

0.49842.  Fragment,  Upper  Maastrichtian,  St.  Pietersberg  near  Maastricht, 
Netherlands.  Voigt  Collection. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN          29 

Genus QUADRICELLARIA  d'Orbigny  1850 

18.    Quadricellaria  elegans  d'Orbigny 

(PI.  6,  figs.  8-10) 

1851  Quadricellaria  elegans  d'Orbigny:  33,  pi.  652,  figs.  1-5. 

1900  Quadricellaria  elegans  d'Orbigny;  Canu:  413. 

1928  Quadricellaria  excavata  d'Orbigny;  Voigt:  112,  text-figs.  1-5. 

1930  Quadricellaria  excavata  d'Orbigny  ;  Voigt  :  489,  pi.  25,  figs.  12-14. 

Two  small  fragments  of  a  Quadricellaria  may  be  determined  as  Q.  elegans  d'Orbigny. 
The  size  of  the  opesia  compared  with  the  length  of  the  zooecia  varies  considerably, 
and  Canu  may  be  right  in  uniting  d'Orbigny's  three  species,  Q.  elegans,  Q.  excavata 
and  Q.  pulchella,  under  the  name  Q.  excavata,  as  I  did  in  1928.  My  specimens  agree 
largely  with  the  figures  of  Q.  elegans  d'Orbigny  which  was  first  recorded  from  Nehou 
(Manche) . 

This  species  is  represented  in  my  material  from  Chef  du  Pont  and  Port  Filiolet. 
One  specimen  from  Chef  du  Pont  has,  on  the  narrow  sides  of  the  segments,  two 
enormous  avicularia  which  are  I  mm.  long  and  have  an  opesia  with  a  length  of 
0-3  mm.  (pi.  6,  fig.  10). 

SPECIMENS.  0.49591-92.  Two  worn  fragments.  From  matrix  of  D.iSi. 
Upper  Maastrichtian,  Cotentin,  Manche,  France. 

A  fragment  with  a  large  avicularium.  Upper  Maastrichtian,  Chef  du  Pont, 
Cotentin,  Manche,  France.  Voigt  Collection,  Hamburg,  Nr.  4133. 


Genus  COSCINOPLEURA  Marsson  1887 
19.     Coscinopleura  sp. 

Two  minute  indeterminable  fragments  of  a  Coscinopleura,  showing  only  a  few 
zooecia  are  present.  They  possibly  belong  to  a  species  of  Coscinopleura  with  small 
zooecia,  like  Coscinopleura  lamourouxi  von  Hagenow,  or  to  a  similar  form.  Common 
in  the  Cotentin  Maastrichtian  near  Port  Filiolet  and  Chef  du  Pont  (Manche). 

SPECIMENS.  0.49593-94.  Two  small  worn  fragments.  From  the  matrix  of 
D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

Genus  SEMIESCHARINELLA  d'Orbigny  1852! 

20.     Semiescharinella  complanata  d'Orbigny 

(PI.  4,  figs.  11-15) 

1840     Cellepora  ricata  von  Hagenow:  616. 

1852     Semiescharinella  complanata  d'Orbigny:  427,  pi.  714,  figs.  1-4. 

1  D'Orbigny  published  his  genera  Semiescharinella  and  Reptescharinella  on  p.  427  and  p.  428  of  his 
work.  According  to  Sherborn  (Geol.  Mag.  1889  :  223-225)  pages  187-472  appeared  in  1852.  I  follow 
here  the  dates  given  by  Sherborn,  although  Lang  (1917  :  172)  states  1853  for  Reptescharinella  and 
d'Orbigny  himself  writes  1851  for  both  genera. 


3o  UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

1852  Escharinella  simplex  d'Orbigny:  205,  pi.  683,  figs.  14-16. 

1900  Rhagasostoma  simplex  (d'Orbigny);  Canu:  433. 

1930  Micropora  subgranulata  (von  Hagenow)  Voigt  (pars):  476,  pi.  24,  fig.  18  (non  19). 

1959  Semiescharinella  complanata  (d'Orbigny)  Voigt:  54,  pi.  6,  fig.  i. 

1962  Semiescharinella  complanata  (d'Orbigny)  Berthelsen:  134,  pi.  14,  figs.  2-3. 

Two  very  small  unilaminar  fragments  agree  very  well  with  some  specimens  from 
Chef  du  Pont  and  with  d'Orbigny's  type-specimen  from  Sainte  Colombe  (Manche). 
The  size  of  the  zooecia  is  a  little  less  in  the  type-specimen  (about  0-56-0-60  mm.) 
compared  with  0-70  mm.  in  0.49595-96;  but  the  abundant  material  in  the  author's 
collection  shows  that  the  size  of  the  zooecia  and  opesiae  is  highly  variable  in  this 
species.  The  "  TYPE  "  is  a  fragment,  with  c.  18  zooecia,  in  which  the  relative 
length  and  width  of  the  zooecia  differ  from  d'Orbigny's  figure  as  noted  by  Canu 
(1900  :  421),  although  in  the  catalogue  of  the  d'Orbigny  Collection  only  one  speci- 
men from  Sainte  Colombe  (Manche)  is  registered.  The  bilaminar  Escharinella 
simplex  d'Orbigny  from  Nehou  (Manche)  belongs  to  the  same  species.  The  distal 
pore  which  was  interpreted  by  Canu  as  the  trace  of  the  ovicell  is  a  true  avicularium. 
Nevertheless,  there  are  some  real  ovicells  which  are  developed  in  place  of  the  avi- 
cularia  (see  pi.  4,  fig.  15). 

Comparison  of  the  photographs  (pi.  4,  figs.  13-15)  shows  that  there  is  no  variation 
in  the  shape  or  size  of  the  zooecia,  nor  of  the  opesiae,  nor  amongst  the  distal  avicu- 
laria.  I  have  figured  the  only  poor  fragment  from  the  d'Orbigny  collection  which 
must  be  regarded  as  the  type  specimen  (pi.  4,  fig.  15).  This  species  is  congeneric 
with  Cellepora  (Discopora)  subgranulata  von  Hagenow  (1851  :  91,  pi.  n,  fig.  15) 
which  was  chosen  by  Lang  (1917  :  172)  as  the  type  species  for  the  genus  Repteschari- 
nella  d'Orbigny  1852.  D'Orbigny  had  united  under  this  name  eight  Cretaceous, 
one  Tertiary  and  two  Recent  species,  of  quite  distinct  systematic  differences,  said 
to  be  characterized  by  an  "  ouverture  mediocre  ",  a  "  pore  special  "  and  encrusting 
zoaria.  The  genus  Semiescharinella  was  not  discussed  by  Lang,  being  represented 
only  by  Semiescharinella  complanata  d'Orbigny.  There  is  no  doubt  what  d'Orbigny 
meant  by  this  name,  and  I  prefer  it,  proposing  to  drop  Reptescharinella  as  a  synonym 
of  Semiescharinella,  the  mode  of  growth  being  no  generic  criterion.  This  form  was 
first  described  by  von  Hagenow  1840  under  the  name  Cellepora  ricata  without  any 
figure  (Voigt  1959  :  54). 

STRATIGRAPHICAL  RANGE.    Maastrichtian-Danian. 

SPECIMENS.  0.49595-96.  Two  small  worn  fragments.  From  the  matrix  of 
D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

0.49847.  A  small  fragment.  Upper  Maastrichtian,  Chef  du  Pont,  Cotentin, 
Manche,  France.  Voigt  Collection. 

A  small  fragment  showing  an  ovicelled  zooecium.  Upper  Maastrichtian,  Chef 
du  Pont,  Cotentin,  Manche,  France.  Voigt  Collection  Hamburg  Nr.  39i8b. 

Bilaminar  fragment,  Holotype  of  Escharinella  simplex  d'Orbigny.  Upper  Maas- 
trichtian, Nehou,  Cotentin,  Manche,  France.  d'Orbigny  Collection,  Paris,  Musee 
d'Histoire  Naturelle  Nr.  7942. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN  31 

Genus  ROSSELIANA  Jullien  1888 
21.     Rosseliana  thomasi  sp.  n. 

(PL  i  fig.  6,  and  PL  4,  figs.  4-6) 

TYPE  SPECIMEN.  0.49597.  Zoarium  encrusting  a  branched  fragment  of  Radu- 
lopora  minor  n.  g.  n.  sp.  Upper  Maastrichtian,  Chef  du  Pont,  Cotentin,  Manche, 
France.  Collection  E.  Voigt. 

DERIVATIO  NOMINIS.  In  honour  of  Dr.  Dighton  Thomas,  London,  who  first 
detected  this  species  in  the  matrix  of  Castanopora  jurassica  (Gregory) . 

DIAGNOSIS.  A  Rosseliana  with  encrusting  zoarium.  Zooecia  very  small,  oval, 
only  0-33  mm.  long,  with  smooth  cryptocyst  and  sharply  raised  rim  in  the  distal 
region  of  the  zooecia.  Opesiae  semicircular,  occupying  nearly  a  third  of  the  length 
of  the  zooecia,  sometimes  with  very  inconspicuous  lateral  processes,  but  never 
trifoliate  in  shape;  its  lower  rim  is  straight  without  opesiules.  Ovicells  globular 
swellings  above  the  opesia  and  occupying  the  proximal  part  of  the  distal  zooecium. 

DESCRIPTION.  In  addition  to  the  British  Museum  specimen,  which  encrusts  a 
branch  of  Multicrescis  laxata  d'Orbigny,  there  are  two  others  from  Chef  du  Pont 
(Manche).  They  show,  although  they  are  more  fragile,  the  same  essential  specific 
characters,  having  no  avicularia  and  no  trifoliate  opesiae.  In  some  zooecia,  very 
minute  lateral  processes  might  indicate  a  Floridina-like  opesia,  but  this  is  so  negli- 
gible that  it  can  be  ignored.  They  are  therefore  classified  as  Rosseliana  and  not 
Floridina  or  Floridinella.  Nevertheless,  it  is  clear  that  the  difference  between 
Rosseliana  and  Floridinella,  based  only  on  the  existence  of  broad  opesiular  indenta- 
tions in  the  latter,  is  slight.  The  specimens  are  very  similar  to  that  figured  by 
Bassler  (1953,  fig.  130.4),  but  he  only  records  Rosseliana  from  the  Oligocene  to 
Recent. 

There  are  very  few  species  with  which  this  form  could  be  compared.  The  shape 
and  size  of  zooecia  are  very  similar  in  Floridina  (or  better  Floridinella)  scutata  Levinsen 
(1925  :  345,  pi.  4,  fig.  39)  from  the  Danish  Maastrichtian  Chalk  and  Danian,  and 
Semieschara  complanata  d'Orbigny  (1852  :  369,  pi.  708,  figs.  5-8).  Apart  from  the 
free  unilaminar  fronds  of  the  latter,  it  is  very  difficult  to  find  any  constant  difference 
between  these  two  forms,  which  are  distinguished  from  our  new  Rosseliana  thomasi 
only  by  their  opesiae,  always  markedly  trifoliate  as  in  Floridina  or  Floridinella. 
PL  4,  fig.  7,  shows  a  specimen  of  Floridinella  scutata  Levinsen  from  a  Maastrichtian 
flint  drift  boulder  from  northern  Germany.  The  difference  of  this  species  from 
Rosseliana  thomasi  n.  sp.  is  clearly  seen  in  the  opesia  which  has  significant  lateral 
indentations. 

STRATIGRAPHICAL  RANGE.     Upper  Maastrichtian. 

SPECIMENS.  0.47323.  Encrusting  zoarium  with  some  ovicelled  zooecia  on 
Multicrescis  laxata  d'Orbigny  (=  0.49575).  In  the  matrix  of  Castanopora 
jurassica  (Gregory)  (D .  180)  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

Zoarium  encrusting  an  echinoid  fragment.  Upper  Maastrichtian,  Port  Filiolet, 
Cotentin,  Manche,  France.  Voigt  Collection,  Hamburg,  Nr.  3932. 


32          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

Genus  MICROPORA  Gray  1848  (non  Eichwald  1855) 
22.     Micropora  transversa  (d'Orbigny) 
(PI.  7,  figs.  9-12) 

1851  Vincularia  transversa  d'Orbigny:  78,  pi.  657,  figs.  7-9. 

1887  Vincularia  rugica  Marsson:  65,  pi.  6,  fig.  8. 

1930  Micropora  rugica  (Marsson)  Voigt:  472,  pi.  21,  fig.  20. 

1951  Micropora  rugica  (Marsson)  Voigt:  63,  pi.  9,  fig.  10  and  pi.  10,  fig.  4. 

HOLOTYPE.  Two  minute  fragments  evidently  2  pieces  of  one  specimen.  Upper 
Maastrichtian,  Nehou,  Cotentin,  Manche,  France.  d'Orbigny  Collection,  Paris, 
Musee  d'Histoire  Naturelle  Nr.  7755. 

There  is  only  a  very  small  fragment  corresponding  to  the  length  of  a  single  zoo- 
ecium  of  this  species  in  the  matrix  of  D .  181.  The  two  symmetrical  opesiules  below 
the  opesia  show  clearly  that  it  must  belong  to  Vincularia  transversa  d'Orbigny,  the 
type  of  which,  from  Nehou,  (pi.  7,  figs.  10-11)  was  studied  by  the  author.  There  is 
only  one  poorly  preserved  fragment  in  d'Orbigny's  collection  which  is  recorded  in  his 
catalogue.  It  is  now  broken  into  two  pieces.  It  does  not  correspond  to  d'Orbigny's 
figure  and  description,  because  the  paired  opesiules,  which  are  clearly  visible, 
are  neither  mentioned  in  his  text  nor  figured  in  his  drawing ;  they  seem  to  be  indicated 
by  the  deep  furrows  accompanying  the  thick  margin  of  the  zooecia,  but  they  do  not 
correspond  to  what  d'Orbigny  called  "  une  depression  lanceolee  "  which  is  situated 
below  the  aperture.  This  depression  must  correspond  to  the  lanceolate  cryptocyst 
of  d'Orbigny's  figure.  I  found  some  well  preserved  fragments  at  Port  Filiolet 
and  Chef  du  Pont  (Cotentin),  see  pi.  7,  fig.  12. 

This  form  is  conspecific  with  Marsson's  Vincularia  rugica  as  is  proved  by  many 
specimens  from  the  Maastrichtian  Chalk  of  Rugen  and  other  localities  in  northern 
Germany.  Marsson  who  published  the  first  good  description  and  figure  of  this 
species,  was  therefore  unable  to  recognize  that  his  species  was  the  same  as  d'Orbigny's, 
and  it  is  understandable  that,  since  Marsson's  description,  this  form  has  been  re- 
corded only  under  the  name  rugica.  Another  similar  species  is  Vincularia  undata 
d'Orbigny  from  the  Santonian  of  Vendome,  which  also  has  true  opesiules  below  the 
opesia,  not  indicated  in  d'Orbigny's  figure  (d'Orbigny  1851  :  75,  pi.  656,  figs.  10-12) 
but  which  can  be  seen  on  the  holotype. 

The  appearance  of  Micropora  transversa  can  vary  considerably  as  shown  in 
Voigt's  figures  (1951).  The  diameter  of  the  rods  is  from  0-5-0 -7  mm.,  and  the 
number  of  zooecial  rows  varies  between  five  and  ten.  The  length  of  the  normal 
zooecia  is  o  -49-0  -54  mm.  The  basal  attenuated  part  of  some  rods  shows  clearly 
that  this  species  was  articulated  or  radicelled  (cellarif orm) .  This  means  that  this 
form  is  not  congeneric  with  all  true  Micropora,  and  it  could  be  suggested  that  it 
belongs  to  a  new  genus.  But  I  hesitate  to  erect  a  new  genus  for  it  because  it  is 
not  impossible  that  this  form  is  conspecific  with  Dimorphostylus  tetrastichtis  Voigt 
1928.  The  genus  Dimorphostylus  was  established  for  articulated  rods,  in  which  the 
zooecia  are  only  developed  on  one  side.  But  the  size  and  other  features  of  the 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN  33 

zooecia  in  Micropora  rugica  and  in  Dimorphostylus  tetrastichus ,  which  are  often 
associated  together  in  the  same  localities,  are  identical,  and  there  is  the  suspicion 
that  Dimorphostylus  tetrastichus  might  represent  specimens  with  the  zooecia  de- 
veloped only  on  one  side,  and  that  the  differentiation  in  a  frontal  and  reversed  side 
may  be  pathological.  The  two  forms  have  not  yet  been  found  united  in  one  speci- 
men, but  if  they  were,  the  generic  name  Dimorphostylus  must  be  applied  to  Micropora 
transversa  d'Orbigny.  Therefore  it  is  provisionally  here  referred  to  Micropora, 
which  also  indicates  that  it  may  belong  to  the  Microporidae. 

STRATIGRAPHICAL  RANGE.  Maastrichtian ;  Lower  Maastrichtian  of  Riigen  and 
Denmark  and  Upper  Maastrichtian  of  the  Cotentin  (Manche),  Maastricht  and  Ilten 
near  Hanover. 

SPECIMENS.  0.49768.  A  minute  fragment.  From  matrix  of  D.iSi.  Upper 
Maastrichtian,  Cotentin,  Manche,  France. 

0.49769.  A  well  preserved  fragment.  Upper  Maastrichtian,  Chef  du  Pont, 
Cotentin,  Manche,  France.  Voigt  Collection. 


Genus  PUNCTURIELLA  Levinsen  1925 

23.     Puncturiella  cf.  superba  Brydone 

(PL  4,  figs.  8-10) 

1936    Puncturiella  superba  Brydone:  84,  pi.  40,  fig.  18. 

The  small  unilaminar  fragment  with  about  a  dozen  poorly  preserved  zooecia, 
shows  the  cryptocyst  pierced  by  two  outer  and  two  inner  rows  of  pores.  These  are 
barely  visible  because  they  are  partly  obscured  by  recrystallization  of  the  calcite, 
and  the  distal  avicularium  which  is  directed  obliquely  is  very  obscure.  However 
the  identity  of  this  specimen  with  a  few  unilaminar  fragments  from  Fresville  and 
Chef  du  Pont  is  evident  (pi.  4,  figs.  8-9),  and  they  are  probably  identical  with  some 
very  similar  specimens  from  Maastricht  which  are  free  or  encrusting,  but  whose 
dimensions  are  a  little  larger. 

The  average  length  of  zooecia  from  the  Cotentin  localities  is  about  0-65-0-70  mm., 
that  from  Maastricht  0-80-1-00  mm.,  although  some  zooecia  from  the  two  localities 
are  the  same  size,  and  it  may  be  that  the  difference  is  ecological.  The  only  form 
with  which  0.49598  can  be  identified  is  Puncturiella  superba  Brydone  from  the 
Lower  Maastrichtian  Chalk  of  Trimingham  (Norfolk),  in  which  the  zooecia  are  about 
0.65-1  mm.  long. 

STRATIGRAPHICAL  RANGE.     Maastrichtian. 

SPECIMENS.  0.49598.  A  small  worn  fragment  embedded  in  matrix  of  D.iSi. 
Upper  Maastrichtian,  Cotentin,  Manche,  France. 

0.49599.  A  small  fragment.  Upper  Maastrichtian,  Chef  du  Pont,  Cotentin, 
Manche,  France.  Voigt  Collection. 

GEOL.   17,   I.  3 


34          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

0.49600.  A  small  well  preserved  fragment  in  matrix.  Upper  Maastrichtian 
Md,  St.  Pietersberg  near  Maastricht.  Voigt  Collection. 

Genus  LUNULITES  Lamarck 
24.     Lunulites  sp. 

One  very  small  fragment  which  is  worn  and  shows  a  few  zooecia  is  indeterminable. 
It  seems  to  be  conspecific  with  one  of  the  numerous  species  of  Lunulites  represented 
in  the  Maastrichtian  of  the  Cotentin. 

SPECIMENS.  0.49601.  A  very  small  indeterminable  fragment.  In  matrix  of 
D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

Genus  ONYCHOCELLA  Jullien  1881 

25.  Onychocella  nysti  (von  Hagenow) 

(PI.  6,  figs.  3-4) 

1851     Eschar  a  nysti  von  Hagenow:  78,  pi.  9,  figs.  15-17. 

1930     Onychocella  nysti  (von  Hagenow)  Voigt:  459,  pi.  16,  figs.  14-16. 

One  small  fragment  belongs  to  this  species  which  is  very  common  at  all  European 
Maastrichtian  localities  and  which  I  found  at  Port  Filiolet  and  Chef  du  Pont  (Manche) . 
It  has  not  previously  been  recorded  from  the  Cotentin  Maastrichtian. 

STRATIGRAPHICAL  RANGE.     Campanian-Maastrichtian. 

SPECIMENS.  0.49602.  A  worn  fragment.  From  the  matrix  of  D.i8i.  Upper 
Maastrichtian.  Cotentin,  Manche,  France. 

0.49603.  A  well  preserved  fragment.  Upper  Maastrichtian,  Chef  du  Pont, 
Cotentin,  Manche,  France.  Voigt  Collection. 

26.  Onychocella  cf.  cepha  (d'Orbigny) 

(PL  6,  figs.  1-2) 

1851     Eschara  cepha  d'Orbigny:  143,  pi.  670,  figs.  8-10. 

Two  fragments  of  a  narrow  Onychocella  belong  to  a  species  which  is  very  abundant 
near  Chef  du  Pont  (Manche)  and  which  is  not  distinguishable  from  an  Onychocella 
from  the  Maastrichtian  of  Archiac  (Gironde).  Of  the  numerous  species  of  Onycho- 
cella described  by  d'Orbigny  from  the  French  Cretaceous,  the  only  one  which  can  be 
compared  with  this  form  is  Eschara  cepha  d'Orbigny  from  Roy  an,  although  the  type- 
specimen  has  more  slender  zooecia  and  thinner  rims  surrounding  the  zooecia.  Com- 
parison with  the  type  alone  would  suggest  a  different  species,  but  when  compared 
with  all  the  material  from  Archiac  and  Chef  du  Pont,  these  differences  are  much 
diminished  and  identity  is  more  justified  than  the  foundation  of  a  new  species. 

STRATIGRAPHICAL  RANGE.     Maastrichtian. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN  35 

MEASUREMENTS. 

Lz  =  0-48-0-55  mm. 

lz  =  0-30-0-37  mm. 

h0  =  0-10-0-17  mm. 

10  =  0-10-0-13  mm. 

Lav  =  0-70-0-75  mm. 

SPECIMENS.  0.49604.  A  branched  fragment.  From  the  matrix  of  D.iSi. 
Upper  Maastrichtian,  Cotentin,  Manche,  France. 

D .  49605 .  A  small  fragment .  From  the  matrix  of  D .  1 8 1 .  Upper  Maastrichtian , 
Cotentin,  Manche,  France. 

0.49606.  A  fragment.  Upper  Maastrichtian,  Chef  du  Pont,  Cotentin,  Manche, 
France.  Voigt  Collection. 

27.     Onychocella  bellona  (d'Orbigny) 
(PI.  6,  figs.  5-7) 

1851     Eschara  bellona  d'Orbigny:  134,  pi.  668,  figs.  7-9. 
1900     Rhagasostoma  bellona  (d'Orbigny)  Canu  (pars):  431. 

Two  fragments  of  this  broad  bilaminar  species  agree  in  all  details  with  the  material 
collected  by  the  author  at  Chef  du  Pont  (Manche)  and  with  as  pecimen  in  d'Orbigny's 
collection  from  Nehou  (Manche).  (d'Orbigny  Collection  Nr.  7812.) 

The  small  difference  between  the  diameter  of  the  apertures  in  our  figured  specimen 
and  that  of  d'Orbigny  is  irrelevant  because  it  is  also  shown  in  the  specimens  from 
Chef  du  Pont.  The  opesiae  of  the  fertile  zooecia  are  a  little  longer  than  the  others. 
This  species  belongs  to  the  group  of  Onychocella  lamarcki  von  Hagenow,  which  is 
closely  allied  to  it;  but  the  latter  always  shows  more  slender  branches  (1-2-2-0  mm. 
diameter),  and  shorter  and  thicker  zooecia,  and  the  difference  in  size  of  the  fertile 
and  non  fertile  zooecia  is  much  more  evident  than  in  Onychocella  bellona  d'Orbigny. 

It  is  the  same  with  the  fertile  zooecia  of  Onychocella  lamarcki  von  Hagenow  from 
Maastricht,  but  the  zoaria  of  that  species  are  always  flat  and  lamellar. 

MEASUREMENTS. 

Lz  =  0-60-0-70  mm. 

lz  =  0-33-0-45  mm. 
h0  =  0-12-0-18  mm. 

10  =  0-12-0-19  mm. 

Lav  =  0-75-1-00  mm. 

lav  =  0-22-0-27  mm. 

This  species  has  been  found  only  in  the  Cotentin  Maastrichtian. 
STRATIGRAPHICAL  RANGE.     Upper  Maastrichtian. 

SPECIMENS.  0.49607.  A  worn  fragment.  From  the  matrix  of  D.iSi.  Upper 
Maastrichtian,  Cotentin,  Manche,  France. 

GEOL.   17,   I.  3§ 


36          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

0.49608.  A  well  preserved  fragment.  Upper  Maastrichtian,  Chef  du  Pont, 
Cotentin,  Manche,  France.  Voigt  Collection. 

A  well  preserved  fragment  with  ovicelled  zooecia.  Upper  Maastrichtian,  Nehou, 
Cotentin,  France.  d'Orbigny  Collection,  Paris,  Musee  d'Histoire  Naturelle  Nr.  7812. 

Genus  ONYCHOCELLARIA  Voigt  1957 
28.     Onychocellaria  caecilia  (d'Orbigny) 
(PL  5,  %s.  1-6) 

1851     Eschara  caecilia  (d'Orbigny):  138,  pi.  669,  figs.  4-7. 

The  zoarium  was  cellariiform  in  growth.  This  is  proved  by  the  tapering  proximal 
ends  of  the  segments  and  by  pits  occasionally  found  on  the  cryptocyst  in  which 
rootlets  are  inserted.  This  criterion,  combined  with  the  straight  avicularia,  puts  it 
in  the  genus  Onychocellaria,  although  the  endozooecial  ovicell,  which  is  characteristic 
for  this  genus,  has  not  yet  been  observed. 

The  zooecia,  the  length  of  which  is  0-4-0-5  mm.,  are  nearly  rectangular  and  are 
very  often  distinguished  by  a  small  horizontal  band  above  the  distal  rim  of  the 
opesia.  The  slit-like  pit  between  the  opesia  and  this  band  is  very  characteristic, 
and  is  shown  in  d'Orbigny's  figures.  The  opesiae,  which  vary  greatly  in  size,  may 
be  rounded  or  oval  to  high-semicircular.  In  some  segments  the  whole  opesia  or  its 
upper  half  is  closed  by  a  calcareous  lamella. 

MEASUREMENTS. 

Lz  =  0-42-0-51  nun. 

lz  =  0-20-0-25  mm. 
h0  =  o-io-0'70  mm. 

10  =  o-io-o-n  mm. 

LAV  =  0-55-0-58  mm. 

1AV  =  0-25-0-30  mm. 

This  species  is  represented  by  many  small  fragments.  It  is  characteristic  of  the 
Maastrichtian  of  the  Cotentin,  and  has  also  been  found  in  abundance  by  the  present 
author  near  Port  Filiolet  (Manche).  The  specimens  agree  entirely  with  the  original 
specimens  of  d'Orbigny  from  the  Maastrichtian  of  Nehou.  These  are  not  "  use  " 
as  stated  by  Canu  (1900  :  420). 

STRATIGRAPHICAL  RANGE.     Upper  Maastrichtian. 

SPECIMENS.  0.49726.  A  small  fragment  with  avicularia.  From  the  matrix  of 
D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 

0.49727.  Another  fragment  with  avicularium.  From  the  matrix  of  D.iSi. 
Upper  Maastrichtian,  Cotentin,  Manche,  France. 

D. 49728-29.  Two  worn  fragments.  From  the  matrix  of  D.iSi.  Upper  Maas- 
trichtian, Cotentin,  Manche,  France. 

0.49767.  A  small  fragment  corresponding  in  preservation  to  0.49730.  From 
the  matrix  of  D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche,  France. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN          37 

0.49849-54.  6  fragments  from  the  matrix  of  D.iSi.  Upper  Maastrichtian, 
Cotentin,  Manche,  France. 

D .  49848.  Eight  very  worn  small  fragments  from  the  matrix  of  D .  181.  Locality 
and  horizon  as  above. 

D. 49730.  A  well  preserved  fragment  with  some  avicularia.  Upper  Maastrich- 
tian, Chef  du  Pont,  Cotentin,  Manche,  France.  Voigt  Collection. 

D. 49731.  A  well  preserved  fragment.  Upper  Maastrichtian,  Chef  du  Pont, 
Cotentin,  Manche,  France.  Voigt  Collection. 

Genus  PLIOPHLOEA  Gabb  &  Horn  1862 
29.     Pliophloea  sp. 

(PI-  7,  ng.  3) 

Of  this  species  there  is  but  one  encrusting  young  zoarium,  which  is  composed 
of  the  ancestrula  and  a  dozen  zooecia.  The  ancestrula  is  0-20  mm.  long  and  the 
other  zooecia  have  a  length  up  to  0-50  mm.  The  smooth  intraterminal  front  wall 
shows  about  twelve  to  thirteen  costae  which  are  barely  visible.  The  orifice  is  cribri- 
line  to  slightly  pliophloean  according  to  Lang's  definition.  There  is  only  one 
avicularium — if  the  interpretation  as  avicularium  is  correct — which  is  c.  0-20  mm. 
long  and  which  is  close  to  the  ovicelled  zooecium  on  the  right  of  our  figure.  Details 
of  the  spines  are  not  visible.  Pliophloea  gluma  Lang  (1921  :  188,  pi.  6,  fig.  3)  from 
the  Danian  shows  some  affinities  in  the  shape  of  the  zooecia  and  the  apertures,  but 
has  clearly  distinct  costae,  and  visible  intercostal  fusions,  and  the  zooecia  are  only 
0-40  mm.  long.  There  is  no  other  species  hitherto  described  with  which  this  form 
can  be  identified  and  therefore  it  may  be  supposed  that  it  is  a  new  one.  But  this 
one  poor  specimen  does  not  suffice  to  found  a  new  species. 

STRATIGRAPHICAL  RANGE.  This  form  has  not  previously  been  observed  in  the 
Maastrichtian  of  the  Cotentin. 

SPECIMENS.  0.47324.  Small  encrusting  zoarium  with  ancestrula  and  an  ovi- 
celled zooecium.  In  the  matrix  of  Castanopora  jurassica  (Gregory)  D .  180.  Upper 
Maastrichtian,  Cotentin,  Manche,  France. 


Genus  DECURTARIA  Jullien  1886 
(=  Prosoporella  Marsson  1887) 

30.     Decurtaria  cf.  cornuta  (Beissel) 
(PI.  7,  figs.  1-2) 

1865  Semiescharipora  cornuta  Beissel:  58,  pi.  7,  figs.  77-81. 

1887  Prosoporella  cornuta  Marsson :  100. 

1922  Decurtaria  cornuta  (Beissel)  Lang:  388,  text-fig.  124,  pi.  8,  fig.  9. 

1925  Barroisina  trifossata  Levinsen:  387,  pi.  8,  fig.  6. 

1930  Decurtaria  cornuta  (Beissel)  Voigt:  516,  pi.  32,  fig.  6. 


38          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

This  species  is  represented  by  a  fairly  large  zoarium  composed  of  about  thirty 
zooecia.  Unfortunately  it  is  covered  by  a  film  of  minute  calcite-crystals  which 
hides  many  of  the  characteristic  minute  details  of  the  costae  and  the  orifices.  The 
small  number  of  costae  (six  to  seven)  and  the  very  stout  distal  shield  indicate  it  to 
be  Decurtaria  cornuta,  although  the  shape  of  the  orifice  is  more  like  that  of  Decurtaria 
allecta  Lang  (1922  :  386,  pi.  8,  fig.  8)  from  the  Upper  Maastrichtian  of  Maastricht. 
D .  49609  corresponds  exactly  to  a  fragment  from  Chef  du  Pont  (Manche) .  In  some 
respects  that  form  might  be  regarded  as  intermediate  between  the  two  species,  which 
are,  indeed,  more  alike  than  is  stated  by  Lang  who  had  only  one  specimen  of  his 
D.  allecta  from  Maastricht.  Decurtaria  cornuta  is  not  confined  to  the  Lower  Maas- 
trichtian as  might  be  concluded  from  Lang's  work.  I  found  a  small  but  typical 
fragment  of  this  species  in  the  Upper  Maastrichtian  of  Biebosch  (South  Limburg, 
Netherlands) . 

It  seems  that  the  shape  of  the  orifice  is  not  so  constant  as  was  assumed  by  Lang. 
According  to  him,  in  D.  cornuta  the  orifice  should  be  "  super-cribriline  "  and  in  D. 
allecta  "  supernormal  ".  But  Beissel  had  already  stated  that  the  different  types  of 
orifice  shape  may  be  observed  in  the  same  zoarium,  and  he  has  figured  both  forms 
in  his  figs.  77  and  78.  The  size  of  the  zooecia  is  between  0-57  and  0-65  mm.  Barroi- 
sina  trifossata  Levinsen  is  a  synonym  of  this  species  which  I  concluded  from  the 
examination  of  Levinsen's  type  specimen. 

STRATIGRAPHICAL  RANGE.     Maastrichtian. 

SPECIMENS.  0.49609.  A  complete  zoarium  with  ancestrula  and  some  ovicelled 
zooecia.  From  the  matrix  of  D.iSi.  Upper  Maastrichtian,  Cotentin,  Manche, 
France. 

An  incomplete  zoarium.  Upper  Maastrichtian,  Chef  du  Pont,  Cotentin,  Manche, 
France.  Voigt  Collection,  Hamburg,  Nr.  4137. 

Genus  FRURIONELLA  Canu  &  Bassler  1926 
30.     Frurionella  fertilis  sp.  n. 

(PI.  8,  figs.  5-7) 

HOLOTYPE.  0.49610.  Upper  Maastrichtian,  Chef  du  Pont,  Cotentin,  Manche, 
France.  Collection  E.  Voigt. 

DERIVATIO  NOMINIS.  Derived  from  fertilis  =  generative,  on  account  of  the 
numerous  ovicells. 

DIAGNOSIS.  A  slender  species  of  Frurionella,  with  bilaminar  branches  consisting 
of  three  to  four  alternating  rows  of  zooecia.  Zooecia  not  distinct,  small,  with 
quadrangular  to  high  triangular  or  semicircular  opesiae  whose  proximal  margin  is 
straight  and  generally  provided  with  an  inconspicuous  proximal  lip  caused  by  the 
swelling  of  the  median  avicularium  below  the  opesia.  Elliptical  or  linear  indistinct 
pores  which  may  be  avicularia  are  developed  in  the  median  axis  of  the  zooecium. 
Ovicells  numerous,  forming  deep  characteristic  cavities  above  the  apertures  if  the 
covering  lamina  has  been  destroyed. 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN  39 

MEASUREMENTS. 

Lz  =  0-50-0-62  mm. 
h0  =  0-10-0-12  mm. 
10  =  o -10-0 -12  mm. 

REMARKS.  A  small  fragment,  showing  a  scarcely  constricted  opesia  with  a 
minute  avicularium  below  it,  has  been  recognized  as  belonging  to  this  new  species 
which  is  represented  from  Chef  du  Pont  by  two  other  similar  specimens.  This 
form  cannot  be  identified  with  any  of  the  few  known  species  of  Frurionella.  It  is 
smaller  in  its  zoarial  and  zooecial  dimensions  than  the  other  species  of  this  genus. 
At  first  it  was  supposed  that  it  might  be  the  slender  distal  branch  of  another  new 
species  of  Frurionella  which  is  much  larger  and  very  common  at  Fresville  (Cotentin, 
Manche).  This  is  improbable  because  there  are  no  intermediate  stages  between 
these  two  very  different  forms  and  no  distal  branches  smaller  than  2-1-5  mm.  seem 
to  exist  in  the  second  species.  This  other  species  of  Frurionella  from  Fresville 
(pi.  8,  fig.  8)  is  conspecific  with  Frurionella  europaea  Voigt  (1951  :  60,  pi.  9,  figs.  1-3) 
from  the  Upper  Maastrichtian  of  Ilten  (Hannover)  and  Kunrade  (Netherlands). 
It  has  now  also  been  found  at  Maastricht  (Netherlands). 

STRATIGRAPHICAL  RANGE.     Maastrichtian. 
SPECIMENS.     0.49610.     Holotype  see  above. 

D .  49611.  A  small  worn  fragment  in  matrix.  From  the  matrix  of  D .  181.  Upper 
Maastrichtian,  Cotentin,  Manche,  France. 

0.49612.  A  fragment  with  broken  ovicelled  zooecia.  Horizon  and  locality  as 
above. 


Genus  BEISSELINA  Canu  1913 
32.     Beisselina  striata  (Goldfuss) 
(PI.  8,  figs.  3-4) 

1826     Eschara  striata  Goldfuss:  25,  pi.  8,  fig.  16. 

1960     Beisselina  striata  (Goldfuss)  Wiesemann:  27,  pi.  i,  figs.  1-3,  pi.  2,  figs.  3-4;  pi.  12,  figs  3-4; 

text-figs.  36,  4  9-10;  5,  Nrs.  12-13  (with  additional  synonymy). 
1967     Beisselina  striata  (Goldfuss)  Voigt:  72,  pi.  25,  fig.  i. 

One  specimen  belongs  to  this  common  Maastrichtian  species  which  is,  according 
to  Wiesemann,  represented  near  Chef  du  Pont  (Manche) .  Nevertheless  the  diameter 
of  the  stem  (1-5  mm.)  and  the  orifices  (peristomicia)  are  a  little  smaller  (0-08-0-12 
mm.)  than  in  most  specimens  from  the  type  locality  of  Maastricht,  and  Kunrade, 
although  the  length  of  the  zooecia  (distance  between  proximal  and  distal  apertures) 
is  the  same.  Specimens  with  such  small  orifices  are  not  lacking,  and  all  intermediate 
sizes  occur  at  these  localities,  but  they  are  rare.  For  comparison  see  pi.  8,  fig.  4, 
showing  the  surface  of  a  worn  specimen  from  Maastricht  itself.  These  small  forms 
are  more  common  in  the  "  Tuffeau  de  St.  Symphorien  "  in  the  Basin  of  Mons  (Bel- 
gium.) 


4o          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

STRATIGRAPHICAL  RANGE.     Maastrichtian. 

SPECIMENS.  0.49721.  A  worn  fragment.  From  the  matrix  of  D.i8i.  Upper 
Maastrichtian,  Cotentin,  Manche,  France. 

0.49722.  A  worn  fragment.  Upper  Maastrichtian,  Geulem,  near  Berg,  Geul 
valley  near  Maastricht  (Netherlands).  Voigt  Collection. 

Genus  BEISSELINOPSIS  Voigt  1951 
33.     Beisselinopsis  flabellata  (d'Orbigny) 
(PL  8,  figs.  1-2) 

1852     Escharifora  flabellata  d'Orbigny:  460,  pi.  715,  figs.  10-12. 
non  1930     Beisselina flabellata  (d'Orbigny)  Voigt:  525,  pi.  34,  fig.  n. 

One  incomplete  young  zoarium  represents  d'Orbigny's  species  which  is  common 
at  Sainte  Colombe  (Manche)  and  the  type  of  which  from  this  locality  was  studied 
by  the  present  author.  There  are  some  specimens  from  Chef  du  Pont  (Manche) 
which  agree  very  well  with  Beisselinopsis  flabellata  d'Orbigny  although  they  are 
more  elongate  than  the  flabelliform  zoarium  of  d'Orbigny's  type  specimen.  The 
species  identified  as  Porina  flabellata  (d'Orbigny)  by  Marsson  (1887  :  85)  is  Beisselin- 
opsis marginata  v.  Hagenow  1839  (cf.  Voigt  1959  :  n,  pi.  9,  figs.  1-2).  The  similar 
Danian  and  Montian  form  determined  as  Porina  or  Beisselina  flabellata  (d'Orbigny) 
by  Levinsen  (1925,  pi.  7,  fig.  83)  and  Voigt  (1930  partim  :  525,  pi.  34,  fig.  n)  is 
neither  conspecific  nor  congeneric  and  corresponds  to  Eschara  oblita  Kade  (1852  :  29, 
pi.  i,  fig.  18)  as  stated  by  Berthelsen  (1962  :  201,  pi.  24,  fig.  6)  and  Voigt  (1964  :  458, 
pi.  8,  fig.  8  and  pi.  14,  figs.  1-3)  and  belongs  to  the  genus  Pavobeisselina  Voigt  1964. 
It  was  formerly  assigned  to  Beisselinopsis  Voigt  1951,  but  this  attribution  was 
incorrect  because  Beisselinopsis  has  no  ascopore  which  can  always  be  observed  in 
Pavobeisselina.  The  inner  structure  of  Beisselinopsis  flabellata  d'Orbigny  does  not 
show  any  ascopore  in  the  frontal  wall  as  is  always  the  case  in  Beisselina  and  flabelli- 
form Pavobeisselina.  Therefore  it  must  be  regarded  as  a  true  Beisselinopsis. 

STRATIGRAPHICAL  RANGE.     Upper  Maastrichtian. 

SPECIMENS.  0.49723.  A  young  zoarium.  From  the  matrix  of  D.iSi.  Upper 
Maastrichtian,  Cotentin,  Manche,  France. 

An  adult  zoarium.  Upper  Maastrichtian,  Chef  du  Pont,  Cotentin,  Manche, 
France.  Voigt  Collection,  Hamburg,  Nr.  3908. 


II.    BRACHIOPODA 

Genus  THECIDEA  Def ranee  1832 

Thecidea  papillata  (von  Schlottheim) 

(PI.  i,  figs.  9-10) 

1959     Thecidea  papillata  von  Schlotheim;  Backhaus:  21,  pi.  i,  figs.  1-4  (see  for  all  references). 


UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN          41 

TABLE  I. 

List  of  the  species  from  the  matrix  of  "  Castanopora  "  jurassica  (Gregory)  (D .  180) 
and  Onychocella  piriformis  (Goldfuss)  (= Onychocella  bathonica  (Gregory)  (D.iSi). 


I.  POLYZOA 


1.  Berenicea  sp. 

2.  Entalophora  benedeniana  (von  Hagenow)  1851 

3.  Idmidronea  macilenta  (von  Hagenow)  1851 

4.  Heterocrisina  communis  (d'Orbigny)  1853 

5.  Crisisina  carinata  (Roemer)  1840 

6.  Osculipora  truncata  (Goldfuss)  1826 

7.  Reteporidea  lichenoides  (Goldfuss)  1826 

8.  Petalopora  sp. 

9.  Multicrescis  laxata  (d'Orbigny)  1854 

10.  Membranipora  unipora  (Marsson) 

11.  Amphiblestrella  elegans  von  (Hagenow)  1851 

12.  Radulopora  minor  n.  sp. 

13.  Hagenowinella  cf.  incrassata  (d'Orbigny)  1853 

14.  Bactrellaria  rugica  Marsson  1887 

15.  Stamenocella  marginata  (d'Orbigny)  1852 

1 6.  Thyracella  cf.  meudonensis  d'Orbigny  1851 

17.  Vincularia  canalifera  von  Hagenow  1851 

18.  Quadricellaria  elegans  (d'Orbigny)  1951 

19.  Coscinopleura  sp. 

20.  Semiescharinella  complanata  d'Orbigny  1852 

21.  Rosseliana  thomasi  n.  sp. 

22.  Micropora  transversa  (d'Orbigny)  1851 

23.  Puncturiella  cf.  superba  Brydone  1936 

24.  Lunulites  sp. 

25.  Onychocella  nysti  (von  Hagenow)  1851 

26.  Onychocella  cf.  cepha  (d'Orbigny)  1851 

27.  Onychocella  bellona  (d'Orbigny)  1851 

28.  Onychocellaria  caecilia  (d'Orbigny)  1851 

29.  Pliophloea  sp. 

30.  Decurtaria  cf.  cornuta  (Beissel)  1865 

31.  Frurionella  fertilis  n.  sp. 

32.  Beisselina  striata  (Goldfuss)  1926 

33.  Beisselinopsis  flabellata  (d'Orbigny)  1852 

II.  BRACHIOPODA 

34.  Thecidea  papillata  (von  Schlottheim)  1813 
*  After  d'Orbigny. 


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42          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

Two  small  valves  of  this  very  common  and  characteristic  brachiopod  of  the 
"  Craie  a  Thecidees  "  were  found  in  the  matrix  of  D.iSi.  They  are  indistinguish- 
able from  those  of  the  Cotentin  Maastrichtian  collected  by  the  author.  The  collec- 
tion of  the  "  Geologisches  Staatsinstitut  "  Hamburg  possesses  about  850  specimens 
from  Chef  du  Pont,  Fresville  and  Port  Filiolet,  which  were  studied  in  the  monograph 
by  Backhaus.  The  locality  "  Port  Fiolet  "  (Backhaus  1959  :  27  and  Hofker  1959  : 
369,  380,  381)  should  be  "  Port  Filiolet ". 

STRATIGRAPHICAL  RANGE.     Maastrichtian,  mainly  Upper  Maastrichtian. 

SPECIMENS.     66.42981.     Pedicle  valve.     From  the  matrix  of  D.iSi.     Upper 
Maastrichtian,  Craie  a  Thecidees,  Cotentin,  Manche,  France. 
66.42982.     Small  dorsal  valve.     Horizon  and  locality  as  above. 

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44          UPPER  CRETACEOUS  POLYZOA  FROM  COTENTIN 

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EXPLANATION    OF   PLATES 

Some  of  the  specimens  of  Bryozoa  and  Brachiopoda  referred  to  and  figured  in 
this  publication  are  housed  in  the  collections  of  the  British  Museum  (Natural  History) 
and  are  prefixed  by  the  letters  D  and  BB  respectively. 


PLATE    i 
Petalopora  sp.  (p.  18) 

FIG.  i.  Worn  branch  with  narrow  ribs  between  the  small  mesopores.  From  the  matrix  of 
D.iSi,  Upper  Maastrichtian  (Cotentin,  Manche).  X25.  0.49573. 

FIG.  2.  Branch  in  similar  condition  to  fig.  i  with  larger  mesopores  and  peristomes.  Upper 
Maastrichtian,  Port  Filiolet  (Cotentin,  Manche).  Presented  by  the  author.  X25.  0.49574. 

FIG.  3.  Well  preserved  branch.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche). 
X25-  Voigt  Collection,  Hamburg,  Nr.  3969. 

Crisisina  carinata  (Roemer)  (p.  17) 

FIG.  4.  Worn  fragment  with  damaged  peristomes,  lateral  view.  From  the  matrix  of  D.  181 
Upper  Maastrichtian  (Cotentin,  Manche).  X25.  0.49566. 

FIG.  5.  Worn  fragment,  frontal  view.  Upper  Maastrichtian,  Port  Filiolet  (Cotentin, 
Manche).  Presented  by  the  author.  X25.  0.49567. 

Multicrescis  laxata  d'Orbigny  (p.  18)  (See  also  pi.  4  figs.  4-6) 

FIG.  6.  Worn  fragment  0.49575  with  encrusting  Rosseliana  thomasi  n.  sp.  (0.47323)  in 
matrix  of  Castanopora  jurassica  (Gregory)  (D.iSo).  Upper  Maastrichtian  (Cotentin,  Manche). 

X20. 

FIG.    7.     Holotype.     Upper   Maastrichtian,    Sainte    Colombe    (Cotentin,    Manche)    in    coll. 
d'Orbigny  Nr.  8416,  Musee  d'Histoire  Naturelle,  Paris,      x  12. 
FIG.  8.     Holotype.      X25. 

Thecidea  papillata  von  Schlottheim  (p.  40) 

FIG.  9.  Pedicle  (ventral)  valve  with  growth-facet  on  the  apex,  with  the  cast  of  a  small 
lamellibranch  on  the  right.  From  the  matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin, 
Manche),  attesting  the  origin  from  the  "  Craie  a  Thecidees  "  of  the  Cotentin-region.  c.  X  12. 
BB.  42981. 

FIG.  10.  Small  dorsal  valve  slightly  damaged.  From  the  matrix  of  D.iSi.  c.  X 12. 
86.42982. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  i 


PLATE i 


8 


GEOL.   17,   I. 


4§ 


PLATE   2 
Hagenowinella  cf.  incrassata  (d'Orbigny)  (p.  23) 

FIG.  i.  Part  of  the  encrusting  zoarium  showing  an  avicularium  in  the  left  upper  corner  and 
some  damaged  zooecia  in  the  proximal  region.  From  the  matrix  of  D.  181.  Upper  Maastrich- 
tian  (Cotentin,  Manche).  x  20.  0.49583. 

FIG.  2.  Worn  specimen  with  broken  ovicells.  The  deeply  immersed  horseshoe-like  processes 
in  the  interior  of  the  zooecia  can  hardly  be  seen.  Upper  Maastrichtian,  Fresville  (Cotentin, 
Manche).  Presented  by  the  author.  X2o.  0.49845. 

FIG.  3.  Unilaminar  fragment  with  large  opesia.  Some  ovicells  and  an  avicularium  in  the 
lower  row  of  zooecia  are  visible.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  x  20 
Voigt  Collection,  Hamburg,  Nr.  3909. 

Thyracella  cf.  tneudonensis  (d'Orbigny)  (p.  26) 

FIG.  4.  Branched  narrow  fragment  with  an  avicularium,  for  comparison  with  fig.  5.  Upper 
Maastrichtian,  Port  Filiolet,  (Cotentin,  Manche).  x  20.  Voigt  Collection,  Hamburg,  Nr.  3919- 

FIG.  5.  Damaged  fragment  of  the  bilaminar  zoarium.  From  the  matrix  of  D.iSi.  Upper 
Maastrichtian  (Cotentin,  Manche).  X2O.  0.49588. 

FIG.  6.  Worn  fragment,  d'Orbigny  Collection,  Paris  Nr.  8040,  Musee  d'Histoire  Naturelle. 
labelled  Meudon  but  likely  Upper  Maastrichtian,  locality  uncertain  (?  Neliou,  Cotentin, 
Manche).  X2o. 

"  Membranipora  unipora  "  Marsson  (p.  19) 

FIG.  7.  Fragment  with  hyperstomial  ovicells  and  median  pores  on  the  gymnocyst  of  the 
undamaged  zooecia.  The  minute  marks  of  spines  are  hidden  by  recrystallization.  From  the 
rock  matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin,  Manche).  x  20.  0.49576. 

FIG.  8.  Holotype,  of  Flustrellaria  simplex  d'Orbigny  Collection,  Nr.  8130,  Musee  d'Histoire 
Naturelle  Paris,  Upper  Maastrichtian,  N6hou  (Cotentin,  Manche).  x  20. 

Bactrcllaria  rugica  Marsson  (p.  24) 

FIG.  9.  Small  worn  fragment  showing  the  pits  of  the  avicularia  below  the  opesium.  From 
the  matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin,  Manche).  X2O.  0.49584. 

FIG.  10.     The  same  specimen,  showing  the  backside,      x  20. 

FIG.  ii.  Fragment  with  well  preserved  avicularia.  Upper  Maastrichtian,  Port  Filiolet 
(Cotentin,  Manche).  Presented  by  the  author,  x  20.  0.49585. 

FIG.  12.  Worn  fragment  in  similar  condition  as  figs.  9-10.  Upper  Maastrichtian,  Port 
Filiolet  (Cotentin,  Manche).  x  20.  Voigt  Collection,  Hamburg,  ^.3914. 

FIG.  13.  Well  preserved  fragment  with  raised  avicularia  whose  rostra  are  damaged.  From 
chalk  of  a  flint-boulder  of  Upper  Maastrichtian  age,  Wulmstorf  near  Harburg-Hamburg, 
Northern  Germany,  x  20.  Voigt  Collection,  Hamburg,  Nr.  4146. 

FIG.  14.  The  same  as  fig.  13,  lateral  view,  showing  the  long  tubular  peduncles  of  the  avicu- 
laria. x  20. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  i 


PLATE  2 


PLATE   3 
Stamenocella  marginata  d'Orbigny  (p.  25) 

FIG.  i .  Fragment  of  the  bilaminar  zoarium  in  the  matrix  of  Onychocella  piriformis  (Goldf  uss) 
(D.iSi).  Upper  Maastrichtian  (Cotentin,  Manche).  X  20.  0.47322. 

FIG.  2.  Similar  fragment.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  Pre- 
sented by  the  author.  X2O.  0.49586. 

FIG.  3.  Holotype  of  Biflustra  tesselata  d'Orbigny.  Upper  Maastrichtian,  Nehou  (Cotentin, 
Manche).  d'Orbigny  Collection,  Nr.  8053,  Paris  Musee  d'Histoire  Naturelle.  X2o. 

FIG.  4.  Holotype  of  Biflustra  marginata  d'Orbigny.  Upper  Maastrichtian,  Sainte  Colombe 
(Cotentin,  Manche).  d'Orbigny  Collection,  Nr.  8134,  Paris  Musee  d'Histoire  Naturelle.  X  20. 

FIG.  5.  Well  preserved  large  fragment  showing  the  different  aspect  and  size  of  zooecia  and 
avicularia.  The  basal  region  represents  Flustrella  convexa  d'Orbigny.  Upper  Maastrichtian, 
Chef  du  Pont  (Cotentin,  Manche).  X20.  Voigt  Collection,  Hamburg,  Nr.  4128. 

Radtilopora  minor  n.  sp.  (p.  22) 

FIG.  6.  Small  worn  bilaminar  fragment  from  the  matrix  of  D.iSi.  Upper  Maastrichtian, 
(Cotentin,  Manche).  X2o.  0.49579. 

FIG.  7.  Fragment  showing  the  gradual  tapering  of  the  zoarium.  Upper  Maastrichtian, 
Chef  du  Pont  (Cotentin,  Manche).  Presented  by  the  author.  X2O.  0.49580. 

FIG.  8.  Another  fragment.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  Pre- 
sented by  the  author.  X2O.  0.49581. 

FIG.  9.  Broad  fragment,  most  zooecia  with  ovicells  on  the  right  side  with  anen  crusting 
Foraminifer.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  Presented  by  the 
author.  X2o.  0.49582. 

FIG.  10.  Holotype,  showing  the  larger  marginal  zooecia  at  the  edge  of  the  zoarium.  Upper 
Maastrichtian,  St.  Pietersberg  near  Maastricht  (Netherlands),  x  20.  Presented  by  the  author. 
0.49844. 

Radulopora  radula  (Marsson)  (p.  21) 

FIG.  ii.  Fragment  of  the  thick  bilaminar  zoarium  for  comparison  with  Radulopora  minor 
n.  sp.,  showing  the  larger  zooecia  and  avicularia  and  some  fertile  zooecia  with  broken  ovicells. 
Upper  Maastrichtian  Chalk  in  drift,  Tornesch  near  Elmshorn  (Schleswig-Holstein,  Northern 
Germany).  X2O.  Voigt  Collection,  Hamburg  Nr.  3974. 

FIG.  12.  Singular  vibraculum,  x  120,  showing  the  thornlike  process  which  is  curved  inwards. 
Horizon,  locality  and  collection  as  for  fig.  n. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  i 


PLATEs 


11 


8 


PLATE   4 
Amphiblestrella  elegans  (v.  Hagenow)  (p.  20) 

FIG.  i.  Small  worn  fragment  from  the  matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin, 
Manche).  x  20.  0.49577. 

FIG.  2.  Holotype  of  Flustrella  baculina  d'Orbigny.  Upper  Maastrichtian,  N6hou  (Cotentin, 
Manche).  d'Orbigny  Collection,  Paris,  Nr.  8127,  Musee  d'Histoire  Naturelle.  x  20. 

FIG.  3.  Branched  fragment.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  Pre- 
sented by  the  author.  X2O.  0.49578. 

Rosseliana  thomasi  n.  sp.  (p.  31)  (see  also  pi.  i,  fig.  6) 

FIG.  4.  Holotype,  showing  the  encrusting  zoarium  upon  a  branched  fragment  of  Radulopora 
minor  n.  sp.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  Presented  by  the 
author,  x  20.  1X49597. 

FIG.  5.  Same  specimen,  part  of  the  zoarium  x  40,  showing  the  form  of  the  opesium  and  some 
fertile  zooecia  with  ovicells. 

FIG.  6.  Zoarium  encrusting  an  echinoid  fragment.  Upper  Maastrichtian,  Port  Filiolet 
(Cotentin,  Manche).  X2o.  Voigt  Collection,  Hamburg,  Nr.  3932. 

Floridina  scut  at  a  (Levinsen)  (p.  31) 

FIG.  7.  Part  of  encrusting  zoarium,  for  comparison  with  Rosseliana  thomasi  n.  sp.  Upper 
Maastrichtian  chalk  boulder  in  drift,  Tornesch  near  Elmshorn  (Schleswig-Holstein,  Northern- 
Germany).  X40.  Voigt  Collection,  Hamburg,  ^.4145. 

Puncturiella  cf .  superba  Brydone  (p.  33) 

FIG.  8.  Small  unilaminar  fragment  in  matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin, 
Manche).  X2O.  0.49598. 

FIG.  9.  Unilaminar  fragment.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche). 
Presented  by  the  author,  x  20.  D .  49599. 

FIG.  10.  Unilaminar  fragment,  well  preserved.  Upper  Maastrichtian,  St.  Pietersberg  near 
Maastricht  (Netherlands) .  Presented  by  the  author,  x  20.  D .  49600. 

Setniescharinella  complanata  d'Orbigny  (p.  29) 

FIGS.  II-I2.  Small  badly  preserved  unilaminar  fragment  from  the  matrix  of  D.  181.  Upper 
Maastrichtian  (Cotentin,  Manche).  x  20.  0.49595  and  49596. 

FIG.  13.  Fragment  with  well  preserved  zooecia  and  distal  avicularia.  Upper  Maastrichtian, 
Chef  du  Pont  (Cotentin,  Manche).  Presented  by  the  author.  X2o.  0.49847. 

FIG.  14.  Small  fragment  showing  one  zooecium  with  ovicell  in  the  right  lower  corner.  Upper 
Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche) .  x  20.  Voigt  Collection,  Hamburg,  Nr. 
39i8b. 

FIG.  15.  Holotype  of  Escharinella  simplex  d'Orbigny.  Upper  Maastrichtian,  N6hou  (Coten- 
tin, Manche).  d'Orbigny  Collection,  Paris,  Nr.  7942,  Mus6e  d'Histoire  Naturelle.  x  20. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  i 


PLATE4 


PLATE   5 
Onychocellaria  caecitia  (d'Orbigny)  (p.  36) 

FIG.  I.  Fragment  of  a  well  preserved  segment  with  some  avicularia  on  the  edges  of  the 
slightly  flattened  rods.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  x  20. 
Presented  by  the  author.  0.49730. 

FIG.  2.     Same  specimen  seen  from  the  narrow  side  with  two  avicularia.      x  20. 

FIG.  3.  Another  fragment  with  closed  zooecia  in  the  proximal  region  and  larger  opesia. 
Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  Presented  by  the  author,  x  20. 

0.49731- 

FIG.  4.  Small  fragment  with  avicularium.  From  the  matrix  of  D.iSi.  Upper  Maastrich- 
tian, Chef  du  Pont  (Cotentin,  Manche).  X2O.  0.49726. 

FIG.  5.  Another  fragment  with  avicularium  in  the  middle  row  of  zooecia.  From  the  matrix 
of  D.iSi.  Upper  Maastrichtian  (Cotentin,  Manche).  X2o.  Photographed  under  water. 
0.49727. 

FIG.  6.  Broad  worn  fragment.  From  the  matrix  of  D .  1 81 .  Upper  Maastrichtian  (Cotentin, 
Manche).  X2O.  Photographed  under  water.  0.49767. 

Vincularia  canalifera  (von  Hagenow)  (p.  27)  (See  also  pi.  8,  figs.  9-12.) 

FIG.  7.  Proximal  part  of  a  well  preserved  segment,  showing  more  oval  opesia  and  some 
zooecia  with  the  characteristic  openings  for  radicell  filaments  at  the  lower  end.  Upper  Maas- 
trichtian, Md.,  St.  Pietersberg  near  Maastricht  (Netherlands).  X  20.  Presented  by  the 
author.  D .  49840. 

FIG.  8.  Fragment  with  some  avicularia  in  the  distal  and  proximal  part,  showing  more 
semicircular  opesia  than  fig.  7.  Upper  Maastrichtian,  Md.,  St.  Pietersberg  near  Maastricht 
(Netherlands).  X2O.  Presented  by  the  author.  0.49841. 

FIG.  9.  Worn  fragment.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  x  20. 
Presented  by  the  author.  0.49734. 

FIG.  10.  Worn  fragment.  From  the  matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin, 
Manche).  x  20.  0.49733. 

Onychocella  pirifortnis  (Goldfuss)  (p.  5) 

FIG.  ii.  Fragment  of  the  unilaminar  frond  with  an  avicularium  in  the  lower  left  corner  for 
comparison  with  fig.  12.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  x  20. 
Voigt  Collection,  Hamburg,  Nr.  3573. 

FIG.  12.  Holotype  of  Onychocella  bathonica  Gregory,  erroneously  labelled  as  "  Bathonian 
Ranville  (Calvados,  France)  ",  but  certainly  from  the  Upper  Maastrichtian  of  the  Cotentin 
(Manche).  X2O.  D.iSi. 


Bull.  Br.  Mm.  nat.  Hist.  (Geol.)  17,  i 


PLATEs 


PLATE   6 
Onychocella  cf.  cepha  d'Orbigny  (p.  34) 

FIG.  i.  Branched  fragment  with  some  avicularia  from  the  matrix  of  D.iSi.  Upper  Maas- 
trichtian  (Cotentin,  Manche).  x  20.  0.49604. 

FIG.  2.  Fragment  for  comparison  with  fig.  i.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin, 
Manche).  X2o.  Presented  by  the  author.  0.49606. 

Onychocella  nysti  (v.  Hagenow)  (p.  34) 

FIG.  3.  Worn  fragment  from  the  matrix  of  D .  181.  Upper  Maastrichtian  (Cotentin,  Manche). 
X20.  0.49602. 

FIG.  4.  Well  preserved  specimen  with  two  ovicells.  Upper  Maastrichtian,  Chef  du  Pont 
(Cotentin,  Manche).  X2O.  Presented  by  the  author.  0.49603. 

Onychocella  bellona  (d'Orbigny)  (p.  35) 

FIG.  5.  Well  preserved  fragment  with  several  ovicells.  Upper  Maastrichtian,  Nehou  (Coten- 
tin, Manche).  x  20.  d'Orbigny  Collection,  Paris,  Nr.  7812. 

FIG.  6.  Fragment  from  the  Upper  Maastrichtian  from  Chef  du  Pont  (Cotentin,  Manche) 
intermediate  between  fig.  5  and  fig.  7.  Presented  by  the  author,  x  20.  0.49608. 

FIG.  7.  Worn  fragment  from  the  matrix  of  D.  181.  Upper  Maastrichtian  (Cotentin,  Manche). 
X40.  0.49607. 

Quadricellaria  elegans  d'Orbigny  (p.  29) 

FIG.  8.  Fragment  of  a  segment.  Upper  Maastrichtian  (Cotentin,  Manche).  From  the 
matrix  of  D.iSi.  x  20.  0.49591. 

FIG.  9.  Another  worn  fragment.  Upper  Maastrichtian  (Cotentin,  Manche).  From  the 
matrix  of  D.iSi.  x  20.  0.49592. 

FIG.  10.  Fragment  with  a  large  avicularium.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin, 
Manche).  X4O.  Voigt  Collection,  Hamburg,  Nr.  4133. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  i 


PLATE6 


10 


PLATE   7 
Decurtaria  cf.  cornuta  (Beissel)  (p.  37) 

FIG.  i.  Zoarium  with  ancestrula  and  one  ovicelled  zooecium.  From,  the  matrix  of  D.iSi. 
Upper  Maastrichtian  (Cotentin,  Manche).  x  20.  0.49609. 

FIG.  2.  Incomplete  zoarium  with  some  ovicelled  zooecia.  Upper  Maastrichtian,  Chef  du 
Pont  (Cotentin,  Manche).  X2O.  Voigt  Collection,  Hamburg,  ^.4137. 

Pliophloea  sp.  (p.  37) 

FIG.  3.  Young  zoarium  with  ancestrula  and  one  ovicelled  zooecium.  In  the  matrix  of 
Castanopora  jurassica  (Gregory)  D.iSo.  Upper  Maastrichtian  (Cotentin,  Manche).  X2o. 
D.47324. 

Castanopora  jurassica  (Gregory)  (p.  7) 

FIG.  4.  Bilaminar  fragment  with  two  ovicelled  zooecia.  Upper  Maastrichtian,  Chef  du 
Pont  (Cotentin,  Manche).  Presented  by  the  author.  X2O.  0.49724. 

FIG.  5.  Same  specimen  from  the  other  side,  showing  one  zooecium  with  preserved  frontal 
wall,  x  20. 

FIG.  6.  Fragment  with  some  zooecia.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin, 
Manche).  x  20.  Voigt  Collection,  Hamburg,  ^.3585. 

FIG.  7.  Part  of  a  young  zoarium  with  smaller  zooecia.  Upper  Maastrichtian,  Chef  du  Pont 
(Cotentin,  Manche).  Presented  by  the  author.  X2O.  0.49725. 

"  Membranipora  "  constricta  d'Orbigny  (p.  13) 

FIG.  8.  Badly  preserved  specimen  encrusting  a  Brachiopod.  Aptian  les  Croutes  (Yonne, 
France)  ?  Holotype,  d'Orbigny  Collection,  Paris,  Mus6e  d'Histoire  Naturelle,  Nr.  5691.  X2O. 

Micropora  transversa  (d'Orbigny)  (p.  32) 

FIG.  9.  Small  worn  fragment  showing  the  two  opesiular  pores  below  the  opesium.  From  the 
matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin,  Manche).  x  20.  0.49768. 

FIGS.  lo-n.  Two  fragments  of  the  Holotype.  Upper  Maastrichtian,  Neliou  (Cotentin, 
Manche).  X2O.  d'Orbigny  Collection,  Paris,  Musee  d'Histoire  Naturelle,  Nr.  7755. 

FIG.  12.  Fragment,  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  X2O.  Pre- 
sented by  the  author.  0.49769. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  i 


PLATE7 


PLATE   8 
Beisselinopsis  flabellat a  (d'Orbigny)  (p.  40) 

FIG.  i.  Adult  zoarium.  Upper  Maastrichtian,  Chef  du  Pont  (Cotentin,  Manche).  x  20. 
Voigt  Collection,  Hamburg,  Nr.  3908. 

FIG.  2.  Young  worn  zoarium.  From  the  matrix  of  D .  181.  Upper  Maastrichtian  (Cotentin, 
Manche).  X2o.  D.  49723. 

Beisselina  striata  (Goldfuss)  (p.  39) 

FIG.  3.  Worn  fragment.  From  the  matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin, 
Manche).  x  20.  0.49721. 

FIG.  4.  Worn  fragment  for  comparison  with  fig.  3.  Upper  Maastrichtian,  Md.,  Geulem 
near  Berg,  Geul-valley  near  Maastricht  (Netherlands).  Presented  by  the  author,  x  20. 
D.  49722. 

Frurionella  fer tilts  n.  sp.  (p.  38) 

FIG.  5.  Holotype,  well  preserved  fragment  with  ovicelled  zooecia.  Upper  Maastrichtian, 
Chef  du  Pont  (Cotentin,  Manche).  Presented  by  the  author,  x  20.  0.49610. 

FIG.  6.  Fragment  showing  zooecia  with  damaged  ovicells.  Upper  Maastrichtian,  Chef  du 
Pont  (Cotentin,  Manche).  Presented  by  the  author.  X2O.  0.49612. 

FIG.  7.  Worn  fragment,  similar  to  fig.  6.  From  the  matrix  of  D .  181.  Upper  Maastrichtian 
(Cotentin,  Manche).  x  20.  0.49611. 

Frurionella  europaea  (Voigt)  (p.  39) 

FIG.  8.  For  comparison  with  Frurionella  fertilis  n.  sp.  Upper  Maastrichtian,  Chef  du  Pont 
(Cotentin,  Manche).  Presented  by  the  author,  x  20.  0.49720. 

Vincularia  canalifera  (v.  Hagenow)  (p  27)  (See  also  pi.  5,  figs.  7-10) 

FIG.  9.  Small  fragment.  From  the  matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin, 
Manche).  x  20.  0.49735. 

FIG.  10.  Badly  preserved  branched  fragment  showing  an  avicularium  in  the  middle  of  the 
right  side.  From  the  matrix  of  D.iSi.  Upper  Maastrichtian  (Cotentin,  Manche).  x  20. 
0.49736. 

FIG.  1 1 .  Fragment  showing  an  avicularium  at  the  right  upper  corner.  Upper  Maastrichtian 
Md.,  Geulem  near  Berg,  Geul-valley  near  Maastricht  (Netherlands) .  x  20.  D .  49842.  E.  Voigt 
Collection. 

FIG.  12.  Part  of  a  branched  stem,  for  comparison  with  fig.  9,  showing  relatively  small  zooecia 
and  opesia.  Upper  Maastrichtian,  locality  as  fig.  n.  0.49732. 

Vincularia  concinna  (d'Orbigny)  (p.  28) 

FIG.  13.  Holotype.  Branched  fragment  for  comparison  with  Vincularia  canalifera  von 
Hagenow.  Upper  Maastrichtian,  Nehou  (Cotentin,  Manche).  x  20.  d'Orbigny  Collection, 
Paris,  Musee  d'Histoire  Naturelle,  Nr.  7756. 

Vincularia  flexuosa  (d'Orbigny)  (p.  28) 

FIG.  14.  Holotype.  Fragment  for  comparison  with  Vincularia  canalifera  von  Hagenow. 
Labelled  as  Santonian  Venddme,  Loir  et  Cher,  France,  x  20.  d'Orbigny  Collection,  Paris, 
Mus6e  d'Histoire  Naturelle,  Nr.  7752. 


Bull.  BY.  Mus.  nat.  Hist.  (Geol.)  17,  i 


PLATES 


PRINTED  IN  GREAT  BRITAIN 
BY  ADLARD  &  SON  LIMITED 
BARTHOLOMEW  PRESS,  DORKING 


THE  CAUDAL  SKELETON  IN 
MESOZOIC  ACANTHOPTERYGIAN 

FISHES 


COLIN  PATTERSON 


BULLETIN  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

GEOLOGY  Vol.  17  No.  2 

LONDON:  1968 


THE  CAUDAL  SKELETON  IN  MESOZOIC 
ACANTHOPTERYGIAN  FISHES 


» 


BY 


COLIN  PATTERSON 

British  Museum  (Natural  History) 


Pp.  47-102;   28  Text-figures 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
GEOLOGY  Vol.  17  No.  2 

LONDON:  1968 


THE  BULLETIN  OF  THE  BRITISH  MUSEUM 
(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they  become 
ready.  Volumes  will  contain  about  three  or  four 
hundred  pages,  and  will  not  necessarily  be  completed 
within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
Papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  17,  No.  2  of  the  Geological 
(Palaeontological)  series.  The  abbreviated  titles  of 
periodicals  cited  follow  those  of  the  World  List  of 
Scientific  Periodicals. 


World  List  abbreviation: 
Bull.  Br.  Mus.  nat.  Hist.  (Geol.) 


Trustees  of  the  British  Museum  (Natural  History)  1968 


TRUSTEES    OF 
THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 

Issued  19  November,  1968  Price 


THE  CAUDAL  SKELETON  IN  MESOZOIC 
ACANTHOPTERYGIAN  FISHES 

By  COLIN  PATTERSON 

CONTENTS 

Page 

I.     INTRODUCTION          .........  49 

II.     SYSTEMATIC  DESCRIPTIONS          .......  50 

Order  Ctenothrissiformes     .......  50 

Order  Beryciformes    .           .          .          .          .          .          .          .  56 

Suborder  Polymixioidei         .          .          .          .          .          .  56 

Suborder  Dinopterygoidei     ......  62 

Suborder  Berycoidei     .......  67 

Order  Lampridiformes          .......  78 

Order  Salmoniformes,  Suborder  Myctophoidei         .          .          .  81 

III.  DISCUSSION      ..........  83 

(a)  The  relationships  of  Ctenothrissiformes,  Myctophoidei  and 

Beryciformes     ........  83 

(b)  The  origin  of  Perciformes       ......  87 

(c)  Intermediate  groups      .......  96 

IV.  CONCLUSIONS  ..........  98 

V.     REFERENCES   ..........  100 

VI.     ABBREVIATIONS  USED  IN  FIGURES         ......         102 

SYNOPSIS 

This  paper  contains  descriptions  of  the  caudal  skeleton  in  all  Ctenothrissiformes  and  Mesozoic 
acanthopterygians,  comparisons  with  living  relatives  being  made  where  possible,  and  a  brief 
account  of  the  caudal  skeleton  in  Mesozoic  Myctophoidei.  These  groups  have  a  basically  similar 
caudal  skeleton  with  the  first  ural  and  pre-ural  centra  fused,  the  second  ural  centrum  free,  six 
hypurals,  three  epurals,  a  stegural  and  a  second  uroneural.  Caudal  scutes  persist  in  Ctenothris- 
siformes and  primitive  Myctophoidei  and  there  is  a  single  urodermal  in  some  Cretaceous 
myctophoids. 

The  bearing  of  caudal  structure  on  the  origin  and  early  evolution  of  acanthopterygians  is 
discussed  and  variations  in  the  second  pre-ural  neural  spine  are  shown  to  be  important.  This 
structure  is  primitively  short  and  slender:  two  opposite  developments  from  this  condition  are 
elongation  of  the  spine  to  support  procurrent  fin-rays  (as  in  Polymixioidei)  and  reduction  to  a 
low  crest  (as  in  Berycoidei  and  Perciformes).  Wherever  a  full  second  pre-ural  neural  spine 
occurs  in  perciform  and  higher  groups  it  is  produced  secondarily  by  fusion  with  the  first  epural. 
The  evidence  of  the  caudal  skeleton  does  not  support  a  polyphyletic  origin  of  Perciformes  from 
different  groups  of  Beryciformes.  The  Berycoidei  appear  to  be  the  beryciform  group  most 
closely  related  to  the  Perciformes. 

The  Danian  Bathysoma  is  shown  to  be  a  lampridiform :  the  lampridiforms  probably  arose  from 
near  Aipichthys  and  Pharmacichthys.  The  Zeiform.es  are  close  relatives  of  the  Berycoidei. 

I.  INTRODUCTION 

IN  an  earlier   paper  (Patterson  1964)  I  gave  detailed    accounts  of   the   skeletal 
anatomy  of  the  known  Mesozoic  acanthopterygian  fishes  (see  also  Patterson  1967) 
but  without  describing  the  caudal  skeleton.     This  was  omitted  because  when  the  bulk 
GEOL.  17,  2  5 


50  THE   CAUDAL   SKELETON   IN 

of  the  work  was  done  there  existed  no  comprehensive  description  or  even  terminology 
of  the  caudal  skeleton  of  living  teleosts.  But  during  the  last  few  years,  following 
from  the  work  of  Gosline  (1960),  the  caudal  skeleton  has  become  recognized  as  an 
important  structure  in  tracing  the  relationships  of  teleost  fishes  and  we  now  have 
both  a  satisfactory  terminology  (Gosline  1960;  Nybelin  1963;  Monod  1967; 
Patterson  1968)  and  a  reasonable  idea  of  the  composition  of  the  caudal  skeleton 
in  most  teleostean  groups  (Gosline  1960,  1961,  ig6ia,  1963,  1965;  Norden  1961; 
Rosen  1962,  1964;  Greenwood,  Rosen,  Weitzman  &  Myers  1966;  Monod  1967; 
Greenwood  1967;  Weitzman  1967).  In  the  light  of  this  new  information  it  seemed 
that  an  investigation  of  the  caudal  skeleton  in  Mesozoic  acanthopterygians  might 
prove  a  useful  check  on  hypotheses  of  the  origins  of  acanthopterygians  and  par- 
acanthopterygians  (Greenwood  et  al.  1966) .  Also  included  are  accounts  of  caudal 
structure  in  the  Ctenothrissiformes  and,  more  briefly,  in  the  Mesozoic  Myctophoidei. 

The  terminology  used  here  is  that  of  Nybelin  (1963)  with  the  distinction  between 
"  urodermal  "  and  "  uroneural  "  introduced  by  Patterson  (1968)  and  the  addition 
from  Monod  (1967)  of  the  terms  "  parhypural  ",  for  the  haemal  arch  of  the  first 
pre-ural  centrum,  the  terminal  structure  perforated  by  the  caudal  vein  and  artery 
and  which  bears  the  hypurapophysis  (Nursall  1963),  and  "  stegural  "  for  the  paired 
structure  which  articulates  with  the  dorso-lateral  surface  of  the  first  pre-ural  centrum 
and  has  a  slender  shaft  extending  postero-dorsally,  lateral  to  the  nerve  cord.  Monod 
considers  the  stegural  to  be  an  element  sui  generis,  not  a  compound  structure. 
In  my  opinion  (Patterson  1968),  the  stegural  represents  the  first  uroneural  fused 
with  neural  arch  material  from  the  first  pre-ural  and  first  ural  vertebrae.  The  term 
stegural  is  nevertheless  useful,  since  it  obviates  repetition  of  the  cumbersome  term 
"  first  uroneural  fused  with  the  neural  arches  of  the  first  ural  and  pre-ural  centra  ". 

The  material  used  is  mainly  in  the  collections  of  the  British  Museum  (Natural 
History) :  these  specimens  are  referred  to  by  a  registered  number  without  prefix 
or  with  the  prefix  "  P  ".  Specimens  from  the  American  University,  Beirut,  the 
American  Museum  of  Natural  History,  New  York,  the  Royal  Scottish  Museum, 
Edinburgh,  and  the  United  States  National  Museum,  Washington,  are  referred  to 
with  the  prefixes  "  AUB  ",  "  AMNH  ",  "  RSM  "  and  "  USNM  "  respectively.  The 
illustrations  of  fossil  species  are  not  reconstructions  but  camera  lucida  drawings  of 
single  specimens. 

I  am  grateful  to  Drs.  P.  H.  Greenwood  and  D.  E.  Rosen,  who  have  read  and 
criticized  parts  of  this  paper  in  manuscript,  and  for  the  loan  of  specimens  to  Prof. 
T.  Raven,  American  University,  Beirut,  Drs.  C.  D.  Waterston  and  R.  S.  Miles,  Royal 
Scottish  Museum,  Dr.  Bobb  Schaeffer,  American  Museum  of  Natural  History,  and 
Drs.  D.  H.  Dunkle  and  D.  M.  Cohen,  United  States  National  Museum. 

II.  SYSTEMATIC  DESCRIPTIONS 

Order  CTENOTHRISSIFORMES  (Patterson  1964  :  218) 

According  to  Marshall  (1961)  the  living  Macristium  chavesi  is  a  surviving  cteno- 
thrissiform  (see  also  Greenwood,  Rosen,  Weitzman  &  Myers  1966;  Patterson  1967  : 
86).  Unfortunately,  the  only  extant  specimen  of  Macristium  is  a  post-larval 


MESOZOIC   ACANTHOPTERYGIAN    FISHES  51 

individual  and  the  caudal  skeleton  is  not  ossified.  But  there  appear  to  be  six  hypurals, 
as  in  Ctenothrissiformes.  Berry  &  Robins  (1967)  have  described  a  second  macristiid, 
Macristiella  perlucens,  known  only  by  a  single  post-larval  specimen.  In  this  fish 
there  are  six  hypurals,  as  in  Ctenothrissiformes,  but  Berry  &  Robins  think  it  unlikely 
that  the  Macristiidae  are  close  to  the  Ctenothrissiformes. 

Family  AULOLEPIDIDAE  Patterson  (1964  :  247) 
Genus  PATEROPERCA  Smith  Woodward  (1942  :  543) 

The  type  and  only  species,  P.  libanica  Smith  Woodward  from  the  Middle  Ceno- 
manian  of  Hajula,  Lebanon,  is  known  only  by  two  specimens,  AUB  108906  (the 
holotype)  and  AUB  108904.  In  108904  the  caudal  region  is  badly  preserved  and 
yields  no  useful  information,  but  in  the  holotype  the  caudal  skeleton  is  very  well 
preserved  (Fig.  i).  The  second  pre-ural  centrum  (pu2]  has  a  fully  developed  neural 
spine  (npu2)  and  an  autogenous  haemal  arch.  It  is  not  possible  to  see  with  certainty 
whether  the  haemal  arch  of  the  third  pre-ural  centrum  (Pu3)  is  autogenous.  The 
first  pre-ural  and  first  ural  centra  are  fused  (pu i  +  ui) ,  these  fused  centra  bearing 
the  parhypural  (ph)  and  the  first  and  second  hypurals  (hi,  h2)  in  the  normal  way. 
The  first  hypural  is  rather  slender,  only  a  little  broader  than  the  parhypural  and 


u2 


npu2 
pu1+u1 


FIG.  i.  Pateroperca  libanica  Smith  Woodward.  Caudal  skeleton  of  the  holotype,  AUB 
108906,  standard  length  86  mm.,  Middle  Cenomanian,  Hajula,  Lebanon.  For  explanation 
of  lettering  see  p.  102.  Arrows  mark  the  outermost  (unbranched)  principal  fin-rays. 


52  THE   CAUDAL   SKELETON   IN 

the  second.  Dorsally,  the  fused  first  pre-ural  and  ural  centra  bear  a  large  stegural 
(st)  and  there  is  a  second  uroneural  (d2)  ending,  as  usual,  in  front  of  the  base  of  the 
first  unbranched  principal  ray  of  the  caudal  fin.  There  are  three  epurals  (ei  —  j), 
the  third  small  and  short.  There  is  a  small  independent  second  ural  centrum  (u2) 
bearing  three  hypurals  (hj  —  5)  decreasing  in  size  upwards,  and  above  these  there 
is  a  small  sixth  hypural  (h6)  which  probably  failed  to  make  contact  with  the  ural 
centrum.  The  foremost  procurrent  rays  of  the  caudal  fin,  which  are  unsegmented 
lepidotrichia,  not  spines,  are  inserted  in  front  of  the  neural  and  haemal  spine  of  the 
third  pre-ural  centrum.  The  uppermost  principal  ray  of  the  fin  (unbranched)  is 
preceded  by  nine  rays,  only  the  last  two  or  three  segmented,  and  the  lowermost  by 
seven  rays,  the  last  two  segmented. 


FIG.  2.  Aulolepis  typus  Agassiz.  Caudal  skeleton  of  47932,  standard  length  c.  165  mm., 
Lower  Chalk,  Burham,  Kent.  /.  r,  foremost  procurrent  fin-ray  (displaced) ;  for  explana- 
tion of  other  lettering  see  p.  102.  The  upper  hypurals  are  displaced  ventrally,  the 
epurals  and  second  uroneural  are  missing. 


MESOZOIC   ACANTHOPTERYGIAN   FISHES 


53 


Genus  AULOLEPIS  Agassiz  (1844  :  109) 

In  the  type  and  only  species,  A.  typus  Agassiz  from  the  Upper  Cenomanian  of 
the  English  Chalk,  only  two  specimens  show  any  details  of  the  caudal  skeleton, 
4033  (the  holotype,  Fig.  3)  and  47932  (Fig.  2),  and  in  both  the  bones  are  disturbed 
to  some  extent.  The  caudal  skeleton  of  Aulolepis  agrees  with  that  of  Pateroperca 
in  most  respects :  fusion  of  the  first  pre-ural  and  first  ural  centra  (pui  +  ui) ,  presence 
of  a  free  second  ural  centrum  (u2],  three  epurals  (ei  —  3),  six  hypurals  (hi  —  6) 
of  which  the  first  and  the  third  are  largest,  an  autogenous  haemal  arch  on  the  second 
pre-ural  centrum  (pu2),  a  stegural  (st)  and  a  free  second  uroneural  (d2).  The  haemal 
arch  of  the  third  pre-ural  centrum  (pu^]  appears  to  be  autogenous  in  47932  but  fused 
with  the  centrum  in  4033.  Aulolepis  differs  clearly  from  Pateroperca  in  having  the 
neural  spine  of  the  second  pre-ural  centrum  developed  as  a  low,  broad  crest  (npu2) 
whose  hind  edge  fits  between  the  front  edges  of  the  stegurals.  In  my  earlier  des- 
criptions of  Aulolepis  and  Pateroperca  (Patterson  1964  :  247)  I  discussed  the  difficulty 
of  separating  the  two  genera  and  considered  the  possibility  that  P.  libanica  is  merely 


npu2 


d2 


pu1+u1 


FIG.  3.  Aulolepis  typus  Agassiz.  Caudal  skeleton  of  the  holotype,  4033,  standard  length 
c.  135  mm.,  Lower  Chalk,  Lewes,  Sussex.  For  explanation  of  lettering  see  p.  102.  The 
epurals  and  upper  hypurals  are  displaced  ventrally,  the  stegural  is  displaced  forwards. 


54 


THE   CAUDAL   SKELETON   IN 


a  species  of  Aulolepis,  but  the  presence  on  the  second  pre-ural  centrum  of  a  fully 
developed  neural  spine  in  Pateroperca  and  of  a  low  crest  in  Aulolepis  clearly  separates 
the  two  genera.  The  caudal  of  47932  shows  one  other  point  of  interest,  a  slender, 
elongated  caudal  scute  (/.  s,  Fig.  2)  in  front  of  the  upper  lobe  of  the  fin.  There  is 
no  conclusive  evidence  of  a  similar  structure  in  front  of  the  lower  lobe  of  the  fin, 
but  it  was  probably  present  since  there  is  a  caudal  scute  in  front  of  each  lobe  of  the 
fin  in  Ctenothrissa  (see  below)  and  in  most  living  teleosts  which  still  retain  these 
structures  (Elops,  Tarpon,  Albula,  Aulopus  and  Chanos,  Gosline  1965  :  192;  Argen- 
tina, Weitzman  1967  :  532).  There  is  no  sign  of  caudal  scutes  in  either  specimen  of 
Pateroperca,  but  again  it  is  probable  that  they  were  present.  No  specimen  of  Aulol- 
epis has  the  fin-rays  of  the  caudal  fin  well  preserved  but  4033  shows  that  the  foremost 
procurrent  rays  of  the  lower  lobe  articulated  with  the  haemal  spine  of  the  third 
pre-ural  centrum,  and  both  4033  and  47932  show  that  these  foremost  rays  were  true 
spines. 

Family  CTENOTHRISSIDAE  Smith  Woodward  (1901) 
Genus  CTENOTHRISSA  Smith  Woodward  (1899  :  490) 

In  Ctenothrissa,  the  only  genus  of  the  Ctenothrissidae,  the  caudal  skeleton  and  fin 
are  well  exposed  in  the  two  species  from  the  Cenomanian  of  the  Lebanon,  C.  vexillifer 


e3 


u2 


pu1+u1 


FIG.  4.  Ctenothrissa  signifer  Hay.  Caudal  skeleton  of  P.  47524,  standard  length  83  mm., 
Middle  Cenomanian,  Hajula,  Lebanon.  For  explanation  of  lettering  see  p.  102.  Arrows 
mark  the  outermost  (unbranched)  principal  fin-rays. 


MESOZOIC   ACANTHOPTERYGIAN    FISHES 


55 


(Pictet  1850),  the  type  species,  and  C.  signifer  Hay  (1903)  (Fig.  4),  and  the  caudal 
skeleton  is  preserved  in  one  or  two  specimens  of  C.  microcephala  (Agassiz)  (Fig.  5) 
and  C.  radians  (Agassiz)  from  the  Upper  Cenomanian  of  the  English  Chalk.  There 
seem  to  be  no  significant  differences  in  caudal  anatomy  between  these  four  species. 
As  in  Aulolepididae,  there  is  a  free  second  ural  centrum  (u2),  the  first  ural  and  first 
pre-ural  centra  are  fused  (pui  +  ui],  there  are  six  hypurals  (hi  —  6),  the  first  and 
third  the  largest  and  the  sixth  failing  to  articulate  with  the  second  ural  centrum, 
there  are  three  epurals  (ei  —  j),  a  stegural  (st)  and  a  second  uroneural  (£2)  and  the 
haemal  arch  of  the  second  pre-ural  centrum  (pu2)  is  autogenous.  The  haemal  arch 
of  the  third  pre-ural  centrum  (pu>3)  appears  partially  or  completely  fused  to  the 
centrum.  As  in  Aulolepis,  but  in  contrast  to  Pateroperca,  the  neural  spine  of  the 
second  pre-ural  centrum  (npu2)  is  represented  by  a  broad  crest,  about  half  as  high 
as  the  preceding  spine.  There  is  a  slender  caudal  scute  (/.  s.)  in  front  of  both  the 
upper  and  lower  lobes  of  the  caudal  fin.  In  C.  vexillifer  and  C.  signifer,  the  only 
species  in  which  the  caudal  fin-rays  are  well  preserved,  both  the  upper  and  lower 
principal  rays  are  preceded  by  six  rays,  the  last  two  segmented,  and  the  foremost 
fin-rays  are  inserted  on  the  neural  and  haemal  spines  of  the  third  pre-ural  centrum. 


e3 


e1 


npu2 


2mm 


FIG.  5.  Ctenothrissa  microcephala  (Agassiz).  Caudal  skeleton  of  49881,  standard  length 
c.  1 15  mm.,  Lower  Chalk,  zone  of  Holaster  subglobosus,  Reigate,  Surrey.  For  explanation 
of  lettering  see  p.  102.  The  upper  hypurals  are  displaced  ventrally.  Of  the  two  elements 
labelled  ?  h6,  that  on  the  left  may  be  the  base  of  a  fin-ray. 


56  THE   CAUDAL   SKELETON   IN 

In  these  small  species  from  the  Lebanon,  the  foremost  procurrent  caudal  rays  are 
unsegmented  lepidotrichia,  with  separate  right  and  left  halves,  but  in  the  larger 
species  from  the  English  Chalk  they  appear  to  be  true  spines. 

Order  BERYCIFORMES 

Suborder  POLYMIXIOIDEI  Patterson  (1964  :  433) 
Family  POLYMIXIIDAE  Gill  (1862) 

The  only  living  genus  of  this  family  and  suborder  is  Polymixia.  The  caudal 
skeleton  of  Polymixia  nobilis  has  been  figured  by  Regan  (1911,  fig.  i)  and  briefly 
discussed  by  Gosline  (1961  :  14).  The  specimen  illustrated  here  (Fig.  6)  is  almost 
certainly  that  used  by  Regan.  The  haemal  arches  of  the  second  and  third  pre-ural 
centra  (pu2,  pus)  are  autogenous,  the  second  pre-ural  centrum  has  a  fully  developed 
neural  spine  (npu2) ,  the  first  pre-ural  and  first  ural  centra  are  fused  (pui  +  ui)  and 
there  is  a  free  second  ural  centrum  (14,2).  There  is  a  large  stegural  (st)  and  a  free 
second  uroneural  (d2).  There  are  three  slender  epurals  (ei  —  j)  and  six  hypurals 
(hi  —  6)  of  which  the  first  and  the  fourth  are  the  largest,  the  third  being  excavated 
posteroventrally  to  give  a  notch  between  the  hypurals  supporting  the  upper  and 
lower  lobes  of  the  fin.  The  second  ural  centrum  has  a  long  posterior  process  and 
makes  contact  with  the  sixth  hypural.  The  first  rays  of  the  caudal  fin  articulate 
with  the  neural  and  haemal  spines  of  the  third  pre-ural  centrum.  In  the  upper  lobe 
of  the  fin  the  first  unbranched  principal  ray  is  preceded  by  four  true  spines  and  two 


e1-3 


npu2 


pu1+u1 


u2 


FIG. 


3mm 


6.     Polymixia  nobilis  Lowe.     Caudal  skeleton   of  a   dried   skeleton,    1895.5.28.1, 
standard  length  220  mm.,  Madeira.     For  explanation  of  lettering  see  p.  102. 


MESOZOIC   ACANTHOPTERYGIAN   FISHES 

segmented  rays,  in  the  lower  lobe  by  three  spines  and  two  segmented  rays, 
caudal  skeleton  of  P.  japonicus  does  not  differ  from  that  of  P.  nobilis. 


57 


Genus  BERYCOPSIS  Dixon  (1850  :  372) 

In  the  type  species,  B.  elegans  Dixon  from  the  Upper  Cenomanian  and  Turonian 
of  the  English  Chalk,  the  caudal  skeleton  is  well  preserved  in  25881  (Fig.  7)  and 
P. 6465.  The  caudal  skeleton  of  B.  elegans  agrees  with  that  of  Polymixia  in  almost 
every  detail  except  that  the  third  hypural  (Aj)  is  not  excavated  postero-ventrally 
so  that  it  is  as  large  as  the  fourth,  the  haemal  arch  of  the  third  pre-ural  centrum, 
probably  autogenous  in  P. 6465,  is  fused  with  the  centrum  in  25881,  and  the  third 
and  fourth  hypurals,  separate  in  25881  (standard  length  c.  13  cm.),  are  completely 
fused  in  the  larger  P. 6465  (standard  length  c.  16  cm.).  In  25881  the  lateral  surface 
of  the  second  ural  centrum  is  covered  by  a  small,  triangular  plate  (x,  Fig.  7). 
P. 6465  is  not  sufficiently  well  preserved  to  see  whether  this  plate  is  present.  The 
plate  appears  to  be  part  of  the  caudal  skeleton,  not  a  dermal  element  which  has 
become  apposed  to  the  centrum  post  mortem.  The  only  record  of  a  similar  structure 
is  in  the  living  myctophoid  Synodus  foetens,  where  Hollister  (193 ja,  figs.  5,  12)  found 
a  "  uroneural  "  in  this  position,  ventral  to  the  normal  uroneurals.  The  foremost 


h6 


h5 


npu2 


pul+ul 


FIG.  7.  Berycopsis  elegans  Dixon.  Caudal  skeleton  of  25881,  standard  length  c.  130  mm., 
Chalk,  Sussex,  x,  unidentified  plate  covering  second  ural  centrum;  for  explanation  of 
other  lettering  see  p.  102. 


58  THE   CAUDAL   SKELETON   IN 

unsegmented  caudal  fin  rays  of  B.  elegans  are  soft  rays,  not  spines  as  they  are  in 
Polymixia. 

Of  P.  1047 1,  a  small  fish  lacking  the  head,  I  rashly  stated  that  it  is  "  certainly 
B.  elegans  "  (Patterson  1964  :  278),  but  on  re-examining  this  specimen  I  find  that  the 
scales  are  larger  than  in  B.  elegans,  the  dorsal  and  anal  spines  are  longer,  there  are 
five  anal  spines  (not  four  as  stated  earlier)  and  in  the  caudal  skeleton  the  neural 
spine  of  the  second  pre-ural  centrum  is  much  reduced.  This  specimen  is  clearly  not 
B.  elegans  and  the  reduced  neural  spine  of  the  second  pre-ural  centrum  shows  that 
it  is  not  even  a  polymixioid,  but  there  is  insufficient  evidence  to  decide  on  its  real 
position. 

The  second  species  of  Berycopsis,  B.  germanus  (Agassiz)  from  the  Campanian  of 
Westphalia,  shows  nothing  in  the  caudal  skeleton  to  distinguish  it  from  B.  elegans 
except  that  the  haemal  arch  of  the  third  pre-ural  centrum  is  clearly  autogenous  and 
there  is  no  sign  of  fusion  between  the  third  and  fourth  hypurals  in  the  two  specimens 
where  this  region  is  visible.  B.  germanus  is  so  preserved  that  it  shows  the  details 
of  the  caudal  fin-rays  much  better  than  B.  elegans.  The  foremost  rays  of  the  caudal 
fin  are  arranged  asymmetrically,  the  upper  rays  articulating  with  the  neural  spine 
of  the  second  pre-ural  centrum,  the  lower  with  the  haemal  spine  of  the  third  pre-ural 
centrum.  The  first  principal  ray  (unbranched)  of  the  upper  lobe  is  preceded  by 
four  unsegmented  and  one  segmented  ray,  the  lower  by  three  unsegmented  and  one 
segmented  ray.  It  is  not  possible  to  see  whether  the  foremost  rays  are  spines  (as 
in  Polymixia)  or  soft  rays,  as  in  B.  elegans  and  Omosoma. 

Genus  OMOSOMA  Costa  (1857  :  Io6) 

Having  now  had  the  opportunity  to  make  a  direct  comparison  between  the  holo- 
types  of  Omosoma  pulchellum  (Davis  1887)  (RSM  1891.59.72)  and  0.  intermedium 
Smith  Woodward  (1901)  (48112)  I  find  that  the  two  are  conspecific,  the  median  fin 
counts  (D  V,  28-30;  A  III-IV,  24)  being  the  same  in  both.  The  apparent  differences 
in  proportions  of  the  two  (Smith  Woodward  1901  :  420)  are  due  to  longitudinal 
compression  of  the  holotype  of  0.  pulchellum,  the  holotype  of  0.  intermedium  showing 
the  true  proportions  of  the  fish.  In  my  earlier  description  of  Omosoma  (Patterson 
1964  :  374)  the  median  fin  counts  given  for  0.  pulchellum  (D  IV-V,  35 ;  A  III-IV,  29) 
were  based  on  distorted  specimens  of  0.  sahelalmae.  0.  intermedium  is  therefore  a 
synonym  of  0.  pulchellum. 

None  of  the  specimens  of  Omosoma  available,  belonging  to  0.  sahelalmae  Costa 
(the  type  species)  and  0.  pulchellum  (Davis),  from  the  Upper  Santonian  of  Sahel 
Alma,  Lebanon  (Ejel  &  Dubertret  1966,  have  recently  found  evidence  of  the  precise 
age  of  these  beds),  has  the  caudal  skeleton  sufficiently  well  preserved  to  be  illustrated, 
but  all  give  a  picture  of  a  caudal  skeleton  which  does  not  differ  significantly  from  that 
of  Berycopsis.  The  foremost  rays  of  the  caudal  fin  are  arranged  asymmetrically, 
those  of  the  upper  lobe  articulating  with  the  neural  spine  of  the  second  pre-ural 
centrum,  those  of  the  lower  lobe  with  the  haemal  spine  of  the  third  pre-ural  centrum, 
just  as  in  B.  germanus.  In  both  the  lobes  of  the  fin  there  are  4  unsegmented 
lepidotrichia  (not  spines)  and  two  segmented  rays  in  front  of  the  principal  rays. 


MESOZOIC   ACANTHOPTERYGIAN   FISHES 


59 


It  is  impossible  to  see  whether  the  haemal  arch  of  the  third  pre-ural  centrum  is 
autogenous. 

Genus  PYCNOSTERINX  Heckel  (1849  :  337) 

Pycnosterinx  is  known  by  four  species,  all  from  the  Upper  Santonian  of  Sahel 
Alma,  Lebanon.  Figure  8  shows  the  caudal  skeleton  of  P.  russeggerii  Heckel,  the 
type  species:  it  is  very  similar  to  those  of  the  other  polymixiids  and  differs  from 
Polymixia  (Fig.  6)  only  in  having  the  postero-ventral  part  of  the  third  hypural  com- 
pletely ossified,  so  that  there  is  no  gap  between  the  hypurals  supporting  the  upper 
and  lower  lobes  of  the  fin.  As  in  Polymixia  the  neural  and  haemal  spines  are  strongly 
inclined  and  the  epurals  slender.  As  in  Berycopsis  and  Omosoma,  but  in  contrast 
to  Polymixia,  the  foremost  caudal  fin-rays  are  arranged  asymmetrically,  the  upper 
ones  articulating  with  the  neural  spine  of  the  second  pre-ural  centrum,  the  lower 
with  the  haemal  spine  of  the  third  pre-ural  centrum.  In  both  the  upper  and  lower 
lobes  of  the  fin  the  principal  rays  are  preceded  by  three  unsegmented  soft  rays  and 
three  segmented  rays.  P.  discoides  Heckel  and  P.  gracilis  Davis  do  not  differ  in 
caudal  structure  from  P.  russeggerii  except  that  in  P.  gracilis  there  appear  to  be 
only  five  procurrent  rays.  I  have  seen  no  specimens  of  P.  dubius  Davis  in  which 
the  caudal  skeleton  and  fin  are  well  preserved,  but  in  the  holotype  of  Pycnosterinx 
latus  Davis  (1887  :  534,  pi.  27,  fig.  2),  RSM  1891.59.77  (referred  to  as  a  "frag- 
mentary fish  of  indeterminable  genus  "  by  Smith  Woodward  1901  :  395),  the  scales 
have  the  same  spinous  surface  as  in  P.  dubius  (Patterson  1964  :  380)  and  it  is  probable 
that  the  specimen  is  a  large,  distorted  individual  of  P.  dubius,  although  the  state  of 


u2 


pu1>u1 


2mm 


FIG.  8.  Pycnosterinx  russeggerii  Heckel.  Caudal  skeleton  of  47820,  standard  length 
78  mm.,  Upper  Santonian,  Sahel  Alma,  Lebanon.  For  explanation  of  lettering  see 
p.  102. 


6o 


THE   CAUDAL   SKELETON   IN 


preservation  of  this  and  the  holotype  of  P.  dubius  is  such  that  this  cannot  be  demon- 
strated with  sufficient  certainty  to  synonymize  the  two  species.  P.  latus  appears 
to  have  one  or  two  more  anal  fin-rays  and  caudal  vertebrae  than  the  holotype  of 
P.  dubius.  The  caudal  skeleton  is  completely  exposed  in  the  holotype  of  P.  latus 
and  does  not  differ  from  that  of  P.  russeggerii  in  any  way,  but  both  in  this  specimen 
and  in  P.  dubius  the  foremost  fin-rays  are  true  spines,  a  difference  from  the  other 
species  of  Pycnosterinx  and  a  resemblance  to  the  living  Polymixia. 

Genus  HOMONOTICHTHYS  Whitley  (1933  :  146) 

All  three  species  of  this  genus  occur  in  the  Upper  Cenomanian — Lower  Turonian 
of  the  English  Chalk.  In  the  type  species,  H.  dorsalis  (Dixon),  only  one  specimen, 
43575,  shows  anything  of  the  caudal  skeleton  and  here  it  is  only  possible  to  see  that 
there  was  a  fully  developed  neural  spine  on  the  second  pre-ural  centrum  and  that 
the  foremost  rays  of  the  upper  caudal  lobe  are  true  spines,  articulating  with  the 
neural  spine  of  the  third  pre-ural  centrum.  In  these  last  two  features  H.  dorsalis 


e2 


e3 


d2 


2mm 


FIG.  9.     Homonotichthys  pulchellus  (Dixon).  Caudal  skeleton  of  P.  11112,  standard  length 
c.  no  mm.,  Lower  Chalk,  Amberley,  Sussex.     For  explanation  of  lettering  see  p.  102. 


MESOZOIC   ACANTHOPTERYGIAN   FISHES  61 

resembles  Polymixia  and  differs  from  other  Cretaceous  polymixiids  except 
Pycnosterinx  dubius. 

In  H.  pulchellus  (Dixon)  the  caudal  skeleton  is  preserved  in  the  holotype,  25886, 
and  in  two  specimens  only  recently  recognized  as  belonging  to  this  species,  P.  11112 
(Fig.  9)  and  P.  10639:  the  latter  are  about  no  and  120  mm.  respectively  in  standard 
length,  the  largest  examples  of  this  species  yet  recorded.  The  caudal  skeleton  of 
P.IIH2  (Fig.  9)  is  abnormal  in  the  partial  doubling  of  the  neural  spine  of  the  second 
pre-ural  centrum  and  in  having  the  second  epural  smaller  than  the  third.  The 
caudal  skeleton  of  H.  pulchellus  is  very  like  that  of  Polymixia,  with  autogenous 
haemal  arches  on  the  second  and  third  pre-ural  centra  (pu2,  pu3)  six  hypurals 
(hi  —  6),  three  slender  epurals  (ei  —  3),  a  free  second  ural  centrum  (u2),  a  stegural 
(st)  and  a  second  uroneural  (d,2}.  The  postero- ventral  corner  of  the  third  hypural 
is  truncated,  but  less  so  than  in  Polymixia.  Broad  flanges  on  the  anterior  margin 
of  the  neural  spines  of  the  second  and  third  pre-ural  centra  are  characteristic  of 
H.  pulchellus.  All  three  specimens  show  that  the  foremost  caudal  rays  are  spines, 
as  in  H.  dorsalis  and  Polymixia.  In  P .  10639  the  caudal  rays  articulate  with  the 
neural  and  haemal  spines  of  the  third  pre-ural  centrum,  as  in  Polymixia.  In  P .  11112 
where  the  second  pre-ural  spine  is  abnormal,  the  foremost  upper  caudal  rays  articu- 
late with  the  neural  spine  of  the  second  pre-ural  centrum.  The  lowermost  principal 
caudal  ray  (unbranched)  is  preceded  by  four  spines  and  three  segmented  rays. 
P.iiii2  shows  the  anal  fin  of  H.  pulchellus,  previously  unknown.  The  fin  contains 
five  spines,  the  first  very  small  and  the  fifth  the  longest  and  thickest,  and  about 
eleven  soft  rays.  The  fifth  anal  spine  is  equal  in  length  to  the  longest  dorsal  spine, 
just  over  one-quarter  of  the  maximum  depth  of  the  trunk. 

H.  rotundus  (Smith  Woodward)  is  known  only  by  the  holotype,  P .  315,  and  P .  5682. 
P .  5682  shows  most  of  the  details  of  the  caudal  skeleton :  there  is  nothing  to  distin- 
guish it  from  the  other  polymixiids  described  here  except  that  there  is  no  flange 
on  the  anterior  margin  of  the  neural  spine  of  the  second  pre-ural  centrum  as  there 
is  in  H.  pulchellus.  P. 315  shows  that  the  foremost  caudal  rays  are  soft  rays, 
longitudinally  divided,  not  spines  as  they  are  in  H.  dorsalis,  H.  pulchellus  and  Poly- 
mixia. These  two  points  may  be  added  to  others  (Patterson  1964  :  299)  indicating 
that  this  species  is  incorrectly  placed  in  Homonotichthys,  but  more  specimens  are 
necessary  before  its  true  position  can  be  decided. 

Family  SPHENOCEPHALIDAE  Patterson  (1964  :  383) 

The  only  member  of  this  family  is  Sphenocephalus  fissicaudus  Agassiz  from  the 
Campanian  of  Westphalia.  Of  the  six  specimens  in  the  British  Museum  (Natural 
History),  three,  P. 2100  (Fig.  10),  P. 8772  and  P. 9059  have  the  caudal  skeleton  well 
preserved.  As  in  Polymixiidae,  the  second  pre-ural  centrum  has  a  fully  developed 
neural  spine  (npu2],  there  is  a  free  second  ural  centrum  (1*2),  a  stegural  (st),  a  second 
uroneural  (d2),  and  six  hypurals  (hi  —  6).  As  in  Polymixia,  the  foremost  procurrent 
rays  articulate  with  the  neural  and  haemal  spines  of  the  third  pre-ural  centrum. 
In  contrast  to  all  Polymixiidae  there  are  only  two  epurals  (ei,  2),  the  first  curved 
forwards  proximally  and  with  a  gap  between  it  and  the  neural  spine  of  the  second 


62 


THE   CAUDAL   SKELETON   IN 


FIG.  10.     Sphenocephalus  fissicaudus  Agassiz.     Caudal  skeleton  of  P. 2100,  standard  length 
92  mm.,  Campanian,  Sendenhorst,  Westphalia.     For  explanation  of  lettering  see  p.  102. 

pre-ural  centrum,  there  is  a  wide  gap  between  the  upper  and  lower  hypurals,  and 
there  is  a  large  number  of  procurrent  rays,  apparently  nine  in  each  lobe,  six 
unsegmented  soft  rays  and  three  segmented  in  the  upper,  five  unsegmented  and 
four  segmented  in  the  lower. 

Suborder  DINOPTERYGOIDEI  Patterson  (1964  :  434) 

This  suborder  contains  four  monotypic  Uppper  Cretaceous  familes,  probably  not 
closely  related.  As  the  caudal  skeleton  in  the  type  family,  the  Dinopterygidae,  is 
poorly  known,  the  best  known  family,  the  Aipichthyidae,  will  be  described  first. 

Family  AIPICHTHYIDAE  Patterson  (1964  :  303) 
Genus  AIPICHTHYS  Steindachner  (1860  :  763) 

I  have  seen  no  material  of  the  type  species,  A.  pretiosus  Steindachner  from  the 
Lower  Cenomanian  of  Dalmatia,  and  the  caudal  region  is  not  preserved  in  A. 


MESOZOIC    ACANTHOPTERYGIAN   FISHES  63 

nuchalis  (Dixon)  from  the  English  Chalk.  The  other  two  species,  A.  minor  (Pictet) 
and  A.  velifer  Smith  Woodward,  both  from  the  Cenomanian  of  Hakel,  Lebanon, 
are  well  represented  in  the  British  Museum  (Natural  History).  They  show  a 
remarkable  range  of  variation  in  the  structure  of  the  caudal  skeleton.  In  most  speci- 
mens of  Aipichthys  the  distal  parts  of  the  hypurals,  epurals  and  uroneurals  are 
obscured  by  the  deeply  cleft  bases  of  the  caudal  fin-rays,  but  in  P. 82  (A.  minor, 
Fig.  uA)  the  fin-rays  are  displaced,  exposing  these  bones.  In  this  specimen  and  in 
all  others  the  neural  and  haemal  spines  of  the  third  pre-ural  centrum  (pu3)  are  broad 
and  elongate  and  the  first  procurrent  rays  of  the  fin  articulate  with  them.  The 
haemal  arches  of  the  second  and  third  pre-ural  centra  are  autogenous.  In  contrast 
to  all  Polymixioidei,  the  neural  spine  of  the  second  pre-ural  centrum  (npu2)  is 
normally  only  about  half  as  long  as  its  predecessor,  which  makes  contact  with  the 
first  epural  distally.  Though  somewhat  expanded,  this  neural  spine  is  more  like 
those  of  Elops  (Nybelin  1963,  figs,  i,  4)  and  Nematonotus  (Fig.  25)  than  the  shorter, 
broader  spine  in  Aulolepis  and  Ctenothrissa  (Figs.  2,  4,  5) :  it  does  not  resemble  the 
very  low  crest  on  this  centrum  in  Berycoidei  and  percoids.  In  one  specimen  of 
Aipichthys  minor,  P. 6183  (Fig.  nC),  the  neural  spine  of  the  second  pre-ural  centrum 
is  fully  developed  and  supports  epaxial  fin-rays,  as  in  Polymixioidei.  This  is 
clearly  an  individual  variation,  comparable  to  those  found  in  certain  individuals  of 
Monocentris  (Fig.  14)  and  Siniperca  (Fig.  26),  and  is  recognizable  as  such  by  the  gap 
between  the  spine  and  the  proximal  part  of  the  first  epural,  which  is  filled  by  flanges 
from  the  posterior  face  of  the  spine  and  from  the  anterior  face  of  the  epural.  In 
all  specimens  of  Aipichthys  there  are  three  epurals  (ei  —  3)  and  the  first  ural  and 
pre-ural  centra  are  fused  (pui  +  ui).  There  is  normally  a  free  second  ural  centrum 
(u2,  Figs.  1 1  A,  C),  but  in  occasional  individuals,  like  the  large  specimen  shown  in 
Fig.  nB,  the  second  ural  centrum  is  fused  into  the  preceding  compound  centrum 
(pui  +  ui  +  U2),  although  the  line  of  fusion  is  visible  in  transparency  under  xylene. 
There  is  always  a  stegural  (st)  and  a  second  uroneural  (d2).  In  A.  minor  there  are 
normally  six  hypurals  (hi  —  6,  Figs.  nB,  C),  as  in  Polymixioidei,  but  in  P. 82  (Fig. 
1 1  A)  the  fifth  hypural  is  partially  divided  distally  suggesting  that  the  sixth  is 
fused  into  it.  In  all  specimens  of  A.  velifer  in  which  the  hypurals  are  visible  (P. 4743, 
P. 4744,  49486,  P. 47862)  there  appear  to  be  only  five  hypurals.  Preceding  the 
principal  rays  of  the  caudal  fin  there  are  four  unsegmented  soft  rays  and  three  seg- 
mented rays  in  the  upper  lobe,  three  unsegmented  and  three  segmented  rays  in  the 
lower. 

Family  PHARMAGICHTHYIDAE  Patterson  (1964  :  398) 

This  family  contains  only  Pharmacichthys  venenifer  Smith  Woodward  (1942)  from 
the  Middle  Cenomanian  of  Hakel,  Lebanon.  None  of  the  five  known  specimens 
of  this  species  has  the  caudal  skeleton  sufficiently  well  preserved  to  be  illustrated, 
but  the  holotype,  AUB  104691/99,  and  AUB  101872  show  most  of  the  caudal  anatomy. 
The  suggestion  (Patterson  1964  :  401)  that  the  nearest  relative  of  Pharmacichthys 
is  Aipichthys  is  borne  out  by  the  structure  of  the  caudal  skeleton  and  fin.  In  almost 
every  respect  the  caudal  skeleton  of  Pharmacichthys  is  identical  with  that  of  Aipi- 
chthys (Fig.  n).  The  foremost  caudal  fin-rays  articulate  with  the  neural  and  haemal 

GEOL.    I  7,   2  6 


64 


THE   CAUDAL   SKELETON    IN 

,e1-3 


d2 


npu2 


h5 


•pu1+u1+u2 


MESOZOIC   ACANTHOPTERYGIAN   FISHES 

e1-3 


d2. 


h6 


puUul 


u2 


FIG.  ii.  Aipichthys  minor  (Pictet).  Caudal  skeleton  of  A,  P. 82,  standard  length  32  mm.; 
B,  RSM  1881 .5.41,  standard  length  c.  50  mm.;  C,  P. 6183,  standard  length  36  mm.  All 
from  Middle  Cenomanian,  Hakel,  Lebanon.  For  explanation  of  lettering  see  p.  102. 
In  B  arrows  mark  the  outermost  (unbranched)  principal  fin-rays. 

spines  of  the  third  pre-ural  centrum,  the  neural  spine  of  the  second  pre-ural  centrum 
is  about  half  as  long  as  its  predecessor,  as  in  Aipichthys,  the  haemal  spine  of  the 
second  pre-ural  centrum  has  a  broad  flange  on  its  anterior  edge,  there  are  three 
epurals  and  the  principal  rays  of  the  fin  are  preceded  by  four  unsegmented  soft  rays 
and  three  segmented  rays  above,  three  unsegmented  and  three  segmented  rays  below. 
It  is  impossible  to  see  whether  there  are  five  or  six  hypurals.  A  further  resemblance 
to  Aipichthys,  not  previously  noticed,  is  that  the  bases  of  the  caudal  rays  are  deeply 
cleft,  overlapping  much  of  the  hypurals  ("  hypurostegy  ",  Le  Danois  &  Le  Danois 
1964).  The  only  difference  from  Aipichthys,  suggested  by  the  holotype,  AUB 
101872  and  AUB  102601,  is  that  the  first  and  second  hypurals  appear  to  be  fused. 

Family  PYGNOSTEROIDIDAE  Patterson  (1964  :  389) 

The  only  member  of  this  family  is  Pycnosteroides  levispinosus  (Hay  1903)  from  the 
Middle  Cenomanian  of  Hajula,  Lebanon.  The  caudal  skeleton  and  fin  are  present 
in  two  specimens,  P.  13900  (Fig.  12)  and  AMNH  45190  (Hay  1903,  pi.  32,  fig.  3). 
Pycnosteroides  differs  from  Aipichthys  and  Pharmacichthys  in  having  the  foremost 
rays  of  the  caudal  fin  inserted  on  the  first  epural  above  and  the  haemal  spine  of 
the  third  pre-ural  centrum  below,  and  in  having  a  fully  developed  neural  spine  on 


66 


THE   CAUDAL   SKELETON    IN 

el 

npu2, 


st 


FIG.  12.  Pycnosteroides  levispinosus  (Hay).  Caudal  skeleton  of  P.  13900,  standard 
length  c.  50  mm.,  Middle  Cenomanian,  Hajula,  Lebanon.  For  explanation  of  lettering 
seep.  1 02. 

the  second  pre-ural  centrum  (npu2).  This  neural  spine  differs  from  those  of  the 
Polymixioidei  in  being  expanded  distally.  The  haemal  arches  of  the  second  and 
third  pre-ural  centra  (pu2,  puj)  are  autogenous.  In  the  fusion  of  the  first  ural  and 
first  pre-ural  centra  (pui  -f-  ui),  the  free  second  ural  centrum  (u2),  and  the  form  of 
the  stegural  (st}  and  second  uroneural  (missing  in  the  figured  specimen  but  present 
in  AMNH  45190;)  Pycnosteroides  agrees  with  Aipichthys  and  the  Polymixioidei.  There 
are  three  epurals  (ei  —  3).  In  P.  13900,  as  in  Aipichthys  velifer,  there  are  only  five 
hypurals  (hi  —  5),  but  here  the  first  and  third  are  the  largest,  the  third  being  much 
larger  than  the  fourth.  In  AMNH  45190  there  are  six  hypurals.  In  contrast  to 
Aipichthys  and  Pharmacichthys  there  are  only  three  soft  rays,  all  unsegmented,  in 
front  of  both  the  upper  and  lower  principal  rays,  and  the  fin-rays  are  not  deeply 
cleft  basally. 

Family  DINOPTERYGIDAE  Jordan  (1923  :  173) 

This  family,  the  type  of  the  suborder,  contains  only  Dinopteryx  spinosus  (Davis) 
from  the  Upper  Santonian  of  Sahel  Alma,  Lebanon. 

The  caudal  region  is  very  imperfectly  preserved  in  the  holotype, 


MESOZOIC   ACANTHOPTERYGIAN   FISHES 


67 


but  is  more  or  less  completely  visible  in  USNM  22217  and  22219.  As  in  other  respects 
(Patterson  1964  :  392),  Dinopteryx  resembles  Pycnosteroides  in  the  caudal  skeleton. 
The  second  pre-ural  centrum  has  a  fully  developed  neural  spine  and  both  this  and 
the  preceding  neural  spine  are  expanded  distally,  as  in  Pycnosteroides  (Fig.  12).  As 
in  Pycnosteroides  there  are  three  epurals,  the  first  ural  and  pre-ural  centra  are  fused, 
there  is  a  free  second  ural  centrum,  a  stegural  and  a  second  uroneural.  USNM  22219 
shows  that  there  were  at  least  three  upper  hypurals,  shaped  as  in  Pycnosteroides, 
but  it  is  impossible  to  be  certain  whether  or  not  a  small  sixth  hypural  was  present. 
The  foremost  procurrent  caudal  rays  articulate  with  the  neural  spine  of  the  second 
pre-ural  centrum  above  and  the  haemal  spine  of  the  third  pre-ural  centrum  below, 
further  forwards  than  in  Pycnosteroides.  There  are  four  spines  and  two  segmented 
rays  in  front  of  the  principal  rays  above,  three  spines  and  two  segmented  rays  below, 
both  the  holotype  and  USNM  22217  showing  that  the  foremost  procurrent  rays  were 
true  spines,  a  difference  from  other  Dinopterygoidei. 

Suborder  BERYCOIDEI  Patterson  (1964  :  433) 

This  suborder,  containing  eight  living  families,  is  represented  in  the  Cretaceous 
only  by  two  families,  the  Trachichthyidae  and  Holocentridae.  A  detailed  discussion 
of  the  relationships  between  these  two  families  in  the  Cretaceous  will  be  found  in 


npu3 


pu1+u1 
npu2 


FIG.  13.  Hoplostethus  mediterraneus  Cuvier  &  Valenciennes.  Caudal  skeleton  of  a  dried 
skeleton,  1878.4.5.8,  standard  length  190  mm.,  Japan.  For  explanation  of  lettering 
seep.  102. 


68  THE   CAUDAL   SKELETON    IN 

Patterson  (1967).  Among  living  Berycoidei,  the  caudal  skeleton  of  Centroberyx 
affinis  (Berycidae)  has  been  illustrated  by  Regan  (1911,  fig.  2),  those  of  Hoplostethus 
(Trachichthyidae),  Holocentrus  and  Myripristis  (both  Holocentridae)  are  briefly 
discussed  by  Gosline  (1961  :  14)  and  a  caudal  skeleton  of  Holocentrus  ascensionis  is 
figured  by  Rosen  (1964,  fig.  23D).  Fig.  13  shows  the  caudal  skeleton  of  the  living 
Hoplostethus  mediterraneus  (Trachichthyidae).  In  most  respects  this  is  typical  of 
primitive  Berycoidei,  with  autogenous  haemal  spines  on  the  second  and  third 
pre-ural  centra,  the  second  pre-ural  centrum  without  a  neural  spine,  bearing  only  a 
low,  broad  crest  (npu2)  which  lies  below  the  proximal  end  of  the  first  epural,  three 
epurals  (ei  —  3),  six  hypurals  (hi  —  6),  a  free  second  ural  centrum  (u2),  a  stegural 
(st)  and  a  second  uroneural  (d2) .  In  the  specimen  illustrated  the  neural  spine  of  the 
third  pre-ural  centrum  (npuj)  is  double  distally  and  the  third  and  fourth  hypurals 
are  fused  proximally:  these  features  are  individual  abnormalities.  The  neural  crest 
on  the  second  pre-ural  centrum  is  autogenous — this  is  a  primitive  feature  which  also 
occurs  in  primitive  myctophoids  (N ematonotus  and  Aulopus,  p.  81).  In  Hoplo- 
stethus the  foremost  procurrent  fin-rays  articulate  with  the  neural  and  haemal  spines 
of  the  third  pre-ural  centrum.  In  living  trachichthyids  (Hoplostethus,  Trachichthys, 
Paratrachichthys,  Gephyroberyx)  the  nineteen  principal  caudal  rays  are  normally 
preceded  by  six  spines  and  one  segmented  ray  above  and  below. 

The  caudal  skeleton  in  living  holocentrids  is  described  below  (p.  75,  Fig.  20). 
In  Berycidae  (Regan  1911,  fig.  2)  the  caudal  skeleton  is  advanced  over  the  trachi- 
chthyid  condition  in  having  both  the  second  ural  centrum  and  the  stegural  fused 
into  the  compound  first  ural  and  pre-ural  centrum.  Nothing  is  known  of  the  caudal 
skeleton  in  Korsogasteridae  and  Anomalopidae.  Dissection  of  single  specimens  of 
Diretmus  (Diretmidae)  and  Anoplogaster  (Anoplogasteridae)  shows  that  both 
resemble  Berycidae  in  having  the  stegural  and  second  ural  centrum  fused  into  the 
preceding  compound  centrum,  while  in  Anoplogaster  the  second  uroneural  is  lost 
and  in  Diretmus  the  sixth  hypural  is  lost  and  there  is  fusion  between  the  first  and 
second  hypurals  and  between  the  third  and  fourth  hypurals.  In  the  Stephano- 
berycoidei,  which  appear  to  be  only  specialized  derivatives  of  the  Berycoidei, 
Gibberichthys  (Gibberichthyidae)  agrees  with  trachichthyids  such  as  Hoplostethus  in 
the  caudal  skeleton  but  Melamphaes  (Melamphaeidae)  has  both  the  stegural  and 
second  ural  centrum  fused  with  the  preceding  centrum,  a  much  reduced  second 
uroneural,  only  five  hypurals  and  fusion  within  the  upper  and  lower  hypurals.  All 
living  berycoids  and  stephanoberycoids  seem  to  be  characterized  by  the  presence 
of  spinous  procurrent  caudal  rays. 

In  Monocentris  (Monocentridae)  the  caudal  skeleton  normally  shows  the  same 
major  features  as  the  figured  specimen  of  Hoplostethus  (even  to  the  partial  doubling  of 
the  neural  spine  of  the  third  pre-ural  centrum),  but  in  one  of  the  available  skeletons 
(Fig.  14)  there  is  a  fully  developed  neural  spine  on  the  second  pre-ural  centrum 
(npu2)  and  the  neural  spine  of  the  third  pre-ural  centrum  is  single.  There  are  three 
epurals  (ei~3),  so  that  in  this  individual  the  neural  spine  on  the  second  pre-ural 
centrum  has  apparently  developed  instead  of  the  normal  doubling  of  the  neural 
spine  of  the  preceding  centrum.  The  caudal  skeleton  of  this  specimen  resembles 
those  of  polymixioids  and  the  dinopterygoids  Pycnosteroides  and  Dinopteryx,  but 


MESOZOIC   ACANTHOPTERYGIAN   FISHES 


69 


FIG.  14.  Monocentris  japonicus  (Houttuyn).  Caudal  skeleton  of  a  dried  skeleton  showing 
a  neural  spine  on  the  second  pre-ural  centrum,  1862.  u  .  i  .47,  standard  length  125  mm., 
Japan.  For  explanation  of  lettering  see  p.  102. 


it  is  questionable  whether  this  is  significant.  However,  Monocentris  seems  to  be 
the  only  living  berycoid  in  which  the  procurrent  caudal  rays  are  sometimes  not  spines 
but  unsegmented  soft  rays :  of  four  specimens  examined,  one  has  no  spines  and  one 
has  only  one  spine  in  front  of  the  lower  caudal  lobe  and  none  above.  Monocentris 
is  a  fish  of  highly  specialized  appearance  and  the  skull  suggests  that  the  Monocent- 
ridae  are  derivatives  of  the  Trachichthyidae,  but  there  is  a  patch  of  teeth  on  the 
endopterygoid  (Starks  1904  :  618),  a  character  otherwise  unknown  in  Berycoidei, 
and  this  and  the  occasional  absence  of  procurrent  caudal  spines  suggest  that  the 
Monocentridae  may  have  had  a  long  independent  history. 

Family  TRACHICHTHYIDAE  Bleeker  (1859) 

The  caudal  skeleton  of  the  living  Hoplostethus  mediterraneus  is  described  above 
(Fig.  13). 

Genus  HOPLOPTERYX  Agassiz  (1838  :  4) 

Hoplopteryx,  with  eight  species  ranging  from  the  Middle  Cenomanian  to  the  Upper 
Senonian,  is  the  longest-ranging  and  largest  genus  of  Cretaceous  Trachichthyidae. 


7o 


THE   CAUDAL   SKELETON   IN 


h6 


h5 


pu1+u1 


FIG.  15.     Hoplopteryx  lewesiensis  (Mantell).     Caudal  skeleton  of  P.  19486,  standard  length 
c.  175  mm.,  Chalk,  Sussex.     For  explanation  of  lettering  see  p.  102. 


In  the  type  species,  H.  antiquus  Agassiz  from  the  Campanian  of  Westphalia,  I  have 
seen  no  specimens  in  which  the  caudal  skeleton  is  preserved.  The  best  known 
species  is  H.  lewesiensis  (Mantell)  which  ranges  throughout  the  English  Chalk: 
several  specimens  show  the  caudal  skeleton  (Figs  15,  16)  which  is  almost  identical 
with  that  of  the  living  Hoplostethus  (Fig.  13),  with  the  neural  arch  of  the  second 
pre-ural  centrum  (npu2)  reduced  (though  not  so  much  as  in  Hoplostethus},  the  haemal 
arches  of  the  second  and  third  pre-ural  centra  autogenous,  three  epurals  (ei~3)  of 
which  the  first  is  especially  large,  a  stegural  (sf)  and  a  second  uroneural  (d2),  a  free 
second  ural  centrum  (u2)  and  six  hypurals  (hi-6),  the  uppermost  very  small.  The 
neural  arch  of  the  second  pre-ural  centrum  is  not  autogenous  as  it  is  in  Hoplostethus. 
The  foremost  caudal  fin-rays  are  inserted  on  the  first  epural  above  and  on  the  haemal 
spine  of  the  third  pre-ural  centrum  below.  There  are  only  three  spines  and  one 
segmented  ray  in  front  of  the  upper  principal  rays  and  two  spines  and  one  segmented 
ray  in  front  of  the  lower. 

The  other  species  of  Hoplopteryx  in  the  English  Chalk  are  H.  simus  Smith  Wood- 
ward, H.  macr acanthus  Patterson  and  H.  gephyrognathus  Patterson.     The  caudal 


MESOZOIC   ACANTHOPTERYGIAN   FISHES 


U2 


2mm 


FIG.  16.  Hoplopteryx  lewesiensis  (Mantell).  Caudal  skeleton  of  P. 5421,  standard  length 
c.  135  mm.,  Lower  Chalk,  Lewes,  Sussex.  For  explanation  of  lettering  see  p.  102.  The 
second  uroneural  is  displaced  ventrally  and  the  fifth  and  sixth  hypurals  are  missing. 


skeleton  of  H.  simus  is  exposed  in  P.  11202,  that  of  H.  macmcanthus  in  P. 30186: 
neither  appears  to  differ  from  H.  lewesiensis  in  any  way.  The  caudal  region  is  not 
preserved  in  the  two  known  specimens  of  H.  gephyrognathus. 

The  earliest  species  of  Hoplopteryx  is  H.  lewisi  (Davis)  from  the  Middle  Ceno- 
manian  of  Hakel,  Lebanon.  In  this  species  the  caudal  skeleton  and  fin  are  exposed 
in  P.  10709  and  partially  shown  in  the  holotype,  P. 4758.  H.  lewisi  seems  to  agree 
with  H.  lewesiensis  in  every  detail,  even  to  the  insertion  of  the  foremost  upper  fin 
rays  on  the  first  epural,  except  that  there  are  four  spines  and  one  segmented  ray  in 
front  of  the  upper  principal  rays,  three  spines  and  one  segmented  ray  in  front  of  the 
lower. 

The  remaining  two  species  of  Hoplopteryx,  H.  syriacus  (Pictet  &  Humbert)  and 
H.  spinulosus  Smith  Woodward,  are  from  the  Upper  Santonian  of  Sahel  Alma, 
Lebanon.  In  H.  spinulosus  I  have  seen  no  specimen  in  which  the  caudal  skeleton 


72 


THE   CAUDAL   SKELETON   IN 


is  preserved.     In  H.  syriacus  parts  of  the  caudal  skeleton  are  preserved  in  49553 
and  they  show  nothing  to  distinguish  the  species  from  H.  lewesiensis. 

Genus  LISSOBERYX  Patterson  (1967  :  73) 

The  type  species,  L.  dayi  (Smith  Woodward  1942),  is  from  the  M.  Cenomanian 
of  Hakel  and  Hajula,  Lebanon.  I  have  briefly  described  the  caudal  skeleton  (Pat- 
terson 1967  :  78)  which  is  preserved  in  AUB  108926  (Fig.  17)  and  AUB  101997. 
The  caudal  skeleton  agrees  well  with  those  of  Hoplostethus  (Fig.  13)  and  Hoplopteryx 
(Figs  15,  16).  The  neural  spine  of  the  second  pre-ural  centrum  (npu2)  is  reduced 
to  about  the  same  extent  as  in  Hoplopteryx,  there  are  three  epurals  (21-3),  the 


d2 


st 


npu2 


pu3 


FIG.  17.  Lissoberyx  dayi  (Smith  Woodward).  Caudal  skeleton  of  AUB  108926,  standard 
length  34  mm.,  Middle  Cenomanian,  Hajula,  Lebanon.  For  explanation  of  lettering 
see  p.  102.  Arrows  mark  the  outermost  (unbranched)  principal  fin-rays. 


first  long  and  closely  applied  distally  to  the  neural  spine  of  the  third  pre-ural  centrum, 
a  stegural  (st)  and  a  second  uroneural  (d,2),  and  six  hypurals  (hi-6).  The  only 
significant  differences  from  Hoplopteryx  and  Hoplostethus  are  that  the  fused  first  ural 
and  pre-ural  centrum  is  longer,  with  clear  signs  in  the  surface  sculpture  of  its  origin 
from  two  centra,  and  that  the  second  ural  centrum  is  also  longer,  so  that  the  caudal 
skeleton  appears  more  upturned.  As  in  Hoplopteryx  the  foremost  caudal  rays 
articulate  with  the  first  epural  above  and  the  haemal  spine  of  the  third  pre-ural 
centrum  below.  The  principal  rays  are  preceded  by  four  spines  and  one  unseg- 
mented  ray  above,  three  spines  and  one  segmented  ray  below. 


MESOZOIC    ACANTHOPTERYGIAN   FISHES  73 

Genus  ACROGASTER  Agassiz  (1838  :  5) 

I  have  seen  no  specimens  of  A.  parvus  Agassiz,  the  type  species,  or  of  A.  brevi- 
costatus  von  der  Marck,  both  these  species  from  the  Campanian  of  Westphalia  being 
poorly  known.  The  remaining  species,  A.  heckeli  (Pictet)  and  A.  daviesi  (Davis), 
from  the  Upper  Santonian  of  Sahel  Alma,  Lebanon,  are  common  and  several  specimens 
in  the  British  Museum  (Natural  History)  show  the  caudal  skeleton  clearly:  I  can 
find  no  differences  between  these  two  species  in  caudal  anatomy.  A  specimen  of 


u2 


FIG.  18.     Acrogaster  heckeli  (Pictet).     Caudal  skeleton  of  P. 4155,  standard  length  c.  55  mm. 
Upper  Santonian,   Sahel  Alma,   Lebanon.     For  explanation  of  lettering  see  p.  102. 


A.  heckeli  is  illustrated  in  Fig.  18.  As  in  the  other  trachichthyids  described  above, 
the  neural  spine  of  the  second  pre-ural  centrum  is  reduced  (npu2),  there  are  three 
epurals  (ei-j),  a  stegural  (st),  a  second  uroneural  (d2)  and  six  hypurals  (hi-6). 
The  fused  first  ural  and  pre-ural  centra  (pui  +  ui)  and  the  second  ural  centrum  (u2) 
are  elongate,  as  in  Lissoberyx,  and  the  caudal  skeleton  appears  strongly  upturned. 
As  in  Hoplopteryx  and  Lissoberyx,  the  foremost  caudal  rays  articulate  with  the  first 
epural  above  and  the  haemal  spine  of  the  third  pre-ural  centrum  below.  The 
principal  caudal  rays  are  preceded  by  four  or  five  spines  and  one  segmented  ray  above, 
three  or  four  spines  and  one  segmented  ray  below. 


74 


THE   CAUDAL   SKELETON   IN 


Genus  TUBANTIA  Patterson  (1964  :  413) 

The  only  species  is  T.  cataphractus  (von  der  Marck),  from  the  Campanian  of 
Westphalia,  in  which  the  caudal  skeleton  is  well  exposed  in  P. 21984  (Fig.  19). 
Tubantia  agrees  with  other  trachichthyids  in  the  reduction  of  the  neural  spine  of  the 
second  pre-ural  centrum  (npu2),  the  three  epurals  (ei-3),  stegural  (st),  second  uro- 


d2 


h6 


npu2 


pul+ul 


FIG.  19.  Tubantia  cataphractus  (von  der  Marck).  Caudal  skeleton  of  P. 21984,  standard 
length  130  mm.,  Campanian,  Baumberg,  Westphalia.  For  explanation  of  lettering 
seep.  102. 

neural  (d,2],  and  six  hypurals  (hi-6).  As  in  Hoplostethus,  the  fused  first  ural  and  pre- 
ural  centrum  (pui  +  ui]  is  only  as  long  as  the  preceding  centrum.  Tubantia  differs 
from  other  Cretaceous  trachichthyids  and  resembles  living  forms  in  having  the 
number  of  procurrent  rays  increased  to  nine  spines  and  two  segmented  rays  above, 
six  spines  and  two  segmented  rays  below,  these  small  rays  extending  forwards  in 
front  of  the  tips  of  the  neural  and  haemal  spines  of  the  third  pre-ural  centrum. 

Genus  GNATHOBERYX  Patterson  (1967  :  81) 

The  type  and  only  species,  G.  stigmosus  Patterson  (1967  :  82),  from  the  Upper 
Santonian  of  Sahel  Alma,  Lebanon,  is  known  by  two  specimens  and  the  caudal 
skeleton  is  preserved  only  in  the  holotype,  AUB  100402,  where  it  is  compressed  and 


MESOZOIC   ACANTHOPTERYGIAN   FISHES 


75 


distorted.  So  far  as  can  be  seen,  the  caudal  skeleton  and  fin  agree  with  other  Creta- 
ceous trachichthyids  such  as  Lissoberyx,  Hoplopteryx  and  Acrogaster,  with  the  neural 
spine  of  the  second  pre-ural  centrum  reduced,  the  stegural  free,  a  free  second  ural 
centrum,  and  the  upper  principal  rays  preceded  by  four  spines  and  one  segmented 
ray,  the  foremost  articulating  with  the  first  epural. 

Family  HOLOCENTRIDAE  Richardson  (1846) 

The  caudal  skeleton  of  the  living  Myripristis  adustus  is  shown  in  Fig.  20 :  it  shows 
no  significant  differences  from  those  of  several  species  of  Holocentrus.  The  neural 
and  haemal  spines  of  the  fourth  (puj)  and  fifth  pre-ural  centra  are  expanded  but 
short.  The  neural  spine  of  the  second  pre-ural  centrum  (npu2)  is  greatly  reduced 


2mm 


FIG.  20.  Myripristis  adustus  Bleeker.  Caudal  skeleton  of  a  dried  skeleton,  1858 .4.21. 239, 
standard  length  155  mm.,  Amboina.  hap,  hypurapophysis ;  for  explanation  of  other 
lettering  see  p.  102. 

(and  not  autogenous  as  it  is  in  Hoplostethus),  with  the  tip  of  the  first  epural  lying 
above  it.  The  haemal  arches  of  the  second  and  third  pre-ural  centra  (pu2,  pu3) 
are  autogenous  and  there  are  three  epurals  (ei~3) .  In  contrast  to  the  trachichthyids 
the  two  ural  centra,  the  first  pre-ural  centrum  and  the  stegural  are  all  fused  into  a 
single  structure  (pui  +  UI  +  U2-}-  st)  and  there  are  only  five  hypurals  (hi-5) 
the  uppermost  hypural  present  in  trachichthyids  having  been  lost.  The  second 
uroneural  (d2)  is  free  and  fits  proximally  into  a  notch  in  the  stegural  rather  than 
lying  below  and  behind  it.  In  Myripristis  the  foremost  caudal  rays  articulate  with 
the  first  epural  above  (as  in  most  Cretaceous  trachichthyids)  and  with  the  haemal 


76 


THE   CAUDAL   SKELETON   IN 


spine  of  the  third  pre-ural  centrum  below.  In  Holocentrus  the  foremost  rays  arti- 
culate with  the  neural  and  haemal  spine  of  the  third  pre-ural  centrum.  In  Myrip- 
ristis  the  principal  caudal  rays  are  preceded  by  four  spines  and  one  segmented  ray 
above  and  below.  Rosen  (1964,  fig.  230)  has  figured  a  caudal  skeleton  of  Holo- 
centrus ascensionis  which  differs  from  all  the  Recent  holocentrid  skeletons  that  I 
have  seen  in  having  a  free  second  ural  centrum.  Rosen  does  not  say  how  large  his 
specimen  was,  but  if  it  was  an  alizarin-stained  juvenile  this  difference  can  be 
explained. 

Genus  CAPROBERYX  Regan  (1911  :  8) 

In  the  type  species,  C.  superbus  (Dixon)  from  the  Turonian  of  the  English  Chalk, 
the  caudal  skeleton  is  exposed  in  P. 3979  (Fig.  21).  The  neural  and  haemal  spines 
of  the  fourth  (Puj)  and  fifth  pre-ural  centra  are  normal,  not  expanded  as  in  living 
holocentrids.  The  haemal  arches  of  the  second  and  third  pre-ural  centra  (pu2, 
PUJ)  are  autogenous,  the  neural  spine  of  the  second  pre-ural  centrum  (npu2)  is 
reduced  and  there  are  three  epurals  (ei-3),  all  as  in  living  holocentrids.  In  contrast 
to  living  holocentrids,  the  stegural  (st)  is  not  fused  to  the  underlying  centrum  (pui  + 
ui)  and  the  second  ural  centrum  (u2)  is  free.  The  second  uroneural  (d2)  is  fused  to 
the  stegural  (st)  distally,  but  this  is  perhaps  only  a  consequence  of  the  very  large 
size  (standard  length  c.  40  cm)  of  this  specimen.  As  in  living  holocentrids  there  are 
only  five  hypurals,  but  the  distribution  of  the  branched  principal  fin-rays  (one  on 


e1-3 


d2 


npu2 


10mm 


ph 


FIG.    21.     Caproberyx  superbus    (Dixon).     Caudal  skeleton   of  P. 3979,   standard   length 
c.  400  mm.,  Chalk,  Sussex.     For  explanation  of  lettering  see  p.  102. 


MESOZOIC   ACANTHOPTERYGIAN   FISHES  77 

the  uppermost  hypural,  two  on  the  one  below  and  six  on  the  next)  shows  that  the 
three  upper  hypurals  are  the  third  and  fourth  fused  (hj  +  4),  the  fifth  (/»5)  and  the 
sixth  (h6),  for  in  living  holocentrids  the  three  upper  hypurals,  the  third,  fourth  and 
fifth,  bear  two,  six  and  one  branched  principal  rays  respectively.  Caproberyx 
therefore  agrees  with  the  trachichthyids  in  retaining  the  small  sixth  hypural,  and 
the  fusion  of  the  third  and  fourth  hypurals  shown  by  P. 3979  is  again  probably 
merely  a  consequence  of  the  large  size  of  the  fish.  As  in  Myripristis  and  most 
Cretaceous  trachichthyids,  the  foremost  caudal  fin-rays  articulate  with  the  first 
epural  above  and  the  haemal  spine  of  the  third  pre-ural  centrum  below,  and  there 
are  probably  four  spines  and  one  segmented  ray  in  front  of  the  principal  caudal 
rays  above  and  below.  Characters  of  the  skull  and  fins  suggest  that  Caproberyx  is  the 
most  primitive  holocentrid  known,  lying  near  to  the  common  stock  of  the  Holo- 
centridae  and  Trachichthyidae  (Patterson  1964  :  359;  1967  :  103).  This  is  con- 
firmed by  the  structure  of  the  caudal  skeleton  in  C.  superbus,  for  in  the  six  hypurals, 
the  free  stegural  and  second  ural  centrum,  and  the  unexpanded  neural  and  haemal 
spines  of  the  posterior  caudal  vertebrae,  this  species  resembles  the  trachichthyids 
rather  than  other  holocentrids. 

The  other  species  of  Caproberyx  are  C.  polydesmus  (Arambourg  1954)  from  the 
Lower  Cenomanian  of  Jebel  Tselfat,  Morocco,  and  C.  pharsus  Patterson  (1967  :  97) 
from  the  Middle  Cenomanian  of  Hakel,  Lebanon.  C.  pharsus  is  known  only  by  a 
specimen  lacking  the  caudal  region.  C.  polydesmus  is  known  only  by  the  holotype: 
Arambourg's  figure  (1954,  pi.  19,  fig.  i)  shows  that  the  neural  and  haemal  spines 
of  the  fourth  and  fifth  pre-ural  centra  are  not  expanded,  as  in  C.  superbus,  and  he 
described  the  principal  rays  as  having  five  or  six  small  rays  in  front  of  them  in  each 
lobe. 

Genus  STICHOCENTRUS  Patterson  (1967  :  88) 

The  type  and  only  species  is  S.  liratus  Patterson  from  the  Middle  Cenomanian 
of  Hajula,  Lebanon.  The  caudal  skeleton  is  well  preserved  in  AUB  108923  (Fig.  22) 
and  is  partially  shown  in  AUB  108927  and  108929.  The  neural  and  haemal  spines 
of  the  fifth  pre-ural  centrum  are  unmodified  but  those  of  the  fourth  (puj)  are 
expanded,  though  not  so  strongly  as  in  living  holocentrids.  The  autogenous  haemal 
arches  on  the  second  and  third  pre-ural  centra,  the  reduced  neural  spine  on  the 
second  pre-ural  centrum  (npu2)  and  the  three  epurals  (ei~3)  are  as  in  Caproberyx 
and  living  holocentrids.  The  stegural  is  fused  with  the  underlying  centrum  (pui  + 
ui  +  U2  -\-  st)  anteriorly.  The  second  uroneural  (d2)  is  free  and  lies  below  and 
behind  the  first,  not  notched  into  the  first  as  it  is  in  living  holocentrids.  The  second 
ural  centrum  is  fused  to  the  compound  first  pre-ural  and  ural  centrum  in  AUB 
108923  (standard  length  c.  75  mm.)  although  the  line  of  fusion  is  clearly  seen,  but  in 
AUB  108929,  a  much  smaller  specimen  (standard  length  c.  35  mm.)  the  centrum 
appears  free.  As  in  living  holocentrids,  there  are  only  five  hypurals  (hi-5),  the 
small  sixth  hypural  present  in  Caproberyx  having  been  lost.  As  in  Myripristis, 
the  foremost  caudal  rays  are  inserted  on  the  first  epural  above  and  the  haemal  spine 
of  the  third  pre-ural  centrum  below,  and  there  are  four  spines  and  one  segmented 
ray  in  front  of  both  the  upper  and  lower  principal  rays. 


THE   CAUDAL   SKELETON   IN 

e3 


2mm 


FIG.  22.  Stichocentrus  liratus  Patterson.  Caudal  skeleton  of  AUB  108923,  standard 
length  c.  75  mm.,  Middle  Cenomanian,  Hajula,  Lebanon.  For  explanation  of  lettering 
see  p.  102.  Arrows  mark  the  outermost  (unbranched)  principal  fin-rays. 

As  in  the  skull  and  fins,  Stichocentrus  is  more  advanced  towards  the  living 
holocentrids  than  Caproberyx  in  the  expanded  neural  and  haemal  spines  of  the  fourth 
pre-ural  centrum,  the  five  hypurals  and  the  partial  fusion  of  the  stegural  and  second 
ural  centrum  with  the  preceding  centrum. 

The  remaining  Cretaceous  holocentrids  are  Trachichthyoides  ornatus  Smith  Wood- 
ward (1902),  known  only  by  an  isolated  head  from  the  English  Chalk,  and  Kansius 
sternbergi  Hussakof  (1929)  known  by  the  two  syntypes  from  the  Niobrara  Formation, 
Gove  Co.,  Kansas.  Nothing  is  known  of  the  caudal  anatomy  of  these  forms. 

Order  LAMPRIDIFORMES 
Suborder  LAMPRIDOIDEI  Berg  (1940  :  463) 

This  suborder  is  used  to  contain  both  the  Lampridoidei  (Lampris  only)  and  the 
Veliferoidei  (Velifer,  etc.]  of  Berg  (Bonde  1966). 

?  Family  VELIFERIDAE  Bleeker  (1860) 
Genus  BATHYSOMA  Davis  (1890  :  424) 

The  type  and  only  species  is  B.  lutkeni  Davis  from  the  Danian  stage  of  southern 
Sweden.  The  caudal  skeleton  is  partially  preserved  in  two  specimens  in  the  British 
Museum  (Natural  History).  P. 9947  (Fig.  23 A)  shows  that  the  upper  hypurals  are 


MESOZOIC   ACANTHOPTERYGIAN    FISHES 


79 


fused  into  a  triangular  plate  and  that  this  plate  is  fused  basally  with  the  second 
ural  centrum  (u2  -\-  uh).  Above  the  second  pre-ural  centrum  (pu2)  and  the  fused 
first  pre-ural  and  ural  centra  (pui  +  ui)  there  are  two  elongate  bones:  the  second 
of  these  (e)  is  certainly  an  epural  but  it  is  impossible  to  be  certain  whether  the  first 
is  an  epural  or  the  neural  arch  and  spine  of  the  second  pre-ural  centrum.  Above 
the  upper  hypural  plate  there  is  a  third  slender  bone  of  uncertain  nature.  P. 9948 


B 


pu1+u1 


u2+uh 


2mm 

FIG.  23.  Bathysoma  lutkeni  Davis.  Caudal  skeleton  of  A,  P. 9947,  standard  length  c. 
95  mm.;  B,  G,  P. 9948  (part  and  counterpart),  standard  length  78  mm.  Both  from 
Danian,  Limhamn,  southern  Sweden.  U2  +  uh,  second  ural  centrum  fused  with  one  or 
more  upper  hypurals;  for  explanation  of  other  lettering  see  p.  102. 

(in  counterpart,  part  and  counterpart  shown  in  Fig.  236,  C)  shows  the  second  pre- 
ural  centrum  (pu2)  with  an  autogenous  haemal  arch  and  no  sign  of  a  neural  spine, 
the  fused  first  pre-ural  and  ural  centra  (pui  +  ui)  bearing  the  parhypural  (ph) 
and  the  first  hypural  (hi)  ,the  distal  part  of  the  second  hypural  (h2),  and  the  upper 
hypural  plate  with  a  fragment  of  the  second  ural  centrum  fused  to  it  (u2  +  uh).  As 
in  P. 9947,  there  is  a  slender  bone  lying  above  the  upper  hypural  plate.  The  bones 
above  the  first  and  second  pre-ural  centra  are  shattered  and  displaced.  Although 

GEOL.   17,  2  7 


8o 


THE   CAUDAL   SKELETON    IN 


these  two  specimens  are  far  from  complete,  they  show  that  in  Bathysoma  the  haemal 
arch  of  the  second  pre-ural  centrum  was  autogenous,  the  first  pre-ural  and  ural 
centra  were  fused,  the  parhypural  and  the  first  and  second  hypurals  were  separate 
and  autogenous,  there  was  at  least  one  epural,  and  the  upper  hypurals  were  fused 
with  each  other  and  with  the  second  ural  centrum.  On  the  available  material  it  is 
difficult  to  interpret  the  upper  hypural  plate  and  the  slender  bone  above  it,  which 
may  be  an  epural,  a  stegural  or  a  free  hypural.  In  the  upper  hypural  plate  at  least 
three  hypurals  can  be  recognized  in  transparency  under  xylene,  presumably  hypurals 
3-5,  but  the  uppermost  part  of  the  plate  is  of  a  different  texture,  suggesting  that  the 
stegural  may  also  be  fused  into  the  structure,  a  most  unusual  condition  which  can 
only  be  confirmed  on  more  complete  material. 


e1-3. 


st>d2 


pti        W  hp 

FIG.  24.  Mene  maculata  (Bloch  &  Schneider).  Caudal  skeleton  of  a  dried  skeleton, 
1866.6.8.59,  standard  length  95  mm.,  Taiwan,  hp,  hypurapophysis ;  for  explanation 
of  other  lettering  see  p.  102. 

The  caudal  skeleton  of  Bathysoma  is  very  different  from  that  of  Mene,  the  only 
genus  of  the  Menidae,  in  which  Bathysoma  was  previously  placed  (Patterson  1964 : 
423).  In  the  living  Mene  maculata  (Fig.  24)  the  first  pre-ural  centrum,  both  the  ural 
centra,  and  all  but  the  uppermost  hypural  are  fused  into  a  symmetrical,  fan-shaped 
plate  (pui  +  ui  +  U2  +  hi-4).  The  parhypural  (ph)  has  a  very  large  hypura- 
pophysis (hp)  and  lies  free  below  this  plate.  The  uppermost  hypural  (h^)  is  also 
free,  articulating  with  a  hook  on  the  upper  edge  of  the  hypural  plate.  There  are 
three  normal  epurals  (ei-3)  and  the  neural  arch  of  the  second  pre-ural  centrum 
(pu2)  is  reduced  to  a  very  low  crest.  The  stegural  (st  +  d2)  is  autogenous,  articu- 
lating with  the  underlying  compound  centrum  by  a  large  and  clearly  mobile  joint. 
The  distal  part  of  the  shaft  of  the  stegural  is  grooved  longitudinally  in  M.  maculata ; 
in  the  Middle  Eocene  species  M.  rhombeus  (Volta)  and  M.  oblongus  (Agassiz),  in 
which  the  caudal  skeleton  is  otherwise  identical  with  that  of  the  living  species,  there 


MESOZOIC    ACANTHOPTERYGIAN    FISHES  81 

is  a  free  second  uroneural,  and  the  groove  on  the  shaft  of  the  stegural  in  M.  maculata 
clearly  marks  the  line  of  fusion  between  the  first  and  second  uroneurals.  Mene 
has  seventeen  principal  caudal  rays  with  fifteen  branched,  the  outermost  two  or 
three  rays  unsegmented  and  the  inner  ones  only  sparsely  segmented,  preceded  by 
five  unsegmented  but  divided  (in  the  median  plane)  rays  above  and  four  below. 
The  bases  of  the  caudal  rays  are  deeply  cleft,  covering  much  of  the  hypural  plate. 
The  caudal  skeleton  of  Mene  can  be  derived  from  the  basal  perciform  type  (p.  87) 
by  fusion  of  the  first  four  hypurals  with  each  other  and  with  the  supporting  centra, 
and  the  caudal  fin  has  the  perciform  number  of  rays.  The  caudal  skeleton  of 
Bathysoma  is  of  a  much  more  primitive  type,  differing  from  that  of  Beryciformes 
mainly  in  the  fusion  of  the  upper  hypurals  with  the  second  ural  centrum.  A  caudal 
skeleton  of  this  type  occurs  in  the  Lampridiformes  Velifer  (Gosline  1961;  fig.  3D), 
Palaeocentrotus  (Kiihne  1941,  fig.  2 ;  Bonde  1966)  and  Lampris,  and  the  known  skele- 
tal features  of  Bathysoma  (Patterson  1964,  fig.  90)  agree  as  well  with  Palaeocentrotus 
and  Velifer  (Regan  1907;  Smith  1951)  as  they  do  with  Mene.  The  holotype  of 
Bathysoma  lutkeni,  in  Copenhagen,  shows  that  the  supraoccipital  crest  is  attached 
to  the  skull  roof  only  at  the  posterior  end,  with  a  gap  between  it  and  the  frontal 
crest  (Bonde,  personal  commn) :  an  exactly  similar  supraoccipital  crest  occurs  in 
Bonde's  (1966)  ?  veliferid  from  the  Lower  Eocene  [Mo-clay],  while  in  Palaeocentrotus 
there  is  a  large  foramen  between  the  supraoccipital  and  frontal  crests  (Kiihne  1941, 
fig.  3).  Provisionally  Bathysoma  may  be  placed  in  the  Veliferidae  but,  like  Bonde's 
Eocene  form,  it  may  well  prove  to  be  closer  to  Palaeocentrotus. 

Order  SALMONIFORMES  (Greenwood  et  al.  1966) 
Suborder  MYCTOPHOIDEI 

The  most  generalized  of  living  myctophoids  is  Aulopus  (Aulopodidae,  Regan 
191  la  :  121 ).  The  caudal  skeleton  of  Aulopus  is  briefly  discussed  by  Gosline 
(1961  :  10),  who  notes  that  this  genus  is  one  of  the  few  living  teleosts  retaining  large 
caudal  scutes  in  front  of  the  caudal  lobes.  The  caudal  skeletons  of  more  advanced 
myctophoids  have  been  figured  by  HoUister  (Synodus,  Trachinocephalus ,  1937^, 
figs  1-14),  Gosline  (C hlorophthalmus ,  1961,  fig.  26),  Rosen  (Myctophum,  1964,  fig.  236) 
Greenwood  et  al.  (Neoscopelus,  1966,  fig.  36)  and  Weitzman  (Parasudis  and  Saurida 
1967,  figs  17,  18).  In  Aulopus  the  caudal  skeleton  is  almost  identical  with  that  of  the 
Cretaceous  Ctenothrissa  (Figs  4,  5),  with  a  free  second  ural  centrum,  a  stegural  and  a 
second  uroneural,  three  epurals  and  six  hypurals,  of  which  the  first  and  third  are 
the  largest.  The  only  differences  from  Ctenothrissa  are  that  the  haemal  arch  of  the 
third  pre-ural  centrum  and  the  neural  arch  of  the  second  pre-ural  centrum  are  auto- 
genous, the  neural  spine  of  the  second  pre-ural  centrum  is  less  expanded,  more 
spine-like,  and  just  over  half  as  long  as  its  predecessor,  and  the  second  ural  centrum 
has  a  long  posterior  process.  In  all  these  characters,  Aulopus  appears  to  be  more 
primitive  than  Ctenothrissa.  In  other  living  myctophoid  families  conditions  are 
much  as  in  Aulopus,  but  the  caudal  scutes  are  lost,  the  neural  spine  of  the  second 
pre-ural  centrum  ceases  to  be  autogenous  and  becomes  shorter  and  expanded,  the 
first  epural  tending  to  move  forwards  above  it,  there  is  often  fusion  within  the  upper 


82 


THE   CAUDAL   SKELETON    IN 


and  lower  hypurals,  the  sixth  hypural  and  one  epural  may  be  lost  (Synodontidae), 
and  the  second  ural  centrum  and  stegural  may  fuse  with  the  compound  first  ural 
and  pre-ural  centrum.  These  trends  are  very  like  those  seen  within  the  Berycoidei 
(p.  68;  cf.  figs.  236,  D  in  Rosen  1964). 

In  the  Upper  Cretaceous  myctophoids  were  abundant,  the  best  known  genera 
being  Sardinioides,  Acrognathus,  Cassandra  (  =  Leptosomus)  and  Nematonotus. 
Nematonotus  appears  to  be  the  most  primitive  of  these  and  will  serve  as  an  example. 
Figure  25  shows  specimens  of  Nematonotus  bottae  (Pictet  &  Humbert),  from  the 
Cenomanian  of  Hakel,  Lebanon,  and  N.  longispinus  (Davis),  from  the  Cenomanian 


e1-3 


d2- 


e1-3. 


h5 


pu1*u1 


pu1*u1 


ph 


ud 


2mm 

FIG.  25.  The  caudal  skeleton  in  A,  Nematonotus  bottae  Pictet  &  Humbert,  49563,  stand- 
ard length  c.  90  mm.,  Middle  Cenomanian,  Hakel,  Lebanon;  B,  Nematonotus  longispinus 
(Davis),  P.  13882,  standard  length  114  mm.,  Middle  Cenomanian,  Hajula,  Lebanon. 
In  G  are  the  bases  of  the  upper  caudal  rays  in  P. 48825,  N.  longispinus,  standard  length 
97  mm.,  to  show  the  urodermal,  ud.  For  explanation  of  other  lettering  see  p.  102. 
In  B  the  second  uroneural  is  missing,  in  A  and  C  arrows  mark  the  outermost  (unbranched) 
principal  fin-rays. 


MESOZOIC   ACANTHOPTERYGIAN   FISHES  83 

of  Hajula,  Lebanon.  There  are  no  significant  differences  between  these  two  species 
in  the  caudal  region.  In  the  specimen  of  N.  bottae  illustrated,  the  haemal  arches 
of  the  third  and  fourth  pre-ural  centra  are  fused  and  the  haemal  spine  of  the  third 
pre-ural  centrum  is  partially  doubled.  This  is  an  individual  abnormality.  As  in 
Aulopus  there  is  a  large  caudal  scute  (/.  s)  above  and  below  the  caudal  skeleton  and 
the  neural  spine  of  the  second  pre-ural  centrum  (npu2)  is  autogenous,  spine-like  and 
just  over  half  as  long  as  its  predecessor.  The  autogenous  haemal  spines  on  the 
second  and  third  pre-ural  centra,  three  epurals  (ei-3),  six  hypurals  (hi-6),  long  second 
ural  centrum  (u2,  Fig.  256)  and  second  uroneural  (d2)  are  as  in  Aulopus.  The 
stegural  (st)  is  forked  proximally,  with  a  process  extending  forwards  on  to  the  second 
pre-ural  centrum.  This  process,  absent  in  Aulopus  and  Ctenothrissiformes,  is  a 
primitive  feature  present  in  Flops  and  some  clupavids  (Patterson  19670,  fig.  n) 
which  indicates  the  double  origin  of  the  first  uroneural  (Regan  1910  :  355 ;  Patterson 
1968  :  226).  In  Nematonotus  there  is  a  single  urodermal  (Fig.  256),  a  structure  absent 
from  living  myctophoids  and  Ctenothrissiformes  but  also  present  in  the  Cenomanian 
Sardinioides  attenuatus.  In  Nematonotus  the  foremost  procurrent  caudal  rays 
articulate  with  the  neural  and  haemal  spine  of  the  third  pre-ural  centrum  and  the 
nineteen  principal  rays  are  preceded  by  four  unsegmented  and  four  segmented  rays 
above,  four  unsegmented  and  two  segmented  rays  below.  In  myctophoids  the 
procurrent  caudal  rays  are  normally  longitudinally  divided,  not  spinous,  but  pro- 
current  caudal  spines  appear  in  advanced  members  of  the  family  Myctophidae 
(Fraser-Brunner  1949  :  1033). 

III.  DISCUSSION 

(a)  The  relationships  of  Ctenothrissiformes,  Myctophoidei  and  Beryciformes. 

Monod  (1967  :  118)  has  remarked  that  the  structure  of  the  caudal  skeleton  is 
rarely  of  value  in  discriminating  between  taxa  at  the  generic  level  and  below,  but 
that  it  becomes  increasingly  valuable  at  the  familial,  subordinal  and  ordinal  level. 
This  observation  is  fully  borne  out  by  the  forms  described  here,  the  only  exception 
being  the  occurrence  of  two  types  of  caudal  skeleton  in  the  ctenothrissiform  family 
Aulolepididae,  Pateroperca  having  the  neural  spine  of  the  second  pre-ural  centrum 
fully  developed,  Aulolepis  having  it  short  and  expanded,  as  it  is  in  Ctenothrissidae. 
This  suggests  that  Aulolepis  and  Pateroperca  belong  to  different  families,  but 
Pateroperca  is  as  yet  so  poorly  known  that  no  conclusion  can  be  reached  on  this 
point  until  more  specimens  are  discovered. 

Among  the  Ctenothrissiformes,  Myctophoidei  and  Beryciformes  caudal  anatomy 
is  very  similar.  The  basal  type  of  caudal  skeleton  in  these  groups  has  the  following 
features:  the  first  pre-ural  and  ural  centra  fused,  a  free  second  ural  centrum,  three 
epurals,  a  stegural,  a  second  uroneural  and  six  hypurals  (two  lower  and  four  upper). 
Caudal  scutes  above  and  below  the  caudal  skeleton  are  present  in  Aulopus,  most 
Cretaceous  myctophoids  (Nematonotus,  Acrognathus,  Sardinioides)  and  Ctenothriss- 
iformes (Aulolepis,  Ctenothrissa) :  they  are  absent  in  Beryciformes  and  higher  groups. 
A  single  urodermal  is  present  in  Nematonotus  and  Sardinioides  attenuatus  among 
Cretaceous  myctophoids.  There  is  no  urodermal  in  Ctenothrissiformes  or  Bery- 
ciformes. The  main  variations  encountered  in  the  caudal  skeletons  of  myctophoids, 

GEOL.  17,  2  7§ 


84  THE   CAUDAL   SKELETON    IN 

ctenothrissiforms  and  beryciforms  involve  reductions  in  the  number  of  caudal 
elements  by  fusion  or  suppression  and  the  condition  of  the  neural  spine  of  the  second 
pre-ural  centrum,  which  may  be  fully  developed  (Pateroperca,  Polymixioidei,  Pycno- 
steroides,  Dinopteryx),  spine-like  and  about  half  as  long  as  its  predecessor  (Nematono- 
tus,  Aulopus,  Aipichthys,  Pharmacichthys) ,  short  and  expanded  (Ctenothrissa, 
Aulolepis,  many  myctophoids)  or  reduced  to  a  low  crest  (Berycoidei) . 

The  basal  teleostean  caudal  skeleton,  seen  in  such  genera  as  Leptolepis,  Allothris- 
sops,  Thrissops,  Ichthyodectes,  Hiodon,  Flops,  Salmo,  etc.,  contains  two  free  ural 
centra,  three  epurals,  two  lower  hypurals  and  five  upper  hypurals  (there  may  be  six 
or  seven  upper  hypurals  in  Leptolepis;  Patterson  1968  :  220;  there  are  only  four  in 
salmonids;  Norden  1961  :  738),  up  to  seven  uroneurals  (Patterson  1968)  which  extend 
forwards  to  the  fourth  pre-ural  centrum  in  Thrissops  and  Ichthyodectes,  to  the  third 
in  Leptolepis  and  Allothrissops  and  to  the  second  pre-ural  centrum  in  Hiodon,  Elops, 
Salmo,  etc.,  and  one  or  two  urodermals  (Patterson  1968  :  230).  The  condition  of  the 
neural  arches  and  spines  in  the  caudal  region  is  often  complicated  by  doubling  of  the 
segmental  structures,  usually  as  individual  variations,  but  in  Leptolepis,  Allothrissops, 
Flops  and  some  salmonids  there  is  normally  a  neural  arch  and  spine  on  the  first  pre-ural 
centrum,  and  in  Leptolepis,  Hiodon,  osteoglossoids  (Greenwood  1967)  and  Alepo- 
cephalus  (Patterson  1968,  fig.  12)  there  may  be  a  more  or  less  well  developed  arch  and 
spine  on  the  first  ural  centrum.  It  is  usually  assumed  (e.g.  Gosline  1961  : 14;  Patterson 
19670  :  104)  that  the  second  pre-ural  centrum  primitively  bears  a  complete  neural 
spine,  supporting  epaxial  fin-rays.  But  among  the  forms  described  here  the  most 
primitive  (Aulopus,  Nematonotus)  have  the  neural  spine  of  the  second  pre-ural 
centrum  about  half  as  long  as  that  of  the  third,  and  this  is  also  true  of  Elops  (Nybelin 
1963,  figs,  i,  4),  which  in  other  respects  seems  to  be  the  most  primitive  living  teleost. 
This  suggests  that  the  short  second  pre-ural  neural  spine  may  be  primitive  for  some 
teleost  groups.  There  are  three  possible  conditions  of  the  second  pre-ural  neural 
spine.  It  may  be  fully  developed,  as  in  Allothrissops  (Patterson  19670,  fig.  6), 
Ichthyodectes  (Cavender  1966,  fig.  i),  Tarpon  (Nybelin  1963,  fig.  7),  and  many  other 
primitive  telosts,  normally  supporting  epaxial  fin-rays  but  in  Tarpon  ending  just 
in  front  of  the  foremost  epaxial  fin-ray;  it  may  be  about  half  as  long  as  its  prede- 
cessor, as  in  Elops,  "  Clupavus  "  (Patterson  19670,  fig.  n),  Nematonotus  and  Aulopus; 
or  it  may  be  represented  only  by  a  low  crest,  as  in  Berycoidei  and  generalized  percoids 
(p.  87).  Intermediates  between  the  second  and  third  of  these  conditions  occur  in 
Aulolepis,  Ctenothrissa  and  many  myctophoids.  Intermediates  between  the  first 
two  conditions  seem  to  occur  only  in  primitive  protacanthopterygian  groups 
(salmonids,  as  in  the  specimen  of  Cristivomer  illustrated  by  Vladykov  1954,  fig.  2; 
characinids,  as  in  the  specimen  of  Brycon  illustrated  by  Weitzman  1962,  fig.  15) 
in  which  the  pre-ural  neural  spines  are  very  variable  and  both  conditions  may  occur 
in  a  single  species.  The  third  condition,  the  spine  reduced  to  a  low  crest,  is 
undoubtedly  advanced  and  may  be  left  out  of  consideration  here.  In  the  ancestors 
of  the  teleosts,  the  pholidophorids,  the  neural  spines  of  the  last  three  pre-ural  verte- 
brae decrease  in  size  progressively  so  that  all  three  end  on  approximately  the  same 
oblique  plane :  the  first  pre-ural  neural  spine  is  very  short,  the  second  is  both  shorter 
and  more  slender  than  the  third  (Patterson  1968,  figs.  1-4).  In  pholidophorids 


MESOZOIC   ACANTHOPTERYGIAN   FISHES  85 

these  neural  spines  do  not  reach  the  dorsal  edge  of  the  trunk  and  do  not  support 
fulcra  or  fin-rays.  This  condition  of  the  pre-ural  neural  spines  seems  to  be  primitive 
for  the  teleosts  as  a  whole  and  it  persists  in  the  Lower  Jurassic  Leptolepis  cory- 
phaenoides  and  L.  normandica  (Nybelin  1963,  figs.  9,  10)  and  the  Upper  Jurassic 
L.  dubia  (Nybelin  1963,  fig.  8;  Patterson  1968,  fig.  10).  In  the  Upper  Jurassic 
two  of  the  three  modern  types  of  second  pre-ural  neural  spine  were  already  in  exis- 
tence. In  Allothrissops  and  Thrissops  (Nybelin  1963,  figs,  u,  12;  Patterson  19670, 
fig.  6)  both  the  second  and  third  preural  neural  spines  have  elongated  so  that  they 
reach  the  dorsal  edge  of  the  trunk,  ending  just  in  front  of  the  foremost  procurrent 
fin-rays:  this  is  essentially  the  condition  in  living  Tarpon.  In  the  Upper  Jurassic 
E lops-like  fish  illustrated  by  Nybelin  (1963,  fig.  6)  the  second  pre-ural  neural  spine 
remains  short  but  the  third  and  fourth  pre-ural  neural  spines  are  elongated,  reaching 
the  dorsal  edge  of  the  trunk  and  supporting  the  foremost  procurrent  fin-rays.  This 
is  essentially  the  condition  in  living  Flops.  There  is  no  a  priori  reason  to  regard 
either  of  these  two  conditions  as  more  primitive,  both  are  a  response  to  a  new  need, 
the  necessity  to  support  the  epaxial  procurrent  rays  as  they  extend  forwards  to 
increase  the  dorso-ventral  symmetry  of  the  tail.  However,  we  know  that  the  Elops 
type,  with  a  short  second  pre-ural  neural  spine,  has  persisted  unchanged  in  elopids 
since  the  Upper  Jurassic  and  that  this  type  of  second  pre-ural  neural  spine  is  primi- 
tive for  the  teleosts  as  a  whole,  and  there  is  no  reason  for  regarding  the  short  neural 
spine  of  Elops  and  its  Jurassic  relative  as  a  secondary  regression  from  a  long  neural 
spine  of  Tarpon  type.  I  conclude,  therefore,  that  when  one  finds  a  second  pre-ural 
neural  spine  resembling  that  of  Elops  in  a  generalized  teleost  one  should  regard  it 
as  a  primitive  feature  unless  there  is  good  evidence  to  the  contrary.  In  support 
of  this  interpretation  is  the  occurrence  of  a  second  pre-ural  neural  spine  of  this  type 
only  in  teleosts  in  which  the  caudal  skeleton  retains  such  primitive  features  as  a 
free  second  ural  centrum  and  nineteen  principal  caudal  rays.  A  generalized  teleost 
having  an  elongate  second  pre-ural  neural  spine  is  to  be  regarded  as  having  developed 
this  from  a  short  spine  of  leptolepid  or  elopid  type :  this  development  can  apparently 
take  place  spontaneously  (see  Aipichthys,  Fig.  n). 

From  the  basal  type  of  teleostean  caudal  skeleton,  the  most  primitive  members  of 
the  myctophoid-ctenothrissiform-beryciform  assemblage  (such  as  Nematonotus} 
differ  in  the  loss  of  one  hypural,  the  seventh,  whether  by  suppression  or  by  fusion 
with  the  sixth  is  as  yet  unknown,  have  reduced  the  number  of  uroneurals  to  two, 
principally  by  loss  of  the  small  posterior  uroneurals  (ural  neural  arches  6-8),  since 
the  forked  first  uroneural  of  Nematonotus  (Fig.  25)  is  clearly  homologous  with  that 
of  Elops,  representing  the  second  and  third  ural  neural  arches,  while  the  second 
uroneural  is  probably  homologous  with  the  second  uroneural  of  Elops,  representing 
the  fourth  and  fifth  ural  neural  arches  (Patterson  1968  :  226),  the  first  pre-ural  and 
ural  centra  have  fused,  and  the  first  uroneural  has  fused  with  the  first  ural  and  pre- 
ural  neural  arches  to  produce  a  stegural,  a  development  which  took  place  very  early 
in  the  protacanthopterygian  lineage  (salmonids,  "  Clupavus  ",  etc.).  These  changes 
raise  the  question  of  the  origin  of  the  myctophoids  and  ctenothrissiforms.  Green- 
wood et  al.  (1966  :  371)  wrote  of  the  ctenothrissiforms  "  we  link  them  with  some 
early  group  of  myctophoid-like  salmoniform  fishes  in  which  the  supramaxillae  were 


86  THE   CAUDAL   SKELETON   IN 

not  reduced,  the  premaxilla  had  not  excluded  the  maxilla  from  the  gape,  and  in 
which  the  adipose  fin  had  disappeared.  "  By  extension,  the  ancestor  of  the  mycto- 
phoids  would  be  such  a  fish  with  an  adipose  fin.  On  the  other  hand,  Gosline  (1961  : 
35 ;  also  Gosline,  Marshall  &  Mead  1966  :  5)  points  out  that  the  large  caudal  scutes 
of  Aulopus  make  it  impossible  to  derive  the  myctophoids  from  any  living  teleost 
except  the  elopoids.  Weitzman  (1967  :  532)  discusses  this  point  and  notes  that 
caudal  scutes  are  present  in  Argentina,  but  the  structures  he  describes  in  the  salmonoid 
Plecoglossus  and  the  galaxioid  Retropinna  do  not  seem  to  resemble  caudal  scutes. 
The  absence  in  all  known  salmonoids  of  large  caudal  scutes  and  of  a  forked  first 
uroneural  of  the  type  found  in  Elops  and  Nematonotus,  together  with  the  absence  of 
recognizable  salmonoids  from  pre-Tertiary  rocks,  make  it  difficult  to  envisage  any 
direct  relationship  between  salmonoids  and  myctophoids  (Greenwood  et  al.  1966,  fig.  i) . 
A  short  second  pre-ural  neural  spine  resembling  those  of  Aulopus  and  Nematonotus 
occurs  in  some  salmonoids  (Coregonus,  which  also  has  a  urodermal),  but  the  last  few 
neural  arches  and  spines  and  the  epurals  are  apparently  very  variable,  and  no  clear 
pattern  emerges  from  Norden's  (1961  :  738)  analysis.  In  Argentina  caudal  scutes  are 
present  (though  they  are  reduced)  and  the  first  pre-ural  and  ural  centra  are  fused 
(Gosline  1960,  fig.  10),  as  they  must  have  been  in  the  common  ancestor  of  Myctophoidei 
and  Ctenothrissiformes,  but  the  neural  spine  of  the  second  pre-ural  centrum  is  elon- 
gate and  supports  procurrent  fin-rays :  in  this  character  Argentina  is  advanced  over  the 
basal  myctophoids  and  ctenothrissiforms.  In  the  Clupavidae,  a  family  with  a  fossil 
record  extending  back  to  the  Upper  Jurassic,  large  caudal  scutes  are  usually  present 
in  the  tail,  the  skull  seems  primitive  enough  to  have  given  rise  to  both  myctophoids 
and  Ctenothrissiformes,  and  the  caudal  skeleton  may  be  strikingly  like  that  of 
Nematonotus  (Patterson  19670,  fig.  n).  It  is  unlikely  that  the  known  clupavids 
were  ancestral  to  the  myctophoids  and  ctenothrissiforms  because  of  their  reduced 
dentition  and  clupeid-like  jaws,  with  a  high  coronoid  process  on  the  dentary,  but 
they  suggest  a  possible  source  for  this  type  of  caudal  skeleton. 

Taking  the  caudal  skeleton  of  Nematonotus  as  the  primitive  condition  for  the  mycto- 
phoids and  ctenothrissiforms,  the  myctophoids  are  characterized  by  shortening 
and  broadening  the  neural  spine  of  the  second  pre-ural  centrum,  and  in  more 
advanced  forms  by  fusion  of  the  hypurals  with  each  other  and  with  the  supporting 
centra.  Within  the  ctenothrissiforms  two  distinct  types  of  caudal  skeleton  occur: 
in  Ctenothrissa  and  Aulolepis  the  neural  spine  of  the  second  pre-ural  centrum  is 
expanded,  as  in  myctophoids,  tending  towards  the  condition  in  Berycoidei  and 
Percoidei;  in  Pateroperca  the  second  pre-ural  neural  spine  is  fully  developed,  as  it 
is  in  Polymixioidei.  In  Aulolepis  and  some  species  of  Ctenothrissa  there  are  pro- 
current  spines  in  front  of  the  caudal  fin :  this  is  specific  evidence  of  evolution  towards 
the  Berycoidei  and  like  the  reduction  of  the  second  pre-ural  neural  spine  in  these 
fishes  it  distinguishes  them  from  the  Polymixioidei  and  Dinopterygoidei,  in  which 
procurrent  caudal  spines  occur  only  in  advanced  forms. 

Within  the  Beryciformes,  the  structure  of  the  caudal  skeleton  gives  some  support 
to  the  division  of  the  order  into  three  suborders.  In  Polymixioidei  there  are  always 
six  hypurals  and  the  neural  spine  of  the  second  pre-ural  centrum  is  fully  developed. 
In  Berycoidei  the  second  pre-ural  spine  is  reduced  to  a  low  crest  (except  in  one 


MESOZOIC   ACANTHOPTERYGIAN   FISHES  87 

individual  of  Monocentris,  Fig.  14),  the  procurrent  rays  are  spinous,  and  there  are 
trends  towards  loss  of  the  sixth  hypural  and  fusion  of  the  stegural  and  second  ural 
centrum  with  the  preceding  centrum.  In  the  Dinopterygoidei,  already  known  to  be 
a  heterogeneous  group,  the  neural  spine  of  the  second  pre-ural  centrum  retains  the 
primitive  short  condition  (Aipichthys,  Pharmacichthys)  or  is  fully  developed  (Dinop- 
teryx,  Pycnosteroides)  and  there  are  trends  towards  loss  of  the  sixth  hypural.  The 
polymixioid  caudal  skeleton  (also  found  in  Dinopteryx  and  Pycnosteroides)  resembles 
that  of  the  ctenothrissiform  Pateroperca,  differing  only  in  having  no  caudal  scutes 
and  in  having  one  less  principal  ray.  The  caudal  skeleton  of  Berycoidei  resembles 
those  of  Aulolepis  and  Ctenothrissa,  differing  only  in  having  lost  the  caudal  scutes 
and  further  reduced  the  second  pre-ural  neural  spine.  Aipichthys  and  Pharma- 
cichthys seem  to  have  the  most  primitive  caudal  skeletons  known  in  Beryciformes, 
with  nineteen  principal  rays,  the  primitive  short  second  pre-ural  spine,  and,  at  least 
in  some  specimens  of  Aipichthys,  six  autogenous  and  separate  hypurals  and  a  free 
second  ural  centrum.  Although  both  Aipichthys  and  Pharmacichthys  are  too 
specialized  in  other  characters  to  have  given  rise  to  Beryciformes,  their  caudal  skele- 
ton could  give  rise  to  both  the  polymixioid  condition  (by  elongation  of  the  second 
pre-ural  neural  spine,  which  occurs  spontaneously  in  some  individuals  of  Aipichthys, 
Fig.  nC)  and  the  berycoid  condition  (by  shortening  of  the  second  pre-ural  neural 
spine). 

(b)  The  origin  ofPerciformes. 

Gosline  (i96ia)  discussed  the  caudal  skeleton  of  Perciformes  and  found  that  the 
most  generalized  type  contains  fifteen  branched  principal  rays,  no  neural  spine  on  the 
second  pre-ural  centrum,  three  epurals,  two  free  uroneurals,  no  free  ural  centra, 
five  autogenous  hypurals,  and  the  haemal  arches  of  the  second  and  third  pre-ural 
centra  autogenous  (see  also  Monod  1967,  fig.  3).  Gosline  mentioned  that  this 
type  of  caudal  skeleton  occurs  in  Kuhlia  (Kuhliidae),  Chaetodon  (Chaetodontidae), 
Polydactylus  (Polynemoidei)  and  juvenile  Sphyraena  (Sphyraenoidei) .  Monod  (1967) 
refers  to  this  type  of  caudal  skeleton  as  "  sciaeno-sparidien  banal  "  and  states  that 
it  occurs  in  many  Perciformes,  mentioning  Sciaena  (Sciaenidae),  Pagrus,  Sparus 
(Sparidae)  and  Gaterin  (Pomadasyidae) .  I  find  that  this  generalized  type  of  caudal 
skeleton  also  occurs  in  Centropomus  (Centropomidae) ,  Lateolabrax,  Polyprion, 
Dicentrarchus,  M  or  one,  Acanthistius  (Percichthyidae,  sensu  Gosline  1966),  Branchio- 
stegus  (Branchiostegidae),  Pomatomus  (Pomatomidae),  Brama  (Bramidae),  Arripis 
(Arripidae),  Lutjanus  (Lutjanidae),  Nemipterus,  Scolopsis  (Nemipteridae),  Lobotes 
(Lobotidae),  Xenocys,  Xenistius,  Xenichthys  (Pomadasyidae),  Lethrinus,  Sphaerodon 
(Lethrinidae),  Monodactylus  (Monodactylidae) ,  Kyphosus,  Medialuna  (Kyphosidae) , 
Ephippus,  Drepane,  Platax  (Ephippidae),  Chelmo,  Heniochus,  Pomacanthus  (Chaeton- 
ontidae),  Histiopterus  (Pentacerotidae),  Cirrhitus  (Cirrhitidae)  and  Schedophilus 
(Stromateoidei).  Many  other  groups,  among  them  the  Serranidae  (sensu  Gosline 
1966),  Cichlidae,  Percidae,  Acanthuroidei,  etc.  differ  from  this  basal  type  only  in 
the  loss  or  incorporation  in  the  stegural  of  the  second  uroneural.  The  occurrence 
of  an  apparently  identical  type  of  caudal  skeleton  in  such  a  wide  range  of  perciform 
groups,  including  forms  with  lunate,  forked,  emarginate  and  rounded  caudal  fins 


88  THE   CAUDAL   SKELETON   IN 

in  habitats  ranging  from  pelagic  to  lacustrine,  suggests  that  caudal  anatomy 
is  unlikely  to  contribute  much  to  the  unravelling  of  lineages  among  generalized 
Perciformes. 

A  caudal  skeleton  very  similar  to  the  basal  perciform  type  occurs  in  some  advanced 
Berycoidei  (living  Holocentridae  and  Diretmidae  differ  only  in  having  the  stegural 
fused  with  the  underlying  centrum  (Fig.  20),  Berycidae  have  the  stegural  fused  with 
the  centrum  and  also  retain  the  sixth  hypural  (Regan  1911,  fig.  i))  but  here  the 
hypurals  support  nineteen  principal  rays.  In  Myctophoidei  a  slightly  different 
sequence  of  fusion  is  followed  in  which  the  second  ural  centrum  partially  retains  its 
individuality.  So  far  as  I  know,  the  basal  perciform  caudal  skeleton  is  not  precisely 
duplicated  elsewhere. 

The  differences  between  the  caudal  skeleton  and  fin  of  basal  Perciformes  and  those 
of  generalized  Beryciformes  (Polymixia,  Aipichthys,  Pycnosteroides,  Monocentris)  are: 

(i)  Reduction  of  the  neural  spine  of  the  second  pre-ural  centrum.  This  has 
already  taken  place  in  all  Berycoidei. 

(ii)  Fusion  of  the  second  ural  centrum  with  the  preceding  two  centra.  This  has 
taken  place  in  most  living  Berycoidei  and  occurs  in  some  individuals  of  Aipichthys. 

(iii)  Loss  of  the  sixth  hypural  (the  development  of  the  caudal  skeleton  in  Mugil 
and  Sphyraena  (Hollister  1937)  suggests  that  Perciformes  have  lost  the  sixth  hypural, 
not  incorporated  it  in  the  fifth).  This  has  already  occurred  in  some  individuals  of 
Aipichthys  and  Pycnosteroides,  and  takes  place  during  the  evolution  of  the  Berycoidei 
(Holocentridae,  Diretmus] . 

(iv)  Reduction  of  the  number  of  principal  caudal  rays  from  nineteen  (Berycoidei, 
Aipichthys  and  Pharmacichthys]  or  eighteen  (Polymixioidei,  Dinopteryx  and  Pycnos- 
teroides) to  seventeen. 

(v)  In  all  living  Beryciformes,  the  foremost  procurrent  rays  in  each  caudal  lobe 
are  true  spines,  but  in  Perciformes  they  are  usually  (?  always)  unsegmented  lepido- 
trichia,  with  the  right  and  left  halves  separate,  and  this  is  true  of  the  earliest  Perci- 
formes (Prolates).  In  this  character  Perciformes  are  more  primitive  than  living 
Beryciformes.  Among  Cretaceous  Beryciformes,  all  Berycoidei,  like  their  living 
relatives,  have  spines  in  front  of  the  caudal  fin;  in  Polymixioidei  procurrent  spines 
occur  only  in  Homonotichthys  and  Pycnosterinx  dubius,  already  known  to  be  evolving 
towards  the  living  Polymixia  (Patterson  1964  :  301,  380),  and  in  Dinopterygoidei 
they  occur  only  in  Dinopteryx.  It  is  striking  to  find  that  spinous  procurrent  caudal 
rays  occur  only  in  those  Cretaceous  genera  (except  Dinopteryx}  already  known  to  be 
closely  related  to  riving  Beryciformes. 

The  first  of  these  five  differences,  the  condition  of  the  neural  spine  of  the  second 
pre-ural  centrum,  is  the  most  interesting.  As  discussed  above  (p.  84)  the  primitive 
condition  of  this  structure  in  teleosts  seems  to  be  as  in  Flops,  Aulopus  and  Nematono- 
tus,  where  the  spine  is  slender  and  about  half  as  long  as  its  predecessor.  This 
type  of  spine  may  elongate  so  that  it  supports  procurrent  fin-rays,  as  in  Polymixioidei 
and  many  primitive  telosts,  or  it  may  become  reduced  to  the  percoid  condition  (Fig. 
28) .  But  apart  from  these  two  simple  alternatives  there  are  other  possibilities  which 
complicate  the  issue.  First,  the  fully  developed  spine  might  become  detached  as  an 
epural,  producing  the  percoid  condition  direct.  Secondly,  from  a  low  neural  crest 


MESOZOIC   ACANTHOPTERYGIAN    FISHES  89 

of  percoid  type  an  apparent  full  neural  spine  might  develop  secondarily  by  fusion 
with  the  first  epural.  Thirdly,  an  apparent  neural  spine  on  the  second  pre-ural 
centrum  might  be  produced  by  fusion  between  the  second  and  third  pre-ural  centra. 
The  last  of  these  possibilities  can  normally  be  recognized  by  the  partial  or  complete 
doubling  of  the  neural  or  haemal  spine  on  the  compound  centrum,  as  in  Pleuronectes 
(Harrington  1937,  fig.  i)  and  the  specimen  of  Saurida  illustrated  by  Weitzman 
(1967,  fig.  18).  The  abnormal  specimen  of  Monocentris  illustrated  in  Fig.  14  is 
evidently  a  special  case  of  this  type  of  fusion,  where  the  neural  and  haemal  spines 
of  the  third  pre-ural  centrum  are  normally  double  and  the  posterior  half  of  the 
neural  spine  has  become  attached  to  the  succeeding  centrum.  This  type  of  fusion 
does  not  seem  of  general  significance  in  the  present  discussion. 

Fusion  of  an  epural  with  the  neural  crest  of  the  second  pre-ural  centrum  to  produce 
a  secondary  neural  spine  (Fig.  28E)  is  a  common  occurrence  in  acanthopterygians : 
this  process  appears  to  account  for  the  complete  neural  spine  on  the  second  pre-ural 
centrum  in  such  groups  as  the  Nandidae  (but  not  Pristolepis ;  Gosline  1968,  fig.  2b) 
among  Percoidei,  the  Channiformes,  Anabantoidei,  Luciocephalus ,  some  scombroids, 
pleuronectoid  and  soleoid  pleuronectiforms,  tetraodontiforms,  etc.  (Monod  1967; 
Liem  1963,  1967;  Gosline  1968).  In  Psettodes,  the  most  primitive  living  pleuronecti- 
form,  the  caudal  skeleton  (Monod  1967,  fig.  13)  is  of  basal  perciform  type,  with  five 
autogenous  hypurals  and  two  uroneurals,  but  there  is  only  one  free  epural  and  there 
appears  to  be  a  neural  spine  on  the  second  pre-ural  centrum.  Monod  identifies  this 
spine  as  the  first  epural,  for  the  element  is  partially  or  completely  autogenous  and 
the  suture  at  the  base  lies  not  between  the  arch  and  the  centrum  but  between  the 
arch  and  the  spine.  Psettodes  demonstrates  clearly  that  the  neural  spine  of  the 
second  pre-ural  centrum  in  Pleuronectoidei  and  Soleoidei  is  an  epural  which  has 
secondarily  regained  contact  with  and  fused  with  a  neural  arch.  That  this  has  also 
occurred  in  scombroids  such  as  Neothunnus  can  be  seen  by  comparing  figs.  15  and 
16  of  Monod  (1967) .  A  further  peculiarity  of  the  caudal  skeleton  of  pleuronectoids 
and  soleoids  is  that  the  parhypural  tapers  proximally  and  fails  to  make  contact  with 
the  centrum  (Monod  1967  :  117).  The  effect  of  this  is  to  give  dorso- ventral  sym- 
metry to  the  caudal  skeleton,  the  free  parhypural  opposing  the  single  epural  just  as 
the  neural  and  haemal  spines  of  the  second  pre-ural  centrum  oppose  one  another. 
A  free  parhypural,  tapering  proximally,  also  occurs  in  acanthopterygians  such  as 
the  Channiformes  (Monod  1967  :  117;  Gosline  1968,  fig.  20),  most  Anabantoidei 
(Liem  1963  :  32),  Luciocephalus  (Liem  1967  :  114)  and  balistoids  (Whitehouse  1910, 
pi.  50,  fig.  33;  Monod  1967  :  117),  all  forms  with  a  neural  spine  on  the  second  pre- 
ural  centrum.  Since  none  of  these  fishes  has  more  than  two  epurals,  all  these  groups 
appear  to  be  cases  of  secondary  fusion  between  the  first  epural  and  the  second  pre-ural 
neural  arch  in  order  to  increase  the  dorso-ventral  symmetry  of  the  caudal  skeleton. 
A  complete  neural  spine  on  the  second  pre-ural  centrum  also  occurs  occasionally  in 
basal  percoids:  Fig.  26  shows  such  a  structure  in  a  large  specimen  of  Siniperca 
(Percichthyidae).  In  this  individual  there  is  a  perfectly  formed  neural  arch  and 
spine  (npu2]  fully  fused  to  the  second  pre-ural  centrum,  and  there  are  only  two 
epurals  (ei,  2]  compared  with  the  three  of  normal  Siniperca,  most  percichthyids  and 
serranids.  This  specimen  is  best  regarded  as  an  abnormality  foreshadowing  the 


go 


THE   CAUDAL   SKELETON    IN 


fusion  of  the  first  epural  with  the  second  pre-ural  neural  arch  in  nandids,  pleuro- 
nectiforms,  etc. 

It  is  thus  well  established  that  in  many  perciform  groups  and  perciform  derivatives 
the  first  epural  can  fuse  with  the  second  pre-ural  neural  arch.  This  raises  the 
question  of  the  homology  of  the  perciform  first  epural:  does  the  frequent  fusion  of 
this  bone  with  the  second  pre-ural  centrum  indicate  that  these  two  structures  were 


h5. 


FIG.  26.  Siniperca  chuatsi  (Basilewsky).  The  caudal  skeleton  of  a  dried  skeleton  showing 
a  complete  neural  spine  on  the  second  pre-ural  centrum,  1888.3.23.3,  standard  length 
340  mm.,  Kiu  Kiang,  China.  For  explanation  of  lettering  see  p.  102. 

originally  part  of  the  same  segment?  If  so,  the  perciform  first  epural  may  have 
appeared  by  detachment  of  a  fully  developed  neural  spine  in  a  caudal  skeleton  of 
polymixiid  type.  Rosen  (1964  :  244)  suggested  that  this  took  place  in  the  evolution 
of  the  exocoetoids :  that  the  ancestral  exocoetoid  had  only  two  epurals  and  a  fully 
developed  neural  spine  on  the  second  pre-ural  centrum  which  became  detached  as 
the  foremost  of  the  three  epurals  in  such  a  fish  as  Dermogenys.  If  this  neural  spine 
became  detached  in  a  fish  which  still  retained  the  original  three  epurals  one  would 
expect  there  to  be  four  epurals.  Such  a  condition  occurs  very  occasionally  in 


MESOZOIC   ACANTHOPTERYGIAN    FISHES 

.d2 


h5 


•puUuUu2 


5  mm 


FIG.  27.  Pentaceropsis  recurvirostris  (Richardson).  Caudal  skeleton  of  a  dried  skeleton 
showing  four  epurals,  1869.2.24.24,  standard  length  405  mm.,  Tasmania.  For  explana- 
tion of  lettering  see  p.  102. 


the   generalized   percoid   caudal   skeleton,  as  in    the   individual    of  Pentaceropsis 
(Pentacerotidae)  shown  in  Fig.  27. 

The  epurals  of  teleosts  are  the  remnants  of  a  more  numerous  set  of  bones  in  primi- 
tive actinopterygians  which  are  serial  homologues  of  the  supraneurals  above  the 
anterior  vertebrae,  and  like  these  bones  they  were  primitively  metameric  (Patterson 
1968  :  221),  but  the  metameric  arrangement  of  the  epurals  has  been  lost  in  living 
chondrosteans.  In  the  most  primitive  teleosts,  such  as  the  Jurassic  Leptolepis  and 
Allothrissops  (Patterson  19670:,  figs.  3,  6),  the  three  epurals  still  show  an  apparent 
metamery,  suggesting  that  they  represent  the  neural  spines  of  the  first  pre-ural 
neural  arch  and  two  ural  neural  arches :  this  seems  to  be  supported  by  conditions  in 
osteoglossoid  fishes,  where  there  are  often  fully-developed  neural  spines  on  both 
the  first  pre-ural  and  the  first  ural  centra,  and  where  there  is  never  more  than  one 
epural  (Greenwood  1967).  But  in  higher  teleosts,  as  in  living  chondrosteans,  a 
metameric  arrangement  of  the  epurals  is  no  longer  recognizable.  At  the  percoid 
level,  for  example,  one  can  find  fishes  with  all  the  epurals  behind  the  neural  crest  on 
the  second  pre-ural  centrum  (Gosline  ig6ia,  fig.  i,  Kuhlia;  1968,  fig.^c.Bathymaster), 
or  with  one  epural  above  the  crest  (Gosline  ig6ia,  fig.  2,  Parupeneus;  Hollister  1937, 


92  THE   CAUDAL   SKELETON    IN 

figs.  12-14,  Sphyraena)  or  with  two  above  it  (Hollister  1937,  fig.  8,  Mugil).  Further, 
in  groups  such  as  the  Berycoidei  and  Percoidei  there  is  good  evidence  that  in  primi- 
tive forms  the  first  epural  lies  behind  the  neural  crest  of  the  second  pre-ural  centrum, 
moving  forwards  above  it  in  more  advanced  forms  (in  berycoids  cf.  Figs.  17,  21  with 
Figs.  13,  20).  Since  there  is  no  evidence  that  strict  metamery  of  the  epurals  is 
maintained  in  acanthopterygians  there  is  no  reason  to  believe  that  the  perciform 
first  epural  is  the  detached  neural  spine  of  the  second  pre-ural  centrum.  The 
individual  of  Pentaceropsis  shown  in  Fig.  27  is  best  interpreted  merely  as  exhibiting 
a  supernumary  epural:  such  a  condition  is  already  known  to  occur  in  the  salmonid 
Oncorhynchus,  where  Vladykov  found  four  epurals  in  three  out  of  1,020  specimens 
(1962,  tableS). 

The  conclusions  drawn  from  this  discussion  of  the  second  pre-ural  neural  arch 
and  spine  are  that  the  low  neural  crest  in  basal  percoids  is  to  be  regarded  as  having 
evolved  by  reduction  of  the  primitive  short  neural  spine,  and  that  the  ancestors  of 
the  perciforms  are  to  be  found  among  fishes  having  a  low  crest  or  short  neural  spine 
on  the  second  pre-ural  centrum,  not  among  those  with  a  full  neural  spine.  Where 
there  is  a  full  neural  spine  on  the  second  pre-ural  centrum  in  Perciformes  and  perci- 
form derivatives,  it  is  to  be  regarded  as  having  arisen  by  fusion  of  an  epural  with  a 
low  neural  crest.  These  conclusions  are  illustrated  in  Fig.  28. 

With  this  background  on  the  perciform  caudal  skeleton,  we  can  now  consider  the 
evidence  of  caudal  structure  in  the  various  beryciform-perciform  lineages  that  have 
been  suggested  (Patterson  1964).  These  were,  in  decreasing  order  of  confidence, 

Polymixiidae  (Omosoma — Berycopsis  lineage)  —  — >     Scorpididae, 

Monodactylidae  and 
Kyphosidae 

Aipichthyidae  — >     Carangidae 

Pharmacichthyidae  — >     Acanthuroidei 

Sphenocephalidae  — >     basal  Percoidei 

(Serranidae,  etc.) 

Pycnosteroididae  — >     Chaetodontidae 

Dinopterygidae  — >     Centrarchidae 

Detailed  study  of  the  caudal  skeleton  cannot  be  said  to  give  support  to  these 
lineages.  Of  the  various  perciform  groups  mentioned,  the  scorpidids,  monodacty- 
lids,  kyphosids,  basal  percoids  (Centropomidae,  Percichthyidae),  Chaetodontidae 
and  Centrarchidae  have  the  generalized  percoid  type  of  caudal  skeleton,  differing 
from  those  of  Beryciformes  in  the  characters  listed  on  p.  88.  The  acanthuroids 
are  more  advanced  only  in  having  lost  the  second  uroneural,  while  the  carangids 
have  enlarged  the  first  epural  and  show  fusion  between  the  first  and  second  hypural 
and  between  the  third  and  fourth  hypurals.  Among  the  beryciform  groups,  the 
polymixiids,  sphenocephalids,  pycnosteroidids  and  dinopterygids  all  have  a  com- 
plete neural  spine  on  the  second  pre-ural  centrum :  for  reasons  given  in  the  discussion 
above,  it  is  unlikely  that  a  percoid  caudal  skeleton  can  be  derived  directly  from  this 
condition.  In  Sphenocephalus  there  are  only  two  epurals,  suggesting  the  possibility 


MESOZOIC   ACANTHOPTERYGIAN    FISHES 


93 


of  arriving  at  a  percoid  arrangement  of  three  epurals  and  a  low  neural  crest  on  the 
second  pre-ural  centrum  by  detachment  of  the  neural  spine  on  this  centrum,  but 
further  study  of  Sphenocephalus  has  yielded  strong  evidence  (to  be  discussed  in 
a  forthcoming  paper  by  the  author  and  D.  E.  Rosen)  that  far  from  being  an  ancestral 
percoid  it  is  related  to  the  percopsiforms.  In  the  aipichthyids  and  pharmacichthyids 


FIG.  28.  Diagrams  to  show  changes  in  the  second  pre-ural  neural  spine  (black)  in  the 
evolution  of  the  myctophoids,  ctenothrissiforms  and  acanthopterygians.  A,  the  primi- 
tive short  neural  spine,  as  in  Nematonotus,  Aulopus,  Aipichthys  and  Pharmacichthys; 
B,  elongation  of  the  neural  spine  to  support  procurrent  fin-rays,  as  in  Pateroperca,  Poly- 
mixiidae,  Dinopteryx  and  Pycnosteroides ;  C,  expansion  of  the  neural  spine  into  a  plate,  as 
in  most  myctophoids,  Ctenothrissa,  A  ulolepis  and  some  primitive  Berycoidei ;  D,  reduction 
to  a  low  crest,  with  the  first  epural  (stippled)  moving  forwards  above  it,  as  in  most 
Berycoidei  and  Perciformes  (a  berycoid  is  illustrated) ;  E,  production  of  a  secondary  neural 
spine  by  fusion  of  the  first  epural  in  fishes  with  no  free  second  ural  centrum  and  five 
hypurals,  as  in  Zeiformes  (except  Caproidae),  Nandidae,  Channiformes,  Anabantoidei, 
Pleuronectiformes,  etc. 


94  THE   CAUDAL   SKELETON   IN 

the  neural  spine  of  the  second  pre-ural  centrum  is  still  short,  as  in  the  most 
primitive  teleosts,  and  could  give  rise  to  the  percoid  condition  by  reduction.  Further, 
in  Aipichthys  (Fig.  n)  there  is  a  tendency  to  reduce  the  number  of  hypurals  to  five 
and  also  to  fuse  the  second  ural  centrum  into  the  preceding  centrum,  as  in  percoids. 
There  seems  to  be  nothing  in  the  caudal  skeleton  of  Aipichthys  to  oppose  the  postu- 
lated link  with  the  carangids.  In  Pharmacichthys  the  caudal  skeleton  is  like  that 
of  Aipichthys,  so  far  as  it  is  known,  and  the  fin-rays  are  deeply  cleft  basally,  as  in 
carangids:  this  character  reduces  the  possibility  of  a  relationship  between  Phar- 
macichthys and  the  acanthuroids  and  balistoids,  in  which  the  caudal  fin-rays  are 
unmodified. 

Except  for  the  Aipichthys-carangid  lineage,  the  evidence  of  the  caudal  skeleton 
indicates  that  Gosline's  (19660)  criticism  of  the  polyphyletic  scheme  of  perciform 
origins  which  I  proposed  are  well  founded.  But  the  evidence  on  which  Gosline 
based  these  criticisms,  the  structure  of  the  supraoccipital  crest,  does  not  in  fact 
oppose  the  various  beryciform-perciform  lineages.  Gosline  (19660  :  412)  contrasted 
the  type  of  supraoccipital  crest  seen  in  the  polymixiids  Homonotichthys  and  Poly- 
mixia,  and  in  the  carangids,  priacanthids,  etc.,  which  extends  forwards  between  the 
frontals,  is  knife-edged  and  buried  in  musculature,  with  the  type  of  crest  seen  in 
Antigonia,  acanthuroids,  chaetodontids,  etc.,  which  is  short,  high  and  thickened 
anteriorly,  extending  "  up  and  back  over  the  nape  as  a  sort  of  protective  shell ". 
Gosline  finds  that  these  two  types  of  supraoccipital  crest  are  "  structurally  and 
functionally  ...  far  apart  ",  and  writes  "  nor  does  it  appear  that  one  could  be 
developed  from  the  other  except  by  going  all  the  way  back  through  some  inter- 
mediate form  with  a  relatively  small,  unspecialized  occipital  crest  ".  Gosline  draws 
the  conclusion  that  Aipichthys  and  Sphenocephalus ,  both  with  a  short  crest,  thickened 
anteriorly,  could  not  have  given  rise  respectively  to  the  carangids  and  serranids, 
which  have  a  long,  knife-edged  crest.  But  the  evolution  of  the  Berycoidei  shows 
(as  clearly  as  such  processes  can  be  shown  by  the  fossil  record)  that  fishes  with  a 
short  "  Aipichthys-type  "  crest  can  give  rise  to  fishes  with  a  long  "  carangid-type  " 
crest.  The  most  primitive  Berycoidei  have  an  "  Aipichthys-type,  "  crest,  moderately 
high  and  thickened  anteriorly :  this  is  true  of  both  the  trachichthyid  lineage  (Lisso- 
beryx,  Patterson  1967,  fig.  2)  and  the  holocentrid  lineage  (Caproberyx,  Patterson 
1964,  fig.  67;  1967,  fig.  10,  Stichocentrus,  Patterson  1967,  fig.  8).  From  this  basal 
type  there  are  in  berycoids  two  divergent  trends  in  the  evolution  of  the  supra- 
occipital  crest.  In  Holocentridae  the  frontals  grow  backwards,  partially  covering 
the  parietals,  eliminating  the  supratemporal  fossa,  and  producing  a  low  supra- 
occipital  crest  which  secondarily  comes  to  resemble  the  small  supraoccipital  crest 
of  primitive  teleosts.  In  Trachichthyidae  (Hoplopteryx,  Hoplostethus)  the  supra- 
temporal  fossa  extends  forwards  and  the  supraoccipital  crest  becomes  elongated, 
thickened  centrally  and  knife-edged.  This  trend  continues  further  in  the  Berycidae, 
and  Beryx  has  a  long  knife-edged  crest,  continued  forwards  by  the  frontals  above  the 
orbit,  which  resembles  those  of  Polymixia  and  the  carangids.  If  a  carangid- 
type  "  supraoccipital  crest  developed  from  an  "  Aipichthys-type. "  within  the 
Berycoidei,  there  is  no  reason  why  these  changes  could  not  have  occurred  in  other 
lineages. 


MESOZOIC   ACANTHOPTERYGIAN   FISHES  95 

It  should  also  be  mentioned  here  that  Gosline  (19660  :  410)  has  indicated  that 
Beryciformes  differ  from  Perciformes  in  the  number  of  infraorbitals  and  the  extent 
of  the  subocular  shelf.  He  finds  that  in  Beryciformes  (except  Holocentridae)  there 
are  four  circumorbitals  behind  the  lachrymal  whereas  in  Perciformes  there  are  five, 
and  that  in  Beryciformes  the  subocular  shelf  extends  over  more  than  one  infra- 
orbital  while  in  Perciformes  it  is  confined  to  the  second  infraorbital.  The  subocular 
shelf  extends  along  all  the  infraorbitals  in  Holocentridae  (living  and  fossil)  and 
in  Polymixiidae  (Polymixia,  Homonotichthys) ,  but  in  Trachichthyidae  (living 
and  fossil),  Berycidae,  Monocentridae,  etc.,  the  shelf  is  restricted  to  the  second 
infraorbital,  as  in  Perciformes.  The  subocular  shelf  also  extends  along  all  the 
infraorbitals  in  Anabantidae  and  Belontiidae  (Liem  1963) :  probably  a  subocular 
shelf  on  all  the  infraorbitals,  as  in  polymixiids,  holocentrids  and  anabantids,  is  the 
primitive  condition  of  the  structure  (Smith  &  Bailey  1962  :  3).  In  the  number  of 
infraorbitals  the  basic  beryciform  condition  is  undoubtedly  as  in  percoids,  with  a 
total  of  six  bones,  a  lachrymal,  four  infraorbitals,  and  a  dermosphenotic  overlying 
the  autosphenotic :  this  condition  occurs  in  Polymixiidae,  living  and  fossil,  and  in 
living  Holocentridae.  In  Trachichthyidae,  Berycidae,  Monocentridae  and  some 
Cretaceous  Holocentridae  (Caproberyx,Stichocentrus,  Patterson  1964  :  347;  1967  :  89) 
the  apparent  reduction  in  number  of  circumorbital  bones  is  caused  by  fusion  of 
the  dermo-  and  autosphenotics,  which  had  already  occurred  in  the  Cretaceous 
trachichthyid  Hoplopteryx  (Patterson  1964,  fig.  55). 

In  summary,  of  the  various  beryciform-perciform  lineages  which  have  been 
proposed,  only  the  Aipichthys — carangid  lineage  emerges  unscathed  from  a  detailed 
examination  of  caudal  structures.  The  caudal  skeletons  of  Polymixiidae,  Sphen- 
ocephalus,  Dinopteryx  and  Pycnosteroides  differ  fundamentally  from  those  of  Perci- 
formes in  having  a  full  neural  spine  on  the  second  pre-ural  centrum.  In  Aipichthys 
and  Pharmacichthys  there  are  tendencies  towards  the  Perciformes  in  the  occasional 
fusion  of  the  second  ural  centrum  with  the  preceding  centrum  and  the  occurrence 
of  five  hypurals,  but  it  is  only  among  the  Berycoidei  that  the  basal  perciform  caudal 
skeleton  is  duplicated.  All  known  Berycoidei,  even  the  very  generalized  Lissoberyx, 
are  more  specialized  than  Perciformes  in  having  procurrent  caudal  spines,  and  this 
and  characters  of  the  skull  (Patterson  1964  :  467)  show  that  no  perciform  could 
have  evolved  from  any  known  berycoid.  But  if  the  Perciformes  and  cognate 
groups  (Channiformes,  Scorpaeniformes,  Pleuronectiformes,  Tetraodontiformes) 
should  prove  to  be  a  monophyletic  group,  an  alternative  to  the  traditional  method — 
"  an  attempt  first  to  define  orders  and  other  higher  taxa  and  then  to  speculate  upon 
their  origin,  albeit  in  the  light  of  the  known  fossils  "  (Greenwood  et  al.  1966  :  346) 
is  to  use  the  criteria  recommended  by  Hennig  (1966  :  88,  120)  and  to  search  among 
the  living  fauna  for  the  sister  group  (Hennig  1966  :  139;  see  also  Brundin  1966  :  17) 
of  this  assemblage.  A  preliminary  analysis  suggests  that  the  Berycoidei,  not  the 
Beryciformes  as  a  whole,  may  fill  this  role.  This  is  indicated  not  only  by  features 
of  the  caudal  skeleton  but  by  the  fact  that  the  Berycoidei  is  the  only  beryciform 
group  showing  such  perciform  features  as  a  subocular  shelf  confined  to  the  second 
infraorbital,  the  absence  of  epineurals,  the  pelvic  girdle  firmly  joined  to  the  cleithra 
(in  Berycidae  especially),  pelvic  fins  containing  a  spine  and  five  soft  rays  (in  Anomalo- 


96  THE   CAUDAL   SKELETON   IN 

pidae  and  Gibberichthyidae,  for  example),  partially  separate  soft  and  spinous  dorsal 
fins,  etc.,  and  in  the  stephanoberycoids,  which  appear  to  be  merely  specialized 
offshoots  of  the  trachichthyid  lineage,  loss  of  the  orbitosphenoid.  While  it  is  clear 
that  many  of  these  perciform  characters  in  Berycoidei  have  arisen  independently 
within  the  group  and  were  not  inherited  from  a  common  ancestor  of  Berycoidei  and 
perciforms,  they  appear  to  be  true  parallelisms  (Simpson  1961  :  78),  and  are 
indicative  of  relationship. 

(c)  Intermediate  groups. 

Between  the  Beryciformes  and  the  basal  Percoidei,  Greenwood  et  al.  (1966  :  398) 
place  the  Zeiformes,  Lampridiformes,  Gasterosteiformes,  Channiformes,  Synbranchi- 
formes,  Scorpaeniformes,  Dactylopteriformes  and  Pegasiformes.  Of  the  Gastero- 
steiformes, Scorpaeniformes,  Dactylopteriformes  and  Pegasiformes  I  have  nothing 
to  say.  The  Channiformes  (see  Gosline  1968)  and  Synbranchiformes  are  probably 
derived  from  the  percoid  level  rather  than  from  the  beryciform  or  pre-beryciform. 

In  Zeiformes  the  caudal  skeleton  resembles  those  of  basal  Perciformes  in  having 
no  free  second  ural  centrum  and  in  having  only  five  hypurals,  and  is  more  advanced 
than  basal  percoids  in  having  lost  the  second  uroneural.  In  Zeidae  (Zeus,  Cyttus], 
Oreosomatidae  (Neocyttus)  and  Grammicolepidae  (Xenolepidichthys]  there  is  a  com- 
plete neural  spine  on  the  second  pre-ural  centrum,  but  this  never  occurs  in  conjunc- 
tion with  three  epurals,  and  in  Caproidae  (Capros,  Antigonia;  Gosline  1961,  fig.  4A), 
which  in  other  respects  appear  to  have  the  most  primitive  caudal  skeletons  of  the 
group  (the  hypurals  autogenous,  the  stegural  autogenous  in  Antigonia)  there  are 
three  epurals  and  there  is  a  low  crest  on  the  second  pre-ural  centrum,  as  in  percoids. 
Conditions  in  the  Caproidae  indicate  that  the  neural  spine  on  the  second  pre-ural 
centrum  in  Zeidae,  Oreosomatidae  and  Grammicolepidae  has  arisen  secondarily  by 
fusion  of  the  first  epural  (see  above,  p.  89,  Fig.  28),  and  that  as  Gosline  (1961)  has 
already  said,  there  is  nothing  in  the  caudal  skeleton  to  distinguish  Zeiformes  from 
Perciformes.  It  has  long  been  recognized  that  the  Zeiformes  are  probably  related 
to  the  Beryciformes,  principally  because  of  the  pelvic  ray  count,  but  they  also  show 
many  perciform  features  (Gosline  1961  :  36)  and  no  conclusion  has  yet  been  reached 
on  whether  they  are  more  closely  related  to  the  beryciforms  or  the  perciforms, 
although  in  most  recent  classifications  they  are  placed  directly  after  the  Beryci- 
formes. Stinton  (1967)  has  recently  shown  that  there  is  a  remarkable  resemblance 
between  the  otoliths  of  Antigonia  and  those  of  Berycoidei  (Berycidae,  Trachichthyi- 
dae,  and  especially  Monocentridae) .  Stinton  interprets  this  as  indicating  that 
Antigonia  is  a  berycoid,  for  he  finds  that  the  otoliths  of  Capros  resemble  those  of 
the  zeids  rather  than  Antigonia.  However,  the  evidence  that  Antigonia  and  Capros 
are  related  can  hardly  be  ignored,  and  in  Stinton's  illustrations  of  zeiform  otoliths 
it  seems  possible  to  recognize  a  trend  in  reduction  and  specialization  of  the  otolith 
in  the  sequence  Antigonia-Capros-Cyttus-Zeus:  the  same  sequence  of  increasing 
specialization  is  also  shown  by  fusion  within  the  caudal  skeleton.  In  my  opinion 
Stinton's  otolith  evidence  indicates  not  that  Antigonia  is  a  berycoid,  but  that  as 
the  most  primitive  living  zeiform  it  retains  the  clearest  evidence  of  a  common  ancestry 
with  the  Berycoidei.  The  percoid-like  caudal  skeleton,  pelvic  spine,  etc.,  of  the 


MESOZOIC    ACANTHOPTERYGIAN   FISHES  97 

Zeiformes  do  not  oppose  such  a  relationship.  In  Hennig's  terminology,  the  Zei- 
formes  appear  to  be  the  apomorph  sister  group  of  the  Berycoidei,  these  two  groups 
together  forming  the  plesiomorph  sister  group  of  the  perciform  assemblage  (see 
above,  p.  95). 

The  Lampridiformes,  previously  unknown  before  the  Oligocene,  have  recently 
acquired  a  respectable  fossil  record  with  Bonde's  (1966)  preliminary  description  of 
a  ?  veliferid  from  the  basal  Eocene  Mo-clay  of  Denmark,  his  opinion  that  Palaeo- 
centrotus  Kuhne  (1941),  from  the  same  beds,  is  a  lampridoid,  not  a  zeiform,  and  the 
suggestion  (p.  81)  that  the  Danian  Bathysoma  is  a  lampridiform,  not  a  menid. 
It  appears  that  the  deep-bodied  Lampridiformes  of  the  suborder  Lampridoidei  were 
an  important  element  of  early  Tertiary  faunas.  The  caudal  skeleton  of  Lampridi- 
formes (known  in  Velifer,  Lampris,  Palaeocentrotus  and  Bathysoma)  is  characterized 
by  fusion  of  one  or  more  of  the  upper  hypurals  with  the  second  ural  centrum,  but 
in  other  respects  it  does  not  differ  from  that  of  Beryciformes  (there  are  six  hypurals 
in  Velifer).  In  Lampris  and  Velifer  the  neural  spine  of  the  second  pre-ural  centrum 
is  reduced,  but  in  Palaeocentrotus  (Kiihne  1941,  fig.  2)  it  is  about  half  as  long  as  its 
predecessor,  a  primitive  condition  only  found  among  Beryciformes  in  Aipichthys 
(Fig.  n)  and  Pharmacichthys.  Further,  in  Lampris  and  Velifer  there  are  seventeen 
branched  caudal  rays  and  the  caudal  rays  are  deeply  cleft  basally,  covering  much  of 
the  hypurals.  These  points  tend  to  confirm  the  suggestion  (Patterson  1964  :  473) 
that  the  Lampridiformes  are  an  offshoot  of  the  Dinopterygoidei.  Within  the  Dino- 
pterygoidei,  the  second  pre-ural  neural  spine,  the  seventeen  branched  principal 
rays,  the  "  hypurostegy  "  and  the  absence  of  a  pelvic  spine  all  point  to  the  Aipich- 
thyidae  and  Pharmacichthyidae  as  ancestral  forms. 

Transference  of  Bathysoma  to  the  Lampridiformes  leaves  unsettled  the  position 
of  Mene,  to  which  I  thought  Bathysoma  was  related  (Patterson  1964  :  424).  The 
similarities  between  Mene  and  Bathysoma  are  numerous,  and  extend  to  the  form  of 
the  fin-rays,  which  are  preserved  in  the  middle  part  of  the  anal  fin  of  the  holotype 
of  B.  lutkeni  in  Copenhagen,  and  are  short,  broad  and  unbranched  (Bonde,  personal 
commn).  The  skull  of  Mene  is  also  strikingly  like  that  of  Velifer  (Regan  1907, 
figs.  167,  169).  But  the  caudal  skeletons  of  Mene  and  the  Lampridiformes  are 
very  different.  The  six  hypurals  (in  Velifer},  fusion  of  the  second  ural  centrum  with 
the  upper  hypurals  rather  than  with  the  preceding  centrum,  and  the  seventeen 
branched  principal  rays  of  Lampridiformes  can  only  be  derived  from  beryciform  or 
pre-beryciform  ancestors,  but  the  caudal  skeleton  of  Mene,  with  fifteen  branched 
principal  rays  and  fusion  of  the  second  ural  centrum  and  first  four  hypurals  with  the 
preceding  centrum  could  have  evolved  from  the  caudal  skeleton  of  basal  percoids 
(p.  81).  Nor  can  I  find  anything  in  the  skull  and  vertebral  column  of  Mene  which 
is  against  perciform  ancestry.  Nevertheless,  Mene  has  an  unusually  long  fossil 
record,  extending  back  to  the  Lower  Palaeocene.  The  earliest  recorded  species, 
M.  phosphaticus  Astre  (1927)  from  the  Montian  of  Tunisia,  seems  to  agree  with 
Mene  rather  than  with  the  Lampridiformes  in  caudal  structure.  Mene  is  very  com- 
mon in  the  Middle  Eocene  of  Monte  Bolca:  with  Mene  at  Monte  Bolca  there  occur 
other  deep-bodied  fishes  of  similar  structure  such  as  Exellia  ( =  Semiophorus) . 
These  fishes,  though  poorly  known,  seem  to  agree  with  Mene  in  the  absence  of  spines 


98  THE  CAUDAL  SKELETON   IN 

in  the  dorsal  and  anal  fins  (see  Blot  1967  on  Exellia}  and  also  in  the  deeply  cleft 
bases  of  the  caudal  fin-rays.  It  is  difficult  to  know  how  much  significance  can  be 
attached  to  this  last  character.  E.  &  Y.  Le  Danois  (1964),  who  coined  the  term 
"  hypurostegy  "  for  it,  give  great  importance  to  it  and  use  it  to  unite  in  an  "  ordre 
des  Scombres  "  fishes  as  diverse  as  the  holostean  Pachycormidae,  the  Cretaceous 
Tselfatiidae,  the  lampridiform  Veliferidae  and  Lampridae,  the  carangids,  scombrids 
and  others.  In  my  opinion  this  assemblage  is  entirely  spurious;  the  occurrence  of 
hypurostegy  in  such  varied  groups  indicates  not  that  the  fishes  are  related  but  that 
hypurostegy  has  arisen  independently  in  a  number  of  lines,  for  reasons  as  yet 
unknown.  E.  &  Y.  Le  Danois  made  a  new  family  Vomeridae  to  include  Mene,  the 
Eocene  Vomeropsis,  and  the  deep-bodied  carangids  Vomer,  Selene,  Alectis  and 
Hynnis,  grouping  this  family  with  the  Lampridae,  Veliferidae,  Ephippidae  and 
Exellia.  Although  the  reasoning  on  which  this  grouping  is  made  is  doubtful,  there 
may  be  some  truth  in  it.  It  seems  possible  that  Mene  and  Exellia  could  represent  an 
independent  attainment  of  the  perciform  grade  from  the  Palaeocene  lampridiform 
stock.  But  the  possibility  that  the  resemblances  between  these  Eocene  forms  and 
the  Lampridiformes  are  due  to  convergence  is  by  no  means  ruled  out:  revisionary 
studies  on  the  Monte  Bolca  fauna  now  in  progress  (Blot  1967)  may  settle  this  question. 

IV.  CONCLUSIONS 

Study  of  a  single  structural  complex  such  as  the  caudal  skeleton  is  unlikely  to 
produce  firm  conclusions  on  matters  of  phylogeny  and  relationships.  Rather  it 
will  serve  as  a  means  of  checking  existing  hypotheses  and  will  raise  questions  to  be 
settled  by  more  comprehensive  work.  The  main  points  arising  from  this  paper  are 
as  follows.  An  asterisk  indicates  that  the  genus  or  group  is  extinct. 

1.  Ctenothrissiformes,*  Myctophoidei  and  Beryciformes  have  a  basically  similar 
caudal  skeleton,  with  the  first  ural  and  pre-ural  centra  fused,  a  free  second  ural 
centrum,  a  stegural  (the  first  uroneural  fused  with  neural  arch  material  from  the 
first  ural  and  pre-ural  centra),  a  second  uroneural,  three  epurals  and  six  hypurals. 
The  primitive  nineteen  principal  caudal  rays  are  retained  in  all  ctenothrissiforms 
and  myctophoids,  and  in  all  beryciforms  except  the  Polymixiidae,  Sphenocephalidae,* 
Dinopterygidae  *  and  Pycnosteroididae,*  which  have  eighteen. 

2.  Ctenothrissiformes  *    (Aulolepis,    Ctenothrissa)    and    Myctophoidei    (Aulopus, 
Nematonotus,*  Sardinioides*  Acrognathus  *)  have  a  large  caudal  scute  above  and 
below   the   caudal   skeleton.     The   myctophoids   Nematonotus  *   and  Sardinioides 
attenuatus  *  have  a  single  urodermal  on  the  base  of  the  upper  caudal  rays.     Both 
caudal  scutes  and  urodermals  are  relict  structures  absent  in  all  higher  groups. 

3.  The  neural  spine  of  the  second  pre-ural  centrum  in  teleosts  is  primitively  slender 
and  about  half  as  long  as  its  predecessor,  as  in  Elops  and  Leptolepis.*    This  type  of 
second  pre-ural  neural  spine  persists  in  the  myctophoids  Aulopus  and  Nematonotus  * 
and  in  the  beryciforms  Aipichthys  *  and  Pharmacichthys.*     In  Ctenothrissiformes,* 
Ctenothrissa  and  Aulolepis  have  this  short  spine  expanded  into  a  plate,  as  it  is  in 
most  myctophoids,  but  in  Pateroperca  the  spine  is  elongate  and  supports  procurrent 
fin-rays.     A  fully  developed  second  pre-ural  neural  spine  also  occurs  among  Beryci- 
formes in  all  polymixioids  (Polymixiidae,  Sphenocephalidae*)  and  in  the  dinop- 


MESOZOIC   ACANTHOPTERYGIAN    FISHES  99 

terygoids  Dinopfayx*  and  Pycnosteroides.*     In  Berycoidei,  as  in  generalized  Perci- 
formes,  the  second  pre-ural  neural  arch  and  spine  are  reduced  to  a  low  crest. 

4.  The  differences  between  the  caudal  skeletons  of  Myctophoidei  and  Cteno- 
thrissiformes*  and  those  of  the  most  primitive  teleosts  are  minor.     Among  primitive 
teleosts  the  Clupavidae  *  come  closest  to  the  Ctenothrissiformes  and  Myctophoidei 
in  caudal  anatomy. 

5.  The  foremost  procurrent  caudal  fin-rays  are  spinous  in  all  living  Beryciformes. 
This  is   a   feature   peculiar   to  Beryciformes:    Perciformes  appear  to   be   without 
procurrent  caudal  spines.     Procurrent  caudal  spines  also  occur  in  some  species  of 
Ctenothrissa  and  in  Aulolepis  (Ctenothrissiformes*),  probably  an  indication  of  rela- 
tionship between  Ctenothrissiformes  and  Beryciformes,  although  procurrent  caudal 
spines  have  also  developed  in  advanced  members  of  the  myctophoid  family  Mycto- 
phidae.     Among  Cretaceous  Beryciformes,  procurrent  caudal  spines  occur  only  in 
forms  already  known  to  be  closely  related  to  living  Beryciformes  (all  Berycoidei  and 
the  polymixiids  Homonotichthys*  and  Pycnosterinx*)  and  in  Dinopteryx.* 

6.  Within  the  Beryciformes,  the  principal  variations  in  the  caudal  skeleton  and 
fin  (apart  from  those  in  the  second  pre-ural  neural  spine,  principal  fin-ray  count  and 
procurrent  fin-rays  already  mentioned)  are  the  presence  of  only  two  epurals  in 
Sphenocephalus*  (Polymixioidei),  the  presence  of  only  five  hypurals  in  Aipichthys 
velifer*  some  specimens  of  Pycnosteroides*  (both  Dinopterygoidei)  and  in  many 
Berycoidei  (living  holocentrids,  Diretmus),  and  the  fusion  of  the  second  ural  centrum 
with  the  preceding  centrum  in  some  specimens  of  Aipichthys*  and  in  many  Bery- 
coidei   (Berycidae,    Diretmidae,    Anoplogasteridae,   living   Holocentridae) .     A   full 
neural  spine  on  the  second  pre-ural  centrum  has  been  found  in  one  individual  of 
Aipichthys*  and  one  of  Monocentris,  simulating  the  polymixoid  condition. 

7.  The  basal  perciform  caudal  skeleton  differs  from  the  basal  beryciform  type  in 
having  only  five  hypurals,  no  free  second  ural  centrum  and  only  seventeen  principal 
rays.     Perciformes  are  primitively  characterized  by  having  a  low  neural  crest  on 
the  second  pre-ural  centrum.     Where  a  complete  second  pre-ural  neural  spine  occurs 
as  a  normal  feature  in  perciform  or  higher  groups  the  condition  is  secondary,  the 
spine  representing  an  epural  which  has  secondarily  fused  with  the  neural  arch. 

8.  Evidence  from  the  caudal  skeleton  does  not  support  the  various  independent 
beryciform — perciform  lineages  which  have  been  proposed.     The  polymixioids  and 
the  dinopterygoids  Dinopteryx*  and  Pycnosteroides*  differ  fundamentally  from  the 
various  perciform  groups  which  they  otherwise  resemble  in  having  a  fully  developed 
neural  spine  on  the  second  pre-ural  centrum.     Pharmacichthys*  which  resembles 
the  acanthuroids  and   balistoids  in  many  ways,  differs  from  them  in  having  the 
bases  of  the  caudal  fin-rays  deeply  cleft,  and  is  therefore  unlikely  to  have  been  ances- 
tral to  these  groups.     Only  the  postulated  link  between  Aipichthys*  and  the  caran- 
gids  is  not  opposed  by  evidence  from  the  caudal  skeleton  and  fin. 

9.  Although  no  known  berycoid  could  have  been  ancestral  to  any  perciform,  the 
Berycoidei  is  the  only  beryciform  group  in  which  the  caudal  skeleton  evolves  towards 
the  percoid  condition.     There  are  many  other  characters  and  evolutionary  trends 
in  which  the  Berycoidei  is  the  only  beryciform  group  to  resemble  the  percoids :  these 
suggest  that  the  Berycoidei  is  the  sister  group  of  the  Perciformes  and  cognate  groups. 


TOO  THE   CAUDAL   SKELETON   IN 

10.  The  caudal  skeleton  shows  that  the  Danian  Bathysoma*  is  a  lampridiform 
(the  earliest  yet  known),  not  a  member  of  the  Menidae.     It  is  suggested  that  the 
Lampridiformes  originated  from  near  the  beryciform  families  Aipichthyidae*  and 
Pharmacicthyidae.*     In  the  Eocene  there  are  fishes  (Mene,  Exellia*}  which  have 
reached  the  perciform  grade  but  resemble  the  Palaeocene  and  Eocene  Lampridi- 
formes :  possibly  these  forms  represent  an  independent  attainment  of  the  perciform 
grade  from  lampridiform  ancestors. 

11.  The  caudal  skeleton  of  the  Zeiformes  is  basically  of  percoid  type,  but  the 
evidence  of  zeiform  otoliths  indicates  that  they  are  closely  related  to  the  Berycoidei. 
The  Zeiformes  is  evidently  the  sister  group  of  the  Berycoidei,  these  two  groups 
together  being  the  sister  group  of  the  perciform  assemblage. 

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102  THE   CAUDAL   SKELETON    IN   AC  ANTHOPTERYGI  AN    FISHES 

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—  191 1.     The  anatomy  and  classification  of  the  teleostean  fishes  of  the  Orders  Berycomorphi 

and  Xenoberyces.     Ann.  Mag.  nat.  Hist.,  London,  (8)  7  :  1-9,  pi.  i. 
—  191 1«.     The  Anatomy  and  Classification  of  the  Teleostean  Fishes  of  the  Order  Iniomi. 

Ann.  Mag.  nat.  Hist.,  London,  (8)  7  :  120-133,  7  figs. 
ROSEN,  D.  E.     1962.     Comments  on  the  Relationships  of  the  North  American  Cave  Fishes  of 

the  Family  Amblyopsidae.     Amer.  Mus.  Novit.,  New  York,  2109  :  1-35,  24  figs. 
1964.     The  relationships  and  taxonomic  position  of  the  halfbeaks,  killifishes,  silversides, 

and  their  relatives.     Bull.  Amer.  Mus.  Nat.  Hist.,  New  York,  127  :  217-268,  pis.  14,  15. 
SIMPSON,  G.  G.     1961.     Principles  of  Animal  Taxonomy,     xii  +  247  pp.,  30  figs.     Columbia 

Univ.,  New  York. 
SMITH  C.  L.  &  BAILEY,  R.  M.     1962.     The  Subocular  Shelf  of  Fishes.     /.  Morph.,  Philadelphia, 

110  :  1-18,  3  pis. 
SMITH,  J.  L.  B.     1951.     The  Fishes  of  the  Family  Veliferidae  from  South  Africa.     Ann.  Mag. 

nat.  Hist.,  London,  (12)  4  :  497-510,  pis.  10-12. 

STARKS,  E.  C.     1904.     The  osteology  of  some  Berycoid  fishes.     Proc.  U.S.  Nat.  Mus.,  Washing- 
ton, 27  :  601-619,  9  figs. 
STINTON,  F.  C.     1967.     The  otoliths  of  the  Teleostean  fish  Antigonia  capros  and  their  taxonomic 

significance.     Bocagiana,  Funchal,  13  :  1-7,  2  pis. 
VLADYKOV,  V.  D.     1954.     Taxonomic  characters  of  the  eastern  North  American  chars  (Salveli- 

nus  and  Cristivomer] .     J.  Fish.  Res.  Bd.  Can.,  Ottawa,  11  :  904-932,  12  figs. 
1962.     Osteological  studies  on  Pacific  salmon  of  the  genus  Oncorhynchus .     Bull.  Fish. 

Res.  Bd.  Can.,  Ottawa,  136  :  1-172,  89  figs. 
WEITZMAN,  S.  H.     1962.     The  osteology  of  Brycon  meeki,  a  generalized  characid  fish,  with  an 

osteological  definition  of  the  family.     Stanford  ichthyol.  Bull.,  Palo  Alto,  8  :  1-77,  21  figs. 

1967.     The  origin  of  the  stomiatoid  fishes  with  comments  on  the  classification  of  salmoni- 

form  fishes.     Copeia,  Washington,  1967  :  507-540,  18  figs. 

WHITEHOUSE,  R.  H.     1910.     The  caudal  fin  of  the  Teleostomi.     Proc.  zool.  Soc.  Lond.,  1910  : 

590-627,  pis.  47-50. 
WOODWARD,  A.  SMITH.     1901.     Catalogue  of  the  fossil  fishes  in  the  British  Museum  (Natural 

History}.     4.     xxxviii  +  636  pp.,  19  pis.     Brit.  Mus.  (Nat.  Hist.),  London. 
1942.     Some  new  and  little-known  Upper  Cretaceous  fishes  from  Mount  Lebanon.     Ann. 

Mag.  nat.  Hist.,  London,  (n)  9  :  537-568,  5  pis. 

VI.  ABBREVIATIONS  USED  IN  FIGURES 

dz  second,  uroneural 

e  1-3  epurals 

f.s  caudal  scute 

h  1-6  hypurals 

hpu2  haemal  spine  of  second  pre-ural  centrum 

npu2,  npu3  neural  spines  of  second  and  third  pre-ural  centra 

ph  parhypural  (haemal  spine  of  first  pre-ural  centrum) 

pui  +  ui  centrum  formed  by  fusion  of  first  pre-ural  and  ural  centra 

puz,  pus,  pu4  second,  third  and  fourth  pre-ural  centra 

st  stegural  (first  uroneural  fused  with  pre-ural  neural  arch  material) 

U2  second  ural  centrum 

Combinations  of  symbols  linked  by  plus  signs  indicate  compound  elements  formed  by  fusion 
of  the  bones  indicated. 


PRINTED  IN  GREAT  BRITAIN 
BY  ADLARD  &  SON  LIMITED 
BARTHOLOMEW  PRESS,  DORKING 


NON-CALCAREOUS 

MICROPLANKTON  FROM  THE 

CENOMANIAN  OF  ENGLAND, 

NORTHERN  FRANCE  AND 

NORTH  AMERICA 

PART  I 


R.  J.  DAVEY 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
GEOLOGY  Vol.  17  No.  3 

LONDON:   1969 


NON-CALCAREOUS  MICROPLANKTON  FROM 

THE  CENOMANIAN  OF  ENGLAND, 
NORTHERN  FRANCE  AND  NORTH  AMERICA 

PART  I 


BY 


ROGER  JACK  DAVEY 


Pp.  103-180;  ii  Plates;  16  Text-figures 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
GEOLOGY  Vol.  17  No.  3 

LONDON:  1969 


THE     BULLETIN    OF    THE    BRITISH    MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they  become 
ready.  Volumes  will  contain  about  three  or  four 
hundred  pages,  and  will  not  necessarily  be  completed 
within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  17,  No.  3  of  the  Geological 
(Palaeontological]  series.  The  abbreviated  titles  of 
periodicals  cited  follow  those  of  the  World  List  of 
Scientific  Periodicals. 


World  List  abbreviation : 
Bull.  Br.  Mus.  nat.  Hist.  (Geol.) 


Trustees  of  the  British  Museum  (Natural  History)  1969 


TRUSTEES    OF 
THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 

Issued  17  January,  1969  Price  £2     i6s, 


NON-CALCAREOUS  MICROPLANKTON  FROM 

THE  CENOMANIAN  OF  ENGLAND, 
NORTHERN  FRANCE  AND  NORTH  AMERICA 

PART  I 

By  ROGER  JACK  DAVEY 


Manuscript  accepted  May  1967 
CONTENTS 


SYNOPSIS 
I.     INTRODUCTION 


Page 
107 
107 


Acknowledgments      .          .          .          .          .          .          .          .  no 

II.     STRATIGRAPHIC  LOCATION  OF  SAMPLES         .          .          .          .          .  112 

Fetcham  Mill  Borehole        .          .          .          .          .          .          .  112 

Compton  Bay    .........  112 

Speeton    ..........  112 

Hunstanton       .........  115 

Devon      ..........  115 

Escalles  Borehole       .          .          .          .          .          .          .          .  115 

Saskatchewan   .........  115 

Texas        ..........  115 

III.     SYSTEMATIC  DESCRIPTIONS         .         .         .         .         .         .         .  120 

Genus  Gonyaulacysta  Deflandre       .          .          .          .          .          .  120 

Gonyaulacysta  cassidata  (Eisenack  &  Cookson)        .          .          .  120 

whitei  Sarjeant      .          .          .          .          .          .  120 

fetchamensis  Sarjeant     .          .          .          .          .  120 

exilicristata  sp.  nov.        .          .          .          .          .  121 

delicata  sp.  nov.  .          .          .          .          .  123 

sp.  A. 124 

Genus  Cribroperidinium  Neale  &  Sarjeant         .          .          .          .  125 

Cribroperidinium  intricatum  sp.  nov.  .          .          .          .  125 

Other  species     .........  128 

Cribroperidinium  orthoceras  (Eisenack)  .          .          .          .  128 

Genus  Carpodinium  Cookson  &  Eisenack  .          .          .          .  129 

Carpodinium  obliquicostatum  Cookson  &  Hughes    .          .          .  129 

Genus  Ellipsodinium  Clarke  &  Verdier     .          .          .          .          .  129 

Ellipsodinium  rugulosum  Clarke  &  Verdier    .          .          .          .  130 

Genus  Apteodinium  Eisenack  .          .          .          .          .          .  130 

Apteodinium  granulatum  Eisenack         .          .          .          .          .  130 

Genus  Trichodinium  Eisenack  &  Cookson          .          .          .          .  131 

Trichodinium  castaneum  (Deflandre)      .          .          .          .          .  131 

Genus  Microdinium  Cookson  &  Eisenack  .          .          .          .  132 

Microdinium  cf.  ornatum  Cookson  &  Eisenack        .          .          .  132 

setosum  Sarjeant     .          .          .          .          .          .  133 

distinctum  sp.  nov.  .          .          .          .          .  133 

variospinum  sp.  nov.        .          .          .          .          .  135 

veligerum  (Deflandre)       .          .          .          .          .  136 

crinitum  sp.  nov.  .          .          .          .          .  137 

GEOL.  17,  3  i 


106  CENOMANIAN   NON-CALCAREOUS   MICROPLAN  KTON,    i 

Genus  Histiocysta  nov.  .          .          .          .          .          .          .  138 

Histiocysta  palla  sp.  nov.     .          .          .          .          .          .          .  138 

Genus  Fromea  Cookson  &  Eisenack  .          .          .          .          .  140 

Fromea  amphora  Cookson  &  Eisenack  .          .          .          .  140 

Genus  Chytroeisphaeridia  Sarjeant  .          .          .          .          .  140 

Chytroeisphaeridia  euteiches  sp.  nov.      .....  141 

Genus  Cassiculosphaeridia  nov.        .          .          .          .          .          .  141 

Cassiculosphaeridia  reticulata  sp.  nov.  .....  142 

Genus  Epelidosphaeridia  nov.  .          .          .          .          .          .  142 

Epelidosphaeridia  spinosa  (Cookson  &  Hughes)       .          .          .  143 

Genus  Hystrichosphaeridium  Deflandre     .          .          .          .          .  143 

Hystrichosphaeridium  tubiferum  (Ehrenberg)  .          .          .  143 

deanei  Davey  &  Williams     .          .          .  144 

readei  Davey  &  Williams      .          .          .  144 

radiculatum  Davey  &  Williams      .          .  144 

mantelli  Davey  &  Williams  .          .  145 

bowerbanki  Davey  &  Williams       .          .  145 

difficile  Manum  &  Cookson  .          .          .  145 

Genus  Oligosphaeridium  Davey  &  Williams       .          .          .          .  146 

Oligosphaeridium  complex  (White)          .          .          .          .          .  146 

reticulatum  Davey  &  Williams     .          .          .  147 

prolixispinosum  Davey  &  Williams      .          .  147 

anthophorum  (Cookson  &  Eisenack)      .          .  147 

reniforme  (Tasch)       .          .          .          .          .  148 

Genus  Litosphaeridium  Davey  &  Williams         .          .          .          .  148 

Litosphaeridium  siphoniphorum  (Cookson  &  Eisenack)    .          .  148 

Genus  Polysphaeridium  Davey  &  Williams        .          .          .          .  151 

Polysphaeridium  pumilum  Davey  &  Williams         .          .          .  151 

laminaspinosum  Davey  &  Williams       .          .  151 

Genus  Tanyosphaeridium  Davey  &  Williams     .          .          .          .  151 

Tanyosphaeridium  variecalamum  Davey  &  Williams        .          .  151 

Genus  Callaiosphaeridium  Davey  &  Williams    .          .          .          .  151 

Callaiosphaeridium  asymmetricum  (Deflandre  &  Courteville)     .  152 

Genus  Cleistosphaeridium  Davey,  Downie,  Sarjeant  &  Williams  152 

Cleistosphaeridium  heteracanthum  (Deflandre  &  Cookson)          .  152 

multifurcatum  (Deflandre)          .          .          .  152 

armatum  (Deflandre)         .          .          .          .  153 

polypes  (Cookson  &  Eisenack)  .          .          .  154 

var.  clavulum  nov.  .          .          .          .          .  154 

huguonioti  (Valensi)  .          .          .          .  155 

var.  pertusum  nov.  .          .          .          .  156 

flexuosum    Davey,     Downie,    Sarjeant     & 

Williams     ......  157 

parvum  sp.  nov.       .....  157 

aciculare  sp.  nov.     .          .          .          .          .  158 

Genus  Surculosphaeridium  Davey,  Downie,  Sarjeant  &  Williams  158 

Surculosphaeridium  longifurcatum  (Firtion)    .          .          .          .  158 

Genus  Hystrichokolpoma  Klumpp    .          .          .          .          .          .  159 

Hystrichokolpoma  ferox  (Deflandre)        .          .          .          .          .  159 

Genus  Prolixosphaeridium  Davey,  Downie,  Sarjeant  &  Williams  160 

Prolixosphaeridium  conulum  sp.  nov.    .          .          .          .          .  160 

Genus  Coronifera  Cookson  &  Eisenack     .....  161 

Coronifera  oceanica  Cookson  &  Eisenack        .          .          .          .  162 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  107 

Genus  Exochosphaeridium  Davey,  Downie,  Sarjeant  &  Williams  162 

Exochosphaeridium  phragmites  Davey  et  al.    .          .  .  .  163 

pseudohystrichodinium  (Deflandre)  .  .  163 

striolatum  (Deflandre)      .          .  .  .  164 

var.  truncatum  nov.          .          .  .  .  164 

Other  species     .........  166 

Genus  Cyclonephelium  Deflandre  &  Cookson      .          .  .  .  166 

Cyclonephelium  distinction  Deflandre  &  Cookson     .  .  .  166 

membraniphorum  Cookson  &  Eisenack  .  .  167 

vannophorum  sp.  nov.  .          .  .  .  168 

paucispinum  sp.  nov.  .          .  .  .  170 

eisenacki  sp.  nov.  .          .          .  .  .  170 

Genus  Adnatosphaeridium  Williams  &  Downie  .  .  .  171 

Adnatosphaeridium  chonetum  (Cookson  &  Eisenack)  .  .  171 

Genus  Hystrichosphaera  O.  Wetzel  .          .          .          .  .  .  172 

Hystrichosphaera  ramosa  (Ehrenberg)    .          .          .  .  .  172 

var.  ramosa  (Ehrenberg)      .          .  .  .  172 

var.  gracilis  Davey  &  Williams    .  .  .  172 

var.  multibrevis  Davey  &  Williams  .  .  173 

var.  reticulata  Davey  &  Williams  .  .  173 

cingulata  (O.  Wetzel)  .          .  .  .  173 

var.  reticulata  Davey  &  Williams  .  .  174 

crassimurata  Davey  &  Williams  .  .  .  174 

crassipellis  Deflandre  &  Cookson  .  .  174 

Genus  Achomosphaera  Evitt.  .          .          .          .  .  .  174 

Achomosphaera  ramulifera  (Deflandre)  .          .  .  .  174 

sagena  Davey  &  Williams    .          .  .  .  174 

Genus  Hystrichodinium  Deflandre   .          .          .          .  .  .  174 

Hystrichodinium  voigti  (Alberti)   .          .          .          .  .  .  175 

dasys  sp.  nov.    .          .          .          .  .  .  175 

IV.     REFERENCES   .          .          .          .          .          .          .          .  .  .176 

SYNOPSIS 

This  paper,  which  will  appear  in  two  parts,  presents  the  results  of  a  detailed  study  of  some 
non-calcareous  microplankton  from  the  Upper  Cretaceous,  and  in  particular  of  assemblages  of 
Cenomanian  age.  The  stratigraphical  potentialities  of  the  fossil  microplankton  are  briefly 
assessed  by  the  analysis  of  samples  from  five  localities  in  England  and  one  in  France.  To 
assess  the  potentialities  of  long-range  correlation,  assemblages  from  Saskatchewan  and  Texas 
have  also  been  examined.  Both  quantitative  and  qualitative  methods  have  been  employed 
and  the  correlations,  both  intra-  and  inter-regional,  are  promising.  Seven  new  genera  and 
thirty-five  new  species  and  varieties  are  described. 

I.    INTRODUCTION 

FOSSIL  non-calcareous  microplankton  consist  mainly  of  cysts  of  dinoflagellates, 
together  with  various  forms  of  unknown  affinity  placed  in  the  Group  Acritarcha 
Evitt  (1963).  The  majority  of  dinoflagellates  are  free-living,  oceanic  and  plank- 
tonic.  They  have  a  complex  life-cycle,  usually  composed  of  four  stages,  in  one  of 
which  (the  motile  stage)  they  are  capable  of  limited  vertical  movement  by  the  use 
of  two  flagella.  During  the  life  cycle,  if  the  organism  is  subject  to  adverse  con- 
ditions, a  resting  cyst  is  formed.  This,  most  palynologists  believe,  is  the  only  stage 


io8 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 


in  the  dinoflagellate  life-cycle  preserved  in  the  fossil  state.  Fossil  non-calcareous 
microplankton  are  useful  as  stratigraphic  indices  because  they  are  planktonic,  of 
relatively  resistant  composition,  abundant  in  most  marine  sedimentary  samples,  and 
easy  to  extract.  Hence  the  principal  object  of  the  study  was  to  assess  how  accurate 
dinoflagellate  cysts  are  for  intra-  and  inter-regional  stratigraphic  correlations. 
The  order  of  description  of  the  dinoflagellate  cyst-families  follows  that  in  Sarjeant 


FIGURE    ONE 

MAP    of  SPECIMEN    LOCALITIES 
in  ENGLAND  and  FRANCE 


FIG.  i.     Map  of  specimen  Localities  in  England  and  France. 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  109 

&  Downie  (1966).     The  adjectives  used  when  describing  the  frequency  of  a  species 
in  an  assemblage  have  been  defined  as  follows: — 

Very  Common  10-100% 

Common  1-10% 

Infrequent  0-1-1% 

Rare  0-01-0-1% 

Very  Rare  under  o  •  o  i  % 

The  first  fossil  microplankton  were  described  and  figured  by  Ehrenberg  (1838, 
1843,  1854)  and  included  forms  embedded  in  flakes  of  Upper  Cretaceous  flint  from 
Germany  and  Denmark.  These  observations  stimulated  English  microscopists  in 
the  mid-nineteenth  century  into  finding  these  organisms  in  English  flints  of  the  same 
age. 

Little  further  research  in  this  sphere  was  published  until  1933  when  O.  Wetzel 
described  assemblages  from  Upper  Chalk  flints  of  the  Baltic  region.  In  1934 
Deflandre  published  the  first  of  a  number  of  well  illustrated  papers  dealing  with 
fossil  microplankton  from  the  flints  of  the  Paris  Basin.  Unfortunately  the  strati- 
graphic  horizons  of  the  flints  are  unknown,  many  being  picked  up  in  the  streets  of 
Paris.  His  two  most  important  papers  describing  Upper  Cretaceous  microplankton 
were  published  in  1936  and  1937.  They  contain  accurate  figures  and  descriptions 
of  many  new  species  and  genera  mainly  from  the  Senonian,  although  some  of  the 
flints  are  probably  of  Cenomanian  and  Turonian  age. 

Between  1936  and  1952  few  papers  dealing  with  microplankton  from  the  Upper 
Cretaceous  were  published.  Firtion  (1952)  described  the  first  definite  Cenomanian 
assemblage,  the  material  having  been  obtained  from  the  Lower  Cenomanian  of 
France.  All  his  species  have  subsequently  been  observed  in  the  Lower  Chalk  of 
England  and  France  except  for  Pareodina  sp.  which  is  unidentifiable.  Firtion's 
Hystrichosphaeridium  cf .  salpingophorum  may  correspond  to  H.  mantelli,  and  Micrhy- 
stridium  ambiguum  is  probably  Cleistosphaeridium  huguonioti. 

Since  1955  a  number  of  publications  have  been  produced  dealing  mainly  with  the 
systematics  of  Cretaceous  microplankton.  Assemblages  have  been  described  from 
Germany  by  Gocht  (1957,  1959),  Eisenack  (1958),  and  Alberti  (1959,  1961),  but  only 
the  latter  author  records  Cenomanian  species.  Alberti  (1961)  described  a  series  of 
assemblages  from  the  Valanginian  to  Turonian  of  northern  Germany.  Eight  species 
were  recorded  from  the  Cenomanian  and  all  of  these,  except  for  Korojonia  dubiosa, 
have  been  found  during  the  present  study.  Hystrichodinium  pulchrum,  identified  by 
Alberti  from  the  Cenomanian,  is  here  included  within  Hystrichodinium  voigti.  Gony- 
aulax  orthoceras,  figured  by  Alberti,  is  undoubtedly  Cribroperidinium  intricatum  sp. 
nov.,  and  Palaeohystrichophora  cf.  paucisetosa  is  probably  a  form  of  P.  infusorioides 
possessing  fewer  spines  than  usual. 

Lower  Cretaceous  assemblages  were  described  by  Neale  &  Sarjeant  (1962),  Pocock 
(1962),  and  Tasch,  McClure  &  Oftedahl  (1964).  Gorka  (1963)  described  nine  species 
of  microplankton  from  the  Cenomanian  of  Poland.  All  of  these  species  have  been 
found  in  the  Cenomanian  deposits  of  England  and  France,  although  some  are  con- 
sidered to  be  incorrectly  identified.  Gonyaulax  orthoceras,  illustrated  by  Gorka 


no  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

appears  to  be  Gonyaulacysta  exilicristata  sp.  nov.  and  her  Gonyaulax  sp.  is  probably 
Cribroperidinium  intricatum ;  Hystrichosphaeridium  asterigerum  is  probably  equivalent 
to  Oligosphaeridium  complex  and  Hystrichosphaeridium  polytrichum  possibly  to 
Cleistosphaeridium  armatum.  Her  Hystrichosphaeridium  cf.  striolatum  may  also 
belong  to  the  latter  species. 

Baltes  (1963)  described  eight  species  of  microplankton  from  the  Cenomanian 
deposits  of  Roumania.  Of  these  seven  have  been  recorded  in  the  present  study. 
The  species  not  recorded,  Ascodinium  hialinium,  probably  belongs  to  the  genus 
Deflandrea.  The  identities  of  three  other  species  are  in  doubt:  some  of  the  specimens 
illustrated  as  Hystrichosphaeridium  longifurcatum  probably  belong  to  this  species 
(transferred  to  Surculosphaeridium  by  Davey  et  al.  1966),  but  one  specimen  (pi.  7, 
fig.  12)  probably  belongs  in  Hystrichosphaera.  Histrichosphaeridium  sp.  22  resembles 
Exochosphaeridium  striolatum  var.  truncatum  nov.  and  Hystrichosphaeridium  sp.  23  is 
possibly  Cleistosphaeridium  multifurcatum. 

Manum  &  Cookson  (1964)  describe  species  of  supposed  lower  Upper  Cretaceous  age 
from  Arctic  Canada  and  of  these,  eight  have  also  been  recorded  from  the  Lower 
Chalk  of  England  and  France;  however,  the  Arctic  Canadian  assemblages  are  more 
comparable  to  those  obtained  from  Saskatchewan. 

Cookson  &  Hughes  (1964)  gave  the  first  account  of  microplankton  from  the 
deposits  of  Upper  Albian  and  basal  Cenomanian  age  in  England.  Thirty- three  species 
were  described  from  the  Cenomanian  and  of  these  only  six  have  not  been  recorded 
from  the  basal  Cenomanian  of  Fetcham  Mill  (sample  FM  840)  and  Compton  Bay 
(CB  i). 

A  number  of  papers  have  been  published  (between  1954  and  1965)  dealing  with  the 
microplankton  of  Australia  and,  in  part,  of  New  Guinea  and  Papua — Deflandre  & 
Cookson  (1954,  1955),  Cookson  (1956,  1965),  Cookson  &  Eisenack  (1958,  19600,  b, 
1961,  19620,  b)  Eisenack  &  Cookson  (1960),  and  Cookson  &  Manum  (1964).  Un- 
fortunately it  is  usually  only  possible  to  give  the  approximate  age  of  the  samples 
and,  therefore,  stratigraphic  conclusions  are  not  as  meaningful  as  one  would  have 
hoped. 

A  number  of  species  from  the  Cenomanian  deposits  of  England  were  described  by 
Davey,  Downie,  Sarjeant  &  Williams  (1966)  and  the  present  paper  published  in 
two  parts  forms  a  natural  continuation  of  that  work. 

Clarke  &  Verdier  (1967)  describe  microplankton  assemblages  of  Cenomanian  to 
Senonian  age  from  the  Isle  of  Wight,  southern  England.  The  Cenomanian  samples 
examined  by  them  were  obtained  from  a  locality  approximately  sixteen  miles  to  the 
east  of  Compton  Bay.  Their  results  indicated  that  the  Upper  Cretaceous  could  be 
divided  into  5  zones  and  5  subzones,  and  also  into  7  "  intervals  "  based  on  extinction 
points.  The  results  concerning  the  Cenomanian  are  mainly  substantiated  in  the 
present  study. 

ACKNOWLEDGMENTS 

The  research  has  been  carried  out  during  the  tenure  of  a  research  studentship  in  the 
Department  of  Geology,  The  University  of  Nottingham. 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 


646'  1 

0) 

1 
s 

0 

-8 

0) 
_Q 

72  7'* 

<D 

5 

N 

,0 

h. 
0 

84l'l 

XX 

Turonian 

plenus    Marls 

XX 

i      i 

•650-51 

-  A  7O  -  71X 

Ligh 
Ch( 

/ 

Grey 
Ch« 

QIC7 

r  grey 
alk 

^ 

marly 
alk 

i      i 

i      i 

i      i 

i      i 

i-  690-91 
-  710-lf 

i      i 

i 

i      i 

i      i 

i 

}" 

-  7*5  f\     *5  1 

/oU    o  I 
-  750-5  f 
-770-71' 

i 

i 

i      i 

i 

i 

i 

r  790-91 
h810-llx 

i 

•815  - 
Grey  -green   Marl 

OO  £.' 

i 

~  836  - 
'Chloritic'Marl 

I 

i      i 

fj^LlJ     *\  \ 

Albion                                     Upper   Greensand 

FIG.  2.     Section  showing  the  Location  of  the  Samples  analysed  from 
Fetcham  Mill,  Surrey.     Scale  i  inch  to  30  feet. 


H2  CENOMANIAN   NON-CALCAREOUS   MICROPLANKTON,    i 

The  author  would  like  to  thank  sincerely  Dr.  W.  A.  S.  Sarjeant  for  considerable 
help  and  encouragement  given  at  all  stages  during  the  course  of  this  work,  and  also 
particularly  acknowledges  the  interest  and  encouragement  given  by  Professor  W.  D. 
Evans  and  Dr.  A.  J.  Rowell.  Thanks  are  due  to  Professor  G.  Deflandrefor  courtesy 
in  entertaining  the  author  and  allowing  him  to  examine  type  material  at  the  Labora- 
toire  de  Micropaleontologie,  Ecole  Practique  des  Hautes  Etudes,  Paris ;  and  to  Dr. 
G.  L.  Williams  for  his  collaboration  when  dealing  with  the  systematics  of  certain 
genera.  The  author  is  indebted  to  Mr.  R.  Hendry  and  his  staff  in  the  Department 
of  Geology,  The  University  of  Nottingham  for  their  assistance  and  provision  of 
necessary  laboratory  equipment. 

Special  thanks  are  due  to  Sir  James  Stubblefield,  former  Director  of  the  Institute 
of  Geological  Sciences,  for  permitting  study  of  chalk  samples  from  the  Fetcham 
Mill  Borehole;  to  the  Bureau  de  Recherches  Geologiques  et  Minieres,  for  providing 
chalk  samples  from  the  Escalles  Borehole ;  to  the  Department  of  Mineral  Resources, 
Saskatchewan,  for  providing  Cretaceous  samples  from  the  International  Yarbo 
Borehole  no.  17;  to  Dr.  J.  D.  Powell,  for  providing  Upper  Cretaceous  samples  from 
Texas;  to  Dr.  W.  E.  Smith,  for  providing  samples  from  the  Cenomanian  of  south 
Devon;  and  to  Dr.  K.  Diebel,  of  the  Institut  fur  Palaeontologie,  Humboldt  Uni- 
versity, East  Berlin,  for  courteously  permitting  the  loan  of  Ehrenberg's  holotypes. 

The  following  abbreviations  are  used  in  the  text:  B.M.  (N.H.) — British  Museum 
(Natural  History) :  G.S.M. — Institute  of  Geological  Sciences,  London. 


II.  STRATIGRAPHIC  LOCATION  OF  SAMPLES 

1.  Fetcham  Mill  Borehole,  Leatherhead,  Surrey  (TQ  15815650). 

This  borehole  has  been  described  by  Gray  (1965).  Ten  samples  at  20-foot  intervals 
were  processed  from  the  Cenomanian  (Lower  Chalk)  succession  (Fig.  2).  In  addition 
one  sample  from  the  Albian  (Upper  Greensand)  and  one  from  the  Turonian  (Middle 
Chalk)  were  analysed  for  comparative  purposes. 

2.  Compton  Bay,  Isle  of  Wight  (SZ  365854) 

The  Cenomanian  is  well  exposed  in  the  cliff  section  at  this  locality  and  has  been 
described  by  Jukes  Browne  (1903)  and  Osborne  White  (1921).  Samples  were 
collected  at  7-8  ft.  intervals  and  eleven  samples,  at  approximately  14  ft.  intervals, 
were  analysed  for  their  organic-shelled  microplankton  content  (Fig.  3). 

3.  Speeton,  Yorkshire  (TA/i6675o) 

The  Cenomanian  succession  is  fully  exposed  in  the  cliffs  at  Speeton.  The  base  of 
this  stage  was  taken  to  coincide  with  the  bottom  of  bed  V  (Wright  1963)  which, 
together  with  the  overlying  bed  U,  is  placed  in  the  Red  Chalk  (Fig.  4).  The  succes- 
sion has  been  described  recently  by  Kaye  (1964).  Seven  samples  were  collected 
and  analysed  for  their  microplankton  content. 


CENOMANIAN  NON-CALCAREOUS  MICROPLANKTON,  i 


1.5  9' 

Turanian                                     plenus  Marls 

>  i 

^  subglobosus  Zone  ' 

i 

•151-CB21 
-  137-       19 

-116-       17 
-  107-       15 

-92-        13 
•  76'-        11 

Ligh 
Ch 

/ 

Gre) 

C 

t   grey 
alk 

t 

/    marly 
:halk 

l 

i 

l 

l 

ES 

l 

I 

I 

I 

l 

I 

l 

I 

i 

i 

1 

l 

I 

l 

1 

1 

i 

i 

l 

l 

1 

I 

l 

i 

1 

1 

1 

i 

i 
\  varians  Zone 

-  59  -        9 
h44-         7 

-  28'-         5 
•  15'-         3 

l 

l 

1 

I 

l 

I 

l 

i 

l 

1 

l       l 

1       1 

i       i 

i      i 

i       i 

•^_     Phosphatic  nodules 

i      i 

I      i 

i      i 

Blue-grey  Chalk 
.11'  A" 

i       i 

i      i 

i      l 

Chloritic  Marl 

i      i 

l      l 

V 

'      Albion 

23' 
21' 6" 


FIG.  3.     Section  showing  the  Location  of  the  Samples  analysed  from 
Compton  Bay,  Isle  of  Wight.     Scale  i  inch  to  20  feet. 


114  CENOMANIAN   NON-CALCAREOUS   MICROPLAN  KTON,    i 


122- 


Zone 


subg 

** 


78'. 


varians  Zone 


Turanian 


T  ,   I 


II 


^1 


II 


TTI 


"115-Sp7 


100-     6 


78-     5 


52'-     4 


28-3 


•12'-     2 


0'    -     1 


plenus  Marls 


Grey  -white 
Chalk 


- 80 

Hard  grey  Chalk 
-. 78' 

Grey  Chalk 

8' 

Marly  Chalk 

•28' 

Pink  Chalk     U 


Green  -white  Chalk    V 


Aibian 


FIG.  4.     Section  showing  the  Location  of  the  Samples  analysed  from 
Speeton,  Yorkshire.     Scale  i  inch  to  20  feet. 


CENOMANIAN   NON-CALCAREOUS   MICROPLANKTON,    i  115 

4.  Hunstanton,  Norfolk.     (TF  675420) 

The  varians  and  subglobosus  zones  are  exposed  in  the  Hunstanton  cliffs  (Peake  & 
Hancock  1961).  Three  samples  were  processed,  two  from  the  lower  zone  and  one 
from  the  base  of  the  Totternhoe  Stone  (Fig.  5). 

5.  Devon,  South  Coast 

The  Cenomanian  is  represented  between  Salcombe  and  Lyme  Regis  by  isolated 
patches  composed  of  a  few  feet  of  sandy  Cenomanian  Limestone.  These  deposits 
were  divided  into  four  beds  by  Jukes-Browne  (1903) — Ar,  A2,  B  and  C.  Bed  C  is 
probably  the  Actinocamax  plenus  Marls.  The  samples  were  collected  from  four 
localities  by  Dr.  W.  E.  Smith  (Fig.  6) : 

(i)  Maynards  Cliff  (see  Smith  1961  :  114) 

(ii)  Beer  Head  (see  Smith  1957  :  123) 
(iii)  Whitediff  (see  Smith  1957  :  118) 
(iv)  Humble  Point  (see  Smith  1965  :  126) 


6.  Escalles  Borehole,  Cap  Blanc-Nez,  Pas  de  Calais 

The  borehole,  drilled  by  the  Bureau  de  Recherches  Geologiques  et  Minieres  in  1958 
has  been  described  by  Destombes  (1961).  Eleven  samples  were  obtained  for  analysis 
at  about  20  ft.  (6m.)  intervals  (Fig.  7). 


7.  Saskatchewan,  south-east 

The  borehole,  from  which  the  Saskatchewan  samples  were  obtained,  was  drilled 
for  the  Department  of  Mineral  Resources,  Saskatchewan  and  is  called  "  International 
Yarbo,  no.  17  ".  It  is  located  east  of  Regina  at  Lsd.  i,  Sec.  24,  Twp.  20,  Rg.  33, 
Wrst  Meridian.  All  depths  are  measured  from  the  Kelly  Bushing  which  is  at  an 
elevation  of  1,690  ft.  above  sea  level.  Six  samples  of  Albian/Cenomanian  age  were 
analysed  for  their  microplankton  content  (Fig.  8). 


8.  Texas,  north 

Two  samples  were  obtained  from  the  Upper  Cenomanian  of  north  Texas  (Tarrant 
County)  for  the  author  by  Dr.  J.  D.  Powell.  The  lower  sample  (T5)  was  obtained 
from  the  Upper  Woodbine  Formation  (Acanthoceras  wintoni  Zone)  and  consists  of  a 
yellowish,  slightly  calcareous  clay.  The  higher  sample  (T4)  is  from  35  ft.  above  the 
base  of  the  Eagle  Ford  Formation  (Eucalycoceras  Zone).  This  is  a  thin-bedded 
yellow  limestone  containing  shelly  fossils  and  plant  debris,  in  particular  leaf 
fragments. 


n6  CENOMANIAN  NON-CALCAREOUS  MICROPLANKTON,    i 

r20' 


JS 


C 

o 

N 


-13' 10"     -       H2 


•18' 6"      -       H3 


HI 


-18' 


Totternhoe  Stone 


Creamy -white 
Chalk 


4  1 


1  5 


Inoceramus  Bed 


Paradoxica  Bed 


Red  Rock 

FIG.  5.     Section  showing  the  Location  of  the  Samples  analysed  from 
Hunstanton,  Norfolk.     Scale  i  inch  to  2  £  feet. 


CENOMANIAN  NON-CALCAREOUS   MICROPLAN KTON,    i 


117 


n8 


CENOMANIAN   NON-CALCAREOUS  MICROPLANKTON,    i 


502 


N 

«/> 

2 
o 

i 


60f- 


0) 

c 

O 
M 


716 


Turonian 

'    -  E153 


-521' 6" 


-     159 


-541      -     165 


-561'     -     171 


-5806-     177 


-600'    -      183 


•620    -     189 


639V-     195 


659'6"-     201 


679'  -     207 


698' 6'-     213 


Grey-  white 
Chalk 


Grey  marly 
Chalk 


Chloritic  Marl 


712'  6* 


Albian 


FIG.  7.     Section  showing  the  Location  of  the  Samples  analysed  from 
Escalles,  France.     Scale  i  inch  to  30  feet. 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  119 


Q_ 

ID 

o 

0£ 

O 

o 

o 


O 
u 


1 

tt 

Shale 

/ 

Sas 

UJ 

Q_ 

Q. 

ID 

Second  White  Speckled  Shale 

007' 

-835V 

-QO/ 

-890' 

Shale 

c960 

ex. 

\*  f  \j  \J 

-967' 

UJ 

Fish  Scale  Zone 

o 

-c1030' 

-1023 

Shale 

~l-IVJ«jw 

-1084' 

Viking 

FIG.  8.     Section  showing  the  Location  of  the  Samples  analysed  from 
Saskatchewan,  Canada.     Scale  i  inch  to  50  feet. 


GEOL.  17,  3 


120  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

III.    SYSTEMATIC  DESCRIPTIONS 

Class  DINOPHYCEAE  Pasher 

Subclass  DINIFEROPHYCIDAE  Bergh 

Cyst-FamUy  GONYAULACYSTACEAE  Sarjeant  &  Downie  1966 

Genus  GONYAULACYSTA  Deflandre  emend.  Sarjeant  1966 
Gonyaulacysta  cassidata  (Eisenack  &  Cookson)  emend.  Sarjeant 

1960     Gonyaulax  helicoidea  subsp.  cassidata  Eisenack  &  Cookson  :  3,  pi.  i,  figs.  5,  6. 

ig66a  Gonyaulacysta  cassidata  (Eisenack  &  Cookson)  Sarjeant  :  125,  pi.  14,  figs.  3,  4,  text-fig.  31 

(see  also  for  earlier  references). 
1967     Gonyaulacysta  cassidata  (Eisenack  &  Cookson)  Clark  &  Verdier:  29,  pi.  4,  figs.  4-6. 

DIMENSIONS.  Range  of  observed  specimens:  overall  length  59  (67-5)  78/4,  overall 
width  40  (46-4)  60  fi.  Number  of  specimens  measured,  14. 

REMARKS.  The  Cenomanian  specimens  examined  are  very  similar  to  the  Austra- 
lian Aptian-Cenomanian  forms  of  Eisenack  &  Cookson  (1960)  and  Cookson  &  Eisenack 
(19626)  except  that  the  former  are  slightly  smaller  in  size. 

OCCURRENCE.  G.  cassidata  is  an  infrequent  species  at  all  horizons  throughout 
the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  has  not  been 
recorded  in  the  North  American  samples. 

Gonyaulacysta  whitei  Sarjeant 
ig66a     Gonyaulacysta  whitei  Sarjeant  :  126,  pi.  14,  fig.  2,  text-fig.  32. 

DIMENSIONS.  Range  of  observed  specimens:  overall  length  55-62/4,  overall 
width  42-50/4.  Number  of  specimens  measured,  5. 

REMARKS.  The  shape  of  the  shell,  the  tabulation  and  the  crests  make  G.  whitei 
an  easily  recognizable  and  distinctive  species.  G.  cf.  ambigua  Cookson  &  Eisenack 
(19606)  from  the  Upper  Jurassic  is  of  the  same  shape  and  possesses  similar  crests  and 
tabulation;  plate  i""  is  absent.  However,  the  apical  horn  of  G.  cf.  ambigua  is 
rudimentary  or  absent,  thus  making  differentiation  easy. 

OCCURRENCE.  Of  five  specimens  of  G.  whitei  observed,  four  are  from  sample  FM 
770  and  one  from  sample  FM  750.  The  restricted  distribution  of  this  species  and  its 
similarity  to  G.  cf .  ambigua  from  the  Upper  Jurassic  suggest  the  possibility  that  this 
is  a  derived  form. 

Gonyaulacysta  fetchamensis  Sarjeant 

19660     Gonyaulacysta  fetchamensis  Sarjeant  :  128,  pi.  15,  figs,  i,  2,  text-fig.  33. 

REMARKS.  G.  fetchamensis  has  a  rather  unusual  tabulation — two  posterior  inter- 
calary plates  and  a  seventh  postcingular  plate — and,  as  pointed  out  by  Sarjeant 
(19660),  may  subsequently  form  the  basis  of  a  new  genus.  As  yet  only  two  specimens 
have  been  studied  and  this  species  has,  therefore,  been  placed  in  Gonyaulacysta.  It 


CENOMANIAN   NON-CALCAREOUS   MICROPLANKTON,    i  121 

appears  to  be  a  transitional  type  from  the  normal  Gonyaulacysta  to  forms  now  placed 
in  Cribroperidinium  Neale  &  Sarjeant.  The  latter  forms  are  of  similar  overall  appear- 
ance but  the  shell  is  divided  by  low  crests  into  an  unusually  large  number  of  areas. 

OCCURRENCE.  G.  fetchamensis  is  a  rare  species  recorded  only  from  the  Chalk  of 
H.M.  Geological  Survey  borehole,  Fetcham  Mill,  Surrey,  at  840  ft.  depth.  Upper 
Cretaceous  (Lower  Cenomanian). 

Gonyaulacysta  exilicristata  sp.  nov. 
(PI.  i,  figs,  i,  2;  Figs.  gA,  B) 

DERIVATION  OF  NAME.  Latin,  exilis,  thin  or  poor;  cristatus,  crested — with  re- 
ference to  the  poorly  defined  sutural  crests. 

DIAGNOSIS.  Shell  subspherical ;  moderately  well  developed  apical  horn.  Shell 
wall  thick,  finely  but  densely  granular.  Reflected  tabulation  3',  la,  6",  6c,  6'" 
(—7"'?),  i  p,  i"".  Plate  boundaries  marked  by  low,  poorly  defined  crests  which 
sometimes  form  small  spines  at  crestal  nodes.  Cingulum  narrow,  weakly  laevo- 
rotatory;  sulcus  of  moderate  width,  widening  slightly  posteriorly. 

HOLOTYPE.  G.S.M.  slide  PF  3987  (i).  Lower  Chalk,  H.M.  Geological  Survey 
Borehole,  Fetcham  Mill,  Surrey  at  730  ft.  depth.  Upper  Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype:  overall  length  83/4,  overall  width  68  ju,,  length  of  horn 
I2/*.  Range:  overall  length  70  (81-6)  98^,  overall  width  58  (64-5)  71/4.  Number 
of  specimens  measured,  24. 

DESCRIPTION.  The  shell  wall  measures  between  2  and  3/4  in  thickness,  the 
endophragm  being  approximately  twice  as  thick  as  the  periphragm.  The  latter 
forms  the  apical  horn  which  is  triangular  in  cross-section  due  to  the  sutural  ridges 
which  delimit  the  apical  plates  extending  along  it.  Lines  of  ornamentation,  some- 
times similar  to  the  sutural  crests,  are  present  on  some  of  the  plates,  particularly 
those  in  the  postcingular  series.  Occasionally  it  appears  that  plate  4'"  is  subdivided 
by  a  low  ridge  so  giving  seven  postcingular  plates. 

The  cingulum  is  fairly  narrow  (3  to  47*)  and  only  slightly  laevo-rotatory.  The 
sulcus  often  possesses  a  posterior  ventral  plate.  In  the  medial-posterior  part  of  the 
furrow  there  is  usually  an  elongate  depression,  as  seen  in  the  holotype  (Fig.  gA).  A 
precingular  archaeopyle  is  typically  present. 

REMARKS.  G.  exilicristata  sp.  nov.  is  distinguished  from  all  previously  described 
species  by  its  overall  shape,  the  type  of  plate  boundaries  and  the  tabulation. 
Apteodinium  granulatum  Eisenack  (1958)  is  similar  but  has  a  stouter  apical  horn 
and  a  tabulation  appears  to  be  absent;  the  cingulum  is  only  rarely  visible. 

Cribroperidinium  orthoceras  (Eisenack)  comb.  nov.  is  also  similar  but  possesses  a 
longer  apical  horn,  and  the  tabulation  differs  and  is  more  clearly  defined. 

OCCURRENCE  G.  exilicristata  is  a  rare  species  recorded  from  a  number  of  horizons 
throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  At  only 
one  horizon  is  this  species  common,  in  sample  FM  730  from  Fetcham  Mill.  Two 
specimens  have  been  recorded  from  Saskatchewan,  both  from  sample  Sas  1084. 


122  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 


FIG.  9.     Gonyaulacysta  exilicristata  sp.   nov.,  A.  Ventral  Surface  of  Holotype   (X  900), 
B.  Dorsal  Surface  of  Holotype,  (X  900).     Gonyaulacysta  Sp.  A.,  C.  Ventral  Surface  slide 
PF.  3987  (2)  (X  900).     D.  Dorsal  Surface  slide  PF.  3987  (2)  (X  900) 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 


123 


Gonyaulacysta  delicata  sp.  nov. 
(PI.  i,  figs.  7,  8;  Figs.  loA,  B) 

DERIVATION  OF  NAME.  Latin,  delicatus,  delicate — with  reference  to  the  delicate 
nature  of  the  shell. 

DIAGNOSIS.  Shell  subspherical,  epitract  and  hypotract  of  similar  size.  Shell 
wall  thin,  smooth.  Apical  horn  absent,  there  being  a  small  circular  apical  plate  in 
this  position  surrounded  by  three  large  apical  plates.  Reflected  tabulation  4',  la, 
6",  6c,  6"',  ip,  i"".  Plate  boundaries  well  defined  by  low  crests.  Cingulum  wide, 
strongly  laevo-rotatory ;  sulcus  broad. 

HOLOTYPE.  B.M.  (N.H.)  V.  51979(1).  Lower  Colorado,  Second  White  Speckled 
Shale,  International  Yarbo  Borehole  No.  17,  Saskatchewan  at  835  ft.  depth. 
Upper  Cretaceous  (Cenomanian). 

PARATYPE.    B.M.  (N.H.)  V.  51979(2). 

DIMENSIONS.  Holotype:  length  of  shell  57/4,  width  51/4.  Paratype:  length  of 
shell  55/4,  width  47/4.  Range:  length  of  shell  52-60/4,  width  47-51/4.  Number  of 
specimens  measured,  4. 

DESCRIPTION.  The  shell  wall  is  very  thin  (less  than  0-5  ju,  thick)  and  only  attains 
a  thickness  of  0-5  /*  when  forming  the  plate  boundaries.  Due  to  the  thinness 
of  the  shell  wall  specimens  are  easily  distorted.  The  tabulation  has,  however,  been 
formulated  after  the  examination  of  a  number  of  specimens. 


A  B 

FIG.  10.     Gonyaulacysta  delicata  sp.  nov.,  A.  Ventral  surface  of  Holotype  (x  15°°)- 
B.  Dorsal  surface  of  Holotype  (X  1500). 


124  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 

Anteriorly  three  apical  plates  (2',  3'  and  4')  abutt  against  a  small  circular  plate, 
plate  i',  which  is  in  the  position  normally  occupied  by  the  apical  horn.  The  pre- 
cingular  and  postcingular  series  of  plates  are  large  and  more  or  less  pentagonal. 
Plate  3"  is  always  lost  in  archaeopyle  formation.  The  first  postcingular  plate  is 
elongate  and  has  a  poorly  defined  sulcal  border. 

The  cingulum  is  wide  (4  to  7  fi)  and  tends  to  be  constricted  at  the  cingular  plate 
boundaries.  The  sulcus  is  broad  and  widens  slightly  towards  the  posterior.  The 
anterior  end  of  the  sulcus  has  a  flat  margin  from  which  arise  plates  i'  and  la.  At 
the  posterior  end  of  the  sulcus  there  may  be  developed  a  posterior  ventral  plate. 

REMARKS.  The  presence  of  very  low  sutural  crests,  the  lack  of  an  apical  horn  and 
the  tabulation  differentiate  G.  delicata  sp.  nov.  from  all  previously  described  species. 
G.  ambigua  Deflandre,  from  the  Kimeridgian  of  France,  is  of  similar  appearance 
but  possesses  a  small  apical  horn  and  differs  in  tabulation  detail. 

OCCURRENCE.  G.  delicata  has  been  found  in  only  one  sample,  Sas  835,  and  it  is 
there  infrequent.  This  restricted  distribution  may  well  indicate  that  this  is  a 
derived  species. 

Gonyaulacysta  sp.  A. 

(PL  i,  figs.  9,  10;  Figs.  gC,  D) 

DESCRIPTION.  Only  one  well  preserved  specimen  of  this  species  has  so  far  been 
observed.  It  possesses  a  subspherical  shell,  bearing  a  moderately  well  developed 
horn  with  a  trifid  termination.  The  shell  wall  (c.  2/1  thick)  is  irregularly  studded 
with  granules  of  varying  shapes  and  sizes.  The  sutural  crests  are  quite  well  defined, 
but  low,  and  indicate  a  reflected  tabulation  of  3',  la,  6",  6c,  6'"  (—7'"?),  ip.,  i"". 
The  crests,  demarcating  the  three  apical  plates,  extend  along  the  apical  horn  and 
give  rise  to  three  small  spines  at  its  distal  termination.  Plates  2",  3"  and  4"  are 
relatively  large,  plates  i"  and  5"  rather  elongate  and  plate  6"  is  reduced  due  to  the 
anterior  intercalary  plate.  In  the  postcingular  series,  plates  i"'  and  2'"  are 
reduced  and  plate  7'"  does  not  have  a  clearly  marked  plate  boundary.  Plate  2'" 
possesses  a  curved  line  of  ornamentation  which  is  characteristic  of  Cnbroperidinium 
Neale  &  Sarjeant.  There  is  a  single  posterior  intercalary  plate  and  a  large  antapical 
plate. 

The  cingulum  is  strongly  laevo-rotatory  and  varies  considerably  in  width  (2-5  (JL), 
being  constricted  at  the  cingular  plate  boundaries  and  expanding  on  either  side. 
The  sulcus  is  broad  and  possesses  a  central  depressed  area  of  elongate  shape.  A 
archaeopyle  is  present. 

FIGURED  SPECIMEN.  G.S.M.  slide  PF.  3987,  specimen  2.  Lower  Chalk,  H.M. 
Geological  Survey  Borehole,  Fetcham  Mill,  Surrey  at  730  feet  depth,  Upper  Cretaceous 
(Cenomanian) . 

DIMENSIONS.     Overall  length  75/11,  overall  width  65^,  length  of  horn  14  p.. 

REMARKS.  The  form  of  the  apical  horn,  the  type  of  granulation  and  the  shape 
of  the  cingular  plates  distinguish  this  species  from  all  described  forms.  The  shape 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  125 

of  the  shell  and  the  tabulation  are  most  comparable  to  G.  exilicristata  suggesting  a 
relationship  between  the  two  species. 

Genus  CRIBROPERIDINIUM  Neale  &  Sarjeant  emend. 

EMENDED  DIAGNOSIS.  Proximate  cysts,  subspherical  to  ovoidal,  thick-walled. 
Crests  numerous  and  well  developed.  Tabulation  ?6',  (i-5a),  8-9",  oc,  g"',  ip, 
i-3p.v.  (5-7  p.c.),  o""  (— ?2"").  Cingulum  laevo-rotatory.  Archaeopyle 
precingular. 

TYPE  SPECIES.     Cribroperidinium  sepimentum  Neale  &  Sarjeant  1962. 

REMARKS.  The  diagnosis  has  been  emended  to  draw  attention  to  the  fact  that 
the  anterior  intercalary  series  and  the  posterior  circle  series  of  plates  are  not  always 
readily  distinguishable.  The  tabulation  attributed  to  this  genus  by  Neale  & 
Sarjeant  was  6',  i~5a,  8",  9'",  ip,  i-2p.v.  6-Pyp.c.,  o"".  In  determining  the  tabula- 
tion they  numbered  all  the  delimited  areas  on  the  shell  surface  in  the  normal  manner. 
This  numbering  procedure  has  been  used  in  the  study  of  C.  intricatum  sp.  nov.  with 
certain  reservations.  Difficulties  arose  due  to  the  large  number  of  delimited  areas 
and  also  to  some  variation  on  the  dorsal  surface  of  the  hypotract.  Some  of  the 
crests  are  rudimentary  and  others,  although  appearing  fairly  normal,  are  unusual 
in  their  position.  It  was  discovered  that  by  the  removal  of  these  unusual  crests  a 
normal  Gonyaulax-type  tabulation  could  be  reconstructed.  This  is  clearly  shown  in 
Figs.  nA,  B,  of  the  ventral  surface  of  the  holotype  of  C.  intricatum.  Thus  Cribro- 
peridinium is  basically  a  form  of  Gonyaulacysta  which  possesses  additional  crests. 
These  additional  crests  may  correspond  to  an  increase  in  the  number  of  thecal 
plates  composing  the  motile  dinoflagellate,  but  this  is  thought  to  be  unlikely  since: 

(a)  the  plates  formed  would  be  of  an  extremely  unusual  shape ; 

(b)  these  crests  subdivide  detached  opercula  demonstrably  composed  of  a  single 
precingular  plate  (3") ; 

(c)  many  of  these  crests  are  poorly  developed  and  show  considerable  positional 
variation  on  the  dorsal  surface  of  the  hypotract;  and 

(d)  that  by  their  removal  a  normal  Gonyaulax-type  tabulation  remains.     Thus, 
these  additional  crests  may  be  regarded  as  an  ornamentation  or  perhaps  a  rather, 
superficial  cyst  strengthening  device.     It  is,  therefore,  considered  more  practical 
and  correct  to  use  a  different  numbering  system  for  the  Cribroperidinium  cyst 
tabulation,  vis,  roman  numerals. 

Three  species,  Gonyaulacysta  orthoceras  (Eisenack),  G.  muderongensis  (Cookson  & 
Eisenack)  and  G.  edwardsi  (Cookson  &  Eisenack),  are  here  transferred  to  Cribroperi- 
dinium. This  genus  appears  to  be  very  limited  in  stratigraphic  range  (Hauterivian— 
Lower  Turanian)  and,  as  such,  is  a  useful  stratigraphic  indicator. 

Cribroperidinium  intricatum  sp.  nov. 
(PL  2,  figs.  1-3;  Figs,  n,  12) 

DERIVATION  OF  NAME.  Latin,  intricatus,  complicated — with  reference  to  the 
complex  crest  arrangement. 


126 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 


DIAGNOSIS.  Shell  subspherical,  epitract  and  hypotract  of  similar  size.  Apical 
horn  of  moderate  length,  subconical.  Shell  wall  moderately  thick,  granular  and 
bears  a  few,  randomly  arranged  tubercles.  Crests  usually  in  form  of  low  ridges,  well 
denned,  sometimes  membranous,  delimiting  a  large  number  of  plates  on  shell 
surface.  Sutural  spines  absent.  Plate  IF"  crossed  diagonally  by  low  crest. 
Operculum  possessing  semi-circular  crest.  Sulcus  possessing  posterior  ventral  plates. 
Cingulum  narrow,  plates  not  defined. 

HOLOTYPE.  B.M.  (N.H.).  V.  51980  (i).  Upper  Lower  Colorado,  Fish  Scale  Zone, 
International  Yarbo  Borehole  No.  17,  Saskatchewan  at  1,023  feet  depth.  Lower 
Cretaceous  (Albian). 

DIMENSIONS.  Holotype:  overall  length  120  /*,  overall  width  114/1,,  length  of  horn 
20 p.  Range:  overall  length  107  (125-1)  142/11,  overall  width  101  (108-2)  126/1.. 
Number  of  specimens  measured,  17. 

DESCRIPTION.     The  shell  wall  is  i  to  1-5  p  in  thickness  and  densely  granular.     The 
crests  are  typically  low  thickenings  of  the  periphragm  (2-3  p  wide),  but  in  the  ant- 
apical  region,  and  occasionally  elsewhere,  the  crests  take  the  form  of  high  flanges 
The  latter  (up  to  6  /z.  in  height)  are  membranous,  thin  and  always  perforate. 

The  number  of  apical  plates  always  appears  to  be  six.  Plate  I',  equivalent  to 
the  first  apical  plate,  is  elongate  and  abuts  against  the  anterior  end  of  the  sulcus. 
The  crest  arrangement  on  the  ventral  surface  appears  to  be  practically  constant  and 
is  characteristic  of  this  species.  The  crests  limiting  the  plates  F,  I",  II",  III",  IV", 
VII"  and  VIII"  are  always  constant  in  position.  Plates  F"  and  IF"  are  reduced 
due  to  the  presence  of  a  posterior  intercalary  plate.  Plate  IF"  always  possesses  a 
crest  passing  diagonally  across  it  and  the  crest  between  plates  IIF"  and  IV"  is  of  a 


A  B 

FIG.  ii.     Cribroperidinium  intricatum  sp.  nov.     A.  Ventral  surface  of  Holotype, 
(X  700).     B.  Reconstruction  to  show  Gonyaulax-type  tabulation  (x  700). 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i  127 

characteristic  right  angle  shape.  The  positions  of  the  crests  on  the  dorsal  surface 
of  the  epitract  are  fairly  constant,  only  varying  in  minor  details.  The  large  plate 
V"  is  usually  detached  in  archaeopyle  formation  and  possesses  a  semi-circular  crest 
from  which  radiate  a  small  number  of  other  crests  (PI.  2,  fig.  3).  These  pass  over 
the  boundary  of  plate  V"  to  continue  on  adjacent  parts  of  the  epitract.  The  posi- 
tions of  the  crests  on  the  dorsal  surface  of  the  hypotract  appear  to  be  less  constant 
in  position.  Their  predominent  direction  is  parallel  to  the  longitudinal  axis  of  the 
shell;  sometimes  a  series  of  posterior  circle  plates  may  be  present.  Antapical 
plates,  if  present  at  all,  are  very  reduced  and  obscured  by  the  crestal  membranes. 

The  cingulum  is  narrow  (c.  6/z  in  width),  slightly  laevo-rotatory,  and  tends  to 
possess  a  thicker  wall  than  is  usual  for  the  remainder  of  the  shell.  The  sulcus  only 
projects  onto  the  epitract  for  a  short  distance,  being  considerably  larger  and  wider 
on  the  hypotract.  It  is  always  divided  into  a  number  of  posterior  ventral  plates  by 
reduced  crests. 

REMARKS.  Figs  nB,  I2B  have  been  prepared  from  the  holotype  and  one 
other  specimen,  omitting  the  additional  crests,  to  show  the  basic  Gonyaulax-type 
tabulation. 

C.  intricatum  may  be  differentiated  from  C.  orthoceras  (Eisenack),  C.  muderongensis 
(Cookson  &  Eisenack),  Gonyaulacysta  apionis  and  G.  diaphanis  by  its  more  spherical 
form,  details  of  crest  arrangement  and  the  absence  of  spines.  C.  edwardsi  (Cookson 
&  Eisenack)  is  most  similar,  being  almost  spherical,  but  possesses  a  very  well 
developed,  stiff  apical  horn  and  the  crests  on  the  ventral  surface  are  arranged  differ- 
ently. In  particular  the  diagonal  crest  on  plate  II'"  is  absent.  Gonyaulax  sp.  (Gorka 
1963)  from  the  Cenomanian  of  Poland  is  very  similar  and  may  be  conspecific 
with  C.  intricatum. 


A  B 

FIG.    12.     Cribroperidinium   intricatum   sp.    nov.,    A.    Dorsal   surface   of   Holotype   with 
archaeopyle  (X  700).     B.  Reconstruction  to  show  Gonyaulax-type  tabulation  (X  700). 


128  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

OCCURRENCE.  C.  intricatum  is  very  rare  in  the  British  Cenomanian  and  has  been 
observed  only  in  samples  FM  840,  CB  I  and  CB  7.  At  Escalles  it  is  infrequent  in 
sample  E  201,  very  rare  in  sample  E  195  and  lacking  elsewhere.  In  the  Saskatche- 
wan sample  Sas  1023  it  is  common,  in  sample  Sas  1084  rare  and  is  lacking  in  all  the 
other  North  American  samples.  It  is  interesting  to  record  that  this  species  was 
observed  in  the  Albian  sample  FM  886  from  Fetcham  Mill.  Thus  C.  intricatum  has 
a  range  from  Albian  to  Lower  Cenomanian  in  the  examined  material. 

OTHER  SPECIES 

The  following  species  are  here  attributed  to  the  genus  Cribroperidinium  on  the 
basis  of  overal  shape,  and  number  and  position  of  periphragm  crests: — 

Cribroperidinium  orthoceras  (Eisenack  1958)  comb,  nov.,  1958  Gonyaulax  orthoceras  Eisenack, 
Neues.  Jb.  Geol.  Palaont.,  Abh.,  106  (3)  388:  pis.  21,  figs.  3-14;  24,  fig.  i;  text-figs.  2,  3. 

Cribroperidinium  edwardsi  (Cookson  &  Eisenack  1958)  comb,  nov.,  1958  Gonyaulax  edwardsi 
Cookson  &  Eisenack:  Proc.  R.  Soc.  Viet.,  70  (i),  32-33;  PL  III,  figs.  5,  6,  text-fig.  7. 

Cribroperidinium   muderongensis    (Cookson    &    Eisenack    1958)    comb,    nov.,    1958    Gonyaulax 

muderongensis  Cookson  &  Eisenack:  Proc.  R.  Soc.  Viet.,  70  (i),  32;  PI.  Ill,  figs.  3,  4,  text-fig.  15. 

Gonyaulacysta  apionis  (Cookson  &  Eisenack  1958)  and  G.  diaphanis  (Cookson  & 
Eisenack  1958),  both  from  the  Lower  Cretaceous  of  Australia,  are  of  similar  appear- 
ance to  forms  included  in  Cribroperidinium  and  may  at  a  later  date  be  transferred 
to  this  genus. 

Eisenack  (1958  text-figs.  2,  3)  figured  the  ventral  surface  tabulation  of  C.  ortho- 
ceras omitting,  or  dotting  in,  some  of  the  crests  which  are  seen  to  be  present  on  the 
photographs  of  the  same  specimens.  Thus  a  true  representation  of  the  crestal 
positions  was  not  given,  somewhat  misleading  later  workers.  These  figures  have 
been  redrawn  (Figs.  I3A,  B)  from  the  photographs  and  show  the  remarkable  similarity 
between  the  ventral  surface  of  C.  orthoceras  and  that  of  C.  intricatum.  For  this 
reason  the  former  species  has  been  transferred  to  Cribroperidinium  and  the  diagnosis 
emended.  Eisenack  does  in  fact  compare  and  contrast  his  species  with  the 
reattributed  Australian  forms  above,  also  with  Gonyaulactysa  wetzeli  (Lejeune 
Carpentier  1939)  and  G.  obscura  (Lejeune-Carpentier  1946),  all  of  which  he  con- 
siders to  be  in  the  same  group.  The  latter  two  species,  however,  have  a  typical 
Gonyaulacysta  tabulation  and  must  remain  in  that  genus. 

Cribroperidinium  orthoceras  (Eisenack)  emend. 
(Figs.  I3A,  B) 

1958  Gonyaulax  orthoceras  Eisenack  :  388,  pi.  21,  figs.  3-14,  pi.  24,  fig.  i ;  text-figs.  2,  3. 

!959  Gonyaulax  orthoceras  Eisenack;  Gocht  :  54,  pi.  5,  figs.  12,  13. 

1961  Gonyaulax  orthoceras  Eisenack;  Alberti  :  6,  pi.  n,  figs.  1-3. 

1963  Gonyaulax  orthoceras  Eisenack;  G6rka  :  30,  pi.  3,  figs.  1-4. 

1965  Gonyaulax  orthoceras  Eisenack;  Baltes  :  12,  pi.  3,  figs.  95-99. 

EMENDED  DIAGNOSIS.  Shell  ovoidal,  moderately  thick-walled,  bearing  strong, 
thorn-like  apical  horn  constituting  approximately  one  quarter  of  shell  length.  Shell 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  129 

wall  granular  bearing  tubercles.  Distinct  tabulation  marked  by  strong,  low  crests 
delimiting  large  number  of  plates.  Plate  IF"  crossed  diagonally  by  low  crest 
and  operculum  possessing  semi-circular  crest.  Cingulum  narrow,  devoid  of  plate 
boundaries. 

HOLOTYPE.  The  specimen  illustrated  by  Eisenack  (1958,  pi.  21,  fig.  5)  from  Pre- 
paration Ob.  Apt.  No.  32.  Aptian  glauconitic  limestone,  Deutschen  Erdol  A.  G., 
Erdolwerke  Holstein  boring  Marne,  Feld  Heide,  North  Germany,  at  7617  metres 
depth. 

REMARKS.  The  emended  diagnosis  excludes  those  forms  described  by  Sarjeant 
(1966)  from  the  Speeton  Clay  which  are  to  be  described  elsewhere  under  a  new  specific 
name. 

Genus  CARPODINIUM  Cookson  &  Eisenack,  1962 

Carpodinium  obliquicostatum  Cookson  &  Hughes 

(PL  i,  figs.  3,  4) 

1964     Carpodinium  obliquicostatum  Cookson  &  Hughes  :  48,  pi.  6,  figs.  1-6. 

1967     Carpodinium  obliquicostatum  Cookson  &  Hughes;  Clarke  &  Verdier:  23,  pi.  2,  figs.  4,  5. 

DESCRIPTION.  The  shell  is  elongate-ovoidal  and  bears  relatively  high  sutural 
crests.  The  latter  are  smooth  or  slightly  granular  and  the  distal  margin  may  be 
entire  or  spinous.  The  short  apical  horn  is  a  prolongation  of  one  of  the  larger  crests 
and  is  typically  asymmetrically  placed.  The  periphragm  of  the  shell  possesses  an 
unusual  ornamentation — small  areas  of  triangular  to  polygonal  shape,  composed 
of  thick  periphragm,  are  separated  by  narrow  anastomosing  "  canals  "  where  the 
periphragm  is  unthickened  or  perhaps  absent  (PL  r,  fig.  4). 

The  cingulum  is  marked  by  indentations  in  some  of  the  crests  and  is  not  apparent 
on  the  shell  surface.  The  sulcus,  delimited  by  crests,  is  occasionally  observable 
and  extends  the  length  of  the  shell,  being  narrow  near  the  apex  and  widening  pos- 
teriorly. The  precingular  and  postcingular  plates,  probably  six  in  each  series,  are 
very  elongate  and  difficult  to  discern  because  of  the  obscuring  nature  of  the  high 
crests.  A  precingular  archaeopyle  is  commonly  present.  Four  apical  plates  and  a 
single  antapical  plate  appear  to  be  present.  Intercalary  plates  were  not  observed. 

DIMENSIONS.  Range  of  observed  specimens:  overall  length  56(69-5)  82 //,,  overall 
width  33(43-8)  57  /*,  height  of  crests  6-15  ^u,.  Number  of  specimens  measured,  16. 

REMARKS.  The  Cenomanian  specimens  studied  resemble  the  type  material  from 
the  Upper  Albian  and  Lower  Cenomanian  of  Cambridgeshire  in  all  respects. 

OCCURRENCE.  C.  obliquicostatum  is  a  rare  to  very  rare  species  found  at  all 
horizons  throughout  the  Cenomanian  of  Fetcham  Mill  and  Compton  Bay  and  in 
three  samples  from  Escalles  (E  195,  E  189  and  E  159). 

Genus  ELLIPSODINIUM  Clarke  &  Verdier  1967 

REMARKS.  A  number  of  microplankton  genera  have  been  described  as  possessing 
a  reticulate  shell  wall,  occasionally  with  an  outer  membrane  but  only  rarely  with  any 


130  CENOMANIAN   NON-CALCAREOUS    MI  CROPL  AN  KTON,    i 

signs  of  tabulation.  In  all  the  described  forms  possessing  a  cingulum  the  archae- 
opyle  is  apical.  Hence  the  combination  of  numerous  crests,  a  cingulum  and  a 
precingular  archaeopyle  differentiates  Ellipsodinium  from  all  previously  described 
genera. 

Ellipsodinium  rugulosum  Clarke  &  Verdier 

(PI.  3.  fig.  i  ;  Figs.  140,  D) 
1967     Ellipsodinium  rugulosum  Clarke  &  Verdier:  69,  pi.  14,  figs.  4-6,  text-fig.  29. 

DIMENSIONS.  Range  of  observed  specimens  :  shell  length  30  (37  .  7)  46  ju.,  shell  width 
25  (33-6)  40  p,  maximum  height  of  crests  1-5  (2-4)  3-5  ju,.  Number  of  specimens 
measured,  20. 

DESCRIPTION.  The  crests  are  thin,  occasionally  perforate  lamellar  structures  which 
thicken  slightly  before  joining  the  shell  surface.  The  cingulum  may  be  delimited  by 
a  pair  of  crests,  or  in  the  absence  of  cingular  crests,  crests  may  terminate  abruptly 
at  its  borders.  Rarely  crests  traverse  the  cingulum.  The  sulcus  is  not  obvious 
because  of  the  nature  of  the  elongate  areas  outlined  by  the  crests.  Apical  and 
antapical  processes  or  horns  are  absent.  The  precingular  archaeopyle  is  sub- 
triangular  in  outline. 

OCCURRENCE.  E.  rugulosum  is  a  rare  to  fairly  common  species  at  all  horizons 
throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  was 
not  observed  in  the  North  American  material. 

Cyst-Family  PAREODINIACEAE  Gocht  emend.  Sarjeant  &  Downie  1966 
Genus  APTEODINIUM  Eisenack  1958 

REMARKS.  Members  of  this  genus  have  occasionally  been  observed  in  the  Euro- 
pean Cenomanian  and  have  been  placed  in  A  .  granulatum.  Rarely  plate  boundaries 
may  be  discernable.  This  fact  together  with  the  overall  shape  and  the  well  developed 
precingular  archaeopyle  indicate  that  this  genus  is  closely  related  to  Gonyaulacysta 
and  at  a  future  date  may  be  transferred  to  the  Cyst-Family  Gonyaulacystaceae. 

Apteodinium  granulatum  Eisenack 
(PI.  3,  figs.  5,  6) 

?I935  Palaeoperidinium  ventriosum  O.  Wetzel;  Deflandre  :  228,  pi.  5,  fig.  5;  pi.  6,  figs.  9,  10. 

?i936&  Palaeoperidinium  ventriosum  O.  Wetzel;  Deflandre  :  27,  pi.  5,  figs.  1-4. 

?i936a  Palaeoperidinium  ventriosum  O.  Wetzel;  Deflandre:  fig.  100. 

1958  Apteodinium  granulatum  Eisenack  :  386,  pi.  23,  figs.  8-14,  text-fig,  i. 

1958  Apteodinium  granulatum  Eisenack;  Gocht  :  64,  pi.  5,  fig.  2. 

1961  Apteodinium  granulatum  Eisenack;  Alberti  :  24,  pi.  4,  figs.  4-6. 

1963  Apteodinium  granulatum  Eisenack;  Baltes.  :  584,  pi.  4,  fig.  u. 


DESCRIPTION.     Shell  subspherical  with  a  moderately  thick  wall  (1-2-5  //,) 
possessing  a  stout  conical  horn.     The  wall  is  densely  granular,  the  granules  often 
being  elongated  into  fine,  short  processes  which  are  linked  laterally  thus  covering 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  131 

the  shell  surface  with  an  intricate  network  of  a  matted  furry  appearance.  The 
cingular  boundaries  are  always  visible  as  slight  thickenings  of  the  periphragm. 
Very  rarely  similar  but  more  reduced  thickenings  indicate  the  presence  of  plate 
boundaries.  A  precingular  archaeopyle  is  commonly  developed. 

DIMENSIONS.  Range  of  observed  specimens:  overall  length  42  (48-2)  53  /z,  width 
31  (38-6)  44  p.,  length  of  apical  horn  5  (6-1)  8  /*.  Number  of  specimens  measured,  9. 

REMARKS.  The  Cenomanian  specimens  are  very  similar  in  appearance  to  the 
type  material  described  by  Eisenack  from  the  Aptian  of  Germany.  The  matted, 
furry  appearance  was  not  described  but  when  this  feature  is  only  slightly  developed 
the  shell  wall  merely  appears  to  be  very  granular.  The  Aptian  and  Cenomanian 
examples  of  A .  granulatum  appear  to  be  comparable  to  Palaeoperidinium  ventriosum 
O.  Wetzel  as  illustrated  by  Deflandre  (1935,  1936^,  b).  The  holotype  of  this  species 
has  been  re-examined  by  Lejeune-Carpentier  (1946)  and  a  distinct  tabulation  des- 
cribed. Deflandres'  forms  which  do  not  possess  a  tabulation  are  hence  here  ten- 
tatively reattributed  to  A .  granulatum. 

OCCURRENCE.  Only  two  specimens  have  been  recorded  from  the  English  Ceno- 
manian, both  from  sample  FM  690.  At  Escalles  two  specimens  have  been  recorded 
from  both  samples  E  183  and  E  153,  and  three  from  sample  E  165.  A.  granulatum 
is  fairly  common  in  sample  FM  886  (Albian)  from  Fetcham  Mill  but  is  absent  in  the 
North  American  material  and  from  sample  FM  520  (Turanian). 

Genus  TRICHODINIUM  Eisenack  &  Cookson  emend.  Clarke  &  Verdier 

REMARKS.  This  genus  differs  from  Exochosphaeridium  Davey,  Downie,  Sarjeant 
&  Williams  (1966)  by  the  presence  of  a  well  developed  cingulum  and  by  the  shorter 
spines. 

Trichodinium  castaneum  (Deflandre) 
PI.  n,  figs.  1-3 

J935     Palaeoperidinium  castanea  Deflandre  :  49,  pi.  6,  fig.  8. 

19366  Palaeoperidinium  castanea  Deflandre;  Deflandre  :  25,  pi.  16,  figs.  1-4. 

19360  Palaeoperidinium  castanea  Deflandre;  Deflandre:  fig.  99. 

19526  Palaeoperidinium  castanea  Deflandre;  Deflandre:  fig.  96. 

19626  Palaeoperidinium  castanea  Deflandre;  Cookson  &  Eisenack  :  489,  pi.  3,  figs.  9-11. 

1964    Palaeoperidinium  castanea  Deflandre;  Cookson  &  Hughes  :  49,  pi.  5,  fig.  14. 

1967     Trichodium  castanea  (Deflandre)  Clarke  &  Verdier:  19,  pi.  i,  figs,  i,  2. 

DESCRIPTION.  The  shell  is  subspherical  with  occasionally  a  small  apical  horn  or 
a  tuft  of  apical  spines.  The  shell  wall  is  slightly  punctate  and  bears  numerous  small 
spines.  These  are  solid,  often  bifurcate  either  proximally  or  distally,  and  typically 
terminate  in  a  small  bifurcation.  Rarely  they  may  be  acuminate.  The  cingulum 
(3-5  fj,  in  width)  is  marked  by  two  parallel  lines  of  thickening  along  which  spines  are 
concentrated.  Lines  of  similar  thickening  sometimes  occur  perpendicularly  to  the 
cingulum  and  are  probably  sutural.  A  sulcus  has  not  been  observed.  A  precingular 
archaeopyle  is  commonly  present. 


132  CENOMANIAN   NON-CALCAREOUS   MICROPLANKTON,    i 

DIMENSIONS.  Range  of  observed  specimens:  shell  diameter  35  (49-0)  64 /n,  length 
of  spines  I  (3-3)  5  /x.  Number  of  specimens  measured,  16. 

REMARKS.  The  Cenomanian  specimens  resemble  the  type  material  from  the 
Upper  Cretaceous  of  France  in  all  respects.  The  presence  of  a  precingular  archaeo- 
pyle,  an  apical  prominence  or  apical  spines,  and  a  well  developed  cingulum  indicate 
that  this  species  should  be  placed  in  Trichodinium.  T.  intermedium  Eisenack  & 
Cookson,  from  the  Aptian  to  Lower  Albian  of  Australia,  is  very  similar  but  is  larger 
(shell  diameter  69-90  /A). 

OCCURRENCE.  T.  castaneum  is  a  rare  species  occurring  at  most  horizons  throughout 
the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  has  not  been 
recorded  from  the  North  American  samples.  This  species  has  a  recorded  strati- 
graphic  range  from  the  Aptian  to  the  Senonian  (Deflandre). 

Cyst-Family  MIGRODINIACEAE  Eisenack  emend.  Sarjeant  &  Downie  1966 
Genus  MICRODINIUM  Cookson  &  Eisenack  emend.  Sarjeant  1966 

REMARKS.  Microdinium  differs  from  Gonyaulacysta  primarily  in  possessing  a 
single  apical  plate  which  is  detached  in  archaeopyle  formation.  Gorka  (1965),  in  des- 
cribing Upper  Jurassic  assemblages,  erected  the  genus  Tetrasphaera  which  is  diagnosed 
as  having  a  feebly  marked  tabulation  and  short  spines  on  the  plate  boundaries.  Gorka 
has  since  stated  (personal  communication  with  Dr.  W.  A.  S.  Sarjeant)  that  this 
genus  differs  from  Microdinium  only  in  these  two  respects.  However,  it  follows 
that  if  the  tabulation  is  not  clear  it  is  difficult  to  compare  this  new  genus  with 
Microdinium  or  any  other  genus  possessing  a  tabulate  cyst,  since  the  diagnosis  of 
these  genera  is  primarily  based  on  the  observed  tabulation.  Also,  spines  have  been 
observed  on  the  plate  boundaries  of  Microdinium,  even  in  the  type  species.  Thus 
it  would  appear  that  Tetrasphaera  at  the  moment  is  not  precisely  defined. 

Microdinium  cf.  ornatum  Cookson  &  Eisenack. 
(PI.  4,  fig.  5;  Figs.  I3C,  F) 

ig66fl  Microdinium  cf.  ornatum  Cookson  &  Eisenack;  Sarjeant  :  149,  pi.  16,  figs.  3-6,  text-fig.  38. 
1967     Microdinium  ornatum  Cookson  &  Eisenack;  Clarke  &  Verdier:  66,  pi.  5,  figs.  11-14. 

DESCRIPTION.  The  shell  is  subspherical  to  ovoidal  possessing  a  smooth  body  wall 
ornamented  by  a  few  large  tubercles.  The  latter  may  be  flat  or  distinctly  concave 
distally.  In  two  specimens  small  tubercles  were  seen  to  delimit  a  plate  in  the 
posterior  portion  of  the  ventral  area.  The  plates  are  bordered  by  short,  broad 
projections  (Fig.  I3F),  which  may  be  isolated  or  united  distally. 

DIMENSIONS.  Range  of  observed  specimens:  shell  length  31-34 //,,  width  23-31  /z, 
height  of  crests  1-2  /*.  Number  of  specimens  measured,  6. 

REMARKS.  M .  cf.  ornatum  differs  from  M.  ornatum  Cookson  &  Eisenack  (19600) 
in  that  (i)  the  small  cingular  plate,  ventral  to  plate  6c,  is  absent  or  has  a  very 
reduced  ventral  suture,  and  (ii)  the  plates  are  not  bordered  by  ledges,  which  are 
sometimes  perforate,  but  by  isolated  tubercles  and  spines. 


CENOMANIAN   NON-CALCAREOUS   MICROPLANKTON,    i  133 

OCCURRENCE.  M.  cf.  ornatum  is  a  rare  species  in  the  material  examined  being 
found  only  in  the  following  samples:  FM  810,  FM  670,  FM  650,  FM  520  (Turonian), 
E  189  and  E  153.  In  Australia  M.  ornatum  has  a  stratigraphic  range  from  the  Albian 
to  the  Lower  Turonian. 


Microdinium  setosum  Sarjeant. 
(PI.  2,  fig.  4;  Fig.  i3H) 

1966  Microdinium  setosum.  Sarjeant  :  151,  pi.  16,  figs.  9,  10;  text-fig.  39. 

1967  Microdinium  echinatum  Clarke  &  Verdier:  64,  pi.  i,  figs.  9,  10,  text-fig.  26. 

DESCRIPTION.  The  shell  is  spherical  to  ovoidal  and  possesses  a  well  developed 
tabulation.  The  shell  surface  is  either  lightly  or  coarsely  granular.  The  two  plates 
observed  by  Sarjeant  in  the  medial  region  of  the  sulcus  are  not  always  delimited. 
The  sutural  crests  are  often  relatively  high  and  give  rise  to  numerous,  well  developed 
thorn-like  spines  (Fig.  I3H). 

DIMENSIONS.  Range  of  type  material:  shell  length  25  (29-3)  37  p.,  width  21  (26-2) 
31  p,  maximum  height  of  crests  1-5  (3-8)  7  /x.  Number  of  specimens  measured,  22. 

REMARKS.  In  general  form  M.  setosum  is  similar  to  M.  ornatum  but  differs  in  the 
presence  of  spiny  crests,  a  dense  granulation  and,  slightly,  in  the  tabulation  exhibited. 

OCCURRENCE.  M.  setosum  is  an  infrequent  to  fairly  common  species  at  most 
horizons  throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles. 
It  has  also  been  recorded  from  the  Albian  of  Fetcham  Mill. 


Microdinium  distinctum  sp.  nov. 
(PI.  2,  figs.  9-11 ;  Figs.  130,  E.  I) 

1967     Microdinium  ornatum  Cookson  &  Eisenack;  Clarke  &  Verdier:  pi.  5,  figs.  11-12. 

DERIVATION  OF  NAME.  Latin,  distinctus,  different — with  reference  to  the 
distinctive  appearance  of  this  species. 

DIAGNOSIS.  Shell  subspherical,  thick-walled,  smooth.  Sutural  crests  well 
developed  bearing  short,  broad,  flat-topped  spines.  Reflected  tabulation  i,  oa,  6" 
6"',  ip,  i"";  plates  i"  and  6"  very  reduced  and  cingular  plates  absent.  Cingulum 
broad,  weakly  laevo-rotatory. 

HOLOTYPE.  G.S.M.,  slide  PF  3989,  (i).  Lower  Chalk,  H.M.  Geological  Survey 
Borehole,  Fetcham  Mill,  Surrey  at  730  ft.  depth.  Upper  Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype:  shell  length  36^,  width  37  p,  height  of  crests  c.  2,\i. 
Range:  shell  length  29-36^,  width  30-37 //.,  height  of  crests  2-2-5^.  Number  of 
specimens  measured,  7. 

DESCRIPTION.  The  shell  wall  is  relatively  thick,  c.  i  /z,  and  develops  pronounced 
ridges  at  the  plate  boundaries.  The  sutural  spines  are  closely  set,  very  broad  and 
flat  distally. 


134 


GENOMANIAN  NON-CALCAREOUS   MICROPLANKTON,    i 


H 


FIG.  13.  Cribroperidinium  orthoceras  (Eisenack).  A.  Specimen  illustrated  by  Eisenack 
(1958,  text-fig.  3)  redrawn  to  show  full  crestal  arrangement.  B.  Holotype  (Eisenack 
1958,  text-fig.  2)  redrawn  to  show  full  crestal  arrangement.  Microdinium  cf.  ornatum 
Cookson  &  Eisenack,  C.  Ventral  surface  of  Holotype  (X  700).  Microdinium  distinctum 
sp.  nov.,  D.  Ventral  surface  of  Holotype  (x  700),  E.  Dorsal  surface  of  Holotype  (X  700). 
Microdinium  cf.  ornatum  Cookson  &  Eisenack,  F.  Sutural  spines.  Microdinium  vario- 
spinum  sp.  nov.,  G.  Sutural  spines.  Microdinium  setosum  Sarjeant,  H.  Sutural  spines. 
Microdinium  distinctum  sp.  nov.,  I.  Sutural  spines. 


CENOMANIAN   NON-CALCAREOUS    MICROPL ANKTON,    i  135 

The  precingular  plates  are  generally  smaller  than  the  postcingular  plates.  Plates 
i"  and  6"  are  small  and  appear  as  slight  projections  in  the  ventral  area.  This  is 
widest  posteriorly  and  is  open  anteriorly.  The  apical  plate  is  six-sided  and  is  lost 
in  archaeopyle  formation.  The  archaeopyle  possesses  slits  extending  posteriorly 
between  the  precingular  plates. 

REMARKS.  M .  distinctum  may  be  distinguished  from  all  other  species  of  Micro- 
dinium  by  the  thick,  smooth  shell  wall,  the  form  of  the  precingular  plates  and  the 
absence  of  cingular  plates.  It  is  considered  that  the  absence  of  the  latter  does  not, 
at  present,  warrant  the  erection  of  a  new  genus. 

OCCURRENCE.  This  species  is  very  rare,  being  recorded  only  seven  times,  from  the 
following  samples:  FM  790,  FM  730,  FM  710,  FM  690,  £153  and  €617.  It  has 
never  been  observed  in  the  lower  horizons  of  the  Cenomanian. 


Microdinium  variospinum  sp.  nov. 
(PI.  2,  figs.  5,  6;  Fig.  i3G) 

DERIVATION  OF  NAME.  Latin,  varius,  different;  spinosus,  spine — with  reference 
to  the  variable  appearance  of  the  spines. 

DIAGNOSIS.  Shell  subspherical  to  ovoidal;  shell  wall  thin,  smooth,  granular  or 
lightly  reticulate.  Sutural  crests  low,  bearing  small  number  of  variably  shaped 
spines.  These  may  be  simple  tubercles  to  complex  bifurcating  protrusions.  Re- 
flected tabulation  i',  oa,  6",  (6c),  6'",  rp,  i"";  cingular  plate  boundaries  very  faint 
or  absent.  Cingulum  weakly  laevo-rotatory. 

HOLOTYPE.  B.M.  (N.H.)  V.  51981  (i).  Lower  Chalk,  Bureau  de  Recherches 
Geologiques  et  Minieres  Borehole,  Escalles,  Pas  de  Calais  at  165  metres  depth. 
Upper  Cretaceous  (Cenomanian) . 

DIMENSIONS.  Holotype:  shell  length  24^1,  width  23/4,  length  of  spines  1-1*5/4 
Range:  shell  length  20  (23-6)  27/4,  width  17  (20-0)  23/i,  length  of  spines  1-3 /u,. 
Number  of  specimens  measured,  9. 

DESCRIPTION.  The  cingular  plate  boundaries  are  usually  absent,  however,  one 
Saskatchewan  specimen  had  them  lightly  defined.  The  sutural  spines  are  few  in 
number  and  tend  to  be  concentrated  at  the  posterior  of  the  shell  (PI.  2,  fig.  5).  They 
are  stout  and  may  be  either  pointed  or  flattened  distally  (Fig.  130). 

REMARKS.  The  characteristic  features  of  M.  variospinum  are  the  nature  of  the 
shell  wall,  the  form  of  the  spines  and  the  lack  of  well  defined  cingular  plate  bound- 
aries. These  characters  together  differentiate  this  species  from  all  other  described 
species. 

OCCURRENCE.  M.  variospinum  is  a  rare  species  found  at  all  horizons,  save  the 
lower  three,  at  Escalles,  and  in  sample  CB  i,  CB  7  and  Sas  1084.  It  has  not  been 
recorded  from  Fetcham  Mill. 

GEOL.  17,  3  IO 


136  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

Microdiniutn  veligerum  (Deflandre)  comb.  nov. 
(PI.  3,  fig-  4i  PL  4,  fig-  4) 

1937     Micrhystridium  veligerum  Deflandre  :  81,  pi.  12,  fig.  9. 

1943     Ceratocorys  veligera  (Deflandre)  Lejeune-Carpentier  :  22,  text-figs.  1-6. 

19520  Ceratocorys  veligera  (Deflandre)  Deflandre  :  120,  text-fig.  102. 

i952b  Ceratocorys  veligera  (Deflandre)  Deflandre:  text-figs.  304  A-C. 

1967     Eisenackia  crassitabulata  Deflandre  &  Cookson;  Clarke  &  Verdier:  64,  pi.  8,  figs.  4-6. 

DESCRIPTION.  The  shell  is  ovoidal,  densely  granular  and  bears  a  number  of  high 
crests  delimiting  a  tabulation.  The  crests  are  distinctive,  being  i  to  5  /u  in  height, 
with  typically  a  smooth  outer  margin.  They  consist  of  two  membranes,  joined 
distally  and  diverging  proximally  to  form  a  broad  base,  I  to  3  p,  wide  to  the  crest. 
There  is  between  the  two  membranes  a  crestal  cavity  which  is  occasionally  sub- 
divided by  septa,  particularly  where  two  crests  diverge.  Here  a  conical  chamber  is 
usually  found.  The  reflected  tabulation  appears  to  be  i',  ?5",  6c,  6'",  ip,  i"". 
The  hypotract  is  considerably  larger  than  the  epitract,  the  latter  being  devoid  of 
crests  and  usually  possessing  a  pentagonal  apical  archaeopyle.  The  shape  of  the 
latter  is  the  only  indication  that  there  are  five  precingular  plates.  The  cingulum  is 
broad  and  does  not  appear  to  be  spiral.  Plate  i'",  and  to  a  lesser  extent  2'",  are 
reduced  to  accommodate  the  posterior  intercalary  plate.  The  remaining  four 
postcingular  plates  are  large  and  there  is  a  single  large  antapical  plate.  The  sulcus 
is  very  narrow  just  posterior  to  the  cingulum  and  then  widens  rapidly  towards  the 
antapex.  The  sulcus  extends  onto  the  epitract  where  it  sometimes  bears  five  small 
sulcal  plates. 

DIMENSIONS.  Range  of  observed  specimens:  shell  length  28  (31-5)  38 /*,  width 
25  (28-2)  32  //,.  Number  of  specimens  measured,  13. 

REMARKS.  Lejeune-Carpentier  (1943)  placed  this  species,  originally  observed  in 
the  Upper  Cretaceous  of  France,  in  the  genus  Ceratocorys  Stein  (1883)  on  the  basis 
of  its  similarity  to  motile  dinoflagellates  contained  in  this  genus.  However,  it  is  a 
cyst  possessing  an  apical  archaeopyle  and  should  not  be  attributed  to  a  motile 
dinoflagellate  genus.  Thus  this  species  is  here  transferred  to  Microdinium  on  the 
basis  of  the  tabulation,  apical  archaeopyle,  reduced  size  of  epitract  compared  with 
the  hypotract  and  overall  small  size.  M.  veligerum  does,  however,  differ  slightly 
from  the  other  species  in  this  genus  by  the  apparent  absence  of  crests  on  the  epitract 
and  the  probable  presence  of  five  precingular  plates  instead  of  six. 

Eisenackia  crassitabulata  as  illustrated  by  Clarke  &  Verdier  (1967)  is  undoubtedly 
M.  veligerum.  The  former,  as  originally  described  from  the  Australian  Lower 
Tertiary,  is  of  different  overall  form  and  is  considerably  larger  (72-78  by  55-67  /*) . 
The  size  of  the  specimen  illustrated  by  Clarke  &  Verdier  is  approximately  30  by  32  /£ 
and  is  thus  comparable  to  the  Cenomanian  specimens  of  M.  veligerum.  E.  crassi- 
tabulata has  been  recorded  from  the  Lower  Tertiary  and  also  from  the  Maestrichtian 
of  South  Africa  by  the  present  author.  M.  irregulare  Clarke  &  Verdier  (1967) 
appears  to  be  very  similar  to  M.  veligerum  and  any  definite  distinction  is  not  apparent. 

OCCURRENCE.  The  Cenomanian  forms  examined  resemble  the  specimens  illus- 
trated by  Lejeune-Carpentier  in  all  respects.  M.  veligerum  is  an  infrequent  to 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i  137 

common  species  at  all  horizons,  save  two,  at  Fetcham  Mill,  Compton  Bay  and 
Escalles.  These  two  horizons  are  basal  Cenomanian,  FM  840  and  CB  i,  and  it  was 
not  recorded  by  Cookson  &  Hughes  (1964)  from  the  Upper  Albian/basal  Cenomanian 
of  Cambridgeshire.  It  is  also  absent  in  the  North  American  material  and  has  not  been 
described  from  Australia.  This  species  is  present  in  sample  FM  520  of  Turonian 
age.  M.  veligerum  thus  appears  for  the  first  time  just  above  the  base  of  the  Ceno- 
manian and  extends  into  the  Turonian  but  is,  apparently,  of  restricted  geographical 
distribution. 

fMicrodinium  crinitum  sp.  nov. 
(PL  2,  figs.  7,  8) 

1967     Cometodinium  obscurum  Deflandre  &  Courteville;  Clarke  &  Verdier:  pi.  10,  fig.  3:  pi.  n, 
fig.  9- 

DERIVATION  OF  NAME.  Latin,  crinitus,  hairy — with  reference  to  the  numerous 
hair-like  spines. 

DIAGNOSIS.  Shell  subspherical,  periphragm  granular  and  giving  rise  to  numerous, 
fine,  flexuous  spines.  Sutural  crests  low,  bearing  numerous  spines.  Cingulum 
wide,  composed  of  elongate  plates.  Epitract  smaller  than  hypotract.  Archaeopyle 
not  normally  visible. 

HOLOTYPE.  G.S.M.  slide  PF  3990(1).  Lower  Chalk,  H.M.  Geological  Survey 
Borehole,  Fetcham  Mill,  Surrey  at  690  feet  depth.  Upper  Cretaceous  (Cenomanian.) 

DIMENSIONS.  Holotype:  shell  diameter  27  by  28  p,  length  of  spines  c.  12  /JL. 
Range:  shell  diameter  24  (30-1)  38/1,,  length  of  spines  6-19 /x.  Number  of  specimens 
measured,  18. 

DESCRIPTION.  The  periphragm  granules,  which  are  evenly  spaced  on  the  shell 
surface,  are  up  to  0-5  /j,  high,  and  often  form  the  bases  of  the  hair-like  spines.  The 
latter  tend  to  be  especially  concentrated  along  the  sutures,  and  because  of  this 
tendency  and  the  spherical  form  of  the  shell,  it  has  not  been  possible  to  fully  for- 
mulate a  tabulation.  However,  precingular,  cingular  and  postcingular  plates  are 
quite  obvious  when  the  orientation  is  favourable,  the  precingular  plates  being  smaller 
than  the  postcingular  plates.  The  cingulum  is  broad,  c.  5  /x.  The  archaeopyle, 
although  it  has  not  been  observed,  is  probably  apical. 

REMARKS.  The  numerous  hair-like  spines  and  the  tabulation  easily  distinguish 
1M.  crinitum  sp.  nov.  from  all  previously  described  forms  of  dinoflagellate  cysts. 
The  overall  shape,  the  small  size  and  the  fact  that  the  epitract  is  smaller  than  the 
hypotract  all  indicate  that  this  species  is  closely  related  to  the  genus  Microdinium. 
However,  plate  spines  have  not  been  recorded  in  Microdinium,  although  a  granula- 
tion has,  and  since  the  tabulation  has  not  been  elucidated  in  the  present  species  it  is 
only  placed  tentatively  in  this  genus. 

OCCURRENCE.  1M.  crinitum  is  infrequent  at  all  horizons  throughout  the  Ceno- 
manian of  Fetcham  Mill,  Compton  Bay  and  Escalles  and  is  also  present  in  the  Albian 
sample  from  Fetcham  Mill,  sample  FM  886.  It  has  only  once  been  recorded  at 
Saskatchewan,  in  sample  Sas  835  (Cenomanian). 


138  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 

Genus  HISTIOCYSTA  nov. 

DERIVATION  OF  NAME.  Greek,  histos,  mesh  or  network;  kystis,  sac  or  cell — with 
reference  to  the  reticulate  ornamentation  on  the  shell  surface. 

DIAGNOSIS.  Proximate  cysts,  spherical  to  subspherical ;  shell  wall  composed  of 
two  layers;  outer  layer  giving  rise  to  low  crests.  Crests  reflecting  Gonyaulax— 
type  tabulation  and  coarse,  subsidiary  reticulation  within  plate  boundaries.  Sutural 
crests  better  denned  than  crests  of  subsidiary  reticulation.  Archaeopyle  apical  with 
angular  margin.  Operculum  probably  single  apical  plate. 

TYPE  SPECIES.     Histiocysta  palla  sp.  nov.     Lower  Chalk  (Cenomanian) ;  England. 

REMARKS.  The  reasonably  well  denned  tabulation,  the  plate  ornamentation  and 
the  apical  archaeopyle  easily  distinguish  Histiocysta  from  all  previously  described 
genera.  The  most  similar  genera  are  Ellipsoidictyum  Klement  (19606)  and  Dic- 
tyopyxidia  Eisenack  (1961),  both  from  the  Upper  Jurassic.  Both  genera  possess 
an  apical  archaeopyle  and  a  cingulum  but  a  distinct  Gonyaulax-type  tabulation  is 
absent.  However,  it  seems  probable  that  Histiocysta  is  genetically  related  to  these 
two  genera.  Although  the  precise  tabulation  of  Histiocysta  cannot  be  elucidated 
the  presence  of  precingular,  cingular  and  postcingular  series  of  plates  together  with 
an  apical  archaeopyle  indicate  that  this  genus  belongs  to  the  Family  Microdiniaceae. 

Histiocysta  palla  sp.  nov. 
(PI.  i,  figs.  5,  6;  Figs.  I4A,  B) 
1939     Micrhystridium  sp.?  Deflandre  &  Courteville  :  pi.  3,  fig.  4. 

DERIVATION  OF  NAME.  Greek,  palla,  ball — with  reference  to  the  more  or  less 
spherical  shape  of  this  species. 

DIAGNOSIS.  Shell  spherical  to  subspherical,  thin-walled,  periphragm  smooth  and 
forming  a  reticulate  network  of  crests.  Network  consisting  of  reflected  dinoflagel- 
late  tabulation  with  central  region  of  each  plate  occupied  by  coarse  but  simple 
reticulation. 

HOLOTYPE.  G.S.M.  slide  PF  3052  (2).  Lower  Chalk,  H.M.  Geological  Survey 
Borehole,  Fetcham  Mill,  Surrey  at  650  feet  depth.  Upper  Cretaceous  (Cenomanian) . 

PARATYPE.  G.S.M.  slide  PF  3991(1).  Lower  Chalk,  H.M.  Geological  Survey 
Borehole,  Fetcham  Mill,  Surrey  at  710  feet  depth.  Upper  Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype :  shell  diameter  30  by  33  p,  height  of  crests  4-5  /*.  Para- 
type:  shell  diameter  26  by  29^,  height  of  crests  c.  2-5 /A.  Range:  shell  diameter 
25  (31-8)  38  /z,  height  of  crests  1-5  /z.  Number  of  specimens  measured,  19. 

DESCRIPTION.  The  precingular  and  postcingular  plates  may  be  observed  on  most 
specimens  but  the  exact  tabulation  has  not,  as  yet,  been  elucidated.  The  central 
region  of  each  plate  is  occupied  by  a  coarse,  subpolygonal  reticulation  which  occasion- 
ally extends  to  the  plate  boundaries  (Figs.  I4A,  B).  The  cingular  region  is  clearly 
defined  by  crests  and  encircles  the  shell.  Cingular  plates  are  not  usually  discernible, 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 


139 


the  entire  region  being  occupied  by  a  coarse  reticulation.  An  apical  archaeopyle 
with  an  angular  margin  is  typically  present,  the  six-sided  operculum  often  remaining 
attached  to  the  shell. 

REMARKS.     The  combination  of  apical  archaeopyle,  tabulation  and  reticulation 
makes  H.  pdlla  an  easily  recognizable  species  distinct  from  all  previously  described 


B 


C  D 

FIG.  14.  Histiocysta  palla  sp.  nov.,  A.  Lateral  view  of  Holotype  showing  attached  oper- 
culum, apical  archaeopyle,  precingular  plates  and  cingulum  (X  13°°)-  B.  Lateral 
view  of  Holotype  showing  well  defined  plate  boundaries  (X  13°°)-  Ellipsodinium 
rugulosum  Clarke  &  Verdier,  C.  Lateral  view  showing  partially  detached  operculum  and 
cingulum  (X  1300).  D.  Lateral  view  (X  1300). 


i4o  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

forms.  This  species  was  figured,  but  not  described,  by  Deflandre  &  Courteville 
(1939)  as  Micrhystridium  sp.?  from  the  Senonian.  Membranilarnax  cf.  pterosper- 
moides  Deflandre  (1937)  is  of  similar  form  but  does  not  possess  a  reticulation. 

OCCURRENCE.  H.  palla  is  found  throughout  the  European  Cenomanian,  except 
for  the  lowermost  horizons.  It  is  fairly  common  at  Fetcham  Mill  but  rare  at  Comp- 
ton  Bay  and  Escalles.  It  has  not  been  observed  in  the  North  American  samples. 

Cyst-Family  FROMEACEAE  Sarjeant  &  Downie  1966 
Genus  FRO  ME  A  Cookson  &  Eisenack  1958 

REMARKS.  A  number  of  specimens  referable  to  the  type  species,  F.  amphora,  do 
not  appear  to  possess  a  cingulum.  The  absence  of  a  cingulum  makes  this  genus 
similar  to  Chytroeisphaeridia  Sarjeant  (1962).  They  differ,  however,  in  that  the 
archaeopyle  of  Fromea  has  a  rounded  margin,  and  in  the  elongate  shape  typical  of 
the  latter  genus. 

Fromea  amphora  Cookson  &  Eisenack 
(PI.  3,  figs.  2,  3) 

1958     Fromea  amphora  Cookson  &  Eisenack  :  56,  pi.  5,  figs.  10,  u. 

19666  Fromea  amphora  Cookson  &  Eisenack;  Sarjeant  :  209,  pi.  22,  fig.  4;  pi.  23,  fig.  3  (see  also 
for  earlier  references). 

DESCRIPTION.  The  shell  is  ovoidal,  thick-walled  (2-3  p,)  and  typically  possesses 
an  apical  archaeopyle  with  a  rounded  margin.  In  one  specimen  (PL  3,  fig.  3)  the 
apical  region  is  still  attached  and  may  be  seen  to  be  perfectly  rounded.  A  cingulum 
was  not  observed  in  any  of  the  specimens. 

DIMENSIONS.  Range  of  observed  specimens:  shell  length  56  (72-5)  85^,  width 
47  (60)  71  /A.  Number  of  specimens  measured,  6. 

REMARKS.  The  Cenomanian  specimens  are  identical  with  the  type  material  from 
the  Aptian-Cenomanian  of  Australia  except  that  the  cingulum  is  absent.  Cookson 
&  Eisenack  (1958)  state,  however,  that  the  cingulum  may  be  rather  faint  and  it  is 
probable  that  the  European  forms  fall  within  the  range  of  variation  for  this  species. 
F.  amphora  has  been  recorded  from  the  Barremian  of  England  by  Sarjeant  (19666). 
The  specimens  described  by  Sarjeant,  like  the  Cenomanian  forms,  do  not  possess  a 
cingulum.  Maliavkina  et  al.  (1961)  describes  some  very  similar,  but  rather  smaller, 
forms  from  the  Maestrichtian  of  Siberia,  calling  them  Chrysomonadinael .  These 
forms  possess  the  typical  rounded  archaeopyle  of  this  genus. 

OCCURRENCE.  Five  specimens  have  been  recorded  from  Fetcham  Mill,  from 
samples  FM  810,  770,  750  and  650,  and  one  specimen  from  Escalles,  sample  E  177. 

Genus  CHYTROEISPHAERIDIA  Sarjeant  1962 

REMARKS.  Chytroeisphaeridia  and  Canningia  Cookson  &  Eisenack  (19606)  are 
similar  and  probably  fairly  closely  related.  The  latter  is  usually  more  polygonal, 
has  an  apical  horn,  and  sometimes  the  vestiges  of  a  cingulum. 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  141 

Chytroeisphaeridia  euteiches  sp.  nov. 
(PL  3,  figs.  8,  9) 

DERIVATION  OF  NAME.  Greek,  euteiches,  well- walled — with  reference  to  the  stout 
wall  of  this  species. 

DIAGNOSIS.  Shell  subspherical;  shell  wall  thick  and  densely  granular.  Angular 
apical  archaeopyle  typically  present. 

HOLOTYPE.  B.M.  (N.H.)  V.  51982  (2).  Lower  Chalk,  Bureau  de  Recherches  Geo- 
logiques  et  Minieres  Borehole,  Escalles,  Pas  de  Calais,  at  159  metres  depth.  Upper 
Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype:  shell  length  53^,  width  59 /*.  Range:  shell  length 
48-60  jit,  width  49-59  p.  Number  of  specimens  measured,  6. 

DESCRIPTION.  The  shell  wall  is  thick  (2-3  /*)  and  may  be  composed  of  two  layers. 
If  the  wall  is  bipartite  then  the  inner  layer  is  thin,  the  outer  layer  making  up  almost 
the  entire  wall  thickness.  This  layer  appears  to  be  composed  of  minute  cellular 
elements  and  is  densely  granular  on  the  surface.  The  apical  archaeopyle,  when 
developed,  is  angular  with  small  slits  passing  posteriorly  from  its  margin  between  each 
precingular  plate.  A  sulcal  notch  is  also  present  (PI.  3,  fig.  8).  Plate  boundaries 
and  cingulum  are  not  discernible. 

REMARKS.  This  species  is  a  simple,  subspherical  shell  possessing  an  apical 
archaeopyle.  One  species,  C.  chytroeides  Sarjeant  (1962),  from  the  Upper  Jurassic 
of  England,  differs  from  C.  euteiches  in  having  a  fairly  thin  and  only  slightly  granular 
shell  wall.  Chytroeisphaeridia  sp.  Sarjeant  (19656)  is  similar  in  size  and  also  granular 
but  does  not  possess  a  thick  shell  wall.  Canningia  rotundata  Cookson  &  Eisenack 
(1961)  is  also  similar  but  tends  to  have  a  polygonal  shell  with  a  small  apical  horn. 

OCCURRENCE.  C.  euteiches  has  been  recorded  from  two  horizons  only,  E  165,  &  E 
159  from  Escalles,  where  it  is  infrequent. 

Genus  CASSICULOSPHAERIDIA  nov. 

DERIVATION  OF  NAME.  Latin,  cassiculus,  hunting-net;  sphaera,  ball — with 
reference  to  the  surface  reticulation  of  the  shell. 

DIAGNOSIS.  Proximate  cysts;  shell  spherical  to  subspherical,  composed  of  two 
layers,  without  apical  or  antapical  protuberances.  Periphragm  giving  rise  to  low 
crests  or  membranes  which  form  a  reticulate  pattern.  Tabulation  absent.  Archae- 
opyle apical  with  angular  margin. 

TYPE  Species.  Cassiculosphaeridia  reticulata  sp.  nov.  Lower  Chalk  (Ceno- 
manian); France. 

REMARKS.  The  surface  reticulation  of  this  genus  is  very  similar  to  that  of 
Ellipsoidictyum  cinctum  Klement  (1960).  Both  possess  an  apical  archaeopyle,  but 
whereas  in  Cassiculosphaeridia  all  signs  of  a  tabulation  are  absent,  in  Ellipsoidictyum 
there  is  an  obvious  cingulum.  Dictyopyxidia  Eisenack  (1961)  is  also  very  similar 
but  possesses  a  cingulum  and  sulcus. 


142  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 

Cassiculosphaeridia  reticulata  sp.  nov. 

(PL  3,  fig-  7:  PL  4,  %•  3) 

DERIVATION  OF  NAME.  Latin,  reticulatus,  net-like — with  reference  to  the 
reticulate  pattern  formed  by  the  periphragm  crests. 

DIAGNOSIS.  Shell  spherical  to  subspherical.  Shell  surface  bearing  low  ridges, 
forming  a  coarse  reticulation,  from  which  arise  fine  membranous  crests.  Shell  wall 
lightly  to  densely  granular. 

HOLOTYPE.  B.M.  (N.H.)  ¥.51981  (4).  Lower  Chalk,  Bureau  de  Recherches 
Geologiques  et  Minieres  Borehole,  Escalles,  Pas  de  Calais,  at  165  metres  depth. 
Upper  Cretaceous  (Cenomanian) . 

DIMENSIONS.  Holotype:  shell  diameter  38  by  38/11,  height  of  crests  4  to  6  p. 
Range :  shell  diameter  33  (43-6)  55  p,,  maximum  height  of  crests  3  (7-3)  II  /n.  Number 
of  specimens  measured,  15. 

DESCRIPTION.  The  areas  delimited  by  the  low  ridges  are  typically  subpolygonal 
but  may  be  of  irregular  shape.  They  vary  considerably  in  size,  from  2  to  io/u,  in 
diameter.  The  membranous  crests  are  very  fine  and  tend  to  be  flexuous  since 
supporting  structures  are  absent. 

REMARKS.  The  surface  reticulation,  the  absence  of  any  tabulation  and  the  apical 
archaeopyle  together  distinguished  this  species  from  all  previously  described  forms. 

OCCURRENCE.  C.  reticulata  is  rare  to  common  in  samples  from  the  Middle  and 
Upper  Cenomanian  of  Fetcham  Mill  (not  found  below  sample  FM  750),  and  from 
the  Lower,  Middle  and  Upper  Cenomanian  of  Escalles  (not  found  below  sample 
E  207).  This  species  was  absent  from  the  samples  from  Compton  Bay  and  from 
North  America. 

Genus  EPELIDOSPHAERIDIA  nov. 

DERIVATION  OF  NAME.  Greek,  epelidos,  cover  or  lid;  sphaera,  ball — with  reference 
to  the  conical  apical  operculum  which  sometimes  remains  attached  to  the  shell. 

DIAGNOSIS.  Shell  subpolygonal;  epitract  conical  with  small  apical  protuberance, 
hypotract  polygonal  with  small  antapical  horn  on  one  side.  Shell  wall  two  layered, 
periphragm  giving  rise  to  a  moderate  number  of  spines,  truncated  or  forked  distally 
Cingulum  and  sulcus  outlined  by  spines.  Cingulum  slightly  laevo-rotatory.  Apical 
archaeopyle. 

TYPE  SPECIES.  Palaeoperidinium  spinosum  Cookson  &  Hughes  1964.  Cambridge 
Greensand  (Cenomanian),  England. 

REMARKS.  The  presence  of  an  apical  archaeopyle,  together  with  a  well  developed 
cingulum  and  sulcus  differentiate  Epdidosphaeridia  from  all  previously  described 
genera.  Doidyx  Sarjeant  (19666)  is  most  similar  but  differs  in  that  the  shell  is  asym- 
metrical, the  hypotract  is  conical  and  a  sulcus  is  absent. 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i  143 

Epelidosphaeridia  spinosa  (Cookson  &  Hughes)  comb.  nov. 
(PI.  3,  figs.  10-12) 

1964     Palaeoperidinium  spinosum  Cookson  &  Hughes  :  49,  pi.  8,  figs.  6-8. 

1967     Palaeoperidinium  spinosum  Cookson  &  Hughes:  Clarke  &  Verdier:  70,  pi.  14,  figs.  10-12. 

DESCRIPTION.  The  shell  possesses  convex  sides,  conical  epitract  and  a  hypotract 
which  is  more  or  less  truncated  posteriorly.  A  small  apical  prominence  is  commonly 
present.  The  periphragm  is  smooth  or  lightly  granular  and  forms  a  moderate  num- 
ber of  small,  stout  spines.  The  spines  appear  to  be  hollow,  closed  proximally, 
parallel  sided,  and  are  oblate  distally  or  terminate  with  a  small  fork.  The  spines 
widen  slightly  before  joining  the  shell  and  they  are  sometimes  joined  proximally. 
This  is  particularly  well  developed  in  the  antapical  region  where  the  processes  are 
joined  medially  and  form  a  slight  projection  on  one  side  of  the  shell.  The  cingulum 
is  clearly  delimited  by  two  parallel  lines  of  closely  set  spines  and  is  5  to  8  p  in  width. 
It  is  only  slightly  helicoid  and  bears  few  spines  on  its  surface.  The  sulcus  in  most 
specimens  is  clearly  defined,  being  slightly  hollowed  and  almost  devoid  of  spines. 
The  spines  on  the  remainder  of  the  shell  surface  are  usually  randomly  arranged,  but 
occasionally  a  vague  alignment  is  present  suggesting  a  tabulation.  An  apical  archa- 
eopyle  is  constantly  developed,  the  margin  being  only  slightly  angular. 

DIMENSIONS.  Range  of  observed  specimens:  shell  length  32  (43)  56  fj,,  width 
27  (42-1)  57  p,  maximum  length  of  spines  2-5  (3-7)  5ju,.  Number  of  specimens 
measured,  21. 

REMARKS.  The  Cenomanian  specimens  examined  strongly  resemble  the  type 
material  from  the  Cambridge  Greensand  and  Chalk  Marl  (Lower  Cenomanian)  of 
Cambridgeshire,  England.  In  the  lower  horizons  of  the  Cenomanian  the  sulcus, 
although  always  present,  is  not  so  clearly  defined  as  in  higher  horizons. 

OCCURRENCE.  E.  spinosa  is  a  rare  to  common  species  in  the  lower  and  middle 
horizons  of  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  is 
absent  from  samples  FM  690,  670,  650;  CB  13,  15,  17,  19,  21;  and  E  159,  153.  It 
has  only  beeen  observed  in  one  North  American  sample — sample  Sas  1023 
(Saskatchewan,  Albian). 

Cyst-Family  HYSTRICHOSPHAERIDIACEAE  Evit  emend. 
Sarjeant  &  Downie  1966 

Genus  HYSTRICHOSPHAERIDIUM  Deflandre  emend.  Davey  &  Williams  1966 

Hystrichosphaeridium  tubiferum  (Ehrenberg) 

(PI.  5,  figs.  5,  8) 

1838     Xanthidium  tubiferum  Ehrenberg  :  pi.  i,  fig.  16. 

19666  Hystrichosphaeridium  tubiferum  (Ehr.)  Davey  &  Williams  :  56,  pi.  6,  figs,  i,  2;  pi.  8,  fig.  5; 
pi.  10,  fig.  2;  text-fig.  13.     (See  also  for  earlier  references). 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  28  (38-1) 
51  /z,  maximum  length  of  processes  15  (25-0)  37  //,.  Number  of  specimens  measured, 
26. 


i44  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

REMARKS.  H.  tubiferum  is  an  infrequent  to  common  species  at  all  horizons 
throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  is  also 
recorded  from  the  Albian  (sample  FM  886)  and  Turanian  (sample  FM  520)  of  Fetcham 
Mill.  This  species  was  not  recorded  in  the  North  American  material.  Thus  the 
earliest  recording  of  H .  tubiferum  is  from  the  Albian ;  it  ranges  throughout  the  Upper 
Cretaceous  and  has  been  recorded  from  the  Eocene  (Ypresian)  by  Davey  &  Williams 
(19666).  It  is  a  long-ranging  species  of  little  stratigraphic  value. 

Hystrichosphaeridium  deanei  Davey  &  Williams 
(PI.  4,  %.  i) 

19666  Hystrichosphaeridium  deanei  Davey  &  Williams  :  58,  pi.  6,  figs.  4,  8. 

1967     Hystrichosphaeridium  stellatum  Maier;  Clarke  &  Verdier:  55,  pi.  12,  figs,  i,  2. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  37  (45-7) 
54  fji,  maximum  length  of  processes  22  (35-8)  45  [i.  Number  of  specimens  measured, 
10. 

REMARKS.  One  specimen,  occurring  in  sample  E  207  (Escalles),  possesses  broad 
processes  and  appears  to  occupy  a  position  midway  between  H.  deanei  and  H. 
tubiferum. 

OCCURRENCE.  H.  deanei  is  a  rare  species  confined  to  the  Middle  and  Upper 
Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles  (Table  25).  The  samples 
in  which  it  first  occurs  are  FM  710  (Fetcham  Mill),  CB  13  (Compton  Bay)  and  E  195 
(Escalles).//.  deanei  has  also  been  recorded  from  the  Turonian  sample  FM520, 
from  Fetcham  Mill.  It  is  absent  from  the  North  American  material. 

Hystrichosphaeridium  readei  Davey  &  Williams 

19666  Hystrichosphaeridium  readei  Davey  &  Williams  :  64,  pi.  6,  fig.  3   (See  also  for  earlier 
references) . 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  31  (42-1) 
57  p,  maximum  length  of  processes  23  (29-7)  35  /t.  Number  of  specimens  measured, 
ii. 

OCCURRENCE.  H.  readei  is  very  rare  in  samples  FM  810,  790,  770  (Fetcham  Mill) 
and  E  207  (Escalles);  and  it  is  rare  to  infrequent  in  samples  FM  690,  670,  E  183,  177, 
X65,  159  and  CB  9  (Compton  Bay).  Two  specimens  were  recorded  in  the  Albian 
sample  (FM  886)  from  Fetcham  Mill.  This  species  was  not  recorded  in  the  North 
American  samples. 

Hystrichosphaeridium  radiculatum  Davey  &  Williams 
(PI.  4,  fig-  8) 

19666  Hystrichosphaeridium  radiculatum  Davey  &  Williams  :  65,  pi.  7,  fig.  9;  pi.  9,  fig.  6. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  27  (38-8) 
43  ju,  maximum  length  of  processes  12  (15-9)  20  /*.  Number  of  specimens  measured, 
II. 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  145 

OCCURRENCE.  H.  radiculatum  is  a  very  rare  to  infrequent  species  occurring  at  a 
number  of  horizons  throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and 
Escalles.  It  has  also  been  recorded  in  the  Albian  sample  from  Fetcham  Mill 
(sample  FM  886),  but  is  absent  from  the  North  American  material. 

Hystrichosphaeridium  mantelli  Davey  &  Williams 
(PI-  4.  ng.  9) 

19666  Hystrichosphaeridium  mantelli  Davey  &  Williams  :  66,  pi.  6,  fig.  6. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  32  (38-6) 
48  fi,  maximum  length  of  processes  12  (21-4)  26^.  Number  of  specimens  measured, 
15- 

REMARKS.  The  reticulate  nature  of  the  central  body  and  the  fibrous  processes 
differentiate  H.  mantelli  from  most  previously  described  species.  H.  radiculatum  is 
the  most  similar  but  differs  from  H.  mantelli  by  the  more  branched  and  deeply 
furcate  processes  and  the  tendency  for  the  fibrils  of  the  processes  to  continue  across 
the  surface  of  the  central  body. 

OCCURRENCE.  H.  mantelli  is  a  very  rare  to  infrequent  species  occurring  in  most 
samples  throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles. 
It  has  also  been  recorded  in  the  Turonian  sample  from  Fetcham  Mill  (sample  FM 
520),  but  is  absent  from  the  North  American  material. 

Hystrichosphaeridium  bowerbanki  Davey  &  Williams 
(PI-  5,  fig-  9) 

19666  Hystrichosphaeridium  bowerbanki  Davey  &  Williams  :  69,  pi.  8,  figs.  1,4. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  25  (31-9) 
40jLt,  maximum  length  of  processes  20  (25-5)  28  p.  Number  of  specimens  measured, 
10. 

REMARKS.  H.  bowerbanki  is  rare  to  infrequent  in  six  Middle  Cenomanian  samples — 
FM  770,  750,  730  and  690  from  Fetcham  Mill  and  CB  9,  and  17  from  Compton 
Bay.  It  has  also  been  recorded  from  the  Albian  sample  (FM  886)  and  the  Turonian 
sample  (FM  520),  both  from  Fetcham  Mill. 

Hystrichosphaeridium  difficile  Manum  &  Cookson 
(PI.  4,  figs.  2,  6,  7) 

1964     Hystrichosphaeridium  difficile  Manum  &  Cookson  :  12,  pi.  3,  figs.  1-3,  7. 

DESCRIPTION.  The  shell  is  subspherical,  sometimes  with  a  small  apical  prom- 
inence; shell  wall  thick  (c.  i  /JL),  smooth  to  lightly  granular.  The  processes,  approxi- 
mately 30  in  number,  are  complex,  usually  broadly  tubiform  or  buccinate  in  shape, 
of  constant  length  on  any  specimen  but  variable  in  width  (3  to  18/11).  Distally  the 


146  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

larger  processes  have  a  rectangular  opening,  with  a  serrate  margin  which  gives  rise 
to  four  broad  spines.  Typically  there  are  four  bands  of  thickening  extending  along 
the  length  of  the  processes  and  passing  onto  the  shell  surface.  There,  each  joins 
with  a  similar  thickening  from  a  neighbouring  process,  thus  forming  a  coarse  reticula- 
tion on  the  shell  surface.  The  large  tubular  processes  are  arranged  in  a  circular 
manner  around  the  shell,  reflecting  the  precingular,  cingular  and  postcingular  series 
of  plates.  Fine  processes  are  uncommon  and  may  be  sulcal  in  position.  An  apical 
archaeopyle  is  typically  developed  and  possesses  an  angular  margin  marked  at 
intervals  with  V-shaped  notches.  Although  often  remaining  attached,  isolated 
operculae  have  been  identified  (PI.  4,  figs.  6,  7)  and  bear  four  moderate-sized  tubular 
processes. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  49  (64-9) 
79  p,,  maximum  length  of  processes  18  (24-5)  31  p.  Number  of  specimens  measured, 
8. 

REMARKS.  The  specimens  studied  are  extremely  similar  to  the  type  material  of 
Lower  Cretaceous  age  described  by  Manum  &  Cookson  (1964)  from  Arctic  Canada. 
The  only  difference  appears  to  be  that  in  the  type  material  the  shell  surface  has  a 
fine  reticulation  which  is  absent  from  the  specimens  studied. 

Two  similar  species  are  H.  costatum  Davey  &  Williams  (19666)  from  the  Oxford 
Clay  of  England  and  H.  readei  Davey  &  Williams  (19666)  from  the  Cenomanian  of 
England.  However,  both  species  are  considerably  smaller  than  H.  difficile  and  possess 
narrower  and  less  complex  processes. 

OCCURRENCE.  H.  difficile  has  only  been  recorded  from  the  Saskatchewan  material. 
It  is  infrequent  in  samples  Sas  1084  (Albian)  and  Sas  890  (Cenomanian),  and  common 
in  sample  Sas  805  (Cenomanian). 

Genus  OLIGOSPHAERIDIUM  Davey  &  Williams  1966 

Oligosphaeridium  complex  (White) 

(PI.  5,  figs.  6,  7) 

1842     Xanthidium  tubiferum  complex  White  :  39,  pi.  4,  div.  3,  fig.  n. 

19666  Oligosphaeridium  complex  (White)  Davey  &  Williams  :  71,  pi.  7,  figs,  i,  2;  pi.  10,  fig.  3; 

text-fig.  14  (See  also  for  earlier  references). 
1967     Hystrichosphaeridium  complex  (White)  Clarke  &  Verdier:  53,  pi.  n,  figs.  10,  n. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  34  (41-1) 
55  /A,  maximum  length  of  processes  22  (34*6)  43  //,.  Number  of  specimens  measured, 
12. 

REMARKS.  Davey  &  Williams  described  examples  of  0.  complex  from  the  Speeton 
Clay  (Barremian),  Lower  Chalk  (Cenomanian)  and  London  Clay  (Ypresian),  all 
from  England. 

OCCURRENCE.  0.  complex  is  a  rare  to  common  species  in  all  samples  from  Fetcham 
Mill,  Compton  Bay  and  Escalles.  It  is  also  present  in  the  Albian  sample  (FM  886) 
and  the  Turonian  sample  (FM  520)  both  from  Fetcham  Mill.  This  species  is  present 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  147 

in  the  Albian  and  the  lowermost  Cenomanian  sample  from  Saskatchewan — Sas  1084 
1023,  967,  and  890 — but  is  absent  from  the  Texas  samples. 

0.  complex  thus  has  a  known  stratigraphic  range  from  the  Neocomian  (Gocht 
1959;  Cookson  &  Eisenack  1958)  to  the  Eocene,  Ypresian  (Davey  &  Williams  19666). 

Oligosphaeridium  reticulatum  Davey  &  Williams 
19666  Oligosphaeridium  reticulatum  Davey  &  Williams  :  74,  pi.  7,  fig.  10. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  29-47/4, 
length  of  processes  14-26/4.  Number  of  specimens  measured,  5. 

OCCURRENCE.  0.  reticulatum  is  an  infrequent  species  occurring  in  the  two  lower 
samples  from  Fetcham  Mill,  samples  FM  840  and  810. 

Oligosphaeridium  prolixispinosum  Davey  &  Williams 
(PI-  5,  %.  4) 

19666  Oligosphaeridium  prolixispinosum  Davey  &  Williams  :  76,  pi.  8,  figs.  2,  3. 

DIMENSIONS.  Range  of  observed  specimens:  length  of  central  body  33  (39-0) 
43 /A,  width  20  (28-7)  34 /u,  maximum  length  of  processes  18  (25-0)  30/4.  Number  of 
specimens  measured,  15. 

OCCURRENCE.  0.  prolixispinosum  is  a  rare  species  confined  mainly  to  the  Lower 
and  Middle  Cenomanian  of  Fetcham  Mill  and  Escalles,  although  it  does  occur  occas- 
ionally in  the  Upper  Cenomanian.  This  species  has  only  been  recorded  twice  in  the 
samples  CB  17  and  21  from  Compton  Bay,  both  from  the  Upper  Cenomanian.  It 
has  not  been  recorded  in  the  North  American  material. 

Oligosphaeridium  anthophorum  (Cookson  &  Eisenack) 
(PI.  5,  figs,  i,  2,  3) 

1958     Hystrichosphaeridium  anthophorum  Cookson  &  Eisenack  :  43,  pi.  n,  figs.  12,  13;  text-figs. 

16-18. 

1958     Hystrichosphaeridium  anthophorum  Cookson  &  Eisenack;  Eisenack,  402,  pi.  26,  figs,  i,  2. 
1961     Hystrichosphaeridium  anthophorum  Cookson  &  Eisenack;  Alberti  :  34,  pi.  9,  fig.  16. 
19666  Oligosphaeridium  anthophorum  (Cookson  &  Eisenack)  Davey  &  Williams  :  77. 

DESCRIPTION.  The  shell  is  subspherical ;  shell  wall  smooth  to  lightly  granular. 
An  apical  archaeopyle  is  typically  developed  and  possesses  an  angular  margin.  The 
processes  are  hollow,  buccinate  to  infundibular,  with  the  distal  flared  portion  per- 
forate. Distally  the  margins  of  the  processes  are  usually  entire,  but  may  bear  one 
or  two  small  spines.  A  complete  specimen  possesses  18  processes,  reflecting  a 
tabulation  characteristic  of  this  genus.  The  processes  are  all  of  equal  size  except 
for  the  first  postcingular  (i"')  and  the  posterior  intercalary  (ip)  which  are  often 
reduced. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  38  (46-3) 
57  /z,  maximum  length  of  processes  18  (34-1)  43  /*.  Number  of  specimens  measured,  8. 


148  CENOMANIAN  NON-CALCAREOUS   MICROPLANKTON,    i 

REMARKS.  The  presence  of  complex  perforate  processes  having  an  entire  distal 
margin  differentiate  this  species  from  all  other  similar  forms. 

OCCURRENCE.  0.  anthophorum  has  been  recorded  from  the  Upper  Jurassic — 
Lower  Cretaceous  (Aptian-Albian)  of  Australia  (Cookson  &  Eisenack  1958),  from 
the  Aptian  of  Germany  (Eisenack  1958)  and  from  the  Upper  Barremian — Albian  of 
Germany  (Alberti  1961).  This  species  has  been  recorded  from  one  sample,  Sas  1023, 
from  the  Albian  of  Saskatchewan  where  it  is  common. 

Oligosphaeridium  reniforme  (Tasch) 
(PI.  6,  fig.  i) 

1964     Hystrichosphaeridium  reniforme  Tasch  :  193,  pi.  2,  fig.  6. 
19666  Oligosphaeridium  reniforme  (Tasch)  Davey  &  Williams  :  77. 

DESCRIPTION.  The  shell  is  subspherical  to  ovoidal,  shell  wall  lightly  granular. 
The  processes  are  hollow,  tubiform,  widening  distally  into  a  broad,  flat-topped  funnel. 
The  distal  margin  of  the  funnel  bears  a  small  number  of  pointed  and  irregularly 
shaped  spines.  Distally  the  processes  sometimes  possess  large,  subcircular  perfora- 
tions. An  apical  archaeopyle  is  typically  developed. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  31  (42-4) 
49  fj,,  maximum  length  of  processes  20  (24-9)  30  p.  Number  of  specimens  measured,  7. 

REMARKS.  The  specimens  appear  to  be  very  similar  to  the  type  material  from 
the  Albian  of  Kansas  although  Tasch  did  not  describe  the  presence  of  distal  perfora- 
tions. 0.  reniforme  differs  from  0.  anthophorum  and  0.  perforatum  (Gocht  1959)  by 
the  presence  of  distal  spines.  It  differs  from  0.  pulcherrimum  Deflandre  &  Cookson 
(1955)  in  that  the  processes  possess  only  a  few  spines,  are  not  so  complexly  perforate 
and  distally  are  flat-topped. 

OCCURRENCE.  0.  reniforme  is  infrequent  in  all  the  Albian-Cenomanian  samples 
from  Saskatchewan.  It  has  not  been  recorded  elsewhere. 

Genus  LITOSPHAERIDIUM  Davey  &  Williams  1966 
Litosphaeridium  siphoniphorum  (Cookson  &  Eisenack) 

(PI.  6,  figs.  3,  4;  Fig.  15) 

1958     Hystrichosphaeridium  siphoniphorum  Cookson  &  Eisenack  :  44,  pi.  n,  figs.  8-10. 

19666  Litosphaeridium  siphoniphorum  (Cookson  &  Eisenack)   Davey  &  Williams  :  80,  pi.   7, 

figs.  7,  8;  text-figs.  16,  17.     (See  also  for  earlier  references). 
1967     Hystrichosphaeridium  siphoniphorum  Cookson  &  Eisenack;  Clarke  &  Verdier:  55,  pi.  n, 

figs,  i,  2. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  21  (34-0) 
47  /u,,  length  of  processes  4-25  //,.  Number  of  specimens  measured,  160. 

DESCRIPTION.  A  statistical  study  was  made  of  this  species  using  20-30  specimens 
from  each  of  six  samples  at  4O-foot  intervals  from  the  Fetcham  Mill  Borehole.  The 
purpose  of  this  study  was  to  see  if  the  variation  in  the  mean  size  of  L.  siphoniphorum 
was  directional,  and  if  there  was  a  significant  difference  in  this  measurement  for 


0) 

-Q 

E 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 


149 


40 


30 


B 


50 


60 


50 


60 


70 


40 


50 


60 


70 


40 


50 


60 


40 


50 


60 


40 


50 


60 


70 


Overall    diameter 

FIG.  15.  Overall  diameter — frequency  histograms  of  Litosphaeridium  siphoniphorum 
(Cookson  and  Eisenack)  at  six  horizons  from  Fetcham  Mill,  Surrey.  A.  Sample  FM  650, 
B.  Sample  FM  690,  C.  Sample  FM  730,  D.  Sample  FM  770,  E.  Sample  FM  810,  F.  Sample 
FM  840. 


150 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 


successive  samples.  The  measurement  taken  was  the  overall  diameter.  The 
position  of  the  archaeopyle  is  always  obvious,  allowing  easy  specimen  orientation. 
Thus  to  make  all  measurements  strictly  comparable  the  overall  diameter  was  always 
taken  in  the  plane  of  the  archaeopyle. 

Histograms  (Fig.  15)  were  drawn  for  each  assemblage.  The  mean  overall  diameter 
for  each  assemblage  varied  for  each  horizon  but,  unfortunately,  the  variation  was 
not  directional  and,  therefore,  was  of  little  stratigraphic  value.  The  Student's 
t-test  was  performed  on  successive  pairs  of  assemblages  to  see  whether  or  not  they  were 
significantly  different  (Table  A).  A  probability  of  0-05  or  less  was  taken  as  being 
significant. 


FM650 
FM6go 

FM6go 
FM730 


FM  770 

FM  770 
FMSio 

FMSio 
FM84o 


(x=  50-0  ft) 
x  = 


(x=  56-6  fj) 
(x=48-3yM) 

(x  =  48-3/0 
(x=  48-10) 

(X  =   48-I  ft) 
(X=   48-1  ft) 

(x=  48-1/0 
(x=  58-0/0 


TABLE  A 
t  =  3-08 

t  =  3-16 
t  =  0-084 

t  =  o 
t=5'4 


(significant  difference) 
(significant  difference) 
(no  significant  difference) 
(no  significant  difference) 
(significant  difference) 


The  results  show  that  there  is  a  significant  difference  at  the  5  %  level  between 
some  of  the  assemblages  with  respect  to  this  character.  However,  all  the  specimens 
measured  were  apparently  morphologically  identical  and  differ  only  in  size.  Speci- 
mens from  one  sample  vary  considerably  in  size  but  were  probably  formed  by  one 
species  of  motile  dinoflagellate.  Thus  the  size  of  L.  siphoniphorum  appears  to  be 
quite  variable  and  should,  at  the  moment,  not  be  used  as  a  diagnostic  feature  for  the 
subdivision  of  this  species.  The  reason  for  the  means  in  successive  samples  to  be 
significantly  different  is  probably  because  of  palaeoecological  changes  in  the 
environment. 

REMARKS.  All  the  specimens  of  L.  siphoniphorum  examined  agree  fairly  closely 
with  the  type  material  from  Australia.  The  Surrey  specimens  appear  to  be  smaller, 
but  the  range  of  the  Australian  forms  was  not  given  so  no  true  size  comparison  can 
be  made. 

OCCURRENCE.  L.  siphoniphorum  is  rare  to  common  at  all  horizons  throughout  the 
Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  is  present  in  the  Albian 
sample  FM  866,  but  absent  from  the  Turonian  sample,  FM  520.  It  is  rare  to 
infrequent  in  the  lower  four  samples  from  Saskatchewan — samples  Sas  1084, 1023, 967 
and  890.  This  species  is  also  present  in  the  Upper  Woodbine  Formation  of  Texas. 
Thus  L.  siphoniphorum  has  a  wide  geographical  distribution  and  a  fairly  restricted 
range.  It  has  been  recorded  from  the  Albian  of  Australia,  Rumania,  Canada  and 
Britain  and  from  the  Cenomanian  of  Australia,  Canada  and  Britain. 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i  151 

Genus  POLYSPHAERIDIUM  Davey  &  Williams  1966 
Polysphaeridium  pumilum  Davey  &  Williams 

?I955     Hystrichosphaeridium  recurvatum  White;  Deflandre  &  Cookson  :  269,  pi.  i,  fig.  12. 
19666  Polysphaeridium  pumilum  Davey  &  Williams  :  93,  pi.  7,  figs.  3,  4. 

DIMENSIONS.  Range  of  observed  specimens :  overall  diameter  30-40  //,,  diameter 
of  central  body  17-25/1,,  length  of  processes  7-10/1,,  width  of  processes  1-1-5 /A, 
number  of  processes  38-44.  Number  of  specimens  measured,  3. 

OCCURRENCE.  Only  three  specimens  of  P.  pumilum  have  been  observed,  one 
from  sample  FM  750  and  two  from  sample  FM  770. 


Polysphaeridium  laminaspinosum  Davey  &  Williams 
(PL  4,  figs.  10,  ii) 

19666  Polysphaeridium  laminaspinosum  Davey  &  Williams  :  94,  pi.  8,  fig.  8. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  20  (26-8) 
29  [L,  maximum  length  of  processes  9  (13-7)  17  p.  Number  of  specimens  measured,  8. 

OCCURRENCE.  P.  laminaspinosum  is  rare  to  very  rare,  occurring  spasmodically 
throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  has 
not  been  recorded  elsewhere. 


Genus  TANYOSPHAERIDIUM  Davey  &  Williams  1966 

Tanyosphaeridium  variecalamum  Davey  &  Williams 

(PI.  6,  figs.  2,  5) 

19666  Tanyosphaeridium  variecalamum  Davey  &  Williams  :  98,  pi.  6,  fig.  7;  text-fig.  20. 

DIMENSIONS.  Range  of  observed  specimens :  length  of  central  body  27  (32-3)  43  /*, 
width  14  (20-1)  24  n,  maximum  length  of  processes  11  (15-0)  24/4.  Number  of 
specimens  measured,  14. 

OCCURRENCE.  T.  variecalamum  is  a  rare  to  infrequent  species  at  most  horizons 
throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  is 
also  present  in  the  Albian  sample,  FM  886,  and  the  Turonian  sample,  FM  520, 
both  from  Fetcham  Mill.  One  specimen  was  located  in  the  Saskatchewan  material, 
from  the  Albian  sample  Sas  1084. 

Genus  CALLAIOSPHAERIDIUM  Davey  &  Williams  1966 

REMARKS.     Hexasphaera  Clarke  &  Verdier  (1967;  42)  is  a  junior  synonym  of 
Callaiosphaeridium . 
GEOL.  17,  3  IX 


152  CENOMANIAN   NON-CALCAREOUS   MICROPLAN  KTON,    i 

Callaiosphaeridium  asymmetricum  (Deflandre  &  Courteville) 

(PI.  6,  fig.  6) 

1939     Hystrichosphaeridium  asymmetricum  Deflandre  &  Courteville  :  100,  pi.  4,  figs,  i,  2. 
19666  Callaiosphaeridium  asymmetricum   (Deflandre  &  Courteville)   Davey  &  Williams  :  104, 

pi.  8,  figs.  9,  10;  pi.  9,  fig.  2. 
1967     Hexasphaera  asymmetrica  (Deflandre  &  Courtville)  Clarke  &  Verdier:  43,  pi.  7,  figs.  1-3 

text-fig.  17. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  37  (45-8) 
58  ju,  maximum  length  of  cingular  processes  10  (25-0)  32/1,.  Number  of  specimens 
measured,  6. 

REMARKS.  The  author  disagrees  with  the  description  of  Clarke  &  Verdier  (1967) 
in  that  the  archaeopyle  is  epitractal,  not  apical,  and  that  the  antapical  plate  is 
five-sided,  not  six-sided,  bearing  a  process  at  each  corner.  The  position  of  the  three 
sutural  crests  separating  the  large  plates  has,  however,  been  verified. 

OCCURRENCE.  C.  asymmetricum  is  rare  to  infrequent  at  most  horizons  throughout 
the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  has  been  recorded 
in  the  Albian  sample  FM  886  and  the  Turonian  sample  FM  520,  both  from  Fetcham 
Mill,  but  is  absent  from  the  North  American  material. 

Genus   CLEISTOSPHAERIDIUM  Davey,   Downie,   Sarjeant   &  Williams   1966 
Cleistosphaeridium  heteracanthum  (Deflandre  &  Cookson) 

(PI.  7,  fig-  8) 

1955     Hystrichosphaeridium  heteracanthum  Deflandre  &  Cookson  :  276,  pi.  2,  figs.  5,  6;  text-figs. 

40,  41. 
1966     Cleistosphaeridium  heteracanthum  (Deflandre  &  Cookson)   Davey,   Downie,   Sarjeant  & 

Williams  :  168,  pi.  2,  figs.  6,  7  (See  also  for  earlier  references). 

DIMENSIONS.  Range  of  observed  specimens:  shell  diameter  42  (53-2)  63^, 
maximum  length  of  processes  9  (13-8)  17  ju.  Number  of  specimens  measured,  9. 

OCCURRENCE.  C.  heteracanthum  is  a  rare  species  restricted  to  the  Upper  Ceno- 
manian— samples  FM  690,  670  and  650  from  Fetcham  Mill,  samples  CB  19  and  21 
from  Compton  Bay  and  sample  E  153  from  Escalles.  It  was  not  recorded  from  North 
America. 

Cleistosphaeridium  multifurcatum  (Deflandre) 
(PI.  8,  figs.  7,  10) 

1937  Hystrichosphaeridium  multifurcatum  Deflandre  :  76,  pi.  16,  figs.  1-3. 

1939  Hystrichosphaeridium   multifurcatum   Deflandre;    Deflandre   &   Courteville  :  102,    pi.    3, 

fig.  2. 

1952  Hystrichosphaeridium  multifurcatum  Deflandre;  W.  Wetzel  :  400,  text-fig.  16. 

1955  Hystrichosphaeridium  multifurcatum  Deflandre;  Valensi  :  588,  pi.  i,  fig.  21;  pi.  5,  fig.  5. 

1960  Baltisphaeridium  multifurcatum  (Deflandre)  Klement  :  59. 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  153 

1963     Hystrichosphaeridium  multifurcatum  Deflandre;  Gorka  :  66,  pi.  9,  figs.  4-6;  text-fig.  8, 
fig.  i. 

1966  Cleistosphaeridium  multifurcatum  (Deflandre)  Davey,  Downie,  Sarjeant  &  Williams:  170. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  39  (55-0) 
68  fj,,  maximum  length  of  processes  8  (ii'2)  13  //..  Number  of  specimens  measured,  8. 

OCCURRENCE.  C.  multifurcatum  is  rare  at  most  horizons  in  the  Cenomanian  of 
Fetcham  Mill,  Compton  Bay  and  Escalles.  In  a  few  Upper  Cenomanian  samples 
(FM  690,  670  and  650;  CB  19,  21  and  E  153)  it  is  completely  absent.  In  these 
samples  it  seems  to  be  replaced  by  C.  heteracanthum.  C.  multifurcatum  was  not 
observed  in  the  North  American  samples. 

Cleistosphaeridium  armatum  (Deflandre)  comb.  nov. 
(PL  8,  figs,  i,  2,  12) 

1937     Hystrichosphaeridium  armatum  Deflandre  :  76,  pi.  16,  figs.  6,  7. 
1947     Hystrichosphaeridium  armatum  Deflandre;  Deflandre:  fig.  i,  No.  10. 
1952^  Hystrichosphaeridium  armatum  Deflandre;  Deflandre:  fig.  14. 
1963     Baltisphaeridium  armatum  (Deflandre)  Downie  &  Sarjeant  :  91. 

1967  Baltisphaeridium  armatum  (Deflandre)  Clark  &  Verdier:  71,  pi.  13,  fig.  3. 

EMENDED  DIAGNOSIS.  Shell  subspherical ;  shell  wall  of  moderate  thickness,  densely 
granular.  Processes  numerous,  fairly  broad,  rigid,  tapering  gradually  distally. 
Proximally  processes  possess  longitudinal  basal  striations;  distally  simple  or  giving 
rise  to  variable  number  of  small  spines.  Apical  archaeopyle  occasionally  developed. 

HOLOTYPE.  Slide  AJ.  54,  Laboratoire  de  Micropaleontologie,  Ecole  Practique  des 
Hautes  Etudes,  Paris.  (Figured  by  Deflandre  1937,  pi.  16,  fig.  6).  Upper  Cretaceous 
flint  from  the  Paris  Basin. 

DIMENSIONS.  Holotype:  shell  length  20/11,  width  18-20  /z,  length  of  processes 
10-15  p.  Range  of  Cenomanian  specimens:  diameter  of  central  body  19  (30-8)  42  /u,, 
maximum  length  of  processes  5  (9-6)  16  //,.  Number  of  specimens  measured,  33. 

DESCRIPTION.  The  granules  are  elongate  (c.  0-1-0-5  P  in  height)  so  giving  the 
shell  surface  the  appearance  of  possessing  a  matting  of  short  hairs.  This  ornamen- 
tation, commented  on  by  Deflandre  in  the  original  description  of  this  species,  was 
verified  by  the  present  author  when  examining  the  type  material  in  Paris. 

The  processes  are  all  of  a  similar  length  on  any  one  specimen  and  may  vary  in 
width  from  i  to  3  JM.  They  are  hollow,  always  closed  distally  terminating  either 
simply  (the  extremity  may  be  recurved)  or  more  commonly  the  distal  one  quarter 
of  the  processes  bear  a  small  number  of  stiff  spines  (pi.  8,  fig.  2).  The  opening,  when 
observable,  possesses  an  angular  margin  characteristic  of  an  apical  archaeopyle. 

REMARKS.  The  specimens  studied  strongly  resemble  C.  armatum  as  described 
by  Deflandre  from  Upper  Cretaceous  flints,  one  of  which  was  of  Cenomanian  age. 
The  appearance  of  the  shell  surface  and  form  of  the  processes  make  this  an  easily 
recognizable  species. 

OCCURRENCE.  C.  armatum  is  common  at  all  horizons  throughout  the  Cenomanian 
of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  is  also  present  in  the  Upper  Wood- 


154  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 

bine  and  Lower  Eagle  Ford  formations  of  Texas,  but  has  not  been  observed  in  the 
Saskatchewan  samples. 

Cleistosphaeridium  polypes  (Cookson  &  Eisenack)  comb.  nov. 
(PI.  6,  figs.  7,  8) 

19626  Hystrichosphaeridium  recurvatum  subsp.  polypes  Cookson  &  Eisenack  :  491,  pi.  4,  figs. 
11-13. 

DESCRIPTION.  The  shell  is  spherical  to  subspherical  and  bears  numerous  slender 
processes.  The  shell  wall  is  thin  and  may  be  smooth  or  slightly  granular.  The 
processes  are  narrow,  usually  less  than  i  p  in  width  for  most  of  their  length,  broaden- 
ing slightly  proximally.  They  are  closed  distally  and  terminate  in  a  number  of 
short,  fine  spines.  The  latter  are  not  so  numerous  in  the  English  and  French  speci- 
mens as  in  those  from  Australia  and  North  America  where  the  distal  spines  are 
numerous.  A  fairly  large  archaeopyle,  probably  apical,  with  an  angular  margin 
is  often  present. 

DIMENSIONS.  Range  of  observed  specimens :  shell  diameter  31  (38-4)  47  //,,  maxi- 
mum length  of  processes  9  (13-4)  18  /A.  Number  of  specimens  measured,  20. 

REMARKS.  C.  polypes  was  originally  described  by  Cookson  &  Eisenack  from  the 
Aptian-Cenomanian  of  Australia  and  was  considered  to  be  a  subspecies  of 
Hystrichosphaeridium  recurvatum  (White) .  The  resemblance  between  the  two  forms  is 
very  slight  and  they  are  not  considered  to  be  closely  related.  C.  polypes  is  placed 
in  this  genus  because  of  the  presence  of  numerous  closed  processes  and  the  probable 
apical  location  of  the  archaeopyle. 

OCCURRENCE.  C.  polypes  is  a  very  rare  species  restricted  to  the  Middle  and  Upper 
Cenomanian  of  Fetcham  Mill  (samples  FM  750,  730),  Compton  Bay  (samples  CB  9, 
17,  21)  and  Escalles  (samples  E  189,  177,  165).  It  is  common  in  the  Upper  Wood- 
bine formation  of  Texas,  and  is  present  in  one  sample  from  Saskatchewan  (sample 
Sas  835). 

Cleistosphaeridium  polypes  var.  clavulum  nov. 
(PI.  6,  figs.  9,  10) 

1964     Hystrichosphaeridium  recurvatum  subsp.  polypes  Cookson  &  Eisenack;  Cookson  &  Hughes  : 
47,  pi.  9,  fig.  14. 

DERIVATION  OF  NAME.  Latin,  clavulus,  small  nail — with  reference  to  the  pin-like 
shape  of  the  processes. 

DIAGNOSIS.  A  variety  of  C.  polypes  possessing  fine,  capitate  processes.  Processes 
terminating  with  fine  spines  are  extremely  rare. 

TYPE.  G.S.M.  slide  PF  3995(1).  Lower  Chalk,  H.M.  Geological  Survey  Borehole, 
Fetcham  Mill,  Surrey  at  840  feet  depth.  Upper  Cretaceous  (Cenomanian). 

DIMENSIONS.  Type:  diameter  of  central  body  29  by  32 /x,  length  of  processes 
12-13 /A.  Range:  diameter  of  central  body  29  (31-5)  39 p,  maximum  length  of 
processes  13  (14-7)  15  p.  Numbers  of  specimens  measured,  10. 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON.    i  155 

REMARKS.  C.  polypes  var.  davulum  differs  from  C.  polypes  in  that  the  processes 
are  capitate,  the  terminal  bulge  being  flattened  and  resembling  the  head  of  a  pin. 
Two  specimens  have  been  observed  each  possessing  one  process  of  the  type  charac- 
teristic of  C.  polypes  thus  indicating  a  relationship  between  the  two  forms.  C. 
polypes  probably  evolved  from  this  variety,  spiny  processes  replacing  the  capitate 
ones. 

OCCURRENCE.  This  variety  has  only  been  recorded  from  the  lowermost  Cenoman- 
ian  samples  at  Fetcham  Mill  (sample  FM  840)  and  Compton  Bay  (sample  CB  i). 
It  was  previously  recorded  by  Cookson  &  Hughes  (1964)  from  the  Upper  Albian — 
basal  Cenomanian  of  England  and  so  appears  to  be  of  stratigraphic  importance  in 
England  for  indicating  the  base  of  this  stage.  It  does  not  occur  in  the  lowermost 
sample  from  Escalles. 

Cleistosphaeridium  huguonioti  (Valensi)  comb.  nov. 
(PI.  7,  fig.  10) 

1955     Hystrichosphaeridium  huguonioti  Valensi  :  38,  fig.  2a. 

ig6oa  Hystrichosphaeridium  ancoriferum  Cookson  &  Eisenack  :  8,  pi.  2,  fig.  n. 

1963  Hystrichosphaeridium  ancoriferum  Cookson  &  Eisenack;  Baltes  :  586,  pi.  6,  fig.  13. 

1964  Hystrichosphaeridium  ancoriferum  Cookson  &  Eisenack;  Cookson  &  Hughes  :  47,  pi.  9, 

fig.  7- 
1964     Chlamydophorella  nyei  Cookson  &  Eisenack;  Cookson  &  Hughes  :  54,  pi.  6,  fig.  12. 

1966  Cleistosphaeridium   ancoriferum    (Cookson    &    Eisenack)    Davey,    Downie,    Sarjeant    & 
Williams:  167,  pi.  9,  fig.  i. 

1967  Hystrichosphaeridium  huguonioti  Valensi;  Clarke  &  Verdier:  54,  pi.  n,  fig.  4,  5. 

DESCRIPTION.  The  shell  is  subspherical;  the  shell  wall  is  smooth  and  gives  rise  to 
numerous  bifurcating  processes  which  are  not  aligned  to  any  noticeable  extent. 
Most  of  the  specimens  possess  an  apical  archaeopyle,  the  shape  of  which  is  usually 
difficult  to  determine  because  of  distortion.  However,  detached  apical  regions  are 
common  and  are  6-sided.  The  processes  are  hollow,  the  central  cavity  often  being 
constricted  to  some  extent  along  its  length,  and  closed  distally  and  proximally. 
The  sides  of  the  processes  are  practically  parallel,  diverging  slightly  proximally  before 
joining  the  shell.  Distally  they  give  rise  to  two  slightly  recurved  spines.  Cookson 
&  Eisenack  comment  on  the  "  transparent  tips  "  of  the  processes.  The  extremities 
of  the  processes  are  in  fact  closed  by  a  thin,  transparent  membrane.  The  processes 
may  be  isolate  or  a  few  may  be  linked  together  distally  by  their  spines.  The  shell 
wall  rarely  forms  a  small  rounded  apical  bulge. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  20  (31-8) 
45  ju,  length  of  processes  up  to  8  p.  Number  of  specimens  measured,  30. 

REMARKS.  Cookson  &  Hughes  (1964)  described  C.  huguonioti  and  Chlamydo- 
phorella nyei  from  the  Upper  Albian  and  Lower  Cenomanian  of  England,  distinguish- 
ing the  latter,  with  difficulty,  by  the  presence  of  an  outer  membrane  and  an  apical 
bulge.  After  a  detailed  examination  of  numerous  Cenomanian  specimens  it  was 
concluded  that  Chlamydophorella  nyei  s.s.,  as  described  from  Australia,  does  not  occur  in 
the  English  and  French  Cenomanian  and  that  C.  nyei  as  described  by  Cookson  & 


156  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 

Hughes  belongs  in  Cleistosphaeridium  huguonioti.  C.  huguonioti  does  occasionally 
appear  to  possess  a  membrane  linking  some  of  the  processes  when  their  spines  are 
joined  distally,  and  an  apical  prominence  may  also  rarely  be  present.  Some  speci- 
mens possess  joined  processes  but  apparently  no  apical  prominence,  whereas  others 
possess  an  apical  prominence  but  no  joined  processes.  The  presence  of  distinctive 
bifurcate  processes  and  an  apical  prominence  indicates  that  C.  huguonioti  is  related 
to  Chlamydophorella,  the  outer  membrane  of  the  latter  being  reduced  to  a  small, 
transparent  membrane  at  the  distal  end  of  each  process. 

OCCURRENCE.  C.  huguonioti  is  common  throughout  the  Cenomanian  of  Fetcham 
Mill,  Compton  Bay  and  Escalles  (Fig.  22).  It  is  absent  from  the  uppermost  sample 
from  Escalles  (sample  E  153)  and  from  the  Turonian  sample  FM  520.  In  the  Upper 
Cenomanian  at  these  localities  C.  huguonioti  tends  to  be  replaced  by  its  variety, 
C.  huguonioti  var.  pertusum  nov.,  which  appears  to  become  more  abundant  as  C. 
huguonioti  declines.  It  is  present  in  the  Albian  sample  FM  886  and  is  common  in 
the  Upper  Woodbine  and  Lower  Eagle  Ford  formations  of  Texas,  but  was  not 
observed  in  the  Saskatchewan  material.  This  species  was  first  recorded  from  the 
Albian-Cenomanian  of  Australia  and  is  also  present  in  the  Albian-Cenomanian 
of  Rumania  (Balte§,  1963).  Thus  C.  huguonioti  appears  to  be  wide-spread 
geographically  and  restricted  to  the  Albian  and  Cenomanian. 

Cleistosphaeridium  huguonioti  var.  pertusum  nov. 
(PL  7,  figs.  6,  7,  9) 

DERIVATION  OF  NAME.  Latin,  pertusum,  perforated — with  reference  to  the  per- 
forate appearance  of  the  processes. 

DIAGNOSIS.  A  variety  of  C.  huguonioti  possessing  spherical  to  subspherical, 
smooth  walled  shell  bearing  numerous  processes.  Processes  broad-based,  tapering 
distally  and  terminating  with  two  small  recurved  spines.  Lumen  of  processes 
restricted  by  transverse  septa. 

HOLOTYPE.  G.S.M.  slide  PF  3040(2).  Lower  Chalk,  H.M.  Geological  Survey 
Borehole,  Fetcham  Mill,  Surrey  at  670  feet  depth.  Upper  Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype :  diameter  of  central  body  36  by  36  p,,  length  of  processes 
7-9  IJL,  maximum  width  of  processes  distally  2  /*.  Range :  diameter  of  central  body 
24  (34-1)  46  //,,  length  of  processes  6  (8-7)  II  p,  maximum  width  of  processes  distally 
i  (2-7)  4-5  p.  Number  of  specimens  measured,  24. 

DESCRIPTION.  The  processes  taper  distally  from  a  fairly  broad  base  (2-5-4^  in 
width)  to  a  narrow  neck  (c.  0-5  p  in  width)  before  bifurcating  to  give  two  short, 
recurved  spines.  The  processes  are  hollow  but  the  lumen  is  traversed  by  a  number 
of  small  septa  which  thus  subdivide  it,  giving  the  processes  a  "  holey  "  or  vacuolated 
appearance.  An  archaeopyle  is  only  rarely  observable. 

REMARKS.  This  variety  differs  from  C.  huguonioti  in  the  form  of  its  processes 
which  are  vacuolated,  tend  to  be  longer  and  bear  relatively  small  distal  spines. 
C.  huguonioti  found  in  the  same  samples  possesses  processes  which  are  considerably 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i  157 

wider  distally  (5-5-7-5  /i).     C.  huguonioti  var.  pertusum  appears  to  have  evolved  from 
C.  huguonioti  in  the  Middle  to  Upper  Cenomanian. 

OCCURRENCE.  C.  huguonioti  var.  pertusum  occurs  only  in  the  Upper  Cenomanian, 
where  it  is  common  at  all  horizons.  It  first  occurs  in  samples  FM  710  (Fetcham  Mill), 
CB  15  (Compton  Bay)  and  E  171  (Escalles).  At  these  horizons  it  is  occasionally 
difficult  to  distinguish  from  C.  huguonioti.  Like  C.  huguonioti,  this  variety  is  absent 
from  sample  E  153  (Escalles)  and  FM  520  (Fetcham  Mill,  Turonian).  It  has  not 
been  observed  in  the  samples  from  North  America. 


fCleistosphaeridium  flexuosum  Davey,  Downie,  Sarjeant  &  Williams 

(PI.  7,  figs.  4) 

1966  ?  Cleistosphaeridium  flexuosum  Davey,   Downie,   Sarjeant  &  Williams:  169,  pi.  2,  fig.  5. 

OCCURRENCE.     This  is  a  very  rare  species  occurring  at  most  horizons  throughout 
the  Cenomanian  of  Fetcham  Mill.     It  has  not  been  recorded  elsewhere. 


?  Cleistosphaeridium  parvum  sp.  nov. 
(PL  7,  figs,  ir,  12) 

DERIVATION  OF  NAME.  Latin,  parvus,  little — with  reference  to  the  small  size  of 
this  cyst. 

DIAGNOSIS.  Shell  ovoidal,  small;  shell  wall  smooth,  bearing  numerous  long,  fine 
spines.  Spines  may  be  aligned  along  upper  and  lower  boundaries  of  cingulum. 
Cingulum  strongly  laevo-rotatory.  Apical  archaeopyle  typically  developed. 

HOLOTYPE.  B.M.  (N.H.)  ¥.51981  (3).  Lower  Chalk,  Bureau  de  Recherches 
Geologique  et  Minieres  Borehole,  Escalles,  Pas  de  Calais  at  165  metres  depth.  Upper 
Cretaceous  (Cenomanian) . 

DIMENSIONS.  Holotype:  length  of  central  body  I2//,,  width  iiyu,,  length  of 
processes  6-n  p.  Range:  length  of  central  body  11-14 /x,  width  10-13 /z,  length  of 
processes  6-12  /*.  Number  of  specimens  measured,  5. 

DESCRIPTION.  The  spines  are  long,  very  fine,  terminate  distally  in  a  point  and 
widen  only  slightly  when  joining  the  shell.  The  cingulum  is  not  always  observable 
but  some  alignment  of  the  spines  parallel  to  the  archaeopyle  margin  is  usually  pres- 
ent. 

REMARKS.  ?C.  parvum  resembles  only  one  previously  described  species,  that  is 
Palaeostomocystis  echinulata  Deflandre  (1937)  from  the  Upper  Cretaceous  of  France. 
This  species  differs  in  that  there  are  fewer  spines  (c.  12),  but  is  similar  in  its  ovoidal 
shape,  apical  archaeopyle,  cingulum  and  small  size  (6-7  //.  long). 

The  presence  of  a  cingulum  has  not  previously  been  observed  in  members  of 
Cleistosphaeridium  and  might  later  be  used  as  a  character  in  generic  subdivision. 
However,  difficulty  of  observation  makes  it  of  dubious  value. 


158  CENOMANIAN    NON-CALCAREOUS   MICROPL ANKTON,    i 

OCCURRENCE.  ?C.  parvum  is  fairly  common  in  three  Upper  Cenomanian  samples 
from  Escalles — E  171,  165  and  159.  It  has  also  been  observed  in  a  single  Lower 
Cenomanian  sample  from  Compton  Bay  (CB  5).  In  all  other  samples  it  appears  to  be 
absent. 

fCleistosphaeridium  aciculare  sp.  nov. 
(PL  6,  figs,  n,  12) 

DERIVATION  OF  NAME.  Latin,  adcularis,  like  a  needle — with  reference  to  the 
acuminate  shape  of  the  processes. 

DIAGNOSIS.  Shell  spherical  to  subspherical ;  shell  wall  of  moderate  thickness, 
densely  granular.  Processes  numerous,  finely  to  broadly  acuminate,  slightly 
flexuous,  less  than  one-third  of  shell  diameter  in  length. 

HOLOTYPE.  B.M.  (N.H.)  slide  V.  51979  (3).  Second  White  Speckled  Shale, 
International  Yarbo  Borehole  No.  17,  Saskatchewan  at  835  feet  depth.  Upper 
Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype :  diameter  of  central  body  43  by  50  /A,  length  of  processes 
12-14 /*•  Range:  diameter  of  central  body  32  (43-0)  54/^1,  maximum  length  of 
processes  8  (13-9)  21  //..  Number  of  specimens  measured,  n. 

DESCRIPTION.  The  processes  may  be  finely  or  broadly  acuminate  but  on  each 
individual  their  width  is  constant.  On  individuals  bearing  fine  processes  these  are 
more  densely  packed  than  in  individuals  with  broad  processes.  All  intergradations 
exist  between  the  fine  and  the  broad  processed  forms.  The  processes  are  always 
pointed  distally  and  occasionally  bear  small  subsidiary  spines  near  their  extremities. 
An  archaeopyle  has  never  been  observed. 

REMARKS.  ?C.  aciculare  is  only  tentatively  placed  in  this  genus,  for  although  it 
resembles  other  members  in  overall  appearance,  an  apical  archaeopyle  has  not  been 
observed.  The  numerous  acuminate  processes  and  the  densely  granular  shell  sur- 
face differentiate  ?C.  aciculare  from  most  previously  described  forms.  Most  similar 
seems  to  be  Exochosphaeridium  (Hystrichosphaeridiurri)  cf.  striolatum  (Deflandre)  as 
illustrated  by  G6rka  (1963,  pi.  10,  fig.  6)  from  the  Cenomanian  of  Poland. 

OCCURRENCE.  1C.  aciculare  is  common  in  the  following  samples  from  Saskat- 
chewan— Sas  1084,  1023  and  967  (all  Albian)  and  Sas  835  (Cenomanian).  It  has 
not  been  recorded  elsewhere. 

Genus  SURCULOSPHAERIDIUM  Davey,  Downie,  Sarjeant  &  Williams  1966 

Surculosphaeridium  longifurcatum  (Firtion) 

(PI.  8,  fig.  9) 

1952     Hystrichosphaeridium  longifurcatum  Firtion  :  157,  pi.  9,  fig.  i  ;  text-fig,  i,  H,  K,  L  and  M 
1963     Baltisphaeridium  longifurcatum  (Firtion)  Downie  &  Sarjeant:  91. 

1966     Surculosphaeridium  longifurcatum  (Firtion)  Davey,  Downie,  Sarjeant  &  Williams  :  163, 
pi.  8,  figs.  7,  ii ;  text-figs.  43,  44. 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i  159 

DIMENSIONS.  Range  of  observed  forms:  lateral  view — diameter  of  central  body 
30  (38-3)  47 /LI;  apical  view — diameter  of  central  body  36  (42-2)  50/1.,  maximum 
length  of  processes  14  (22-2)  29  /*.  Mean  diameter  of  archaeopyle,  20  //,.  Number  of 
specimens  measured,  24. 

OCCURRENCE.  This  is  a  rare  species  at  all  horizons  throughout  the  Cenomanian 
of  Fetcham  Mill,  Compton  Bay  and  Escalles.  The  only  exception  is  sample  FM  730 
from  Fetcham  Mill,  where  this  species  is  very  common,  composing  22-5  %  of  the  total 
microplankton  present.  The  reason  for  this  unusual  abundance  is  unknown.  5. 
longifurcatum  is  also  present  in  the  Lower  Eagle  Ford  formation  of  Texas,  but  was 
not  observed  in  the  Saskatchewan  samples.  It  has  not  been  observed  in  the  Albian 
sample  (FM  886)  or  the  Turonian  sample  (FM  520)  from  Fetcham  Mill,  and  thus 
appears  to  be  characteristically,  Cenomanian. 

Genus  HYSTRICHOKOLPOMA  Klumpp  emend.  Williams  &  Downie  1966 

Hystrichokolpoma  ferox  (Deflandre) 

(PL  9,  figs.  5-7) 

1937     Hystrichosphaeridium  ferox  Deflandre  :  72,  pi.  14,  figs.  3,  4. 
ig66a  Hystrichokolpoma  ferox  (Deflandre)  Williams  &  Downie  :  181. 

1967     Baltisphaeridium  ferox  (Deflandre)  Clarke  &  Verdier:  73,  pi.  15,  fig.  4  (see  also  for  earlier 
references). 

EMENDED  DIAGNOSIS.  Shell  subspherical,  densely  granular  or  reticulate.  Pro- 
cesses thin  walled,  granular,  often  striated,  of  three  kinds :  (i)  6  large  precingular  and 
4  large  postcingular,  possessing  wide  bases  and  tapering  distally  giving  rise  to  2  or 
more  tubules,  typically  open;  (ii)  a  single  long  tubular  antapical  process  and  (iii) 
cingular  and  sulcal  processes,  of  moderate  length,  slender  and  tubular,  only  joining 
proximally  if  at  all.  Archaeopyle  apical. 

HOLOTYPE.  Slide  AH  72,  Laboratoire  de  Micropaleontologie,  Ecole  Practique 
des  Hautes  Etudes,  Paris.  (Figured  by  Deflandre  1937^,  pi.  14,  fig.  3).  Senonian 
flint  from  the  Paris  Basin. 

DIMENSIONS.  Holotype:  length  of  shell  46  p,  width  36/4,  overall  length  78^, 
length  of  processes  15-17  //..  Range  of  Cenomanian  specimens :  diameter  of  central 
body  39  (46-1)  56  /z,  maximum  length  of  processes  27  (30-2)  36/11.  Number  of 
specimens  measured,  13. 

DESCRIPTION.  H.  ferox  appears  to  be  a  fairly  variable  species.  In  some  speci- 
mens the  tabulation  is  not  clearly  indicated  by  the  processes  which  tend  to  be  smaller 
and  may  even  be  closed  distally.  However,  the  processes  usually  are  well  developed, 
their  bases  covering  an  area  of  the  shell  surface  which  is  often  slightly  raised  and 
of  the  same  shape  as  a  thecal  plate.  The  precingular  and  postcingular  processes  are 
largest  and  give  rise  distally  to  as  many  as  10  tubules.  Each  cingular  process 
divides  proximally  into  2  or  3  long  slender  tubules  aligned  along  the  cingulum.  The 
sulcal  processes  may  proximally  divide  into  two  tubules  or  may  consist  of  a  single 
tubule  which  is  sometimes  reduced  and  closed  distally.  The  antapical  process  is 
long,  tubular,  open  distally,  and  terminates  with  a  smooth  or  serrate  margin. 


160  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 

REMARKS.  The  Cenomanian  specimens  closely  resemble  the  holotype  of  H .  ferox, 
which  was  examined  in  Paris  by  the  author,  by  kind  permission  of  Professor 
Deflandre.  The  processes  of  the  holotype  are  of  the  same  form  as,  and  similarly 
positioned  to,  those  of  the  Cenomanian  forms.  The  antapical  process  of  the  holotype, 
not  shown  in  Deflandre's  illustration  (1937,  pi.  14,  fig.  3),  is  long  and  tubular. 

OCCURRENCE.  H .  ferox  is  rare  to  very  rare  at  most  horizons  throughout  the  Ceno- 
manian of  Fetcham  Mill  and  Escalles,  but  has  not  been  observed  at  Compton  Bay 
or  in  the  North  American  samples.  It  is  also  present  in  the  Turonian  sample 
(sample  FM  520)  from  Fetcham  Mill.  Hence  the  stratigraphic  range  is  from  Aptian 
(Eisenack  1958)  to  Upper  Cretaceous,  probably  Senonian  (Deflandre  1937). 


Genus  PROLIXOSPHAERIDIUM  Davey,  Downie,  Sarjeant  &  Williams  1966 

DIAGNOSIS.  Shell  elongate  ovoidal  to  ellipsoidal,  one  pole  (apical)  typically  lost 
in  archaeopyle  formation.  Opposite  pole  occupied  by  one  or  two  antapical  processes. 
Remaining  processes  arranged  in  distinct  rows,  encircling  shell  and  slightly  offset 
at  a  position  corresponding  to  sulcus.  Number  of  processes  exceeds  30.  Processes 
closed  proximally,  typically  but  not  constantly  closed  distally;  their  distal  termina- 
tions simple,  faring  in  varied  fashion,  or  briefly  furcate.  Shell  surface  sometimes 
bears  cover  of  coarse  granules  or  very  short,  simple  spinelets. 

REMARKS.  The  diagnosis  has  been  changed  slightly:  '  typically  but  not  constantly 
closed  distally  '  being  inserted  in  place  of  '  closed  or  open  distally ',  with  respect  to  the 
processes.  The  processes  of  Prolixosphaeridium  usually  taper  distally  and  are 
closed.  Specimens  with  open  tubular  processes  belong  to  Tanyosphaeridium  Davey 
&  Williams  (19666). 

Prolixosphaeridium  conulum  sp.  nov. 
(PI.  8,  figs.  5,  6) 

DERIVATION  OF  NAME.  Latin,  conulus,  cone — with  reference  to  the  rather  conical 
shape  of  the  processes. 

DIAGNOSIS.  Shell  elongate  ovoidal;  shell  wall  densely  granular  and  bearing 
moderate  number  of  processes.  Processes  acuminate  to  subconical,  smooth  walled, 
typically  rigid,  pointed  distally.  Processes  tend  to  be  aligned  in  circular  manner 
around  shell.  Archaeopyle  apical. 

HOLOTYPE.  B.M.  (N.H.)  ¥.51981  (5).  Lower  Chalk,  Bureau  de  Recherches  Geo- 
logiques  et  Minieres  Borehole,  Escalles,  Pas  de  Calais,  at  165  metres  depth.  Upper 
Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype :  shell  length  47  //,,  width  27  /u,  length  of  processes  11-16  /*. 
Range :  shell  length  38  (43-8)  50  \i,  width  20  (25-9)  29  //,,  maximum  length  of  processes 
ii  (15-1)  i8/i.  Number  of  specimens  measured,  8. 

DESCRIPTION.  The  length  of  the  shell  is  slightly  less  than  twice  the  width.  The 
surface  granules  are  relatively  large  (c.  0-2-0-4 /j,  in  width  and  height);  they  are 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  161 

equidistant  from  one  another.  The  processes  (45-60  in  number)  have  broad  bases 
(up  to  6  fj,)  and  taper  distally  to  terminate  in  a  point.  They  are  approximately 
half  the  shell  width  in  length,  hollow  and  typically  rigid,  only  occasionally  being 
bent  near  the  distal  end.  In  the  central  region  of  the  shell  the  processes  are  aligned, 
the  rows  encircling  the  shell.  However,  in  one  longitudinal  portion  in  this  region  the 
processes  are  generally  smaller  and  haphazard  in  arrangement.  This  region  probably 
corresponds  to  the  sulcus. 

REMARKS.  The  distinctive  shape,  number  and  size  of  the  processes  distinguish 
P.  conulum.  A  rather  similar  but  longer  form  was  illustrated  by  Cookson  &  Eisenack 
(1958,  pi.  8,  fig.  IT )  as  Hystrichosphaeridium  parvispinum  Deflandre.  This  specimen 
comes  from  the  Aptian  of  Australia  and  is  said  to  grade  into  forms  possessing  more 
numerous  processes  similar  to  P.  conulum.  Deflandre's  species  was  placed  in 
Prolixosphaeridium  by  Davey,  Downie,  Sarjeant  &  Williams  (1966). 

Two  other  species  are  rather  similar.  The  holotype  of  P.  granulosum  (Deflandre) 
measures  18  by  33  /*  and  possesses  20-30  long  processes,  the  latter  being  approxi- 
mately equal  to  the  shell  width.  The  number  of  processes  and  the  relative  lengths 
of  the  processes,  therefore,  differentiate  this  species  from  P.  conulum.  P.  granulosum 
as  described  by  Valensi  (1955)  from  the  Upper  Cretaceous,  and  Sarjeant  (1962)  from 
the  Upper  Jurassic,  are  more  similar  to  P.  conulum  but  do  not  possess  conical 
processes. 

P.  mixtispinosum  (Klement)  differs  from  P.  conulum  by  possessing  two  kinds  of 
processes — (i)  approximately  50  processes  of  moderate  length  and  (ii)  numerous  fine 
hairs  covering  the  shell  surface. 

OCCURRENCE.  P.  conulum  has  only  been  observed  at  certain  horizons  in  the 
Upper  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles — in  samples  FM  690, 
CB  19,  E  165,  E  159  and  E  153. 


Genus  CORONIFERA  Cookson  &  Eisenack  emend. 

EMENDED  DIAGNOSIS.  Shell  subspherical  to  ovoidal,  bearing  numerous  simple  or 
bifurcating  processes.  Processes  solid  or  hollow,  closed  distally,  and  joined  proxi- 
mally  by  low  crests  or  membranes.  Apical  process  distinctive,  simple  or  branched. 
Antapical  process  large,  tubular,  often  terminating  with  denticulate  margin.  Archae- 
opyle  apical. 

TYPE  SPECIES.  Coronifera  oceanica  Cookson  &  Eisenack  1958.  Lower  Cretaceous 
(Albian) ;  Australia. 

REMARKS.  The  diagnosis  has  been  emended  to  include  the  presence  of  an  apical 
archaeopyle  and  low  crests  joining  the  processes,  and  the  positions  of  the  two 
distinctive  processes. 

Coronifera  differs  from  Diphyes  Cookson  (1965)  in  that  the  processes  are  never 
tubular  and  open  distally,  and  by  the  presence  of  a  reticulum  joining  the  basal 
portion  of  the  processes.  However,  both  genera  possess  a  large  tubular  antapical 
process  and  an  apical  archaeopyle,  and  are  probably  closely  related. 


162  CENOMANIAN   NON -C  ALC  AREOU  S   MICROPL  ANKTON,    i 

Coronifera  oceanica  Cookson  &  Eisenack 
(PI.  8,  figs.  8,  ii) 

1958  Coronifera  oceanica  Cookson  &  Eisenack  :  45,  pi.  12,  figs.  5,  6. 

1958  Coronifera  oceanica  Cookson  &  Eisenack;  Eisenack  :  407,  pi.  25,  fig.  i. 

1964  Coronifera  oceanica  Cookson  &  Eisenack;  Cookson  &  Hughes  :  56,  pi.  9,  figs.  8,  9. 

1967  Coronifera  oceanica  Cookson  &  Eisenack;  Clarke  &  Verdier:  77,  pi.  17,  fig.  7. 

DESCRIPTION .  The  shell  is  subspherical  to  ovoidal,  thin-walled,  and  bears  numerous 
processes  of  length  between  one-quarter  and  one-third  of  the  shell  diameter. 
There  is,  occasionally,  a  slight  apical  prominence  beneath  the  apical  process.  Fine 
fibres  radiate  from  the  bases  of  the  processes  over  the  shell  surface  reminiscent  of 
those  on  the  shell  surface  of  Exochosphaeridium  striolatum  (Deflandre).  The  pro- 
cesses are  weak,  fairly  flexuous  and  commonly  joined  to  each  other  by  a  network  of 
low  crests  or  fine  membranes.  The  latter  may  be  proximal  or  may  extend  along  the 
entire  length  of  the  processes.  Distally  the  processes  are  closed  and  may  be  simple, 
bifurcate  or  trifurcate.  A  large  tubular  process  is  present  at  the  antapex;  it  is  open 
distally  and  terminates  with  a  denticulate  margin.  At  the  apex,  when  attached, 
there  is  a  process  which  is  only  slightly  larger  than  the  typical  processes  but  is 
usually  branched  and,  therefore,  distinctive.  The  large  archaeoplye,  developed  in 
the  majority  of  specimens,  has  an  angular  margin  and  forms  opposite  the  antapical 
process. 

A  number  of  specimens  of  C.  oceanica  were  observed  in  the  Albian  sample  from 
Fetcham  Mill  (FM  886).  They  resemble  the  specimens  of  Eisenack  (1958)  from  the 
Aptian  of  Germany  in  that  the  processes  are  fewer,  more  solid  and  are  usually  simple. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  31  (40) 
54  /x,  maximum  length  of  processes  11  (15-3)  22  /z.  Number  of  specimens  measured,  15. 

REMARKS.  C.  oceanica  has  previously  been  recorded  from  the  Upper  Aptian  of 
Germany  (Eisenack,  1958),  Albian  of  Australia  (Cookson  &  Eisenack,  1958)  and 
basal  Cenomanian  of  England  (Cookson  &  Hughes,  1964).  The  surface  reticulation 
was  not  described  in  the  Australian  type  material  but  appears  to  be  present  on  the 
photographed  specimens.  It  was  first  commented  on  by  Cookson  &  Hughes. 

OCCURRENCE.  In  addition  to  the  German  and  Australian  records,  C.  oceanica  is 
infrequent  to  common  at  all  horizons  throughout  the  Cenomanian  of  Fetcham  Mill, 
Compton  Bay  and  Escalles,  and  is  also  common  in  the  Albian  sample  (FM  886)  from 
Fetcham  Mill.  It  was  not  present  in  the  Turonian  sample  (FM  520).  One  specimen 
was  observed  in  the  North  American  material,  in  sample  Sas  1084  (Albian)  from 
Saskatchewan.  Thus  the  known  stratigraphic  range  is  from  Upper  Aptian  to 
Cenomanian. 

Cyst-Family  EXOCHOSPHAERIDIACEAE  Sarjeant  &  Downie  1966 
Genus  EXOCHOSPHAERIDIUM  Davey,  Downie,  Sarjeant  &  Williams  1966 

REMARKS.  Exochosphaeridium  differs  from  Trichodinium  Eisenack  &  Cookson 
(1960)  in  that  the  latter  possesses  a  well  developed  cingulum. 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i  163 

Exochosphaeridium  phragmites  Davey,  Downie,  Sarjeant  &  Williams 

(PI.  7>  %.  5) 

1966     Exochosphaeridium  phragmites  Davey,  Downie,   Sarjeant  &  Williams:   165,  pi.  2,  figs. 
8-10. 

DIMENSIONS.  Range  of  observed  specimens:  maximum  diameter  of  central 
body  41  (51-9)  67  IJL,  minimum  diameter  of  central  body  32  (46-7)  57  //,,  maximum 
length  of  processes  10  (18-5)  40  p,.  Number  of  specimens  measured,  18. 

REMARKS.  Superficially  E.  phragmites  resembles  E.  striolatum  (Deflandre)  which, 
however,  has  a  definitely  striated  periphragm.  Trichodinium  paucispinum  Eisenack 
&  Cookson  (1960)  is  also  similar  but  has  fewer  processes  and  a  well  developed 
cingulum. 

OCCURRENCE.  E.  phragmites  is  rare  at  most  horizons  throughout  the  Cenomanian 
of  Fetcham  Mill,  Compton  Bay  and  Escalles,  and  is  recorded  from  the  Albian 
sample,  FM  886.  It  has  not  been  recorded  in  the  North  American  material. 

Exochosphaeridium  pseudohystrichodinium  (Deflandre) 
(PI.  ii,  figs.  4,  5) 

1937     Hystrichosphaeridium  pseudohystrichodinium  Deflandre  :  73,  pi.  15,  figs.  3,  4. 

1966  ? Exochosphaeridium  pseudohystrichodinium    (Deflandre) ;    Davey,    Downie,    Sarjeant    & 
Williams  :  166. 

1967  Baltisphaeridium  pseudohystrichodinium  (Deflandre);  Clarke  &  Verdier:  75,  pi.  15,  fig.  7. 
(see  also  for  earlier  references). 

EMENDED  DIAGNOSIS.  Shell  spherical  to  ovoidal;  shell  wall  thick,  with  pitted 
surface.  Processes  numerous,  slightly  fibrous,  occasionally  bifurcating  medially, 
slender,  broadening  slightly  proximally ;  distally  truncated  or  terminated  with  small 
bifurcation.  Cingular  processes  rarely  aligned.  Apical  process  sometimes  branched 
and  slightly  larger  than  normal.  Archaeopyle  precingular,  formed  by  loss  of  one 
or  two  plate  areas. 

HOLOTYPE.  Slide  AH.  55,  Laboratoire  de  Micropaleontologie,  Ecole  Practique 
des  Hautes  Etudes,  Paris.  (Figured  by  Deflandre  1937,  pi.  15,  fig.  3).  Upper 
Cretaceous  flint  from  the  Paris  Basin. 

DIMENSIONS.  Range  of  type  material:  shell  length  49  to  54/11,  shell  width  38  to 
45 /i,  overall  length  80-90 ju,.  Range  of  observed  specimens:  diameter  of  central 
body  35  (43-8)  54  p,  maximum  length  of  processes  13  (16-8)  21 /u,.  Number  of 
specimens  measured,  12. 

DESCRIPTION.  The  shell  wall  is  moderately  thick  (c.  1-5  ^),  pitted  (never  striated), 
and  bears  a  large  number  of  broad-based  processes.  All  the  processes  appear  to  be 
basically  the  same  except  for  the  apical  process  which  is  typically  branched  and  is 
usually  larger  than  normal. 

The  author  was  permitted,  through  the  courtesy  of  Professor  Deflandre,  to  make  a 
detailed  examination  of  the  holotype  and  paratype.  In  the  holotype  the  cingular 
processes  were  arranged  in  a  definite  circular  manner  around  the  shell.  Such  an 


164  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

alignment  was  not  observed  with  certainty  in  any  of  the  Cenomanian  specimens 
studied.  The  archaeopyle,  present  in  the  paratype,  is  precingular  and  usually 
formed  by  the  loss  of  two  plate  areas. 

REMARKS.  The  Cenomanian  specimens  differ  from  the  holotype  only  in  that 
aligned  circular  processes  were  not  observed.  This  may  be  due  to  unfavourable 
preservation  or  orientation  of  the  specimens.  The  diagnosis  of  E.  pseudohystricho- 
dinium  has  been  emended  to  include  a  description  of  the  apical  process  and  the 
archaeopyle. 

The  overall  form  of  E.  pseudohystrichodinium  resembles  species  included  in  the 
"  hirsutum  "  group,  but  the  typical  fibrous  shell  periphragm  is  absent.  The  forma- 
tion of  the  archaeopyle  by  the  loss  of  two  precingular  plates  has  also  been  observed 
in  E.  striolatum  var.  truncatum  nov.,  indicating  a  relationship. 

OCCURRENCE.  This  species  is  rather  restricted,  being  common  in  samples  FM  690, 
E  165  and  E  159  and  rare  in  samples  FM  750,  E  207  and  FM  520  (Turonian).  It 
appears,  therefore,  to  be  most  common  in  the  Upper  Cenomanian.  It  was  not 
recorded  in  the  samples  from  North  America.  The  recorded  stratigraphic  range 
is  from  Cenomanian  to  Eocene  (Pastiels  1948). 


Exochosphaeridium   striolatum    (Deflandre)   comb.    nov. 
1937     Hystrichosphaeridium  striolatum  Deflandre  :  72,  pi.  15,  figs,  i,  2. 

DIAGNOSIS.  Shell  subspherical  to  ovoidal.  Processes  numerous,  variable, 
fibrous,  often  bifurcate  medially  and  sometimes  terminating  with  small  fork.  Pro- 
cesses distally  may  be  pointed  or  blunted.  Fibres  pass  down  length  of  processes 
onto  shell  surface  and  there  join  with  similar  fibres  from  adjacent  processes.  Apical 
process  and  precingular  archaeopyle  may  be  present. 

HOLOTYPE.  Slide  AH.  89,  Laboratoire  de  Micropaleontologie,  Ecole  Practique 
des  Hautes  Etudes,  Paris.  (Figured  by  Deflandre  1937,  pi.  15,  fig.  i).  Upper 
Cretaceous  flint  from  the  Paris  Basin. 

REMARKS.  Through  the  courtesy  of  Professor  Deflandre,  I  was  able  to  examine 
the  holotype  and  paratype  of  E.  striolatum.  On  the  holotype  one  process  appeared 
to  be  unusually  thick  and  may  have  been  apical  in  position.  Neither  holotype  nor 
paratype  was  observed  to  possess  an  archaeopyle  but  this  may  have  been  due  to  the 
fact  that  the  lower  surfaces  of  the  specimens  were  extremely  dark  and  could  not  be 
studied. 

Exochosphaeridium  striolatum  (Deflandre)  var.  truncatum  nov. 

(PI.  7,  figs.  1-3) 

DERIVATION  OF  NAME.  Latin,  truncatus,  shorten  by  cutting  off — with  reference 
to  the  truncated  extremities  of  the  processes. 

DIAGNOSIS.  A  variety  of  E.  striolatum  possessing  subspherical  to  ovoidal  shell; 
shell  wall  fibrous  or  lightly  pitted.  Processes  numerous,  typically  fibrous  and  blunted 


CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i  165 

distally,  slender  or  subtriangular,  rarely  branched.  Apical  process  large,  often 
foliate.  Precingular  archaeopyle,  commonly  present,  formed  by  loss  of  two  plate 
areas,  rarely  one. 

HOLOTYPE.  B.M.  (N.H.)  ¥.51982  (i).  Lower  Chalk,  Bureau  de  Recherches  Geo- 
logiques  et  Minieres  Borehole,  Escalles,  Pas  de  Calais,  at  159  metres  depth.  Upper 
Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype :  diameter  of  central  body  66  by  67  /z,  length  of  processes 
17-22^.  Range:  diameter  of  central  body  34  (56-1)  81  ^,  maximum  length  of 
processes  6  (17-8)  27  p.  Number  of  specimens  measured,  24. 

DESCRIPTION.  The  shell  possesses  a  moderately  thick  wall  (c.  i  /*)  but  it  is  quite 
often  distorted,  especially  when  an  archaeopyle  is  developed.  The  shell  surface  (peri- 
phragm)  is  typically  fibrous;  the  fibres  pass  down  the  length  of  the  processes  onto 
the  shell  surface  and,  there,  join  up  with  similar  fibres  from  adjacent  processes. 
Some  specimens  are  less  conspicuously  fibrous,  the  fibres  being  apparent  near  the 
bases  of  the  processes  and  only  extending  a  little  way  onto  the  shell  surface.  The 
remainder  of  the  shell  surface  in  these  forms  is  lightly  pitted.  The  processes  may  be 
fairly  slender  to  subtriangular  and  are  occasionally  joined  proximally.  A  small 
number  of  processes  are  subdivided  medially.  The  processes  are  typically  trun- 
cated distally  but  may  be  slightly  bulbous.  Process  alignment  was  not  observed. 

The  apical  process  is  larger  than  the  other  processes  and  often  foliate;  the  endo- 
phragm  occasionally  forms  a  small  apical  bulge  beneath  it.  The  archaeopyle  is 
typically  formed  by  the  removal  of  two  precingular  plates,  as  is  apparent  by  its  shape. 
Rarely  only  one  plate  is  lost.  Detached  opercula  consisting  of  two  precingular 
plates  have  been  located  (pi.  7,  fig.  3). 

REMARKS.  E.  striolatum  var.  truncatum  nov.  is  a  member  of  the  "  hirsutum  " 
group,  a  group  which  is  characterized  by  having  fibres  radiating  from  the  bases  of 
the  processes  over  the  shell  surface.  The  truncate  nature  of  the  processes,  although 
obvious,  has  not  previously  been  remarked  on.  Exochosphaeridium  spinosum  var. 
deflandrei  (Lejeuner-Carpentier  1941)  is  similar,  the  processes  sometimes  being 
bulbous  distally,  but  they  are  very  slender  and  appear  never  to  be  truncated.  Two 
other  forms,  E.  striolatum  (Deflandre)  (illustrated  by  Lejeune-Carpentier  1941  as 
Hystrichosphaeridium  hirsutum  text-figs.  1-4)  and  E.  (Hystrichosphaeridium)  cf. 
hirsutum  (Cookson  &  Eisenack  1958),  are  also  comparable,  differing  in  that  their 
processes  are  not  truncated.  Baltisphaeridium  bifidum  Clarke  &  Verdier  (1967) 
is  similar  but  possesses  fewer  and  more  slender  processes  which,  however,  are 
bifid  distally.  The  above  forms  all  belong  to  the  "  hirsutum  "  group  and  are  rather 
similar,  differing  only  in  the  detail  form  of  the  processes  and  perhaps  in  archaeopyle 
formation,  which  is  noted  here  for  the  first  time. 

Although  the  precingular  archaeopyle  is  typically,  but  not  always,  formed  by 
the  loss  of  two  plate  areas,  this  is  not  considered  reason  enough  for  the  erection  of  a 
new  genus.  This  species  may  indicate  a  trend  towards  the  genus  Lingulodinium  Wall 
(1967)  where  four  or  five  precingular  plate  areas  are  lost. 

OCCURRENCE.  E.  striolatum  var.  truncatum  is  infrequent  to  fairly  common  at 
all  horizons  throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles. 


166  CENOMANIAN   NON-CALCAREOUS   MICROPLANKTON,    i 

At  Fetcham  Mill  it  has  also  been  recorded  from  the  Albian  (sample  FM  886)  and 
Turonian  (sample  FM  520).  A  single  specimen  was  obtained  from  the  Upper  Wood- 
bine of  Texas,  but  the  species  was  absent  from  the  Saskatchewan  samples. 

OTHER  SPECIES 

The  following  species  and  variety  are  here  included  in  Exochosphaeridium  on  the 
basis  of  similarity  in  structure : 

Exochosphaeridium  spinosum   (White   1842)   comb.   nov.    1842   Xanthidium  spinosum  White: 

Microsc.  J.,  11,  35-40,  pi.  4  fig.  6. 
Exochosphaeridium  spinosum  var  deflandrei  (Lejeune-Carpentier  1941)  comb.  nov.  1941  Hystrichos- 

phaeridium  spinosum  var.  defiandrei  Lejeune-Carpentier:  Annls.  Soc.  geol.  Belg.,  63  (bull,  3), 

684,  figs.  6,  7. 

Cyst-Family  AREOLIGERAGEAE  Evitt  emend.  Sarjeant  &  Downie  1966 
Genus  CYCLONEPHELIUM  Deflandre  &  Cookson  emend.  Williams  &  Downie  1966 

REMARKS.  Four  species  are  described  from  the  Cenomanian,  C.  distinctum  Cook- 
son  &  Eisenack  being  the  only  one  fairly  common  throughout  this  stage.  C.  mem- 
braniphorum  Cookson  &  Eisenack,  C.  vannophorum  sp.  nov.  and  C.  paucispinum  sp. 
nov.  are  restricted  in  occurrence.  One  species  C.  eisenacki  sp.  nov.,  is  described  from 
the  Albian  of  Saskatchewan.  In  the  Cenomanian,  the  genus  is  most  abundant  at 
Escalles,  rarest  at  Saskatchewan  and  absent  from  Texas,  possibly  indicating  a 
preference  for  an  open  water  environment. 

Cyclonephelium  distinctum  Deflandre  &  Cookson 
(PI.  n,  figs.  6-8,  10 ;  Figs.  16  C,  D,  F) 

1955     Cyclonephelium  distinctum  Deflandre  &  Cookson  :  285,  pi.  2,  fig.  14. 
1963     Circulodinium  deflandrei  Alberti,  Baltes:  587,  pi.  6,  figs.  9-11. 

1967     Cyclonephelium  distinctum  Deflandre  &  Cookson;  Clarke  &  Verdier:  22,  pi.  i,  figs.  6,  7. 
(see  also  for  earlier  references) . 

DESCRIPTION.  This  species  is  extremely  variable,  as  pointed  out  by  Cookson  & 
Eisenack  (1962),  and  a  member  of  most  Cenomanian  assemblages  studied.  The 
shell,  which  is  always  somewhat  flattened,  may  be  subcircular  to  ovoidal  in  outline 
and  has  a  smooth  or  lightly  granular  shell  wall.  The  regular  outline  is  sometimes 
broken  by  an  apical  protuberance  and  more  rarely  by  two  reduced,  antapical  horns. 
The  latter  are  of  unequal  size  and  when  present  the  antapical  region  is  slightly  con- 
cave (Fig.  i6F).  The  bald  areas,  typical  of  this  genus,  may  occupy  almost  all  of  the 
ventral  and  dorsal  surfaces  of  the  shell  leaving  only  the  peripheral  regions  to  bear 
processes,  or  may  be  practically  invisible  beneath  the  encroaching  processes. 
Usually,  however,  these  bald  areas  are  circular  to  ovoidal  in  shape  and  occupy 
approximately  thirty  per  cent  of  each  side  of  the  shell.  The  processes  are  usually 
abundant,  extremely  variable  in  form  and  typically  under  one-third  of  the  shell 
width  in  length.  They  are  solid  and  usually  widen  distally  and  proximally,  may  be 
fine  or  taeniate,  and  are  often  distally  bifurcate.  The  bases  of  the  processes  are 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i  167 

occasionally  thickened  and  rarely  a  line  of  thickening  on  the  shell  surface  joins  one 
process  with  a  neighbouring  one.  Equally  rarely,  and  only  when  the  processes  are 
broad,  they  may  be  joined  distally. 

A  large  apical  archaeopyle  is  always  present.  This  is  angular,  possesses  a  pro- 
nounced zigzag  margin  and  on  the  ventral  surface  usually  a  relatively  deep  sulcal 
notch.  Although  the  archaeopyle  is  always  discernible,  the  operculum  is  sometimes 
still  attached.  It  then  behaved  as  a  lid,  returning  to  its  original  position  after  the 
escape  of  the  encysted  organism. 

DIMENSIONS.  Range  of  observed  specimens:  height  of  shell  (operculum  absent) 
41  (54-5)  82  ju,  width  48  (61-8)  81  \L,  maximum  length  of  the  processes  4  (10-7)  21  /z. 
Number  of  specimens  measured,  50. 

REMARKS.  C.  distinctum  appears  to  be  extremely  variable,  and  specimens  showing 
extremes  in  variation  sometimes  appear  to  be  transitional  to  other  species. 

OCCURRENCE.  Found  throughout  the  Cenomanian  of  Fetcham  Mill,  Compton 
Bay  and  Escalles,  and  rare  to  common  at  all  horizons.  Two  samples  from  Sas- 
katchewan, samples  Sas  890  and  Sas  805,  also  contain  C.  distinctum.  The  species 
had  a  wide  geographical  distribution  throughout  most  of  the  Cretaceous  and  is  of 
little  value  in  detailed  stratigraphy. 

Cyclonephelium  membraniphorum  Cookson  &  Eisenack 
(PL  ii,  fig.  9) 

1958     Cyclonephelium  compactum  Deflandre  &  Cookson  :  48,  pi.  12,  fig.  8  only. 

19626  Cyclonephelium  membraniphorum  Cookson  &  Eisenack  :  495,  pi.  6,  figs.  8-14. 

1964     Cyclonephelium  membraniphorum  Cookson  &  Eisenack;  Cookson  &  Hughes  :  44,  pi.  10, 

figs.  5,  6. 
1967     Cyclonephelium  membraniphomm  Cookson  &   Eisenack;   Clarke  &   Verdier:   23,   pi.   2, 

figs,  i,  2. 

DESCRIPTION.  The  shell  is  always  dorso-ventrally  flattened,  both  the  apex  and 
antapex  being  circular  to  subcircular  in  outline.  An  apical  archaeopyle  is  always 
developed,  with  a  zigzag  margin  and  sulcal  notch.  The  bald  areas,  typical  of  this 
genus,  may  be  relatively  large  or  quite  small  and  are  surrounded  by  high  membranes. 
The  latter  are  braced  by  stout  supports  which  arise  from  lines  of  thickening  on  the 
shell  surface.  These  lines  of  thickened  periphragm  may  be  curved  and  may  form 
semi-circular  rings.  When  the  latter  are  present,  the  membranes  are  in  the  form 
of  wide  tubular  projections.  The  periphragm  of  the  shell  wall  and  the  membranes 
is  granular  or  pitted.  The  membranes  may  sometimes  be  fenestrate. 

In  some  specimens  attributed  to  C.  membraniphorum  the  bald  areas  are  practically 
non-existent.  The  membranes  in  these  forms  are  particularly  well  developed  and 
commonly  are  in  the  form  of  wide  tubes  or  funnels.  Distally  they  have  a  sub- 
circular  to  polygonal  outline  and  may  possess  a  thickened  outer  rim  bearing  irregular 
pieces  of  periphragm  (PI.  n,  fig.  9). 

DIMENSIONS.  Range  of  observed  specimens:  length  of  shell  (operculum  missing) 
37  (50-6)  79  ft,  width  41  (54-1)  82 /u,  maximum  height  of  membranes  6  (13-3)  26^. 
Number  of  specimens  measured,  16. 

GEOL.    17,   3  12 


168  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 

REMARKS.  Many  of  the  specimens  observed  appear  identical  with  the  Albian  to 
Cenomanian  forms  from  Australia,  except  that  the  former  are  considerably  smaller, 
the  largest  of  them  only  falling  within  the  size  range  of  the  type  material.  It  is 
not  thought  practical  to  separate  the  European  forms  solely  on  the  size  distinction 
and  they  have,  therefore,  been  placed  in  the  Australian  species.  An  interesting 
variation  is  the  apparent  absence  of  the  bald  areas,  although  the  overall  shape  and 
the  apical  archaeopyle  typical  of  Cyclonephelium  are  present. 

OCCURRENCE.  In  addition  to  the  Australian  records,  C.  membraniphorum  is  rare 
and  spasmodic  throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and 
Escalles,  though  absent  elsewhere.  This  species  has  also  been  recorded  from  the 
Turonian  and  Lower  Senonian  by  Clarke  &  Verdier  (1967). 

Cyclonephelium  vannophorum  sp.  nov. 
(PL  9,  fig.  3:  PI.  ii,  figs,  n,  12;  Fig.  i6E) 

DERIVATION  OF  NAME.  Latin,  vannophorum,  fan-bearer — with  reference  to  the 
fan-shaped  processes. 

DIAGNOSIS.  Shell  subcircular  in  outline,  with  slight  apical  prominence  and  one 
or  two  reduced  antapical  horns.  Shell  wall  coarsely  granular  with  areas  devoid  of 
processes  of  moderate  size.  Processes  numerous,  short,  solid,  of  irregular  shape  and 
often  confluent  distally.  Apical  archaeopyle  with  zigzag  margin. 

HOLOTYPE.  B.M.  (N.H.)  slide  ¥.51986  (i).  Lower  Chalk,  Compton  Bay,  Isle  of 
Wight  at  15  feet  6  inches  above  the  base  of  the  Cenomanian.  Upper  Creatceous 
(Cenomanian) . 

DIMENSIONS.  Holotype:  length  of  shell  (including  operculum)  65  /*,  width  62 /u, 
length  of  processes  up  to  8//,.  Range:  length  of  shell  (without  operculum)  47  (57-8) 
70 /*,  width  56  (65-5)  78  p,  maximum  length  of  processes  3  (5-3)  8/u,.  Number  of 
specimens  measured,  11. 

DESCRIPTION.  When  two  antapical  horns  are  present  the  portion  of  the  shell 
between  them  is  concave  and  from  this  region  a  broad  furrow  passes  towards  the 
apex.  The  furrow  or  sulcus  decreases  in  width  and  depth  in  this  direction  and 
disappears  just  posterior  to  the  archaeopyle  margin.  The  processes  vary  greatly 
in  size  from  mere  enlarged  granules  (0-5  ju,)  to  8  p,  in  length.  In  the  larger  processes 
the  stem  is  quite  narrow,  the  distal  third  widening  rapidly  (PI.  9,  fig.  3)  and  is  some- 
times bifurcate.  The  processes  are  often  joined  to  form  a  short  line  on  the  shell 
surface.  Rarely  the  cingulum  is  just  discernible  by  a  concentration  of  small 
processes  along  its  borders. 

REMARKS.  C.  vannophorum  sp.  nov.  is  most  closely  comparable  to  ?C.  attadalicum 
Cookson  &  Eisenack  (19626)  from  the  Aptian /Albian  of  Australia.  The  processes 
are  similar  in  form  but  the  shell  of  ?C.  attadalicum  is  more  polygonal  and  the  cingulum 
is  always  well  defined. 

OCCURRENCE  .  C .  vannophorum  has  only  been  recorded  from  the  Lower  Cenomanian 
sample  CB  3  (Compton  Bay)  and  it  there  comprises  about  2%  of  the  micro- 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 


169 


E  F 

FIG.  16.  Cyclonephelium  eisenacki  sp.  nov.,  A.  Complete  specimen  (x  700).  B.  Specimen 
illustrating  apical  archaeopyle  (  x  700).  Cyclonephelium  distinctum  Deflandre  &  Cookson, 
C.  D.  and  F.  Illustrating  variation  in  the  shape  of  the  shell  of  this  species  (processes  re- 
moved) (X  700).  Cyclonephelium  vannophorumsp.  nov.,  E.  Specimen  with  well  developed 
apical  archaeopyle  (processes  removed)  (X  700). 


iyo  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

plankton  content.     The  unusual  distribution  and  the  similarity  to  a  Lower  Cretaceous 
species  indicate  that  C.  vannophorum  is  possibly  a  derived  form. 


Cyclonephelium  paucispinum  sp.  nov. 
(PI.  9,  figs,  i,  2) 

DERIVATION  OF  NAME.  Latin,  paucus,  few;  spina,  thorn — with  reference  to  the 
scarcity  of  the  processes. 

DIAGNOSIS.  Shell  subpolygonal  in  outline  with  well  defined  antapical  horn. 
Shell  wall  lightly  to  coarsely  granular,  bearing  small  number  of  irregularly  distri- 
buted processes  confined  to  peripheral  region  of  shell.  Processes  of  moderate  size, 
solid  and  widening  distally.  Apical  archaeopyle  with  a  zigzag  margin  and  sulcal 
notch. 

HOLOTYPE.  B.M.  (N.H.)  ¥.51981  (2).  Lower  Chalk,  Bureau  de  Recherches  Geo- 
logiques  et  Minieres  Borehole,  Escalles,  Pas  de  Calais  at  165  metres  depth.  Upper 
Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype :  length  of  shell  (operculum  missing)  54  //,,  width  81  p, 
length  of  processes  1-5-12 /x.  Range:  length  of  shell  (operculum  missing)  40  (72-4) 
92  /z,  width  54  (87-0)  112  /A,  maximum  length  of  processes  5  (10-2)  19  p.  Number  of 
specimens  measured,  12. 

DESCRIPTION.  The  shell  is  subpolygonal  in  outline,  the  cingular  region  being  the 
widest  portion  of  the  shell.  Posteriorly  there  is  a  pointed  antapical  horn.  The 
few  processes  present  are  of  moderate  size  and  if  close  together  tend  to  anastomose 
both  distally  and  proximally. 

REMARKS.  The  large  size,  the  paucity  of  processes  and  the  subpolygonal  shape 
of  the  shell  distinguish  C.  paucispinum  sp.  nov.  from  all  previously  described  species. 
The  processes  most  closely  resemble  those  of  C.  distinctum  but  are  fewer  in  number. 

OCCURRENCE.  C.  paucispinum  is  rare  in  samples  E  165,  CB  3,  and  common  in 
sample  CB  n.  The  distribution  is,  therefore,  rather  erratic  in  the  Lower,  Middle 
and  Upper  Cenomanian. 

Cyclonephelium  eisenacki  sp.  nov. 

(PI.  8,  figs.  3,  4;  PL  9,  fig.  4;  Figs.  I7A,  B) 
1960     Aptea  cf.  polymorpha  Eisenack  &  Cookson  :  9,  PI.  3,  figs.  2-4. 

DERIVATION  OF  NAME.     The  species  is  named  after  Professor  A.  Eisenack. 

DIAGNOSIS.  Shell  subtriangular,  flattened,  with  convex  sides.  Apical  horn  well 
developed;  antapical  horns,  if  present,  very  reduced.  Greater  part  of  shell  surface 
ornamented  by  complex  network  of  low  crests  and  short,  capitate  processes.  In 
centre  of  both  dorsal  and  ventral  sides  there  is  a  circular  area  devoid  of  ornamenta- 
tion. Apical  archaeopyle  always  present. 


CENOMANIAN   NON-CALCAREOUS   MICROPLANKTON,    i  171 

HOLOTYPE.  B.M.  (N.H.)  V.  51980(2).  Upper  Lower  Colorado  (Fish  Scale 
Zone?),  International  Yarbo  Borehole  No.  17,  Saskatchewan  at  1023  feet  depth. 
Lower  Cretaceous  (Albian). 

DIMENSIONS.  Holotype:  shell  length  70 /z,  width  66  p,  height  of  crests  1-5  p. 
Range:  shell  length  70-92/11  (4  complete  specimens  measured);  shell  length  (oper- 
culum  missing)  55  (63-1)  71 /z,  width  66  (72-1)  84 /z,  height  of  crests  2  (4-2)  7  /z. 
Number  of  specimens  measured,  12. 

DESCRIPTION.  The  shell  surface  is  ornamented  by  a  very  characteristic  network 
of  low  crests  or  lamellae  which  are  often  broken,  so  forming  isolated  processes.  A 
poorly-marked  cingulum,  outlined  by  the  crests,  is  occasionally  discernible.  The 
apical  operculum  is  often  in  position. 

REMARKS.  The  illustrations  of  Aptea  cf.  polymorpha  indicate  that  these  speci- 
mens belong  in  the  genus  Cydonephelium  and  may  well  be  conspecific  with  the 
Canadian  specimens  here  described.  This  species  differs  from  Aptea  polymorpha 
Eisenack  (1958)  by  the  absence  of  the  numerous  fine  processes  and  the  outer  mem- 
branous structure  which  they  support.  Pseudoceratium  turneri  Cookson  &  Eiesnack 
(1958),  from  the  Aptian /Albian  of  Australia,  possesses  a  similar  but  stronger  orna- 
mentation. Also  the  apical  and  antapical  horns  are  usually  much  better  developed, 
although  the  specimen  of  P.  turneri  illustrated  (loc.  cit.  pi.  5,  fig.  5),  approaches  a 
number  of  the  C.  eisenacki  examples. 

OCCURRENCE.  C.  eisenacki  sp.  nov.  is  fairly  common  in  two  samples  from 
Saskatchewan,  samples  Sas  1084  and  1023.  It  has  not  been  recorded  elsewhere  in 
the  material  examined. 


Cyst-Family  ADNATOSPHAERIDIAGEAE  Sarjeant  &  Downie  1966 
Genus  ADNATOSPHAERIDIUM  Williams  &  Downie  1966 
Adnatosphaeridium  chonetum  (Cookson  &  Eisenack)  comb.  nov. 

(PL  10,  figS.  II,  12.) 
19626  ICannosphaeropsis  choneta  Cookson  &  Eisenack  :  493,  pi.  4,  figs.  8-10. 

DESCRIPTION.  The  shell  is  spherical  to  subspherical  and  bears  a  number  of  more 
or  less  tubular  processes  which  possess  bands  of  thickening  for  support.  The  pro- 
cesses are  usually  joined  to  their  neighbour  either  along  their  entire  length,  or  only 
distally  by  membranes.  However,  isolated  tubular  processes  are  occasionally 
visible.  The  impression  obtained  from  most  specimens  is  of  a  complex  membranous 
network  perpendicular  to  the  shell  surface.  A  well  developed  apical  archae- 
opyle  is  typically  present. 

DIMENSIONS.  Range  of  observed  specimens:  shell  diameter  27  (34-1)  46^,  length 
of  processes  6  (10-4)  24 /z.  Number  of  specimens  measured,  15. 

REMARKS.  The  Cenomanian  specimens  examined  strongly  resemble,  but  are 
slightly  smaller  than,  the  type  material  from  the  Cenomanian  of  Australia.  This 


172  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 

species,  tentatively  placed  in  Cannosphaeropsis  by  Cookson  &  Eisenack,  is  here 
transferred  to  Adnatosphaeridium  because  of  the  presence  of  interconnecting 
processes  and  an  apical  archaeopyle. 

OCCURRENCE.  Apart  from  the  Australian  occurrence,  it  is  infrequent  to  common 
in  the  Middle  and  Upper  Cenomanian  of  Fetcham  Mill  and  Escalles,  not  occurring 
below  samples  FM  790  and  E  195.  Only  one  specimen  was  recorded  from  Compton 
Bay,  in  sample  CB  19. 


Cyst-Family  HYSTRICHOSPAERAGEAE  O.  Wetzel  emend.  Sarjeant 

&  Downie  1966 

Genus  HYSTRICHOSPHAERA  O.  Wetzel  emend.  Davey  &  Williams  1966 
Hystrichosphaera  ramosa  var.  ramosa  (Ehrenberg) 

(PI.  10,  figS.  I,  2,  5) 

1838     Xanthidium  ramosum  Ehr. :  pi.  i,  figs,  i,  2,  5. 

19660  Hystrichosphaera  ramosa  (Ehr.)  var.  ramosa  Davey  &  Williams  :  33,  pi.  i,  figs,  i,  6;  pi.  3, 

fig.  i;  text-fig.  8.     (See  also  for  earlier  references). 
1967     Hystrichosphaera  furcata  (Ehr.)  Clarke  &  Verdier:  48,  pi.  8,  figs.  12,  13. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  29  (40-7) 
56  fi,  maximum  length  of  processes  13  (19-8)  27  jii.  Number  of  specimens  measured, 
32. 

OCCURRENCE.  This  variety  has  a  known  stratigraphic  range  from  the  Oxfordian 
(Jurassic)  to  the  Ypresian  (Eocene).  It  is  a  common  variety  throughout  the  Ceno- 
manian of  Fetcham  Mill,  Compton  Bay  and  Escalles,  and  is  rare  to  infrequent  in  the 
material  from  Saskatchewan  and  Texas. 


Hystrichosphaera  ramosa  (Ehrenberg)  var.  gracilis  Davey  &  Williams 

1955     Hystrichosphaera  ramosa  (Ehr.)  Deflandre  &  Cookson  :  263,  pi.  5,  fig.  8. 
1963     Hystrichosphaera  ramosa  (Ehr.)  Gorka  :  48,  pi.  6,  figs.  6,  7. 

1966^  Hystrichosphaera  ramosa  (Ehr.)  var.  gracilis  Davey  &  Williams  :  34,  pi.  i,  fig.  5;  pi.  5, 
fig.  6. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  28-35 //,, 
maximum  length  of  processes  16-22 /u,.  Number  of  specimens  measured,  5. 

OCCURRENCE.  The  known  stratigraphic  range  of  this  variety  is  from  the  Ceno- 
manian (England)  to  the  Miocene  (Australia).  It  is  rare  to  infrequent  at  most  hori- 
zons throughout  the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles,  but 
has  not  been  recorded  in  the  North  American  material. 


CENOMANIAN   NON-CALCAREOUS   MICROPLAN  KTON,    i  173 

Hystrichosphaera  ramosa  (Ehrenberg)  var.   multibrevis  Davey  &  Williams 

(PL  10,  figs.  3,  4) 

I955     Hystrichosphaera  furcata  (Ehr.)  Valensi  :  586,  pi.  4,  fig.  4;  pi.  5,  fig.  12. 
1958     Hystrichosphaera  furcata  (Ehr.)  Eisenack  :  406,  pi.  25,  figs.  4-8. 

19660  Hystrichosphaera  ramosa  (Ehr.)  var.  multibrevis  Davey  &  Williams  :  35,  pi.   i,  fig.  4; 
pi.  4,  fig.  6;  text-fig.  9. 

DIMENSIONS.  Range  of  observed  specimens :  diameter  of  central  body  31  (39-7) 
51  n,  maximum  length  of  processes  10  (14-3)  21  p.  Number  of  specimens  measured,  13. 

OCCURRENCE.  H.  ramosa  var.  multibrevis  has  a  stratigraphic  range  from  the 
Lower  Cretaceous  (Hauterivian)  to  the  Eocene  (Ypresian).  It  is  rare  to  infrequent 
in  all  samples  examined  from  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  has 
also  been  recorded  from  the  Lower  Cretaceous  (Albian)  of  Saskatchewan,  in  samples 
Sas  967, 1023  and  1084. 

Hystrichosphaera  ramosa  (Ehrenberg)  var.  reticulata  Davey  &  Williams 

19660  Hystrichosphaera  ramosa  (Ehr.)  var.  reticulata.  Davey  &  Williams  :  38,  pi.  i,  fig.  2. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  33  (45-2) 
59  p,  maximum  length  of  processes  13  (15-3)  17  //,.  Number  of  specimens  measured, 
13- 

OCCURRENCE.  This  variety  is  rare  to  infrequent  at  most  horizons  throughout  the 
Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  has  also  been 
recorded  in  two  Albian  samples  from  Saskatchewan,  samples  Sas  967  and  1084. 

Hystrichosphaera  cingulata  var.  cingulata  (O.  Wetzel) 

J933     Cymatiosphaera  cingulata  O.  Wetzel  :  28,  pi.  4,  fig.  10. 

19660  Hystrichosphaera  cingulata  (O.  Wetzel)  Davey  &  Williams  :  38,  pi.  i,  fig.  9.     (See  also 

for  earlier  references). 
1967     Hystrichosphaera  cingulata  var.  cingulata  (O.  Wetzel)  Clarke  &  Verdier:  45,  pi.  8,  figs.  9,  10. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  26  (36-8) 
48  ju,  maximum  height  of  crests  5  (7-0)  13  /z.  Number  of  specimens  measured,  16. 

OCCURRENCE.  H.  cingulata  is  infrequent  at  all  horizons  throughout  the  Ceno- 
manian of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  has  not  been  observed  in 
the  North  American  material.  The  recorded  stratigraphic  range  is  from  Albian  to 
Middle  Miocene. 

Hystrichosphaera  cingulata  (O.  Wetzel)  var.  reticulata  Davey  &  Williams 

19660  Hystrichosphaera  cingulata  var.  reticulata  Davey  &  Williams  :  39,  pi.   i,  fig.   10;  pi.  2, 

fig-  4- 
1967     Hystrichosphaera  cingulata  var.  perforata  Clarke  &  Verdier  :  46,  pi.  9,  figs.  2-4,  text-fig.  19. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  33  (42-8) 
59  ft,  maximum  height  of  crests  7  (10-2)  17  /u..  Number  of  specimens  measured,  14. 


174  CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i 

OCCURRENCE.  H.  cingulata  var.  reticulata  is  rare  to  infrequent  throughout  the 
Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  appears  to  be  slightly 
more  common  in  the  Middle  and  Upper  Cenomanian  at  these  localities.  It  has  not 
been  observed  in  the  North  American  samples.  Clarke  &  Verdier  (1967)  also  record 
this  variety  from  the  Turonian  and  Senonian. 


Hystrichosphaera  crassimurata  Davey  &  Williams 

19663  Hystrichosphaera  crassimurata  Davey  &  Williams  :  39,  pi.  i,  fig.  u. 

1967     Hystrichosphaera  cingulata  var.  polygonalis  Clarke  &  Verdier:  47,  pi.  8  figs.  7,  8,  text-fig. 
20. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  36-46 /z, 
height  of  crests,  up  to  14  jit.  Number  of  specimens  measured,  4. 

OCCURRENCE.  H.  crassimurata  is  very  rare  to  rare  in  the  Middle  and  Upper 
Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles,  the  lowest  samples  in 
which  it  is  found  being  FM  790,  E  195  and  CB  9.  It  has  also  been  recorded  from  the 
Upper  Woodbine  Formation  of  Texas  but  is  absent  from  the  Saskatchewan  material. 


Hystrichosphaera  crassipellis  Deflandre  &  Cookson 

1954     Hystrichosphaera  crassipellis  Deflandre  &  Cookson:  text-fig.  5. 

ig66a  Hystrichosphaera  crassipellis  Deflandre  &  Cookson;  Davey  &  Williams  :  40,  pi.  i,  figs.  7, 

8.     (See  also  for  earlier  references). 
1967     Hystrichosphaera  crassipellis  Deflandre  &  Cookson;  Clarke  &  Verdier:  48,  pi.  8,  fig.  n; 

pi.  9,  fig.  i. 

DIMENSIONS.  Range  of  observed  specimens:  diameter  of  central  body  34  (47-9) 
68  p,,  maximum  length  of  processes  10  (17-8)  29  p.  Number  of  specimens  measured,  19. 

OCCURRENCE.  H.  crassipellis  is  rare  at  most  horizons  throughout  the  Cenomanian 
of  Fetcham  Mill,  Compton  Bay  and  Escalles;  it  has  not  been  recorded  in  the  North 
American  material.  The  recorded  stratigraphic  range  is  from  Cenomanian  to  Middle 
Miocene  (Gerlach  1961). 


Genus  ACHOMOSPHAERA  Evitt  1963 

Achomosphaera  ramulifera  (Deflandre) 

(PL  10,  fig.  7) 

1935     Hystrichosphaera  cf.  ramosa  (Ehr.)  Deflandre  :  pi.  5,  fig.  n. 

1966^  Achomosphaera  ramulifera  (Deflandre)  Davey  &  Williams  :  49,  pi.  2,  fig.  3.     (See  also  for 

earlier  references) . 
1967     Achomosphaera  ramulifera  (Deflandre);  Clarke  &  Verdier:  40,  pi.  8,  fig.  i. 

DIMENSIONS.     Range  of  observed  specimens:  diameter  of  central  body  36  (44-7) 
61  /A,  maximum  length  of  processes  16  (25-8)  36  p.   Number  of  specimens  measured,  14. 


CENOMANIAN   NON-CALCAREOUS   MICROPL  AN  KTON,    i  175 

OCCURRENCE.  This  species  has  a  known  stratigraphic  range  from  the  Cenomanian 
to  the  Eocene  (Pastiels  1948).  It  is  a  very  rare  to  infrequent  species  at  most  horizons 
in  the  Cenomanian  from  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  is  absent 
from  the  North  American  samples. 

Achomosphaera  sagena  Davey  &  Williams 

ig66a  Achomosphaera  sagena  Davey  &  Williams  :  31,  pi.  2,  figs,  i,  2. 

1967     Achomosphaera  reticulata  Clarke  &  Verdier:  41,  pi.  8,  figs.  2,  3,  text-fig.  16. 

DIMENSIONS.  Range  of  type  material:  diameter  of  central  body  35  (48-4)  59^, 
maximum  length  of  processes  17  (20-8)  28  ^.  Number  of  specimens  measured,  12. 

OCCURRENCE.  A.  sagena  is  a  very  rare  to  rare  species  occurring  spasmodically  in 
the  Cenomanian  samples  from  Fetcham  Mill,  Compton  Bay  and  Escalles.  Clarke  & 
Verdier  (1967)  have  also  recorded  it  (as  A .  reticulata)  from  the  Turonian  and  Senonian. 

Genus  Hystrichodinium  Deflandre  emend.  Clarke  &  Verdier  1967 

Hystrichodinium  voigti  (Alberti) 

(PI.  10,  figs.  6,  10) 

1961     Heliodinium  voigti  Alberti  :  33,  pi.  8,  figs.  1-5. 

1966^  Heliodinium  voigti  Alberti;  Sarjeant  :  142,  pi.  16,  fig.  2;  text-fig.  36. 

1967     Hystrichodinium  voigti  (Alberti)  Clarke  &  Verdier:  38. 

DESCRIPTION.  The  shell  is  ovoidal  to  subpolygonal.  The  periphragm  is  smooth 
or  slightly  granular,  and  forms  sutural  crests  and  ribbon-like  processes.  The  former 
are  variably  developed,  may  occasionally  be  absent,  but  when  present  give  rise  to 
long,  flexuous  processes  along  their  length.  When  the  crests  are  absent  the  pro- 
cesses arise  directly  from  the  shell  surface.  The  processes  are  not  confined  to  the 
plate  boundaries  and  may  be  distributed  at  random  over  the  shell  surface.  The 
processes  are  thin-walled,  typically  simple  and  occasionally  terminate  with  bifurcate 
or  trifurcate  extremities.  The  cingulum  (4-5^  wide)  is  always  delimited  by  low 
crests  and  is  strongly  helicoid.  A  precingular  archaeopyle,  formed  by  the  loss  of 
plate  3",  is  typically  present.  One  detached  operculum  has  been  located  and  bears 
five  processes. 

DIMENSIONS.  Range  of  observed  specimens:  shell  length  40  (49-6)  62 /*,  width 
40  (48-4)  58  ^,  length  of  processes  27  (35-6)  48  //,.  Number  of  specimens  measured,  11. 

OCCURRENCE.  H.  voigti  is  rare  to  fairly  common  and  found  spasmodically  through- 
out the  Cenomanian  of  Fetcham  Mill,  Compton  Bay  and  Escalles.  It  was  originally 
described  by  Alberti  (1961)  from  the  Lower  Barremian  to  ?  Lower  Aptian  of  Germany. 
The  specimens  described  by  Sarjeant  (19660)  are  from  the  Cenomanian  of  Fetcham 
Mill. 

Hystrichodinium  dasys  sp.  nov. 

(PI.  10,  figs.  8,  9) 

DERIVATION  OF  NAME.  Latin,  dasys,  hairy  or  shaggy — with  reference  to  the 
abundant  hair-like  processes. 


176  CENOMANIAN   NON-CALCAREOUS   MICROPL ANKTON,    i 

DIAGNOSIS.  Shell  subspherical,  thin-walled,  smooth  or  slightly  granular.  Tabula- 
tion very  faintly  marked ;  cingulum,  and  more  rarely,  precingular  and  postcingular 
plate  boundaries  visible.  Processes  numerous,  short,  fine  and  flexuous,  aligned 
along  plate  boundaries,  also  intratabular.  Archaeopyle  not  observed. 

HOLOTYPE.  B.M.  (N.H.)  ¥.51982  (3).  Lower  Chalk,  Bureau  de  Recherches  Geo- 
logiques  et  Minieres  Borehole,  Escalles,  Pas  de  Calais  at  159  metres  depth.  Upper 
Cretaceous  (Cenomanian). 

DIMENSIONS.  Holotype :  diameter  of  central  body  58  by  61  p,  length  of  processes 
10-14  p.  Range:  diameter  of  central  body  42  (53-0)  70^,  maximum  length  of 
processes  7  (10-6)  14  p.  Number  of  specimens  measured,  13. 

DESCRIPTION.  The  shell,  being  thin-walled,  is  always  found  deformed.  Only 
two  specimens  possess  a  faint  tabulation.  This  is  best  seen  in  the  holotype  which 
has  a  clear  but  lightly  defined  cingulum  and  two  or  three  precingular  and  post- 
cingular plate  boundaries  visible.  The  numerous  processes  are  hair-like,  approxi- 
mately 0-2 11  wide  for  most  of  their  length,  probably  hollow  and  terminate  with  a 
point.  The  most  characteristic  feature  of  the  processes  is  their  extreme  flexibility. 

REMARKS.  The  characteristic  processes  and  the  presence  of  a  tabulation  differ- 
entiate H.  dasys  sp.  nov.  very  easily  from  all  previously  described  microplankton 
species. 

OCCURRENCE.  This  species  is  rare  to  common  in  all  samples  from  Escalles  except 
the  lower  three  (E  201,  E  207,  and  E  213).  One  specimen  was  recorded  from  Comp- 
ton  Bay  (sample  CB  7)  and  two  from  Fetcham  Mill  (sample  FM  520,  Turonian). 
The  species  was  not  recorded  from  the  North  American  material.  The  distribution 
of  H.  dasys  possibly  indicates  an  environmental  difference  between  Escalles  and 
Fetcham  Mill/Compton  Bay,  perhaps  in  the  depth  of  water  or  distance  from  land. 

IV.  REFERENCES 

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28  :  93-105,  pis.  8,  9. 

1961.     Zur    Kenntnis    mesozoischer   und    alttertiarer   Dinoflagellaten    und   Hystrichos- 

phaerideen  von  Nord-  und  Mitteldeutschland  sowie  einigen  anderen  europaischen  Gebieten. 
Palaeontographica,  Cassel,  Stuttgart,  116,  A  :  1-58,  pis.  1-12. 

BALTES,  N.  1963.  Dinoflagellate  si  Hystrichosphaeride  cretacice  din  Platforma  moezica. 
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1965.     Observatii  asupra  microflorei  cretacice  inferiare  din  zona  R.  Bacaz.     Petrol  Gaze, 

Bucuresti,  16  :  3-17,  pis.  3,  4. 

BARROIS,  C.     1876.     Recherches  sur  le  terrain  Cretac6  Superieur  de  1'Angleterre  et  de  1'Irlande. 

Mem.  Soc.  geol.  N.,  Lille,  3,  189-205. 

BRISTOW,  H.  W.     1889.     The  Geology  of  the  Isle  of  Wight.     Mem.  geol.  Surv.  U.K.,  London. 
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CLARKE,  R.  F.  A.  &  VERDIER,  J.  P.     1967.     An  investigation  of  microplankton  assemblages 

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24,  3  :  1-96,  pis.  1-17. 


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Nomenclature  of  some  Upper  Cretaceous  Dinoflagellate  Taxa  (in  press) . 

COOKSON,  I.  C.  1956.  Additional  microplankton  from  Australian  late  Mesozoic  and  Tertiary 
sediments.  Aust.  J.  mar.  Freshwat.  Res.  Melbourne,  7,  i  :  183-191,  pis.  i,  2. 

—  1965.     Cretaceous  and  Tertiary  Microplankton  from  South-Eastern  Australia.     Proc.  R. 
Soc.  Viet.,  Melbourne,  78,  i  :  85-93,  pis.  9-11. 

COOKSON,  I.  C.  &  EISENACK,  A.  1958.  Microplankton  from  Australian  and  New  Guinea  Upper 
Mesozoic  sediments.  Proc.  R.  Soc.  Viet.,  Melbourne,  70,  i  :  19-79,  pis.  1-12. 

—  igdoa.     Microplankton     from     Australian     Cretaceous     sediments.     Micropaleontology , 
New  York,  6,  i  :  1-18,  pis.  1-3. 

-  19606.     Upper  Mesozoic  microplankton  from  Australia  and  New  Guinea.     Palaeontology, 
London,  2,  2  :  243-261,  pis.  37-39. 

-  1961.     Upper   Cretaceous  microplankton  from  the   Belfast  No.  4  bore,  South- Western 
Australia.     Proc.  R.  Soc.  Viet.,  Melbourne,  74,  i  :  69-76,  pis.  n,  12. 

-  19620.     Some  Cretaceous  and  Tertiary  microfossils  from  Western  Australia.     Proc.  R. 
Soc.  Viet.,  Melbourne,  75,  2  :  269-273,  pi.  37. 

—  19626.     Additional  microplankton  from  Australian  Cretaceous  sediments.     Micropalaeon- 
tology,  New  York,  8,  4  :  485-507,  pis.  1-7. 

COOKSON,  I.  C.  &  HUGHES,  N.  F.  1964.  Microplankton  from  the  Cambridge  Greensand 
(mid-Cretaceous).  Palaeontology,  London,  7,  i  :  37-59,  pis.  5-11. 

COOKSON,  I.  C.  &  MANUM,  S.  1964.  On  Deflandrea  victoriensis  n.  sp.,  D.  tripartita  Cookson  & 
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PLATE    i 

Gonyaulacysta  exilicristata  sp.  nov. 
Fetcham  Mill  Borehole  (depth,  730  feet). 

FIG.  i.     Dorsal  surface  of  holotype.     Slide  PF.  3987  (i).      x  500. 
FIG.  2.     Ventral  surface.     Slide  PF.  3987.      X  500. 

Carpodinium  obliquicostatum  Cookson  &  Hughes 

FIG.  3.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  750  feet).  Lateral  view.  Slide 
PF.  3988.  x  500. 

FIG.  4.  Lower  Chalk,  Escalles  Borehole  (depth,  159  metres) .  Dorsal  surface  with  archaeopyle. 
Slide  E  195/3.  X  I25° 

Histocysta  pa  I  la  sp.  nov. 
Fetcham  Mill  Borehole. 

FIG    5.     Holotype;  view  of  archaeopyle  and  attached  operculum.    Slide  PF.  3052.     x  500. 
FIG.  6.     Paratype;  Antapical  view.     Slide  PF.  3991  (i).     X  500. 

Gonyaulacysta  delicata  sp.  nov. 

FIG.  7.     Ventral  surface  of  holotype.     V.  51979  (i).      X  500. 
FIG.  8.     Paratype;  V. 5 1979(2).      X  500. 

Gonyaulacysta  sp.  A.     Figured  specimen. 
Lower  Chalk,  Fetcham  Mill  Borehole. 

FIG.  9.     Ventral  surface.     Slide  PF.  3987  (2).      x  500 
FIG   10.     Medial  view.     Slide  PF.  3987  (2).      x  500. 


Butt.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 

•       * 


PLATE  i 


m 


4 


GEOL.   17,  3. 


PLATE   2 
Cribroperidiniutn  intricatum  sp.  nov. 

FIG.  i.     Ventral  surface  of  holotype.     V.  51980  (i).      x  500. 

FIG.  2.     Lower  Colorado,  Saskatchewan  (depth,  1,023  feet).     Dorsal  surface  with  operculum 
in  situ.     ¥.51980.     x  500. 

FIG.  3.     Lower  Colorado,  Saskatchewan.     Detached  operculum.     Slide  Sas  1023/3.     X  500. 

Microdinium  setosum  Sarjeant. 

FIG.   4.     Lower  Chalk,   Fetcham  Mill  Borehole   (depth,   840  feet).     Lateral  view.     Slide 
PF.  3036.      x  640. 

Microdinium  variospinum  sp.  nov. 

FIG.  5.     Lower  Chalk,  Escalles  Borehole  (depth,   195  metres).     Antapical  sutural  spines 
visible.     Slide  E  195/3.      X  640. 

FIG.  6.     Lateral  view  of  holotype.     V.  51981  (i).      x  640. 

IMicrodinium  crinitutn  sp.  nov. 

FIG.  7.     Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  770  feet).     Slide  FM  770/15.      x  640. 
FIG.  8.     Dorsal  surface  of  holotype     Slide  PF.  3990  (i).      x  640. 

Microdinium  distinctum  sp.  nov. 

FIG.  9.     Ventral  surface  of  holotype.     Slide  PF.  3989  (i).     X  640. 
FIG.  10.     Dorsal  surface  of  holotype.     X  640. 

FIG.  ii.     Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  790  feet).     Ventral  surface.     Slide 
FM  790/16.      x  640. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 


PLATE  2 


f 


8 


v  '    - 


V 


W 


10 


11      t    V 


PLATE   3 
Ellipsodinium  rugulosum  Clarke  &  Verdier 

FIG.  i.  Lateral  view  of  specimen  showing  operculum  partially  detached.  Slide  PF.  3988. 
X  1250. 

Fromea  amphora  Cookson  &  Eisenack 
Lower  Chalk,  Fetcham  Mill  Borehole. 

FIG.  2.     Slide  FM  770/6  (depth,  770  feet).     X  500. 

FIG.  3.     Specimen  with  operculum  attached  (depth,  650  feet).     Slide  PF.  3041.     x  500. 

Microdinium  veligerum  (Deflandre). 

FIG.  4.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  650  feet).  Antapical  view  to  show 
crestal  cavities.  Slide  FM  650/5.  x  1250. 

Apteodinium  granulatum  Eisenack. 

FIG.  5.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  690  feet).  Ventral  surface.  Slide 
FM  690/12.  x  640. 

FIG.  6.  Lower  Chalk,  Escalles  Borehole  (depth,  165  metres).  Lateral  view.  V.  51981. 
X  640. 

Cassiculosphaeridia  reticulata  sp.  nov. 

FIG.  7.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  730  feet) .  Operculum  partially  detached. 
Slide  FM  730/9.  X  500. 

Chytroeisphaeridia  euteiches  sp.  nov. 

FIG.  8.     Holotype  showing  archaeopyle.     V.  51982  (2).      x  500. 

FIG.  9.  Lower  Chalk,  Escalles  Borehole  (depth,  159  metres).  Archaeopyle  with  detached 
operculum.  V.  51982.  x  500. 

Epelidosphaeridia  spinosa  (Cookson  &  Hughes) 
Lower  Chalk,  Fetcham  Mill  Borehole. 

FIG.  10.     Dorsal  surface.     Slide  PF.  3992,  (depth,  770  feet),      x  500. 

FIG.  ii.     Ventral  surface  with  sulcus.     Slide  PF.  3992  (depth,  770  feet).      X  500. 

FIG.  12.     Complete  specimen.     Slide  PF.  3048  (depth,  770  feet),      x  500. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 


PLATE  3 


m  f 

* 

i 


•A 


\ 


I 


8 


•x- 

-JW 


-  •••"  ': 


10 


11 


12 


PLATE   4 
Hystrichosphaeridium  deanei  Davey  &  Williams 

FIG.  i.     Lower  Chalk,  Compton  Bay.     Complete  specimen  (137  feet  above  base  of  Chalk). 
Slide  CB  19/2.      x  500. 

Hystrichosphaeridium  difficile  Manum  &  Cookson 
Upper  Colorado,  Saskatchewan. 

FIG.  2.     Complete  specimen,  archaeopyle  partially  detached.     ¥.51983  (depth,  805  feet). 
X  500. 

FIG.  6.     Detached  operculum  bearing  4  processes.     Slide  Sas  805/3  (depth,  805  feet).     X  500. 

FIG.  7.     Detached  operculum,  lateral  view.     Slide  Sas  805/3  (depth,  805  feet).     X  500. 
X  500. 

Cassiculosphaeridia  reticulata  sp.  nov. 
FIG.  3.     Holotype;  view  of  archaeopyle.     V.  51981  (4).      x  500. 

Microdinium  veligerum  (Deflandre). 

FIG.  4.     Lower  Chalk,  Fetcham  Mill  Borehole  (depth,   750  feet).     Dorsal  surface.     Slide 
PF.  3988.      x  640. 

Microdinium  cf .  ornatum  Cookson  &  Eisenack 

FIG.  5.     Lower  Chalk,  Escalles  Borehole  (depth,  189  metres).     Dorsal  surface.     Slide  E  189/4. 
X  640. 

Hystrichosphaeridium  radiculatum  Davey  &  Williams 

FIG.  8.     Lower  Chalk,  Escalles  Borehole  (depth,  159  metres).     Lateral  view  showing  archaeo- 
pyle.    V.  51982.      x  500. 

Hystrichosphaeridium  mantelli  Davey  &  Williams. 

FIG.  9.     Lower  Chalk,  Escalles  Borehole  (depth,  153  metres).     Lateral  view  to  show  archaeo- 
pyle.    Slide  E  153/3.      X  500. 

Polysphaeridium  laminaspinosum  Davey  &  Williams 
Lower  Chalk,  Fetcham  Mill  Borehole. 

FIG.  10.     Apical  view  showing  archaeopyle.     Slide  PF.  3035  (depth,  840  feet),      x  500. 
FIG.  ii.     Antapical  view.     Slide  PF.  3035  (depth,  840  feet),      x  500. 


Bull.  Br.  Mus.  not.  Hist.  (Geol.)  17,  3 


PLATE  4 


8 


10 


11 


PLATE   5 
Oligosphaeridium  anthophorurn  (Cookson  &  Eisenack) 

Lower  Colorado,  Saskatchewan. 
FIG.   i.     Enlargement  to  show  process  extremities.     Slide  Sas   1023/3   (depth,   1023  feet). 

X  975- 

FIG.  2.     Detached  operculum.     Slide  Sas  1023/1  (depth,  1023  feet).      X  500. 
FIG.  3.     V.  51980  (boring  depth,  1023  feet),      x  500. 

Oligosphaeridium  prolixispinosum  Davey  &  Williams 

FIG.  4.  Lower  Chalk,  Compton  Bay  (116  feet  above  the  base  of  the  Chalk).  Specimen 
showing  "  bald  "  cingular  region.  Slide  CB  I7/C.  X  500. 

Hystrichosphaeridium  tubiferum  (Ehrenberg) 
Lower  Chalk,  Fetcham  Mill  Borehole. 

FIG.  5.     Detached  operculum.     Slide  FM  690/12  (depth,  690  feet).      X  500. 
FIG.  8.     Slide  PF.  3987  (depth,  730  feet),      x  500. 

Oligosphaeridium  complex  (White) 

FIG.  6.  Lower  Chalk,  Escalles  Borehole  (depth,  159  metres).  Detached  operculum  composed 
of  4  plates.  V.  51982.  x  500. 

FIG.  7.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  750  feet).  Specimen  illustrating  the  6 
precingular  processes.  Slide  FM  750/13.  x  500. 

Hystrichosphaeridium  bowerbanki  Davey  &  Williams 

FIG.  9.  Lower  Chalk,  Compton  Bay  (116  feet  above  the  base  of  the  Chalk).  Lateral  view, 
archaeopyle  to  the  north.  Slide  CB  I7/C.  x  500. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 


PLATE  5 


. 


'/ 

5 


GEOL.  17,  3. 


PLATE   6 

Oligosphaeridiutn  reniforme  (Tasch) 
FIG.  i.     Lower  Colorado,  Saskatchewan  (depth,  1023  feet).     Slide  Sas  1023/1.     x  500. 

Tanyosphaeridiutn  variecalamum  Davey  &  Williams 

FIG  2      Holotype.     Slide  PF  3035  (2)       x  500 

FIG  5      Lower  Chalk,  Compton  Bay  (28  feet  above  base  of  Chalk).     Precingular,  cingular  and 
postcingular  processes  aligned  into  three  series.     Slide  CB  5/C       x  500 

Litosphaeridium  siphoniphorum  (Cookson  &  Eisenack) 
Lower  Chalk,  Fetcham  Mill  Borehole. 

FIG.  3.     Lateral  view  showing  precingular  and  postcingular  processes,  and  antapical  process. 
Slide  PF.  3987  (depth,  730  feet),      x  500. 
FIG.  4.     Detached  operculum.     Slide  FM  690/14  (depth,  690  feet).     X  975. 

Callaiosphaeridium  asymmetricum  (Deflandre  &  Courteville) . 

FIG.  6.     Upper  Greensand,  Fetcham  Mill  Borehole  (depth,  886  feet).     Lateral  view  illustrating 
epitractal  archaeopyle.     Slide  FM  886/2.      x  500. 

Cleistosphaeridiutn  polypes  (Cookson  &  Eisenack) 
FIG.   7.     Upper  Woodbine  Formation.     Enlargement  to  show  process  extremities.     Slide 

T5/3-      X  975- 

FIG.  8.     Lower  Chalk,  Compton  Bay  (151  feet  above  base  of  Chalk).     Specimen  possessing 
apical  archaeopyle.     Slide  CB  2i/C.      x  500. 

Cleistosphaeridiutn  polypes  var.  clavulum  nov. 

FIG.  9.     Type.     Slide  PF.  3995  (i)   x  500. 

FIG.  10.     Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  840  feet).     Enlargement  to  show 
process  extremities.     Slide  PF.  3035.      x  975. 

fCleistosphaeridium  aciculare  sp.  nov. 

FIG.    ii.     Upper   Colorado,    Saskatchewan    (depth,    835    feet).     Specimen   with   numerous 
fine  processes.     V.  51988.      x  500  (phase  contrast). 
FIG.  12.     Holotype.     V.  51979  (3).      X  500. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 


PLATE  6 


8 


12 


,  *t 


10 


PLATE   7 
Exochosphaeridium  striolatutn  var.  truncatum  nov. 

FIG.  i.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  750  feet).  Apical  view  showing  two 
partially  detached  precingular  plates  to  the  north.  Slide  PF.  3988.  x  500. 

FIG.  2.  Type;  precingular  archaeopyle  clearly  shown  to  be  formed  by  the  removal  of  two 
plates.  V.  51982  (i).  x  500. 

FIG.  3.  Lower  Chalk,  Escalles  Borehole  (depth,  165  metres).  Detached  operculum  composed 
of  two  precingular  plates.  ¥.51981.  x  500. 

fCleistospaeridium  flexuosum  Davey  et  al. 

FIG.  4.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  690  feet).  Slide  PF.  3030.  x  500 
(phase  contrast). 

Exochosphaeridium  phragmites  Davey  et  al. 

FIG.  5.  Lower  Chalk,  Escalles  Borehole  (depth,  165  metres).  Apical  process  with  precingular 
archaeopyle  to  the  north-west.  V.  51981.  x  500. 

Cleistosphaeridium  huguonioti  var.  pertusutn  nov. 

FIG.  6.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  690  feet).  Enlargement  showing 
vacuolated  processes  with  small  distal  bifurcations.  Slide  FM  690/14.  x  975. 

FIG.  7.     Type.     Slide  PF.  3040  (2).      x  500. 

FIG.  9.  Lower  Chalk,  Escalles  Borehole  (depth,  159  metres).  Specimen  possessing  apical 
archaeopyle.  Slide  E  159/4.  X  500. 

Cleistosphaeridium  heteracanthum  (Deflandre  &  Cookson) 
FIG.  8.     Lower  Chalk,  Compton  Bay  (137  feet  above  base  of  Chalk).     Slide  CB  19/2.      x  500. 

Cleistosphaeridium  huguonioti  (Valensi) 

FIG.  10.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  730  feet).  Detached  operculum. 
Slide  FM  730/19.  x  975. 

? Cleistosphaeridium  parvum  sp.  nov. 

FIG.  ii.     Holotype.     V.  51981  (3).      x  975. 

FIG.  12.  Lower  Chalk,  Escalles  Borehole  (depth,  159  metres).  Cingular  region  devoid  of 
processes  clearly  visible.  Slide  E  159/1.  X  975. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 


PLATE  7 


/ 

k 
* 


'  *  7 


PLATE   8 

Cleistosphaeridiutn  armatum  (Deflandre) 

FIG.  i.     Lower  Chalk,  Escalles  Borehole  (depth,  189  metres).     View  of  apical  archaeopyle. 
Slide  E  189/4.      x  975- 

FIG.  2.  Lower  Chalk,  Compton  Bay  (116  feet  above  base  of  Chalk).  Enlargement  to  show 
process  extremities.  Slide  CB  iy/C.  x  975. 

FIG.  12.     Lower  Chalk,  Compton  Bay  (i  16  feet  above  base  of  Chalk).     Slide  CB  I7/C.      x  500. 

Cyclonephelium  eisenacki  sp.  nov. 

FIG.  3.  Lower  Colorado,  Saskatchewan  (depth,  1,023  feet).  Specimen  with  archaeopyle 
developed.  ¥.51980.  X  500. 

FIG.  4.     Holotype.     V.  51980  (2).      x  500. 

Prolixosphaeridium  conulurn  sp.  nov. 

FIG.  5.     Holotype;  complete  specimen.     V.  51981  (5).      x  500. 

FIG.  6.  Lower  Chalk,  Compton  Bay  (59  feet  above  base  of  Chalk).  Apical  archaeopyle  to 
the  north.  Slide  CB  9/2.  X  500. 

Cleistosphaeridium  multifurcatum  (Deflandre) 
FIG.  7.     Lower  Chalk,  Escalles  Borehole  (depth,  207  metres).     View  of  apical  archaeopyle. 

Slide  E  207/5.      X  500. 

FIG.  10.     Lower  Chalk,  Compton  Bay  (15  feet  above  base  of  Chalk).     Complete  specimen. 

V.  51986.      x  500. 

Coronifera  oceanica  Cookson  &  Eisenack 

FIG.  8.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  730  feet).  Specimen  possessing  an 
apical  archaeopyle  and  an  antapical  process.  Slide  FM  730/13.  x  500. 

FIG.  ii.  Lower  Chalk,  Escalles  Borehole  (depth,  189  metres).  Complete  specimen.  Slide 
E  189/4.  x  500. 

Surculosphaeridiutn  longifurcatum  (Firtion) 

FIG.  9.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  730  feet).  Lateral  view  to  show  apical 
archaeopyle,  deeply  furcate  precingular  processes  and  completely  subdivided  cingular  processes. 
Slide  PF.  3987.  X  500. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 


PLATE  8 


•**  •••• 


10 


12 


PLATE   9 
Cyclonephelium  paucispinum  sp.  nov. 

FIG.  i.     Holotype.     V.  51981  (2).      x  500. 

FIG.  2.  Lower  Chalk,  Compton  Bay  (15  feet  above  base  of  Chalk).  Complete  specimen  with 
operculum  partially  detached.  V.  51986.  x  500. 

Cyclonephelium  vannophorum  sp.  nov. 

FIG.  3.  Enlargement  of  holotype  to  show  apical  archaeopyle,  and  the  shape  of  the  processes. 
V.  51986  (i).  X  975- 

Cyclonephelium  eisenacki  sp.  nov. 

FIG.  4.  Lower  Colorado,  Saskatchewan  (depth,  1,023  feet).  Complete  specimen.  Slide 
Sas  1023/3.  X  500. 

Hystrichokolpoma  ferox  (Deflandre) 
Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  840  feet).     Slide  FM  840/11. 

FIG.  5.  Lateral  view  (bottom  of  specimen  by  transparency)  showing  precingular,  cingular 
and  postcingular  processes ;  large  antapical  process  to  the  south  and  fine  sulcal  processes  to  the 
west,  x  500. 

FIG.  6.     Medial  section,      x  500. 

FIG.  7.  Lateral  view  (top  of  specimen)  showing  precingular,  cingular  and  postcingular 
processes,  x  500. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 


PLATE  9 

3 


v 


PLATE    10 

Hystrichosphaera  ramosa  var.  ramosa  (Ehr.) 
Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  750  feet). 
FIG.  i.     Ventral  surface.     Slide  PF.  3988.      x  500. 

FIG.  2.     Dorsal  surface  with  precingular  archaeopyle.     Slide  PF.  3988.      x  500. 
FIG.  5.     Detached  operculum  (boring  depth,  770  feet).     Slide  FM  770/15.      x  500. 

Hystrichosphaera  ramosa  var.  multibrevis  Davey  &  Williams 
Lower  Chalk,  Fetcham  Mill  Borehole. 

FIG.  3.     Slide  PF.  3988  (depth,  750  feet),      x  500. 
FIG.  4.     Slide  PF.  3988  (depth,  750  feet),      x  500. 

Hystrichodiniutn  voigti     (Alberti) 
Lower  Chalk,  Escalles  Borehole. 

FIG.  6.     Archaeopyle  to  the  north-east.     V.  51982  (depth,  159  metres). 

FIG.  10.  Detached  operculum  bearing  4  processes.  Slide  E  165/1  (depth,  165  metres). 
X  500. 

Achomosphaera  ramulifera  (Deflandre) 

FIG.  7.  Lower  Chalk,  Escalles  Borehole  (depth,  159  metres).  Specimen  showing  precingular 
archaeophyle  and  apical  process.  V.  51982.  x  500. 

Hystrichodinium  dasys  sp.  nov. 

FIG.  8.     Holotype  illustrating  cingulum.     V.  51982  (3).      x  500. 

FIG.  9.  Middle  Chalk,  Fetcham  Mill  Borehole  (depth,  520  feet).  Specimen  with  unusually 
stout  processes  proximally.  Slide  FM  520/7.  x  500. 

Adnatosphaeridium  chonetum  (Cookson  &  Eisenack) 

FIG.  ii.  Lower  Chalk,  Escalles  Borehole  (depth,  165  metres).  Complete  specimen,  V.  51981. 
X  500. 

FIG.  12.  Lower  Chalk,  Fetcham  Mill  Borehole  (depth,  730  feet).  Apical  archaeopyle  present. 
Slide  PF.  3987.  x  500. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 


PLATE  10 


8 


10 


--• 


12 


PLATE    ii 

Trichodinium  castaneum  (Deflandre). 
Lower  Chalk,  Escalles  Borehole. 

FIG.  i.  Lateral  view  showing  precingular  archaeopyle  and  cingulum,  V.  51989  (depth, 
195  metres),  x  500. 

FIG.  2.     Dorsal  view  showing  small  apical  horn.     Slide  E  159/2  (depth,  159  metres),      x  500. 

FIG.  3.  Complete  specimen  possessing  a  cingulum  and  possibly  a  sulcus,  Slide  E  189/4  (depth, 
189  metres),  x  500. 

Exochosphaeridium  pseudohystrichodinium  (Deflandre) 
Lower  Chalk,  Escalles  Borehole. 

FIG.  4.     Complete  specimen  with  operculum  partially  detached,  V.  51982  (depth,  159  metres). 
X  500. 
FIG.  5.     Complete  specimen  illustrating  pitted  surface,  V.  51981  (depth,  165  metres),      x  500. 

Cyclonephelium  distinctum  Deflandre  &  Cookson 

FIG.  6.  Lower  Chalk,  Escalles  Borehole  (depth,  159  metres).  Specimen  with  unusually 
long  processes,  V.  51982.  x  500. 

FIG.  7.  Lower  Chalk,  Escalles  Borehole  (depth,  159  metres).  Complete  specimen;  archaeo- 
pyle in  the  act  of  developing.  V.  51989.  x  500. 

FIG.  8.  Lower  Chalk,  Compton  Bay  (15  feet  above  base  of  Chalk).  Typical  specimen  with 
archaeopyle  developed.  V.  51986.  X  500. 

FIG.  10.  Lower  Chalk,  Escalles  Borehole  (depth,  165  metres).  Detached  operculum. 
V.  51981.  x  500. 

Cyclonephelium  membraniphorum  Cookson  &  Eisenack 

FIG.  9.  Lower  Chalk,  Escalles  Borehole  (depth,  153  metres).  Apical  archaeopyle  well 
illustrated.  Slide  E  153/3.  X  500. 

Cyclonephelium  vannophorum  sp.  nov. 

FIG.  ii.     Holotype  with  operculum  partially  detached.     V.  51986  (i).      x  500. 
FIG.  12.     Lower  Chalk,  Compton  Bay  (15  feet  above  base  of  Chalk).     Specimen  with  apical 
archaeopyle  developed.     V.  51986.      x  500. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  3 


PLATE  ii 


\    \ 


• 


w? 


10 


PRINTED  IN  GREAT  BRITAIN 
BY  ADLARD  &  SON  LIMITED 
BARTHOLOMEW  PRESS,  DORKING 


A  REDESCRIPTION  OF 

W.  CARRUTHERS'  TYPE 

GRAPTOLITES 


ISLES  STRACHAN 


BULLETIN  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

GEOLOGY  Vol.  17  No.  4 

LONDON:  1969 


A  REDESCRIPTION  OF  W.  CARRUTHERS' 
TYPE  GRAPTOLITES 


BY 

ISLES  STRACHAN 


Pp.  181-206;  5  Plates;  8  Text-figures 


BULLETIN  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

GEOLOGY  Vol.  17  No.  4 

LONDON :  1969 


THE  BULLETIN  OF  THE  BRITISH  MUSEUM 
(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they  become 
ready.  Volumes  will  contain  about  three  or  four 
hundred  pages,  and  will  not  necessarily  be  completed 
within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  17,  No.  4  of  the  Geological 
(Palaeontological)  series.  The  abbreviated  titles  of 
periodicals  cited  followthose  of  the  World  List  of 
Scientific  Periodicals. 


World  List  abbreviation: 
Bull.  Br.  Mus.  nat.  Hist.  (Geol.). 


Trustees  of  the  British  Museum  (Natural  History)  1969 


TRUSTEES    OF 
THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 

Issued  17  January,  1969  Price  £i  45. 


A  REDESCRIPTION  OF  W.  CARRUTHERS' 
TYPE  GRAPTOLITES 

By  ISLES  STRACHAN 

CONTENTS 

Page 

I.     INTRODUCTION          .........  183 

II.     DESCRIPTIONS  .........  185 

Leptograptus  capillaris          .  .          .          .          .          .          .  185 

Pleurograptus  linearis  .          .          .          .          .          .          .  186 

Dicellograptus  elegans  .          .          .          .          .          .          .  187 

Dicellograptus  moffatensis     .          .          .          .          .          .          .  189 

Dicranograptus  clingani        .          .          .          .          .          .          .  190 

Climacograptus  minimus      .......  191 

Climacograptus  minutus       .          .          .          .          .          .          .  193 

Cvyptograptus  tricornis         .          .          .          .          .          .          .  194 

Monograpius  clingani  .          .          .          .          .          .          .  195 

Monograptus  intermedius     .          .          .          .          .          .          .  196 

Diver sograptus?  capillaris     .          .          .          .          .          .          .  198 

Rastrites  maximus       ........  200 

Cyvtograptus  murchisoni       .......  202 

"  Dendrograptus  "  lentus      .          .          .          .          .          .          .  204 

III.     REFERENCES   ..........  204 

SYNOPSIS 

The  fourteen  species  of  graptolites  described  and  figured  by  William  Carruthers,  mainly  in 
his  paper  of  1868,  are  redescribed  from  the  type  material  where  possible.  Two  neotypes  are 
proposed.  Some  of  the  species  appear  to  be  unsatisfactory  by  modern  standards,  but  it  is 
hoped  that  refiguring  of  the  types  will  assist  future  work. 

I.  INTRODUCTION 

IN  their  monograph  of  British  Graptolites  (1901-18)  Elles  &  Wood  described  and 
figured  most  of  the  previously  known  species  as  well  as  numerous  new  forms.  In 
many  cases,  however,  they  did  not  use  the  original  type  material  either  because  it 
was  not  available  or  because  later  collections  had  provided  other,  sometimes  better, 
material.  In  the  fifty  years  which  have  elapsed  since  then,  discrimination  of  grapto- 
lite  species  has  become  keener,  particularly  by  workers  overseas  who  have  frequently, 
in  their  subdividing,  formally  designated  type  specimens  without  consideration  of 
the  availability  or  value  of  the  early  material.  It  seems  appropriate  that  the  species 
described  in  the  nineteenth  century  should  be  re-examined  so  that  these  workers 
should  be  able  to  recognize  the  old  species  accurately  with  the  aid  of  modern 
descriptions  and  illustrations.  The  work  could  be  tackled  systematically  by  genus 
and  species  but  this  involves  consideration  of  generic  and  specific  limits  in  deciding 

GEOL.    IJ,  4  15 


184  W.    CARRUTHERS'   TYPE   GRAPTOLITES 

the  scope  of  a  paper.  It  is  easier  to  treat  the  species  for  redescription  by  discussing 
one  author  at  a  time  and  I  hope  to  undertake  further  papers  of  a  similar  scope  to  this. 

William  Carruthers  (1830-1922)  was  born  at  Moffat,  Dumfriesshire,  and  early 
developed  an  interest  in  the  local  geology,  particularly  in  the  graptolites  for  which 
the  area  had  then  recently  become  famous  through  the  researches  of  Harkness  and 
Salter.  His  own  publications  on  graptolites  are  comparatively  few  in  number  but 
it  is  clear  that  by  1867  he  was  regarded  as  an  expert  on  the  group,  since  he  contributed 
the  article  on  graptolites  to  the  fourth  edition  of  Murchison's  "  Siluria  ".  In  the 
following  year  he  described  more  fully  the  new  species  erected  in  that  memoir. 
He  strongly  disagreed  with  Nicholson  on  the  interpretation  of  a  number  of  graptolite 
features  and  they  carried  on  an  acrimonious  correspondence  in  the  pages  of  the 
Geological  Magazine  during  1867.  At  the  same  time  he  was  also  describing  fossil 
plants,  to  which  he  later  devoted  all  his  research.  In  1870-72  he  was  assisting  the 
young  amateur  John  Hopkinson  with  his  descriptions  of  new  species  of  graptolites 
from  the  Moffat  Shales  but  they  both  appeared  to  realize  that  the  publication  in 
1876  of  Lap  worth's  paper  on  "  Scottish  Monograptidae  "  marked  a  new  and  more 
critical  approach  to  graptolite  systematics  and  stratigraphy  and  neither  of  them 
published  further  papers  on  the  subject.  It  is,  however,  clear  from  Lapworth's 
later  papers  that  he  maintained  a  friendly  association  with  Hopkinson  and  Carruthers 
and  the  latter's  manuscript  notes  on  graptolites  are  in  the  Lapworth  Library, 
Birmingham  University.  These  notes  contain  no  indication  of  how  or  when  Lap- 
worth  acquired  them  but  there  are  no  references  to  papers  after  1872  and  it  is  possible 
that  Carruthers  gave  them  to  Lapworth  before  taking  up  more  administrative 
duties  at  the  British  Museum  (Natural  History)  in  1880  and  confining  his  research 
to  palaeobotany. 

The  new  species  described  by  Carruthers  are  found  in  two  main  papers,  in  1858 
and  1868.  The  first  of  these  was  reprinted  the  following  year  in  a  more  accessible 
journal  and,  as  already  mentioned,  the  1868  paper  described  properly  the  species 
which  were  only  illustrated  in  the  appendix  to  "  Siluria  ".  Carruthers  had  some 
harsh  remarks  to  make  about  Nicholson's  drawings  of  graptolites  and,  on  the  whole, 
his  own  are  generally  sufficiently  accurate  to  allow  recognition  of  the  type  specimens. 
Most  of  these  were  catalogued  at  once  in  the  British  Museum  (Natural  History) 
collections  where  they  can  be  traced.  Almost  all  his  species  can  be  recognized 
fairly  easily.  The  type  of  Rastrites  maximus  is  the  only  one  which  has  not  been 
found  and  a  neotype  is  proposed  for  that  species. 

It  is  a  pleasure  to  acknowledge  the  ready  co-operation  which  I  have  had  from  those 
at  the  British  Museum  (Natural  History)  who  have  had  charge  of  the  graptolite 
collections  and  to  thank  their  photographers  who  have  provided  most  of  the  illus- 
trations for  the  plates.  I  must  also  express  my  thanks  to  the  Geological  Survey  & 
Museum  and  to  the  Royal  Scottish  Museum  for  the  loan  of  specimens  over  a  period 
which  has  been  much  longer  than  I  had  at  first  intended. 

Specimens  with  Q  numbers  are  in  the  British  Museum  (Natural  History);  with 
GSM,  in  the  Institute  of  Geological  Sciences,  London;  with  BU,  in  the  Geology 
Department,  University  of  Birmingham;  and  with  SM,  in  the  Sedgwick  Museum, 
Cambridge. 


W.    CARRUTHERS'    TYPE   GRAPTOLITES  185 

II.  DESCRIPTIONS 

Leptograptus  capillaris  (Carruthers) 
PL  i 

1868  Cladograpsus  capillaris  Carruthers:  130,  pi.  5,  figs.  7,  ya. 

1876  Leptograptus  capillaris (Carr.)  Lapworth  :  9,  pi.  3,  fig.  72. 

1903  Leptograptus  capillaris  (Carr.);  Elles  and  Wood  :  112,  pi.  XV,  figs.  4a-d. 

?I954  Leptograptus  capillaris  (Carr.);  Sherrard  195,  pi.  n,  fig.  8. 

Original  description.  "  Extremely  slender  polypary,  with  remote  branches  and 
very  minute  hydrothecae;  about  twenty-four  in  an  inch.  It  is  not  so  abundant  as 
C.  linearis,  and  is  easily  distinguished  by  its  capillary  appearance.  It  is  probably 
the  same  species  as  that  figured  and  described  by  Emmons  in  his  American  Geology, 
vol.  i,  p.  109,  pi.  i,  fig.  7,  under  the  name  of  Nemagrapsus  capillaris.  Loc.  Moffat  ". 

Comments  on  usage.  There  appears  to  have  been  little  confusion  over  this  species. 
Although  he  quoted  Emmons'  species,  Carruthers  clearly  intended  his  own  form 
to  rank  as  a  new  species  and  the  American  form  is  now  referred  to  Thamnograptus. 
The  original  description  refers  to  "  remote  branches  "  which  are  not  shown  by  the 
type  specimens  or  original  figure  but  one  illustration  of  Cladograpsus  linearis  of  the 
previous  year  (Carruthers  1867,  pi.  2,  fig.  I7a)  shows  the  characteristic  curved 
stipes  with  secondary  branches.  The  original  of  this  has  not  however  been 
recognized. 

TYPE  MATERIAL.  The  syntypes,  British  Museum  (Natural  History)  Q-30,  were 
refigured  by  Elles  &  Wood.  As  lectotype  I  select  the  large  rhabdosome  on  the 
upper  right  of  Carruthers'  original  figure  (see  PI.  i,  fig.  2).  The  precise  locality  and 
horizon  are  not  known  but  Elles  &  Wood  give  it  as  Hartfell  Spa?,  Hartfell  Shales. 

DIAGNOSIS.  Very  slender  Leptograptus  with  markedly  curved  stipes,  maximum 
breadth  0-5  mm.  thecae  eight  to  ten  per  cm. 

REVISED  DESCRIPTION.  The  species  normally  occurs  crowded  on  the  slabs  so  that 
details  of  the  stipes  are  not  readily  seen.  The  sicula  is  inconspicuous  but  the  stipes 
appear  to  grow  initially  horizontally  or  even  somewhat  downward  before  curving 
gracefully  upwards  and  inwards,  forming  loops  on  occasion.  The  stipes  widen  from 
an  initial  breadth  of  0-2  mm.  to  a  maximum  of  0-5  mm.  over  a  distance  of  at  least 
4  or  5  cm.  and  appear  to  be  twisted  so  that  the  thecae  can  appear  on  either  side  of 
the  curve.  The  sicula  is  2  mm.  long  and  about  0-2  mm.  wide  but  details  of  the  origin 
of  the  stipes  are  not  known.  The  thecae  appear  to  be  simple  leptograptid  tubes 
but  the  preservation  of  the  distal  parts  of  the  stipes  is  such  as  to  make  measurements 
of  their  length  and  overlap  impossible. 

GEOLOGICAL  HORIZON.  Lapworth  (1878)  recorded  the  species  only  from  the 
zone  of  Pleurograptus  linearis  but  Elles  &  Wood  (1903)  also  recorded  it  from  the  zone 
of  Dicranograptus  clingani.  The  type  slab  shows  no  associates,  nor  do  most  of  the 
specimens  in  the  Lapworth  Collection.  Lithologically,  however,  the  slabs  agree 
with  the  upper  part  of  the  Lower  Hartfell  Shales. 

DISTRIBUTION.  This  species  is  known  from  only  a  few  localities  in  the  South  of 
Scotland  and  is  not  recorded  from  beds  of  a  similar  age  in  South  Wales.  Ruedemann 


i86  W.    CARRUTHERS'   TYPE   GRAPTOLITES 

and  Decker  recorded  it  from  the  Viola  Limestone  in  Oklahoma  but  gave  no  figures 
(Ruedemann  1947).  Sherrard  (1954)  recorded  it  from  New  South  Wales  but  her 
figure  shows  stipes  which  appear  to  be  too  broad  and  not  curved  enough.  They 
may  be  one  of  the  forms  of  Leptograptus  flaccidus  which  occurs  about  the  same  hori- 
zon, e.g.  L.f.  arcuatus  Elles  &  Wood.  Thomas  (1960)  illustrated  a  form  which  is 
clearly  not  L.  capillaris  as  the  stipes  are  too  straight,  and  this  then  casts  doubt  on 
the  range  of  the  species  as  given  by  Thomas,  which  goes  up  to  the  Ashgill  Series. 

Pleurograptus  linearis  (Carruthers) 
PL  2 ;  Figs.  la-c 

1858  Cladograpsus  linearis  Carruthers  :  467,  fig.  i. 

1859  Cladograpsus  linearis  Carruthers :  24,  fig.  i. 

non  1867     Cladograpsus  linearis  Carruthers;  Carruthers  :  369,  pi.  2,  fig.  17. 
?i8&7a  Cladograpsus  linearis  Carruthers;  Carruthers  :  540,  fig.  8. 

1867  Pleurograpsus  linearis  (Carruthers)  Nicholson  :  257,  pi.  n,  figs.  1-5. 

1868  Cladograpsus  linearis  Carruthers ;  Carruthers  :  129. 

1876     Pleurograptus  linearis  (Carruthers) ;  Lapworth,  pi.  3,  fig.  69. 

1904     Pleurograptus  linearis  (Carruthers);  Elles  &  Wood  :  119,  pi.  16,  fig.  7,  pi.  17,  fig.  i. 

ORIGINAL  DESCRIPTION.  "  From  a  short  and  very  slender  base  the  zoophyte 
divides  into  two  stems,  each  supporting  the  cells  on  their  upper  sides.  Branches 
are  given  off  at  irregular  intervals  from  these  principal  stems.  The  length  of  the 
polypidom  is  very  great;  one  specimen  I  have  been  able  to  trace  for  nearly  three 
feet  .  .  .  The  polypidom  at  its  origin,  near  to  the  slender  base,  is  very  narrow, 
being  little  more  than  a  fine  line;  as  it  increases  in  length  it  increases  in  breadth, 
until  it  is  fully  two-fifths  of  a  line  broad.  The  cells  are  very  remote  from  each 
other,  and  are,  at  first  sight,  from  the  slight  indentation  they  make  in  the  stem, 
scarcely  perceptible,  giving  the  Graptolite  the  appearance  as  if  it  were  a  clear  line. 
The  mouth  of  the  cell  is  straight  and  at  right  angles  to  the  axis ;  it  makes  an  inden- 
tation equal  to  about  one-sixth  of  the  breadth  of  the  polypidom.  The  number  of 
cells  in  an  inch  is  about  eighteen.  Type  locality  Hartfell." 

COMMENTS  ON  USAGE.  Carruthers'  later  figures  (?  1867,  i867a)  are  different  from  his 
original  but  there  has  been  no  difficulty  in  the  recognition  of  his  species.  Carruthers 
accepted  Nicholson's  specimen  of  Pleurograpsus  linearis  although  he  was  very  critical 
of  the  detail  of  the  figures.  Nicholson's  specimen  was  refigured  by  Elles  &  Wood 
who  also  figured  a  variety  simplex  in  which  the  branches  are  much  more  widely 
spaced. 

TYPE  MATERIAL.  Carruthers'  original  specimen  is  in  the  British  Museum  (Natural 
History),  (^.848,  and  the  counterpart  is  amongst  material  presented  to  the  Royal 
Scottish  Museum,  Edinburgh,  by  Carruthers  in  1858  (RSM  1858 . 10 .4).  Nicholson's 
specimen,  Q .  27,  was  said  by  Elles  &  Wood  to  be  the  type  specimen  but  it  is  clearly 
not.  The  lower  of  the  two  large  specimens  on  the  type  slab  has  four  lateral  branches 
and  is  recognizable  as  the  specimen  figured  in  1858  (see  PL  2,  figs,  i,  2). 

REVISED  DESCRIPTION.  Rhabdosome  consisting  of  two  or  three  main  stipes 
arising  from  the  sicula  with  secondary  branching  on  one  or  both  sides.  Stipes  up 


W.    CARRUTHERS'   TYPE   GRAPTOLITES 


187 


to  I  mm.  broad;  thecae  simple  leptograptid,  about  eight  per  cm.,  with  apertural 
excavations  occupying  about  one-third  of  the  rhabdosome  breadth.  The  sicula 
is  not  seen  on  any  of  the  figured  material  and  is  probably  associated  in  older  rhabdo- 
somes  with  one  of  the  branches,  a  feature  also  seen  in  species  of  Leptograptus  where 
a  third  branch  may  be  developed.  Young  specimens  on  the  type  slab,  however, 
suggest  that  the  sicula  is  about  2  mm.  long.  The  stipes  are  initially  very  slender 
but  show  some  secondary  thickening  in  older  specimens  where  the  thecae  on  the 
main  stipes  are  difficult  to  distinguish.  There  is  considerable  variation  in  the 
branching,  Nicholson's  specimen  (PL  2,  fig.  3)  being  much  stiffer  than  Carruthers' 
or  the  other  specimen  figured  by  Elles  &  Wood  in  which  the  tertiary  branched  stipes 
are  aligned  in  flowing  curves.  This  suggests  that  the  variation  in  attitude  of  the 
stipes  is  simply  a  depositional  effect. 

fl 


^ 


FIG.  i.  Pleurograptus  linearis  Carr.  a.  Enlargement  of  proximal  part  of  main  stipe  showing 
2  lateral  branches  and  possible  central  branch  from  the  sicula.  Counterpart  of  type  slab, 
R.S.M.  1858.10.4.  The  2  parallel  distal  parts  of  branches  are  the  left  hand  pair  from  the 
holotype  (PI.  2,  fig.  i).  b,  c.  Young  specimens  showing  sicula,  type  slab,  Q.8^8.  All  X5- 

DISTRIBUTION.  P.  linearis  appears  to  be  confined  to  the  one  horizon  in  the  south 
of  Scotland  where,  however,  it  is  quite  common  if  all  the  fragments  showing  this 
type  of  branching  really  belong  to  it.  Ruedemann  (1908;  1947)  figured  a  single 
specimen  from  the  Utica  Shale  of  New  York  State  which  shows  similar  branching 
at  rather  more  distant  intervals,  approximating  in  that  respect  to  var.  simplex 
which  is  also  recorded  by  Thomas  (1960)  from  Australia. 

Dicellograptus  elegans  (Carruthers) 
PI.  3,  fig.  i. ;  Figs  2a,  b. 


Didymograpsus  elegans  Carruthers  :  369,  pi.  2,  fig.  i6a. 
Didymograpsus  elegans  Carruthers;  Carruthers  :  129,  pi.  5,  figs.  8a,  ?d. 
Dicellograpsus  elegans  (Carruthers)  Hopkinson  :  24,  pi.  i,  fig.  3. 
Dicellograptus  elegans  (Carruthers) ;  Lapworth,  pi.  4,  fig.  87. 
Dicellograptus  elegans  (Carruthers) 
Dicellograptus  elegans  (Carruthers) 


1867 
1868 
1871 
1876 
1877 
1904 
?I947 
1954 

ORIGINAL   DESCRIPTION.     "  Branches  of  the  polypary  divaricating  at  various 
angles,  and  with  a  slight  curve  within  a  short  distance  of  the  proximal  origin  of  the 


Dicellograptus  elegans  (Carruthers) 

Dicellograptus  elegans  (Carruthers) ;  Sherrard,  pi.  10,  fig.  6. 


Lapworth  :  141,  pi.  7,  fig.  8. 

Elles  &  Wood  :  159,  pi.  23,  figs.  2a-e. 

Ruedemann  :  380,  pi.  63,  fig.  i. 


i88 


W.   CARRUTHERS'   TYPE   GRAPTOLITES 


polypary.  The  hydrothecae  are  rounded  at  the  apex,  and  free  throughout  a  con- 
siderable portion  of  their  length  and  the  intervening  spaces  are  rounded  at  the  base ; 
about  twenty-two  cells  in  an  inch.  The  initial  process  is  obvious  in  young  specimens, 
but  I  have  not  been  able  to  detect  it  in  old  individuals ;  the  outer  apex  of  the  angle 
ornamented  with  three  short  strong  spines.  Loc.  Moffat  ". 

COMMENTS  ON  USAGE.  Since  Elles  &  Wood  refigured  the  type  specimen  there 
has  been  no  difficulty  with  this  species.  The  originals  of  Carruthers'  figures  8b 
and  8c  (BM(NH)  Q-54)  were  recognized  by  Hopkinson  (1871)  as  being  a  distinct 
species  (D.  morrisii  Hopkinson)  and  not  merely  young  stages  of  elegans  as  Carruthers 
thought. 

TYPE  MATERIAL.  The  holotype,  Q.850  (PL  3,  fig.  i),  is  from  the  Hartfell  Shales, 
Dobbs  Linn.  The  species  appears  to  be  commonest  in  the  P.  linearis  Zone  but  the 
associates  on  the  type  slab  (Dicellograptus  pumilus  and  Climacograptus  spp.)  give 
no  direct  confirmation  of  horizon. 

DIAGNOSIS.  Dicellograptus  with  markedly  introverted  thecae  and  with  distinct 
sigmoid  curvature  of  stipes  near  proximal  end,  so  that  the  stipes  curve  first  upwards, 
then  outwards,  then  upwards  again. 


FIG.  2.  Dicellograptus  spp.  a.  D.  elegans  Carr.  Detail  of  thecal  aperture  in  proximal  part 
of  stipe.  Q.850.  x  30.  b.  D.  elegans  Carr.  Distal  thecae.  (3.850.  X5-  c.  D.  moffatensis 
Carr.  Distal  thecae.  (3.843.  X5- 

REVISED  DESCRIPTION.  The  proximal  double  curvature  is  quite  distinctive  and 
the  stipes  have  an  almost  uniform  width  of  i  mm.  throughout  their  length.  The 
first  two  thecae  have  prominent  apertural  spines  which,  with  the  virgella,  give  the 
three-spined  proximal  end  noted  by  Carruthers.  The  thecae  number  eight  to  ten 
per  cm.  and  are  of  the  strongly  introverted  type  with  marked  ventral  curvature 
(group  IV  of  Elles  &  Wood,  which  however  are  not  introtorted,  see  Bulman  1944  :  37). 

GEOLOGICAL  HORIZON.  Lapworth  (1878)  and  Elles  &  Wood  (1904)  recorded  the 
species  only  from  the  P.  linearis  zone,  but  in  the  summary  range  chart  at  the  end 
of  the  Monograph,  Elles  &  Wood  also  recorded  it  as  common  in  the  underlying  zone 
of  D.  clingani.  Elles  (1925)  also  recorded  it  from  both  zones  so  its  precise  range 
must  await  a  revision  of  the  Hartfell  Shales.  Foreign  records  are  surprisingly 
scanty  but  it  seems  to  occur  in  Australia  at  the  same  horizon. 


W.    CARRUTHERS'    TYPE   GRAPTOLITES  189 

Dicellograptus  moffatensis  (Carruthers) 
PL  3,  figs.  5,  6;  Fig.  2c 


1858  Didymograpsus  Moffatensis  Carruthers  :  469,  fig.  3. 

1859  Didymograpsus  Moffatensis  Carruthers;  Carruthers 
1868     Didymograpsus  Moffatensis  Carruthers;  Carruthers 


26,  fig.  3. 

129. 

25,  pi.  i,  fig.  4. 


1871  Dicellograpsus  Moffatensis  (Carruthers)  Hopkinson 

1875  Dicellograptus  moffatensis  (Carruthers);  Hopkinson  &  Lapworth  :  654,  pi.  34,  fig.  5a. 

1877  Dicellograptus  Moffatensis  (Carruthers);  Lapworth  :  141,  pi.  7,  fig.  9. 

1904  Dicellograptus  moffatensis  (Carruthers);  Elles  &  Wood  :  157,  pi.  23,  figs.  la-f. 

ORIGINAL  DESCRIPTION.  '  The  base  terminates  in  three  distinct  spinous  pro- 
cesses. The  zoophyte  bifurcates  from  the  base.  The  general  appearance  is  like 
the  figure;  or  occasionally  the  lines  form  an  acute  angle  for  about  a  quarter  of  an 
inch,  then  suddenly  expand  in  slight  curves,  almost  at  right  angles,  for  a  short  dis- 
tance, when  they  again  recur  to  their  original  direction.  The  branches  are  united 
for  about  a  quarter  of  a  line  by  a  slight  web,  which  in  some  specimens  is  terminated 
in  a  fine  process  of  short  length,  taking  the  direction  of  a  line  bisecting  the  angle. 
The  cells  are  arranged  in  the  outer  margins;  they  are  very  remote,  and  penetrate 
the  polypidom  to  scarcely  one-fourth  of  its  breadth;  they  form  slight  openings  on 
the  margin  of  the  polypidom,  first  entering  at  a  right  angle,  and  then  suddenly 
turning  downwards.  These  openings  are  lengthened  ovate  pouches,  answering 
exactly  in  shape  and  size  to  the  cell-serratures  of  the  margin.  The  number  of  cells 
in  an  inch  is  about  twenty.  The  breadth  of  the  polypidom  is  about  two-thirds  of  a 
line.  Type  locality  Hartf ell ." 

COMMENTS  ON  USAGE.  In  1868,  Carruthers  noted  D.  divaricatus  (Hall)  and 
D.  anceps  (Nicholson)  as  synonyms  of  his  own  species  but  this  is  merely  a  reflection  of 
the  confused  state  of  graptolite  systematics  at  the  time.  Elles  &  Wood  refigured 
Carruthers'  type  but  the  species  does  not  seem  to  have  been  widely  recognized. 

TYPE  MATERIAL.  This  is  one  of  the  few  species  of  which  Elles  &  Wood  made  any 
discussion  of  type  specimens.  They  considered  that  Carruthers'  specimen  was  not 
a  satisfactory  type  and  that  a  specimen  from  the  Lapworth  Collection  should  be 
taken  as  the  type.  Carruthers'  original  figure  is  admittedly  poor  (PL  3,  fig.  5)  but 
the  type  slab  shows  three  specimens  which  agree  fairly  well  with  it  and  show  the 
essential  characters  of  the  species.  The  abrupt  widening  of  the  stipes  on  Lap  worth's 
specimen,  which  Elles  &  Wood  wanted  as  a  character  of  the  species,  appears  to  be 
the  result  of  slight  shearing. 

It  is  impossible  to  decide  which  of  the  three  specimens  was  the  original  of 
Carruthers'  figure  so  the  specimen  (Q .  843)  figured  by  Elles  &  Wood  is  here  selected  as 
lectotype  (see  PL  3,  fig.  6).  It  is  from  the  Hartf  ell  Shales  of  Hartf  eU  and  is  associa- 
ted on  the  slab  with  Orthograptus  cf .  whitfieldi.  This  probably  indicates  a  low  horizon 
in  the  Hartf  ell  Shales.  Lapworth  (1878)  records  it  no  higher  than  the  zone  of 
Climacograptus  wilsoni. 

REVISED  DESCRIPTION.  Stipes  sub-parallel  initially,  then  diverging  making  an 
angle  of  about  45°,  sometimes  later  converging.  Stipes  widening  from  an  initial 
breadth  of  0-4  mm.  to  a  maximum  of  about  1-5  mm.  Thecae  eight  to  ten  per  cm., 


igo  W.    CARRUTHERS'    TYPE   GRAPTOLITES 

with  markedly  curved  ventral  walls  and  introverted  apertures.  The  proximal  end 
usually  shows  a  membrane  in  the  axil  of  the  stipes,  obscuring  the  sicula.  The 
virgella  and  first  thecal  spines  variably  developed.  The  thecae  are  poorly  preserved 
but  distally  appear  to  be  very  similar  to  those  of  D.  elegans  (Fig.  2c). 

It  is  unfortunate  that  both  Carruthers  and  Hopkinson  included  D.  divaricatus 
(Hall)  in  their  synonymy  of  this  species  as  it  makes  it  difficult  to  establish  what  they 
regarded  as  the  diagnostic  features  of  the  species.  The  general  shape  of  the  rhab- 
dosome  appears  to  be  the  most  characteristic  feature,  particularly  the  narrow  axil 
and  later  divergence.  In  this  respect  the  specimen  figured  by  Elles  &  Wood  in 
their  Monograph  (pi.  23,  fig.  ic)  is  not  at  all  typical  but  its  only  associates  are  the 
pair  of  specimens  figured  on  the  same  plate  (pi.  23  fig.  ib)  which  have  the  typical 
shape.  As  the  rhabdosomes  were  flexible  in  life  to  some  extent,  this  poses  the  prob- 
lem of  how  much  reliance  should  be  placed  on  rhabdosome  shape  as  a  specific 
character  in  Dicellograptus  and  other  forms  with  long  slender  stipes.  D.  moffatensis 
can  be  readily  separated  from  other  British  Dicellograptus  by  size  and  shape  of  rhab- 
dosome, D.  morrisii  Hopkinson  being  the  most  similar  but  with  a  more  open  axil. 
D.  moffatensis  var.  alabamensis  Ruedemann  1908  is  not  related  at  all  and  is  clearly 
a  Dicranograptus,  close  to  D.  brevicaulis  Elles  &  Wood  1904. 

DISTRIBUTION.  Elles  &  Wood  recorded  the  species  widely  throughout  the  British 
Isles  but  it  seems  doubtful  now  if  it  occurs  in  the  Lake  District.  The  specimen 
figured  by  Hopkinson  &  Lapworth  from  Llanvirn  (SM .  Ai740o)  is  a  poorly-preserved, 
bent  dichograptid  (O.  M.  B.  Bulman,  personal  communication).  The  specimen 
from  Abereiddy  Bay  refigured  by  Elles  &  Wood  may  well  come  from  the  locality 
referred  to  the  "  Dicranograptus  Shales  "  (Cox,  1915  :  304)  and  not  from  the 
D.  murchisoni  Shales  for  which  the  area  is  best  known.  If  this  is  so,  the  species  ranges 
from  PLlandeilo  Series  to  Caradoc  Series  (wilsoni  Zone).  It  has  been  recorded  from 
Australia  but  Thomas  (1960)  does  not  list  it  in  his  range  chart  and  so  presumably 
does  not  consider  it  to  be  present.  Linnarsson  recorded  it  from  Scania  but  Hadding 
(1913)  transferred  this  form  to  his  new  species  D.  vagus. 


Dicranograptus  clingani  Carruthers 
PI.  3,  figs.  2-4;  Fig.  3a 

1868  Dicranograptus  Clingani  Carruthers  :  132,  pi.  5,  figs.  6a-c. 

1870  Dicranograptus  Clingani  Carruthers;  Hopkinson  :  358,  pi.  16,  figs.  4a-c. 

1876  Dicranograptus  Clingani  Carruthers;  Lapworth,  pi.  3,  fig.  76. 

1877  Dicranograptus  Clingani  Carruthers;  Lapworth  :  141,  pi.  6,  fig.  43. 

1904     Dicranograptus  Clingani  Carruthers;  Elles  &  Wood  :  165,  pi.  24,  figs.  ia-1. 
1915     Dicranograptus  Clingani  Carruthers;  Hadding  :  22,  pi.  3,  figs.  1-8. 

ORIGINAL  DESCRIPTION.  "  Polypary  with  a  short  diprionidian  portion,  the  proxi- 
mal end  furnished  with  three  very  delicate  spines;  hydrothecae  forming  a  slight 
serration  along  the  margin;  twenty-one  cells  in  the  inch.  Loc.  Moffat." 

COMMENTS  ON  USAGE.  Elles  &  Wood  put  this  species  in  a  group  by  itself  on  the 
basis  of  the  thecal  characters — approximately  straight  ventral  walls  and  horizontal 


W.    CARRUTHERS'    TYPE   GRAPTOLITES  191 

apertures.  These  characters  are  clearly  seen  in  Hopkinson's  figures  and  serve  to 
differentiate  the  species  from  other  forms  with  a  short  biserial  portion. 

TYPE  MATERIAL.  Elles  &  Wood  refigured  as  "  type  specimen  "  the  original  of 
Carruthers'  fig.  6a  and  this  can  be  taken  as  a  designation  of  a  lectotype.  The 
specimen,  Q-55  (PI.  3,  figs.  3,  4),  is  from  the  Hartfell  Shales,  Hartfell  Spa. 
Carruthers'  fig.  6b,  Q.842  (PL  3,  fig.  2),  is  also  from  this  locality. 

REVISED  DESCRIPTION.  Dicranograptus  with  short  biserial  portion  consisting  of 
three  or  four  pairs  of  thecae,  and  short,  straight  uniserial  stipes  diverging  at  about 
40°.  The  biserial  portion  has  a  uniform  breadth  of  about  i  mm.  and  the  branches 
are  similarly  uniform  at  about  0-8  mm.  The  virgella  is  usually  prominent  as  a 
short  spine  and  the  first  two  thecae  may  have  sub-apertural  spines.  The  ventral 
walls  of  the  later  thecae  are  straight  and  the  apertures  are  horizontal  in  excavations 
which  occupy  about  one-third  of  the  breadth.  The  uniserial  stipes  appear  to  be 
rarely  more  than  2  cm.  long  although  Hadding  figures  a  specimen  with  stipes  nearly 
4  cm.  long.  The  distal  thecae  number  eight  to  ten  per  cm.  but  the  stipes  are  usually 
twisted  so  that  the  thecae  are  in  scalariform  view  (Fig.  3a)  and  it  is  difficult  to  decide 
the  precise  thecal  shape.  The  apertural  excavations  are  clearly  marked  by  lists  and 
it  is  probable  that  the  straight  ventral  walls  of  the  proximal  thecae  continue  in  the 
distal  ones. 


FIG.  3.  a.  Dicranograptus  clingani  Carr.  Proximal  end  of  paratype.  (3.842.  X5-  The 
right  hand  side  of  the  specimen  is  poorly  preserved,  b.  Climacograptus  minutus  Carr.  Lectotype. 
Q8o.  X5-  c.  Cryptograptus  tricornis  Carr.  Proximal  end  of  lectotype  showing  basal  spines. 
X5- 


DISTRIBUTION.  The  species  is  common  in  the  Lower  Hartfell  Shales  of  the  Moffat 
area  and  is  also  found  at  Conway.  Elles  &  Wood  recorded  it  from  equivalent  beds 
in  Ireland.  It  is  found  in  Scandinavia  but  has  not  been  recorded  from  North 
America.  Thomas  (1960)  gives  records  but  omits  it  from  his  range  chart,  so  casting 
doubt  on  the  records.  The  type  specimen  has  no  associates  but  Elles  &  Wood 
recorded  it  only  from  its  own  zonal  association. 


Climacograptus  minimus  (Carruthers) 
PI.  4,  fig.  3;  Fig.  4c 

1868     Diplograpsus  minimus  Carruthers  :  74,  130,  pi.  5,  figs.  iaa,  b. 
?igo6  Climacograptus  minimus  (Carruthers)  Elles  &  Wood  :  191,  pi.  27,  figs.  xa-g. 


192  W.    CARRUTHERS'    TYPE   GRAPTOLITES 

ORIGINAL  DESCRIPTION.  "  This  agrees  with  D.  pristis  in  general  appearance,  and 
in  the  form  and  arrangement  of  the  cells,  except  that  the  whole  polypary  and  all  its 
parts  are  so  very  small.  Had  I  met  with  only  a  few  specimens,  I  would  have  con- 
sidered it  as  merely  an  accidental  variety,  but  I  have  seen  so  many,  all  agreeing  in 
size,  that  I  cannot  doubt  that  it  is  a  good  species,  especially  as  young  specimens 
of  D.  pristis  early  attain  their  full  breadth,  and  the  increase  of  the  polypary  is  by 
addition  to  its  distal  end,  and  not  to  the  size  of  the  already  formed  hydrothecae, 
just  as  in  the  living  Sertulariadae.  About  thirty-eight  cells  to  one  inch.  Loc.  Moffat." 


FIG.  4.  Figures  to  show  Carruthers'  clear  distinction  between  his  two  small  species  of 
diplograptid.  a,  Copy  of  original  MS  drawing  of  "  Diplograptus  minimus  "  which  has 
appended  note  "38  to  inch  ".  b,  Similar  copy  of  Climacograptus  minutus  which  has  notes 
"  32  to  40  to  an  inch,  nearly  opposite  ". 

COMMENTS  ON  USAGE.  In  his  1868  paper,  Carruthers  clearly  distinguished  two 
small  biserial  species  and  Fig.  4,  taken  from  his  notebook,  illustrates  his  ideas  of 
the  differences,  one  clearly  climacograptid,  the  other  what  he  called  diplograptid 
(now  orthograptid) .  His  type  slab  however  shows  a  large  number  of  small,  poorly- 
preserved  rhabdosomes  which  appear  to  be  almost  all  climacograptid  and  Elles  & 
Wood  certainly  regarded  his  species  in  this  light.  In  his  description  of  the  species 
(1868  :  130)  he  called  it  Diplograpsus  minutus  although  elsewhere,  in  the  explanation 
of  the  plate  published  in  the  first  part  of  the  paper  (1868  :  74),  he  called  it  D.  mini- 
mus, agreeing  with  his  MS  notes,  and  usage  of  this  prior  name  avoids  the  homonym 
which  results  from  the  transference  of  the  species  to  Climacograptus. 

Elles  &  Wood  did  not  re-figure  Carruthers'  material,  and  their  specimens,  from 
the  Hartfell  Shales  (clingani  and  linearis  Zones),  reach  a  breadth  of  2  mm. 
Carruthers'  specimens  on  the  type  slab  are  only  about  I  mm.  wide  and  are  associated 
with  some  slender  uniserial  stipes,  which  look  very  like  Monograptus  sp.  (atavus  or 
acinaces  type).  It  thus  seems  probable  that  Elles  &  Wood's  species  is  not  the  same 
as  Carruthers'  but  it  is  clear  from  the  foreign  references  (e.g.  Ross  &  Berry  1963, 
Obyt  &  Sobolevskaya  1964)  that  it  is  Elles  &  Wood's  species  which  is  nowadays 


W.   CARRUTHERS1   TYPE   GRAPTOLITES  193 

recognized  under  this  name.  Carruthers'  original  notes  are  of  little  use  since  the 
only  measurements  he  gives  are  the  thecal  numbers  per  inch.  A  sketch  of  "  Diplogr. 
minimus  "  clearly  shows  thecae  of  an  orthograptid  type  in  accordance  with  his 
comparison  of  it  with  D.  pristis  but,  as  mentioned  above,  the  common  form  on  the 
type  slab  appears  to  have  climacograptid  thecae.  The  species  must  remain  for  the 
moment  in  an  unsatisfactory  state  since  a  proper  appraisal  of  Elles  &  Wood's  form 
must  await  critical  re-examination  of  the  Upper  Ordovician  climacograptids. 

TYPE  MATERIAL.     Q.82,  a  slab  crowded  with  poorly-preserved  specimens. 

REVISED  DESCRIPTION.  The  rhabdosome  is  about  i  mm.  wide  and  up  to  10  mm. 
long.  Thecal  details  are  obscure. 

HORIZON.  As  noted  above,  the  type  slab  shows  slender  uniserial  stipes  very 
similar  to  monograptids  of  the  vesiculosus  and  cyphus  Zones,  in  contrast  with  the 
description  by  Elles  &  Wood  of  the  species  from  the  Hartfell  Shales  (dingani  and 
linearis  Zones) . 

Climacograptus  minutus  Carruthers 
PL  4,  fig.  i,  ;  Figs  3b,  4b 

1868     Climacograptus  minutus  Carruthers  :  132,  pi.  5,  fig.  loa. 
?i9o6  Climacograptus  minutus  Carruthers;  Elles  &  Wood  :  211,  pi.  27,  figs.  i2a-c. 

ORIGINAL  DESCRIPTION.  '  This  is  a  very  minute  but  well-marked  species,  never 
attaining  a  greater  size  then  represented  on  the  Plate.  There  are  at  the  rate  of  from 
thirty-two  to  forty  cells  in  the  space  of  an  inch.  Loc.  Moffat." 

COMMENTS  ON  USAGE.  This  species  does  not  seem  to  have  been  widely  recorded, 
probably  because  of  its  unsatisfactory  nature.  Carruthers'  original  specimens  are 
of  widely  differing  widths  and  Elles  &  Wood  based  their  account  of  the  species  on 
other  material  from  the  British  Museum  (Natural  History).  Packham  (1962) 
revised  some  of  the  British  Silurian  diplograptids  but  did  not  discuss  this  species. 
His  C.  tangshanensis  linearis  appears  to  be  close  to  Carruthers'  fig.  lob  (1868). 

TYPE  MATERIAL.  Carruthers'  original  specimens  can  be  recognized  and  appear  to 
be  of  two  different  forms.  His  fig.  loa  (Q.8o)  is  here  selected  as  lectotype  (PL  4, 
fig.  i) .  The  original  of  fig.  lob  (Q .  1372)  is  a  narrower  form  and  fits  well  with 
Climacograptus  scalaris  miserabilis  Elles  &  Wood.  Elles  &  Wood's  figured  material 
(Q .  849)  is  all  somewhat  distorted  and  it  is  clear  that  they  did  not  use  Carruthers' 
original  specimens  for  their  description,  probably  through  some  confusion  with 
"  Diplogr aptus  minutus  ". 

REVISED  DESCRIPTION.  Rhabdosome  7  mm.  long.  1-2  mm.  broad;  thecae  thirteen 
per  cm.,  with  large  excavations  occupying  about  one  quarter  of  the  breadth  of  the 
rhabdosome  and  about  equal  to  the  length  of  the  free  ventral  wall.  The  apertures 
are  nearly  opposite  each  other  (as  recorded  in  Carruthers'  notes)  and  not  alternate 
as  stated  by  Elles  &  Wood.  The  proximal  end  of  the  type  specimen  is  poorly 
preserved  but  there  appears  to  be  a  stout  virgula  which  is  prolonged  beyond  the 
distal  end  of  the  rhabdosome.  The  type  specimen  is  completely  flattened  and  it  is 
not  possible  to  make  out  any  details  of  a  median  septum. 


194  W.    CARRUTHERS'   TYPE   GRAPTOLITES 

The  uniform  breadth  of  the  rhabdosome  and  the  large  opposite  excavations  com- 
bine to  make  this  form  quite  distinct  from  other  British  climacograptids.  Un- 
fortunately the  range  of  variation  is  not  known  and  since  the  type  specimen  has  no 
associates,  its  precise  horizon  is  also  doubtful.  It  remains  therefore  an  unsatis- 
factory species. 

Cryptograptus  tricornis  (Carruthers) 

PL  4,  figs.  4-6;  Fig.  30 
1858     Diplograpsus  tricornis  Carruthers  :  468,  fig.  2. 


1859     Diplograpsus  tricornis  Carruthers;  Carruthers 

1867  Diplograpsus  tricornis  Carruthers;  Carruthers 

1868  Diplograpsus  tricornis  Carruthers;  Carruthers 


25,  fig-  2. 

290,  pi.  i,  figs.  loa-d. 

131,  pi.  5,  fig.  ua,  b. 


1880  Cryptograptus  tricornis  (Carruthers)  Lapworth  :  171,  pi.  5,  figs.  27a-e. 

1908  Cryptograptus  tricornis  (Carruthers);  Elles  &  Wood  :  296,  pi.  32,  figs.  I2a-d. 

1908  Cryptograptus  tricornis  (Carruthers) ;  Ruedemann  :  443,  pi.  28,  figs.  1-4. 

1913  Cryptograptus  tricornis  (Carruthers);  Hadding  :  40,  pi.  2,  figs.  13-14. 

1915  Cryptograptus  tricornis  (Carruthers);  Hadding  :  325,  pi.  6,  fig.  15. 

1934  Cryptograptus  tricornis  (Carruthers);  Hsu  :  87,  pi.  6,  figs.  I3a-m. 

1937  Cryptograptus  tricornis  (Carruthers) ;  Bulman  :  5,  t-fig.  8. 

1945  Cryptograptus  tricornis  (Carruthers);  Bulman  :  29,  pi.  2,  figs.  r-8. 

1960  Cryptograptus  tricornis  (Carruthers);  Thomas,  pi.  6,  fig.  69. 

ORIGINAL  DESCRIPTION.  "  This  species  can  be  readily  distinguished  by  the  three 
spines  which  adorn  its  base,  and  which  are  almost  always  preserved.  The  central 
spine  is  a  continuation  of  the  line  of  the  axis ;  it  is  shorter  than  the  lateral  ones  .  .  . 
The  polypidom  is  more  slender  than  in  D.  foliaceus,  which  in  general  outline  it  some- 
what resembles.  The  axis  is  slender,  and  produced  beyond  the  other  parts  of  the 
fossil.  The  cell-walls  are  well  marked,  extending  upwards  from  the  axis  to  the 
boundary  of  the  fossil.  Each  cell  forms  a  rhomb  whose  outer  border  is  slightly 
indented,  giving  the  boundary  of  the  fossil  a  faintly  serrated  aspect.  When  the 
fossil  is  preserved  so  as  to  show  the  serratures,  the  spines  are  so  compressed  that  the 
central  one  is  almost  or  altogether  lost.  When  the  spines  are  well  preserved  and  in 
the  position  described,  no  traces  of  the  individual  cells  are  discernible ;  the  boundary 
of  the  fossil  is  an  unbroken  line  ". 

In  1868,  Carruthers  added  "  When  I  described  this  species  I  had  not  detected  the 
mouths  of  the  cells  in  those  specimens  in  which  they  should  have  been  shown  on  the 
upper  surface.  In  more  perfectly  preserved  specimens  since  obtained  these  have 
been  beautifully  shown  ". 

COMMENTS  ON  USAGE.  This  species  has  been  widely  recognized  as  it  is  easily 
identified  from  the  original  description  and  figures.  Lapworth  (1880)  discussed  the 
varying  appearance  of  the  thecae  at  some  length  and  introduced  the  genus  Crypto- 
graptus for  this  and  allied  species.  Hadding  (1915)  showed  by  comparison  with 
Glossograptus  that  the  two  stipes  were  in  lateral  contact  (the  monopleural  arrange- 
ment of  Jaanusson  1960)  and  removed  the  genus  from  the  Diplograptidae.  The 
structure  of  the  proximal  end  was  not,  however,  elucidated  until  1938  when  Bulman 
described  isolated  specimens  from  the  Balclatchie  Limestone  and  the  two  different 


W.    CARRUTHERS'    TYPE   GRAPTOLITES  195 

aspects  of  the  basal  spines  noted  by  Carruthers  was  explained.     A  number  of  varieties 
have  been  described,  differing  mainly  in  the  breadth  of  the  rhabdosome. 

DIAGNOSIS.  Rhabdosome  biserial,  monopleural,  2-4  cm.  long  widest  at  base 
when  preserved  in  lateral  view,  maximum  width  1-8  mm.  but  typically  narrower. 
Thecae  ten  to  twelve  per  cm.  Basal  spines  conspicuous,  but  short. 

TYPE  MATERIAL.  BM(NH)  (^.1299,  presented  to  the  museum  in  1860,  is  almost 
certainly  Carruthers'  original  slab.  It  is  crowded  with  specimens  up  to  35  mm. 
long  and  1-6  mm.  broad  but  it  is  not  possible  to  recognize  the  original  figured  speci- 
mens. Accordingly,  one  of  the  better  specimens  has  been  selected  as  lectotype 
(PI.  4,  fig.  4).  The  association  on  the  slab  includes  abundant  Corynoides  calicularis 
Nich.  Similar  pieces  in  the  Lap  worth  Collection  (Birmingham  University)  are 
labelled  "  The  Cornice,  Hartfell  "  and,  although  this  is  not  marked  on  Lapworths' 
published  map  of  Hartfell,  it  would  appear  from  the  text  that  the  horizon  is  lowest 
Hartfell,  zone  of  Climacograptus  wilsoni. 

DESCRIPTION.  No  detailed  description  is  needed  since  that  in  the  Monograph 
is  satisfactory  and  has  been  recently  supplemented  by  Bulman's  detailed  account 
of  isolated  specimens.  There  seems  to  be  some  variation  in  the  thecal  count,  speci- 
mens from  Girvan  having  consistently  higher  (twelve  to  sixteen  per  cm.)  counts. 
Elles  &  Wood  described  the  variety  schaeferi  which  Lapworth  had  figured  earlier 
and  claimed  that  it  was  "  somewhat  wider  than  the  typical  form  ".  Examination 
of  the  material  in  Lapworth's  collection  does  not  confirm  this  difference  but  there 
seems  to  be  a  difference  in  thecal  shape  and  the  basal  spines  are  not  conspicuous. 

DISTRIBUTION.  This  species  has  been  recorded  from  all  continents  and  some  of 
the  more  recent  records  are  given  in  the  synonymy.  The  stratigraphic  range  was 
given  by  Elles  &  Wood  as  Arenig  Series  (extensus  Zone)  to  Caradoc  Series  (clingani 
Zone),  a  longer  range  than  any  other  Ordovician  species,  and  it  is  possible  that 
critical  examination  of  a  large  number  of  specimens  from  the  lower  horizons  might 
show  them  to  be  distinct  from  the  typical  form  from  the  early  Caradoc.  Sherrard 
(1954)  records  it  from  the  zone  of  P.  linearis  but  no  illustration  is  given  to  confirm 
this. 

Monograptus  clingani  (Carruthers) 

PL  5,  figs.  1-5 

1867  Graptolithus  Clingani  Carruthers  :  369,  pi.  2,  fig.  8. 

1868  Graptolithus  Clingani  Carruthers;  Carruthers  :  127,  pi.  5,  figs,  iga,  b. 
1876     Monograptus  Clingani  (Carruthers);  Lapworth,  pi.  i,  fig.  24. 

18760  Monograptus  Clingani  (Carruthers);  Lapworth  :  501,  pi.  20,  figs.  sa-c. 

non   1897  Monograptus  Clingani  (Carruthers);  Perner,  pi.  n,  figs.  15-17. 

1913  Monograptus  Clingani  (Carruthers);  Elles  &  Wood  :  463,  pi.  46,  figs.  na-f. 

1951  Monograptus  clingani  (Carruthers);  Bulman  :  322,  t-fig.  5. 

1956  Monograptus  clingani  (Carruthers);  Bondarenko  &  Keller  :  91,  t-fig.  2. 

ORIGINAL  DESCRIPTIONS.  1867.  "...  a  beautiful  small  species,  which  at  first 
I  referred  to  G.  millepeda,  M'Coy,  but  that  species  is  certainly  the  proximal  end  of 
G.  Becki,  and  this  differs  from  it  in  having  a  very  broad  common  base,  from  which 
the  hydrothecae  rise  ."  1868.  "  Polypary,  small  and  arcuate,  with  a  broad  common 


I96  W.    CARRUTHERS'   TYPE   GRAPTOLITES 

canal,  and  slender  somewhat  recurved  hydrothecae.  This  beautiful  little  graptolite 
I  long  supposed  to  be  only  the  proximal  portion  of  some  other  species,  but  the  large 
number  I  have  met  with,  all  equally  perfect,  none  larger  than  fig.  iga,  and  many 
showing  the  prolongation  of  the  axis  beyond  the  distal  end,  together  with  the  great 
breadth  of  the  common  canal  (forming  two-thirds  of  the  breadth  of  the  whole 
polypary),  unlike  the  early  portion  or  proximal  fragment  of  any  graptolite  with 
which  I  am  acquainted,  have  induced  me  to  consider  it  a  good  species  ..." 

COMMENTS  ON  USAGE.  Carruthers'  descriptions  gave  no  real  details  of  the  species. 
From  the  syntypes  it  appears  that  two  different  species  may  be  confused  but  as 
both  are  young  rhabdosomes  it  is  not  possible  to  be  sure.  The  characters  of  the 
proximal  thecae  are  not  determinable  from  the  lectotype,  chosen  by  Pfibyl  (1948), 
which  is  completely  flattened  but  they  appear  to  be  of  the  priodon  type  with  well 
marked  hooks  to  the  apertures  (PI.  5,  fig.  5).  The  distal  thecae  are  similar.  Lapworth 
recorded  a  "  Clingani  Band  ",  some  6  inches  thick,  in  his  Dobbs  Linn  sections  and 
numerous  specimens  in  his  collection  bear  this  label.  In  a  recent  resurvey  of  the 
section,  Toghill  (1965)  recognized  the  unit  again  but  recorded  M.  dingani  from  a 
wider  horizon. 

TYPE  MATERIAL.  The  specimen  (Q.87)  figured  by  Carruthers  as  fig.  iga  was 
referred  to  as  "  type  specimen  "  by  Elles  &  Wood  in  the  explanation  of  their  plate 
and  the  second  specimen  called  "  co-type  "  (Q.8-4),  leaving  them  perhaps  of  equal 
status.  Pfibyl  (1948)  however  clearly  stated  that  Carruthers'  fig.  iga  is  the  lecto- 
type which  is  unfortunate  as  this  specimen  has  a  broader  common  canal  than  in 
most  of  the  other  specimens  referred  to  the  species.  This  may  be  only  a  preserva- 
tional  feature  as  specimens  in  the  Lapworth  collection  from  DufTkinnel,  the  type 
locality,  are  all  of  the  common  type  in  which  the  hooks  occupy  at  least  half  of  the 
breadth  of  the  rhabdosome. 

REVISED  DESCRIPTION.  Rhabdosome  dorsally  curved  at  the  proximal  end,  dis- 
tally  becoming  more  or  less  straight;  widening  from  an  initial  breadth  of  0-6  mm. 
to  a  maximum  of  about  1-5  mm.  within  the  first  6  or  7  mm.  The  thecae  are  of 
priodon  type  with  about  one-third  of  the  length  involved  in  the  hook  which  occupies 
half  of  the  breadth  of  the  stipe.  The  shape  of  the  thecae  varies  a  good  deal  with 
the  type  of  preservation  but  there  appears  to  be  no  overlap  of  the  thecae  (Bulman 
1951,  fig.  5),  the  broad  prothecal  portion  occupying  the  whole  of  the  breadth  of  the 
rhabdosome. 

DISTRIBUTION.  Lapworth  recorded  the  species  in  abundance  from  a  6-inch  band 
within  the  sedgwickii  Zone  at  Dobbs  Linn,  and  Marr  &  Nicholson  (1888)  similarly 
had  a  M.  dingani  Band  above  their  convolutus  Zone  in  the  Lake  District.  Elles  & 
Wood  apparently  extended  the  convolutus  Zone  to  include  these  horizons  and  also 
recorded  the  species  from  the  gregarius  Zone  below  in  their  summary  table. 

Monograptus  intermedius  (Carruthers) 
PI.  5,  fig.  6;  Fig.  5 

1868     Graptolithus  intermedius  Carruthers  :  126,  pi.  5,  fig.  18. 
?i8y6a  Monograptus  intermedius  (Carruthers)  Lapworth  :  316,  pi.  10,  figs.  loa,  d. 


W.    CARRUTHERS'    TYPE   GRAPTOLITES  197 

non   1913     Monograptus  intermedius  (Carruthers) ;  Elles  &  Wood  :  485,  pi.  49,  figs.  sa-c. 
?I952     Spirograptus  intermedius  (Carruthers);  Miinch,  pi.  37b,  figs,  ya,  b. 

ORIGINAL  DESCRIPTION.  "  Polypary  slender;  proximal  end  composed  of  a  slender 
canal  with  distant,  isolated,  and  very  small  hydrothecae ;  adult  hydrothecae,  short, 
triangular,  the  upper  margin  of  the  cell  forming  an  acute  angle  with  the  common 
canal.  About  twenty-six  cells  to  an  inch.  This  species  differs  from  G.  Nilssoni, 
G.  tenuis  and  G.  Hisingeri  in  the  form  of  the  cells,  and  from  the  last  also  in  the  slender 
common  canal.  Perhaps  Portlock's  figure  6a.  pi.  19  of  his  Report  belongs  to  this 
species.  Loc.  Moffat.  " 

COMMENTS  ON  USAGE.  It  is  clear  from  the  specimens  in  Lap  worth's  collection 
that  he  included  under  this  name  a  number  of  slender  forms  which  do  not  agree  with 
Carruthers'  type  specimen.  Elles  &  Wood  used  only  one  slab  from  the  Lap  worth 
Collection  for  their  redescription  of  this  species  and  the  crowded  fragments  on  it 
show  thecae  which  are  much  more  slender  than  those  of  the  type  specimen  in  which 
the  thecae  are  nearer  those  of  M.  involutus.  Lapworth  considered  that  M.  acutus 
(Hopkinson  1872)  was  the  same  as  M.  intermedius  but  Hopkinson  quite  clearly 
differentiated  them.  Most  later  workers  have,  however,  apparently  relied  on  Elles 
&  Wood's  account. 

TYPE  MATERIAL.     Holotype,  BM(NH)  Q.88,  Birkhill  Shales,  Moffat. 

REVISED  DESCRIPTION.  Rhabdosome  arcuate,  with  very  slender  proximal  end, 
widening  from  about  0-3  mm.  to  a  maximum  of  about  i-o  mm.  in  the  distal  part. 
Thecae  of  the  spiralis  type,  the  proximal  ones  with  a  slender  prothecal  portion  and 
an  abruptly  widened  apertural  region,  distal  ones  more  uniformly  widening,  about 
ten  thecae  per  cm.  The  proximal  end  of  the  type  specimen  is  poorly  preserved  but 
appears  to  be  comparatively  straight  so  that  the  rhabdosome  is  not  truly  spirally 


c 


Fig.  5.     Sketches  of  proximal,  medial  and  distal  thecae  of  Monograptus  intermedius,  holotype, 
Q.88,  x  loapprox. 


curved  in  the  way  shown  by  M.  communis,  etc.  On  thecal  shape,  however,  Miinch's 
assignation  of  the  species  to  Spirograptus  is  quite  possible.  In  general  shape  and 
the  slender  proximal  end,  the  species  is  close  to  Spirograptus  planus  (Barrande)  as 
figured  by  Pfibyl  (1946,  pi.  8,  figs.  6-8)  but  that  species  rapidly  reaches  a  breadth 


GEOL.  17,  4 


16 


ig8  W.    CARRUTHERS'    TYPE   GRAPTOLITES 

of  more  than  1-5  mm.  and  the  rhabdosome  continues  to  spiral  distally.     The  species 
is  here  retained  in  Monograptus  sensu  lato. 

DISTRIBUTION.  The  type  specimen  is  associated  with  a  proximal  end  of  M. 
triangulatus-type  and  seems  to  be  from  the  zone  of  M.  gregarius.  Most  of  the  records 
appear  to  be  based  on  Elles  &  Wood's  description  and  hence  are  misidentified  so  that 
no  distribution  can  be  given  at  present. 

Diversograptus?  capillaris  (Carruthers) 
Fig.  6a,  b. 

1867  Rastrites  capillaris  Carruthers  :  368,  pi.  2,  fig.  10. 

1868  Rastrites  capillaris  Carruthers;  Carruthers  :  126,  pi.  5,  fig.  16. 
non   i8y6a  Rastrites  capillaris  Carruthers;  Lapworth  :  314,  pi.  10,  fig.  4. 

i8j6a.  Monograptus  attenuatus  Hopkinson;  Lapworth  :  317,  pi.  10,  fig.  9. 
?i8Q7     Monograptus  (Rastrites)  gemmatus  (Barrande);  Perner,  t-fig.  26. 
1913     Monograptus  gemmatus  (Barrande);  Elles  &  Wood  :  436,  pi.  43,  figs.  5a-d,  ?e. 

1952  Monograptus  (Psubgenus)  capillaris  (Carruthers) ;  Boudek  &  Pfibyl  :  206,  t-fig.  4d-f. 

1953  Diversograptus  capillaris  capillaris  (Carruthers)  Bou£ek  &  Pfibyl  :  496,  558,  pi.  i, 

figs.  1-3. 

ORIGINAL  DESCRIPTION.  "  Common  tube  very  slender,  with  short  isolated  tri- 
angular hydrothecae,  their  base  of  attachment  to  the  common  canal  as  long  or  longer 
than  their  depth.  About  sixteen  cells  to  an  inch.  Loc.  Moffat.  Richter  figures 
this  specimen  in  Zeitschr.  Deutsch.  Geol.  Gesellsch.,V,  1853,  Tab.  xii,  fig.  34a,  referring 
it  to  R.  gemmatus  Barr.,  which  is  very  different,  and  of  which  his  fig.  34b  is  a  good 
representation  ". 

There  has  been  considerable  confusion  over  this  slender  species.  Carruthers' 
original  description  is  not  clear  and  Hopkinson  (1872)  described  M.  attenuatus  as 
another  slender  species  without  making  detailed  comparison  with  capillaris. 
Lapworth  took  the  generic  assignment  to  Rastrites  as  correct  in  spite  of  Carruthers' 
statement  that  the  base  of  the  triangular  theca  was  longer  than  its  height.  From 
his  own  larger  and  more  detailed  collections  from  Dobbs  Linn,  Lapworth  (i876a) 
redescribed  M.  attenuatus  and  a  "  R.  capillaris  "  which  is  a  true  rastritid  (see  Fig.  6d). 
Perner  (1897)  redescribed  M .  gemmatus  (Barrande)  which  has  distinctly  hooked 
thecae  (Fig.  6c)  but  while  he  gave  only  a  natural  size  figure  of  the  holotype  he  also 
gave  an  enlarged  drawing  under  the  same  name  of  a  form  with  simpler  thecae. 
It  was  apparently  this  enlarged  figure  which  Elles  &  Wood  took  as  typical  of  M . 
gemmatus  and  accordingly  included  M .  capillaris  Carr.  and  M.  attenuatus  Hopkinson 
in  its  synonymy.  They  do  not  seem  to  have  considered  Lapworth's  drawings  or 
specimens  of  "  R.  capillaris  "  (which  are  clearly  distinct  from  Carruthers'  types) 
although  they  refigured  his  specimens  of  M.  attenuatus.  Boucek  &  Pfibyl  (1952) 
re-examined  the  slender  monograptids  and  showed  clearly  that  Carruthers'  and  Hop- 
kinson's  species  were  distinct  from  M.  gemmatus.  In  the  following  year  they  assigned 
diversograptid  forms  to  D.  capillaris  and  the  species  has  been  widely  recorded  as 
such  in  Central  European  work  since  1952.  Unfortunately  British  material  has  so 
far  provided  very  few  examples  of  diversograptid  rhabdosomes  while  the  German 
material  (e.g.  Manck  1924)  is  usually  poorly  preserved  as  far  as  thecal  characters 


W.    CARRUTHERS'   TYPE   GRAPTOLITES 


199 


are  concerned.     It  is  thus  doubtful  if  the  equivalence  of  the  British  and  Continental 
specimens  can  be  regarded  as  proven. 

TYPE  MATERIAL.     Holotype  Q .  86,  from  the  Birkhill  Shales,  Moffat. 

REVISED  DESCRIPTION.  Rhabdosome  only  known  as  fragments,  up  to  5  cm.  long, 
very  slender,  maximum  breadth  0-4  mm. ;  thecae  elongate  triangular,  about  seven 
per  cm.,  widening  gradually  from  the  initial  prothecal  portion,  probably  with  no 
overlap.  Apertures  of  spiralis-type  but  no  spines  seen. 

DISCUSSION.  The  type  material  is  very  poorly  preserved  and  other  slender 
British  specimens  are  comparatively  rare  in  collections,  generally  only  found  as 
small  fragments.  It  is  difficult  to  match  these  satisfactorily  with  the  German  and 
Bohemian  material  which,  although  more  frequently  showing  complete  rhabdosomes, 
is  usually  quite  flattened  and  does  not  show  thecal  characters  well.  The  slender 


\ 


FIG.  6.  Diversograptus?  capillaris  Carr.  a.  Copy  from  original  drawing  for  Elles  &  Wood, 
pi.  43,  fig.  5d.  Holotype,  Q.86.  X5-  b.  DP  capillaris  Carr.  Two  thecae  for  comparison  with 
M.  gemmatus.  Q86.  X  10.  c.  M.  gemmatus  Barr.  Fragment  of  stipe  with  2  thecae  showing 
hooked  apertures.  R.  maximus  Beds,  Belcraig  Burn,  Moffat.  BU  304.  x  10.  d.  Rastrites 
"capillaris"  Lapw.  (?  =  R.  spina  Richter)  Duffkinnell.  611305.  Xio. 

GEOL.  17,  4  l6§ 


200  W.   CARRUTHERS'   TYPE  GRAPTOLITES 

stipes  appear  to  have  been  fairly  flexible  and  the  shape  of  the  thecae  is  variable 
along  the  stipe,  depending  probably  on  the  twisting  of  the  stipe.  Although  it  is 
quite  clear  that  D.P  capillaris  is  distinct  from  M.  gemmatus  (Barrande),  its  relation- 
ship to  M.  attenuatus  Hopkinson  remains  doubtful.  The  specimens  which  Lapworth 
figured  as  M.  attenuatus  are  obviously  close  to  Carruthers'  capillaris  but  these  may 
be  different  from  Hopkinson's  types  which  have  not  yet  been  traced. 

DISTRIBUTION.  The  type  material  is  associated  with  Rastrites  cf.  peregrinus, 
indicating  a  Middle  to  upper  Birkhill  age  which  agrees  with  the  records  from  the 
rest  of  Europe. 


Rastrites  maximus  Carruthers 
PL  5,  fig.  7;  Fig.  ya 

i86ya  Rastrites  maximus  Carruthers  :  540,  fig.  90(6) 

1868     Rastrites  maximus  Carruthers  :  126,  pi.  5,  fig.  14. 

i8y6a  Rastrites  maximus  Carruthers;  Lapworth,  pi.  i,  fig.  i. 

1907     Rastrites  maximus  Carruthers;   Tornquist  :   15,   pi.    2,   figs.    28,   29,    ?2y,    ?3O~33, 

?pl.  3,  fig.  i. 

1914     Rastrites  maximus  Carruthers;  Elles  &  Wood  :  494,  pi.  50,  figs.  6a-d,  ?6e. 
1941     Rastrites  maximus  Carruthers ;  Pfibyl  :  15. 
?i955     Rastrites  maximus  Carruthers;  Malinowska  :  57,  pi.  n,  fig.  4. 
1967     Rastrites  maximus  Carruthers;  Schauer  :  184,  pi.  6,  fig.  i. 

ORIGINAL  DESCRIPTION.  "  Common  tube  slender,  supporting  very  large  hydro- 
thecae  at  wide  intervals.  Hydrothecae  nearly  half  an  inch  long,  somewhat  enlarged 
towards  the  apex,  and  furnished  at  the  base  with  a  triangular  corneus  membrane 
extending  a  short  distance  up  the  margin  of  the  cells.  About  six  cells  in  an  inch. 
Loc.  Moffat.  " 

COMMENTS  ON  USAGE.  This  species  has  been  recorded  from  a  few  places  in  Europe 
but  appears  to  be  replaced  by  R.  linnaei  Barrande  in  Central  Europe.  The  large 
rastritids  are  usually  found  in  a  very  fragmentary  state  owing  to  the  extremely 
slender  common  canal,  and  isolated  thecae  cannot  always  be  identified  specifically 
since  the  thecal  length /interspace  ratio  appears  to  be  one  of  the  most  useful  characters 
for  discrimination.  All  the  large  forms,  however,  appear  to  occur  about  the  same 
horizon  and  so  are  useful  for  stratigraphic  purposes. 

Although  the  species  has  been  widely  known  and  figured  by  Tornquist  (1907) 
and  other  authors,  no  proximal  ends  appear  to  have  been  conclusively  demonstrated. 
Most  of  the  so-called  "  young  specimens  "  have  short,  well-spaced  thecae  which 
may  be  broken  or  represent  other  species  such  as  R.  distans.  Both  Tornquist  and 
Elles  &  Wood  gave  measurements  for  the  first  few  thecae  and  the  interspaces  be- 
tween them  but  examination  of  a  large  number  of  specimens  from  the  Moffat  area 
and  from  the  Crossfell  Inlier,  Northern  England,  shows  clearly  that  the  long  thecae 
of  the  adult  develop  within  the  first  centimetre  much  more  rapidly  than  was  generally 
thought.  Unfortunately  few  specimens  are  found  with  more  than  a  few  proximal 
thecae  and  it  is  difficult  to  separate  R.  maximus  from  R.  linnaei  and  R.  distans 
prior  to  the  four  or  five  theca  stage. 


W.    CARRUTHERS'   TYPE   GRAPTOLITES 


FIG.  7.     Proximal  ends  of  Rastrites  spp. 
linnaei  Barr.,  BU  1700.     Both,  x  5  aprox. 


a,  R.  maximus  Carr.,  neotype,  BU  1692.      b,  R. 


Table  I  gives  the  dimensions  of  some  of  these  rhabdosomes  but  the  data  are  as 
yet  insufficient  for  a  statistical  analysis.  The  positions  of  theca  i  and  th.2  are  the 
same  in  relation  to  the  sicula  in  the  three  species,  with  th2  leaving  the  common 
canal  at  the  apex  of  the  sicula. 


R.  maximus 
Tornquist  1907 
Elles  &  Wood  1913 
Neotype,  BU  1692 
Q.i39oBM(NH) 

R.  linnaei 

BU  1700  (Elles  &  Wood, 
pi.  5 1,  fig.  ib) 

R.  carnicus 
Seelmeier  1936 


TABLE  I 

length  of  theca 

r 

I 

2             3 

4 

I  -O 

2-7         6-2 

1-7 

J'5 

3'5         5-o 

•  —  • 

I-4 

5-3       10-2 

12-0 

i  -i 

5'3       I0'4 

10  + 

thecal    interspace 

. A 

I 

1/2          2/3          3/4 
i -o          1-4          1-7 


i • i          3-2          4-8         6-0 


1-2  5-0 


0-8 
0-8 


0-6 


1-6 


4'5 
4-0 


(0-1—0-3) 


TYPE  MATERIAL.  Carruthers'  type  specimen  cannot  be  traced  in  the  British 
Museum  (Natural  History)  collections  and  although  Q .  1390  bears  some  resemblance 
to  Carruthers'  figure  there  are  no  details  of  locality  or  horizon  for  it.  Accordingly 
a  neotype  is  here  proposed,  chosen  from  the  specimens  figured  by  Elles  &  Wood. 
The  specimen,  6.11.1692  (Elles  &  Wood,  pi.  50,  fig.  6b)  is  from  the  Upper  Birkhill 
Shales  of  Riskinhope  Burn,  Yarrow,  Peeblesshire.  (For  stratigraphic  details  see 
LapworthiSyS  :  272). 

REVISED  DESCRIPTION.  Adult  thecae  more  than  10  mm.  long  (maximum  observed 
1-8  cm.)  with  interspaces  about  5  mm;  common  canal  very  slender;  base  of  theca 
characteristically  enlarged  with  the  proximal  side  at  right  angles  to  the  common 
canal  and  the  distal  side  somewhat  oblique.  Thecal  apertures  with  a  well-defined 
hook. 

DISTRIBUTION.  The  species  is  known  from  many  localities  in  the  Moffat  area 
and  was  recorded  by  Elles  &  Wood  from  the  Lake  District  and  North  Wales. 


202  W.    CARRUTHERS'   TYPE   GRAPTOLITES 

Records  from  Sweden  and  Central  Europe  are  doubtful  although,  as  noted  above, 
it  is  often  impossible  to  determine  specificially  the  fragmentary  large  rastritids. 
Amongst  the  Bohemian  material  in  the  Lapworth  Collection  there  are  no  specimens 
approaching  the  typical  form  in  length  and  spacing  of  the  adult  thecae.  Recent 
records  from  Belgium  (Legrand  1962)  and  Portugal  (Romariz  1962)  give  no  illustra- 
tions or  dimensions  and  require  confirmation.  The  specimen  figured  by  Malinowska 
(1955)  has  a  broad  common  canal  although  the  thecae  appear  to  be  long  enough. 
Schauer  (1967)  has  figured  a  good  specimen  from  Germany. 

Cyrtograptus  murchisoni  Carruthers 
PI.  5,  fig.  8,  9;  Fig.  8 

i867a  Cyrtograpsus  Murchisonii  Carruthers  :  540,  foss.  90,  fig.  i. 

1868  Cyrtograpsus  Murchisonii  Carruthers;  Carruthers  :  128,  pi.  5,  figs.  lya,  b. 

PiSgg  Cyrtograptus  Murchisoni  Carruthers;  Perner  :  21,  t-fig.  28. 

1900  Cyrtograptus  Murchisoni  Carruthers;  Elles,  pi.  24,  fig.  6. 

1914  Cyrtograptus  Murchisoni  Carruthers;  Elles  &  Wood  :  505,  pi.  51,  figs.  3a-c. 

?I933  Cyrtograptus  murchisoni  Carruthers;  Boucek  :  30,  pi.  2,  figs.  1-3. 

non   1939  Cyrtograptus  murchisoni  Carruthers;  Chang  &  Sun,  pi.  i,  figs.  4-8. 

ORIGINAL  DESCRIPTION.  "  Hydrothecae  triangular  apiculate,  furnished  with  a 
spine.  The  upper  margin  of  the  cell  at  right  angles  to  the  axis,  about  twenty-eight 
cells  to  the  inch.  The  polypary  is  considerably  incurved  at  its  proximal  end,  and 
as  it  grows  it  gradually  opens  into  a  larger  curve.  The  branches  spring  from  the 
celluliferous  surface  of  the  polypary,  but  as  there  is  no  break  in  the  continuity  of 
the  hydrothecae,  they  must  arise  from  the  periderm  covering  the  common  canal. 
The  branches  also  curve  in  the  same  direction  as  the  main  portion  of  the  polypary. 
Loc.  Pencerrig,  Builth.  I  have  associated  the  name  of  the  author  of  '  Siluria  '  with 
this  remarkable  species.  The  only  British  specimens  I  have  seen  are  in  the  Geological 
Museum,  Jermyn  Street,  but  among  the  specimens  obtained  by  the  British  Museum 
from  M.  Barrande  there  is  a  specimen  from  Listice,  labelled  G.  priodon,  which  belongs 
to  this  species.  " 

COMMENTS  ON  USAGE.  Carruthers'  figure  is  somewhat  idealized  but  there  is  no 
difficulty  in  recognizing  the  species.  The  syntypes  are  not  well  preserved  and  do  not 
show  the  proximal  end  well  on  the  large  specimens  although  this  may  be  the  result 
of  damage  to  the  specimens  which  have  at  some  stage  been  broken.  This  in  unfor- 
tunate as  Boucek,  in  his  revision  of  the  Cyrtograptidae  (1933),  distinguished  forms 
with  excentric  proximal  coiling  from  those  with  central  (and  more  open)  coiling. 
He  had  no  forms  of  the  first  type  with  secondary  branches  (a  characteristic  of  mur- 
chisoni) but  he  separated  the  simply-branched  forms  (centrifugus  and  murchisoni 
bohemicus)  on  the  proximal  ends.  The  type  slabs  of  murchisoni,  however,  show 
proximal  curvature  of  the  centrifugus  type  and  this  was  also  figured  by  Elles  & 
Wood. 

It  is  reasonable  to  separate  the  simply-branched  forms  as  a  subspecies  as  there 
appears  to  be  some  stratigraphic  value  in  this  (see  Rickards  1965)  but  if  the  curvature 
is  also  admitted  as  a  diagnostic  character,  a  new  name  is  required  for  the  Bohemian 


W.    CARRUTHERS'    TYPE   GRAPTOLITES  203 

specimens  of  "  murchisoni  ".  If  the  proximal  end  is  regarded  as  a  variable  feature 
within  the  species,  then  there  is  no  good  criterion  for  separating  centrifugus  from 
m.  bohemicus  and  these  may  both  be  regarded  as  junior  synonyms  of  Tullberg's 
C.  murchisoni  var.  crassiusculus.  Pending  a  revision  with  full  stratigraphic  details 
of  these  Lower  Wenlock  cyrtograptids,  it  seems  best  to  restrict  the  present  account 
to  the  redescription  of  Carruthers'  syntypes  and  the  numerous  topotypes. 

TYPE  MATERIAL.  Carruthers  noted  that  his  types  were  in  the  Jermyn  Street 
Museum  and  the  specimen  and  counterpart  are  now  numbered  GSM  10717-8. 
They  are  from  the  Wenlock  Shale,  Pencerraig,  Builth,  Radnorshire. 

REVISED  DESCRIPTION.  Rhabdosome  stout,  proximal  end  forming  a  helical 
spiral  with  triangular  thecae  of  the  spiralis  type.  First-order  cladia  curved,  arising 
at  well-spaced  intervals  (about  five  to  eight  thecae  between  each)  and  bearing 
secondary  branches  which  are  generally  curved  in  the  same  direction.  Thecae  on 
the  distal  part  of  the  main  stipe  and  on  the  cladia  have  a  tubular  proximal  part  and 
a  short,  hooked  aperture.  The  change  takes  place  on  the  main  stipe  after  the  third 
or  fourth  cladium.  Thecae  ten  to  fourteen  per  cm. 


FIG.  8.  Cyrtograptus  murchisoni  Carr.  Poorly  preserved  proximal  end  showing  excentric 
coiling.  The  most  proximal  thecae  are  underneath  the  later  part  of  the  main  stipe.  GSM 
10717.  X3. 

DISTRIBUTION.  As  noted  above,  there  is  some  difficulty  in  matching  Boucek's 
description  of  the  Bohemian  material  with  the  British  types,  although  Carruthers 
recognized  the  species  in  material  from  Listice.  The  stout,  secondarily  branched 
forms  appear  near  the  base  of  the  Wenlock  throughout  Europe  and  it  appears  con- 
venient to  regard  them  all  as  one  species.  Further  geographic  variation  may  be 
noticed  when  well-preserved  material  from  other  areas  is  studied.  The  species  has 
been  recorded  from  Canada  but  so  far  without  confirmatory  illustrations.  The 
form  recorded  from  China  appears  to  be  the  earlier  species,  Monograptus  spiralis. 
Russian  records  (Obut  1964;  Obut,  Sobolevskaya  &  Bondarev  1965)  appear  to  be 
of  the  species  group  but  not  the  typical  form.  The  relationship  of  C.  murchisoni 
to  the  forms  with  only  first  order  branches  is  still  unknown  but  the  latter  seem 
slightly  earlier  stratigraphically  (Rickards  1965)  and  could  well  be  ancestral.  The 
absence  of  cyrtograptids  from  the  succeeding  zone  of  M.  riccartonensis  suggests  that 
the  later  C.  rigidus  and  its  allies  are  not  phylogenetically  related  to  C.  murchisoni, 
since  this  would  involve  a  reversal  of  the  trend  towards  more  numerous  branches. 

In  my  notes  on  British  zones  (Strachan  1960)  I  observed  that  C.  murchisoni  was 
"  apparently  unknown  outside  the  type  locality  ".  Since  then  I  have  seen  many- 
branched  specimens  from  the  Wenlock  of  the  South  of  Scotland  and  Rickards  (1965) 
has  also  found  numerous  specimens  in  the  Lake  District. 


204  W.    CARRUTHERS'    TYPE   GRAPTOLITES 

Dendrograptus  lentus  Carruthers 

i86ya  Dendrograptus  lentus  Carruthers:  541,  foss.  90,  fig.  5. 
1868     Dendrograptus  lentus  Carruthers:  130,  pi.  5,  fig.  5. 

The  specimen  described  under  this  name  is  preserved  in  the  Geological  Survey 
&  Museum,  GSM  10691  (and  counterpart  10692).  The  species  was  noted  by  Wood 
(in  Elles  &  Wood,  p.  Ivii)  as  belonging  to  Clonograptus  and  a  note  on  the  specimens 
says  "  PShineton  Shales,  Shrops.  "  Carruthers  recorded  the  species  as  "  Caradoc, 
Co.  Fermanagh  "  and  the  specimens  are  listed  as  such  in  the  Survey  Catalogue  of 
1878.  However  there  is  a  collection  number  with  the  drawing  of  this  species  in 
Carruthers'  notes  and  this  appears  in  the  Survey  Catalogue  of  1864  as  "  Graptolithus 
sp.  Cambrian,  Wrekin.  "  There  thus  is  some  confusion  over  this  species  and  since 
the  name  has  not  been  used,  so  far  as  I  can  ascertain,  since  the  original  descriptions 
it  is  best  regarded  as  a  nomen  dubium,  and  an  appropriate  recommendation  to  the 
ICZN  has  been  made.  If  the  species  were  to  be  regarded  as  valid,  the  name  lentus 
would  have  priority  over  the  widely  used  name  tenellus,  and  these  fragments  are 
too  poor  to  be  the  basis  of  this  well-known  species. 


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94  :  1-58,  pis.  1-15. 
TOGHILL,  P.     1965.     Stratigraphical  variations  in  the  Lower  Silurian  of  the  South  of  Scotland. 

Ph.D.  thesis,  Univ.  of  Birmingham. 
TOMCZYK,  H.     1962.     [Statigraphical  problems  of  the  Ordovician  and  Silurian  in  Poland  in 

the  light  of  recent  Studies].     Pr.  Inst.  Geol.,  35  :  134,  4  pis.,  13  figs.     [In  Polish  with  Engl. 

transl.].     Warsaw. 
TORNQUIST,  S.  L.     1907.     Observations  on  the  genus  Rastrites  and  some  allied  species  of  Mono- 

graptus.     Acta.  Univ.  lund.,  Handl.  K.  Fysiogr.  Sallsk.  N.F.,  18,  5  :  1-22,  pis.  1-3. 
TULLBERG,    S.    A.     1883.     Skanes    Graptoliter    II.     Graptolitfaunorna  i  Cardiolaskiffern    och 

Cyrtograptusskiffrarne.     Sver.  geol.  Unders.  Abh.  C,  55  :  1-44,  4  pis.     Stockholm. 


EXPLANATION  OF  PLATES 


Most  of  the  specimens  are  in  the  British  Museum  (Natural  History)  and  their 
numbers  bear  the  prefix  Q.  Those  with  prefixes  GSM  and  BU  are  in  the  Institute 
of  Geological  Sciences  and  Birmingham  University  respectively. 


PLATE  i 
Leptograptus  capillaris  (Carruthers) 

FIG.  i.     Copy  of  original  figure,  Carruthers  1868,  fig.  ya. 

FIG.  ?..     Type  slab,  Q.$o,  natural  size.     Hartfell  shales.     Hartfell  Spa. 


Butt.  BY.  Mus.  nat.  Hist.  (Geol.)  17,  4 


PLATE  i 


GEOL.  17,  4. 


PLATE  2 
Pleurograptus  linearis  (Carruthers) 

FIG.  i.     Holotype,  Q.848,  natural  size.     Hartfell  Shales.     Hartfell  Spa. 

FIG.  2.  Copy  of  page  from  Carruthers'  MS  notes  showing  drawing  of  type  slab  and  the 
original  published  figures  (1858-59)  half  original  size. 

FIG.  3.  Specimen  figured  by  Nicholson,  1867,  fig.  i,  refigured  Elles  &  Wood,  pi.  16,  fig.  7. 
Q.27,  natural  size.  Hartfell  Shales.  Hartfell  Spa. 


Bull.  BY.  Mus.  nat.  Hist.  (Geol.)  17,  4 


PLATE  3 

Diceltograptus  elegans  (Carruthers) 

FIG.  i.     Holotype,  Q.850,  natural  size.     Hartfell  Shales.     Hartfell  Spa. 
Dicranograptus  clingani  Carruthers 

FIG.  2.  Specimen  figured  Carruthers  1868,  fig.  6b,  Q.842,  natural  size.  Hartfell  Shales. 
Hartfell  Spa. 

FIG.  3.  Lectotype,  Carruthers  1868,  fig.  6a,  refigured  Elles  &  Wood,  pi.  24,  fig.  la,  Q-55, 
natural  size.  Hartfell  Shales.  Hartfell  Spa. 

FIG.  4.     Same,  x  5. 

Dicellograptus  moffatensis  (Carruthers) 

FIG.  5.  Page  from  Carruthers'  MS  notes  showing  original  drawings  and  copy  of  published 
figure  (1858-59)  half  original  size. 

FIG.  6.  Type  slab,  (3.843,  natural  size.  The  lectotype,  figured  by  Elles  &  Wood,  pi.  23, 
fig.  la,  is  marked  with  the  arrow.  Hartfell  Shales.  Hartfell  Spa. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  4 


PLATE  3 


. 

':       J  -** 


:'•       £V        *'•          \      i 

\i 

s<        IF -I:*.  ^ 


PLATE  4 
Climacograptus  minutus  Carruthers 

FIG.  i.     Lectotype,  Q.8o,   X5.     Moffat,  horizon  and  locality  uncertain. 
FIG.  2.     Specimen  figured  Carruthers  1868,  fig.  job.  (^.1372,  as  C.  minutus  but  probably 
C.  scalaris  miserabilis  E.  &  W.      X5.     Moffat,  horizon  and  locality  uncertain. 

Climacograptus  minimus  (Carruthers) 

FIG.  3.  Type  slab,  Q .  82,  natural  size,  showing  poorly  preserved  biserial  graptolites.  Horizon 
and  locality  uncertain. 

Cryptograptus  tricornis  (Carruthers) 

FIG.  4.  Slab  with  numerous  specimens,  (^.1299,  lectotype  marked  with  arrow,  natural  size. 
Hartfell  Shales.  Hartfell  Spa. 

FIG.  5.  Drawings  from  Carruthers'  MS  notes  showing  the  different  appearances  of  the 
rhabdosome  half  original  size. 

FIG.  6.     Copy  of  original  figures  (1858-59)  half  original  size. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  4 


PLATE  4 


PLATE  5 
Monograptus  clingani  (Carruthers) 

FIG.  i.     Specimen  showing  distal  part  of  the  rhabdosome,  BU  1667,  figured  Lapworth  1876, 
fig.  3a,  refigured  Elles  &  Wood,  pi.  46,  fig.  nd,  natural  size.     Birkhill  Shales.     Dobbs  Linn. 

FIGS.  2  and  4.     Specimen  figured  Carruthers  1868,  fig.  igb.  (3.84,  natural  size  and    X5. 
MofEat. 

FIGS.  3  and  5.     Lectotype,  figured  Carruthers  1868,  fig.  iga,  Q.87,  natural  size  and    X5- 
Mofiat. 

Monograptus  intertnedius  (Carruthers) 

FIG.  6.     Holotype,  Q.88,  figured  Carruthers  1868,  fig.  18,  natural  size.     Birkhill  Shales. 
Moffat. 

Rastrites  tnaximus  Carruthers 

FIG.  7.     Neotype,  BU  1692,  figured  Elles  &  Wood,  pi.  50,  fig.  6b,    x  2^.     Upper  Birkhill 
Shales.     Riskinhope  Burn,  Yarrow. 

Cyrtograptus  murchisoni  Carruthers 

FIGS.  8  and  9.     Specimen  and  counterpart,  GSM  10717-8,  half  natural  size.     These  are  the 
slabs  from  which  Carruthers  made  his  description  and  idealized  figure.    Wenlock  Shales.    Builth 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  4 


PLATE  5 


«-~      ~  -^ ; 
~*  *        v  ^~ 

,    A--- 

7 


PRINTED  IN  GREAT  BRITAIN 
BY  ADLARD  &  SON  LIMITED 
BARTHOLOMEW  PRESS,  DORKING 


A  REVISION  OF  THE  ENGLISH 
WEALDEN  FLORA,  I 

CHARALES— GINKGOALES 


J.  WATSON 


BULLETIN    OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
GEOLOGY  Vol.  17  No.  5 

LONDON:  1969 


A  REVISION  OF  THE  ENGLISH  WEALDEN 

FLORA,  I 

CHARALES— GINKGOALES 


BY 

JOAN  WATSON 


Pp. 207-254;   6  Plates;    64  Text-figures 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
GEOLOGY  Vol.  17  No.  5 

LONDON :  1969 


THE    BULLETIN    OF    THE    BRITISH    MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they  become 
ready.  Volumes  will  contain  about  three  or  four 
hundred  pages,  and  will  not  necessarily  be  completed 
within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  17,  No.  5  of  the  Geological 
(Palaeontological)  series.  The  abbreviated  titles  of 
periodicals  cited  follow  those  of  the  World  List  of 
Scientific  Periodicals. 


World  List  abbreviation 
Bull.  Br.  Mus.  nat.  Hist.  (Geol.). 


Trustees  of  the  British  Museum  (Natural  History)  1969 


TRUSTEES    OF 
THE    BRITISHfMUSEUM    (NATURAL    HISTORY) 

Issued  21  January,  1969  Price  £i  155. 


A  REVISION  OF  THE  ENGLISH  WEALDEN 

FLORA,  I 

CHARALES— GINKGOALES 

By  JOAN  WATSON 

CONTENTS 

I.     INTRODUCTION          .........  210 

II.     SYSTEMATIC  DESCRIPTIONS          .......  214 

THALLOPHYTA       .          .          .          .          .          .          .          .          .214 

CHLOROPHYCEAE        .         .         .         .         .         .         .         .  214 

CHARALES      .          .          .          .          .          .          .          .          .214 

CIRCONITELLA  gen.  nov 214 

Circonitella  knowltoni  (Seward)  n.  comb.            .          .  215 

GYROGONA  Lamarck 217 

Gyrogona  medicaginula  Lamarck      .          .          .          .  217 

BRYOPHYTA           .........  218 

Musci             .........  218 

?  Bryophyte  leaf             ......  218 

HEPATICAE    .........  218 

HEPATICITES  Walton 219 

Hepaticites  zeilleri  (Seward)  n.  comb.        .          .          .  219 

Hepaticites  ruffordi  sp.  nov.    .....  220 

TH A  LUTES  Walton 220 

Thallites  valdensis  (Seward)  n.  comb.        .          .          .  221 

Thallites  catenelloides  (Seward)  n.  comb.  .          .          .  223 

PTERIDOPHYTA      .........  224 

LYCOPODIALES        ........  224 

SELAGINELLA 224 

Selaginella  dawsoni  (Seward)  .          .          .          .          .  224 

EQUISETALES           ........  227 

EQUISETITES  Sternberg 227 

FlLICALES         .........  228 

MATONIACEAE     ........  228 

MATONIDIUM  Schenk            ...  228 

Matonidium  goepperti  (Ettingshausen)      .          .          .  228 

DlPTERIDACEAE  ........  228 

HAUSMANNIA  Dunker 228 

Hausmannia  dichotoma  Dunker        ....  228 

SCHIZAEACEAE       ........  229 

RUFFORDIA  Seward  ....  229 

Ruffordia  goepperti  (Dunker)  ....  229 

PELLETIERIA  Seward  ...  232 

Pelletieria  valdnensis  Seward  .....  232 

POLYPODIACEAE                .......  235 

ASPIDISTES  Harris         .          .  235 

Aspidistes  sewardi  sp.  nov.      .....  236 

GEOL.   17,  5  J8 


2io  A   REVISION    OF   THE 

UNCLASSIFIED  FERNS   .......  237 

CLADOPHLEBIS  Brongniart  .  .  .  .  .  237 

Cladophlebis  longipennis  Seward  ....  237 

Cladophlebis  albertsii  (Dunker)  ....  237 

Cladophlebis  browniana  (Dunker)  ....  238 

Cladophlebis  dunkeri  (Schimper)  .  .  .  .  238 

SPHENOPTERIS  Sternberg 238 

Sphenopteris  ruffordi  (Seward)  n.  comb.  .  .  .  238 

Sphenopteris  fittoni  Seward  .....  238 

Sphenopteris  fontainei  Seward  ....  238 

LECKENBYA  Seward 238 

Leckenbya  valdensis  (Seward)  .  .  .  .  238 

TEILHARDIA  Seward  238 

Teilhardia  valdensis  Seward        .          .          .          .          .  238 

ONYCHIOPSIS  Yokoyama 238 

Onychiopsis  psilotoides  (Stokes  &  Webb)  .  .  238 

WEICHSELIA  Stiehler 239 

Weichselia  reticulata  (Stokes  &  Webb)      .          .          .  240 

SPERMATOPHYTA  ........  240 

PTERIDOSPERMAE  ........  240 

PACHYPTERIS  Brongniart 240 

Pachypteris  lanceolata  Brongniart     ....  240 

GYMNOSPERMAE          ........  242 

CYCADALES  .........  242 

NILSSONIA  Brongniart 242 

Nilssonia  schaumburgensis  (Dunker)  .  .  .  242 

BECKLESIA  Seward 244 

Becklesia  anomala  Seward  .....  244 

Becklesia  sulcata  sp.  nov.  .....  247 

GlNKGOALES  ........  248 

PSEUDOTORELLIA  Florin 248 

Pseudotorellia  heterophylla  sp.  nov.  .          .          .          248 

III.     REFERENCES   ..........         251 

SYNOPSIS 

The  English  Wealden  flora  described  by  Seward  (1894,  1895)  is  here  revised  using  modern 
techniques,  in  particular  in  the  study  of  the  plant  cuticles.  Much  of  the  work  has  been  carried 
out  on  the  existing  collections  in  the  British  Museum  (Natural  History),  but  a  number  of  new 
species  in  the  form  of  isolated  leaf  cuticles  have  been  obtained  by  bulk  maceration  of  "  coaly- 
shales  "  collected  by  the  author  from  the  Fairlight  Clay  at  Hastings,  Sussex. 

The  present  work  covers  the  Thallophyta,  Bryophyta,  Pteridophyta,  Pteridospermae,  Cy- 
cadales  and  Ginkgoales.  A  second  part  will  deal  with  the  Bennettitales  and  Coniferales  together 
with  a  special  section  devoted  to  a  complete  revision  of  the  Wealden  Equisetales. 

I.  INTRODUCTION 

English  Wealden  Flora. 

THIS  work  is  a  revision  of  part  of  the  English  Wealden  Flora,  using  modern  tech- 
niques. It  has  been  carried  out  largely  from  a  study  of  the  Wealden  plants  in  the 
British  Museum  (Natural  History).  Most  of  this  material  was  collected  from  the 
coast  in  the  neighbourhood  of  Hastings,  Sussex  in  the  latter  part  of  the  last  century 


ENGLISH   WEALDEN   FLORA   I  211 

and  belongs  to  the  Beckles,  Dawson,  Mantell  and  Rufford  Collections.  The  majority 
was  collected  by  Mr.  P.  Rufford  but  unfortunately  his  specimens  are  merely  labelled 
"  Ecclesbourne,  Nr.  Hastings  "  and  the  exact  localities  are  not  recorded.  Attempts 
were  made  to  find  fresh  material  along  the  coast  between  Hastings  and  Pett  Level 
but  unfortunately  no  fossils  were  found.  However,  material  was  obtained  from 
coaly  lenses  and  partings  which  occur  in  the  Hastings  Beds.  Bulk  maceration  of 
these  "  coaly-shales  "  yielded  quantities  of  spores,  isolated  leaves  and  even  small 
shoots.  Two  of  the  species  thus  obtained  are  described  here,  including  the  first 
record  of  a  member  of  the  Ginkgoales  from  the  English  Wealden.  Similar  coaly- 
shales  from  the  Wealden  in  the  Isle  of  Wight  yield  identical  plant  fragments. 

The  flora  was  previously  described  by  Seward  (1894,  1895).  His  identifications 
of  the  fossils  were  based  solely  on  macroscopic  features  and  were  figured  by  means  of 
woodcuts  and  lithographs.  A  re-examination  of  the  fossils  using  modern  techniques 
shows  that  many  of  Seward's  determinations  are  wrong.  However,  where  the  type 
material  of  a  particular  species  (usually  from  the  German  Wealden)  has  not  been 
redescribed  or  examined  it  is  impossible  to  say  if  the  identification  is  truly  accurate. 

Seward  arranged  the  flora  into  about  seventy-five  species.  From  more  than  a 
dozen  of  these  I  was  unable  to  get  any  significant  new  information,  and  unless  better 
material  is  found  it  seems  that  only  a  redescription  of  their  gross  morphology  would 
be  possible.  Most  of  the  ferns  come  into  this  category  and  only  a  few  species  are 
dealt  with  here  in  any  detail.  There  are  very  few  fertile  specimens  available  and 
one  such  fern,  Onychiopsis  psilotoides  (Stokes  &  Webb),  has  recently  been  redes- 
cribed by  Tattersall  (1961).  The  practice  of  coating  specimens  with  varnish  in  the 
past  has  caused  the  destruction  of  the  cuticle  in  many  specimens  and  has  greatly 
hindered  revision. 

The  Thallophyta,  Bryophyta,  Pteridophyta,  Pteridospermae,  Cycadales  and 
Ginkgoales  are  described  in  this  paper  with  the  exception  of  the  Equisetales.  A 
recent  find  of  more  petrified  material  belonging  to  the  Equisetales  renders  necessary 
a  further  revision  of  the  species  (see  Watson  1964),  and  this  will  be  published  in  a 
later  volume  together  with  the  Bennettitales  and  Coniferales. 

Other  Wealden  Floras 

The  Wealden  floras  of  Germany,  France  and  Belgium  were  all  described  in  the 
last  century  in  the  same  way  that  Seward  described  the  English  flora.  Since  then 
little  progress  has  been  made  and  work  as  detailed  as  that  presented  here  scarcely 
exists.  Cuticles  have  been  described  for  a  few  species  from  time  to  time  (Carpentier 
1939)  but  the  amount  of  work  to  be  done  remains  enormous.  The  German  Wealden 
flora  in  particular  should  be  thoroughly  revised  as  it  contains  so  many  type  specimens. 
A  number  of  English  fossils  are  referred  to  German  species  merely  on  external  charac- 
ters. When  a  species  under  revision  can  only  be  compared  with  the  inadequate, 
original  nineteenth  century  description  of  the  type  an  impossible  situation  arises. 
It  has  hitherto  been  thought  that  the  English  and  German  floras  are  more  or  less 
identical  but  I  suspect  that  this  may  not  be  so.  In  some  cases  fairly  recent  work 
on  foreign  material  has  shown  that  a  number  of  English  specimens  previously  re- 


A    REVISION    OF   THE 


f erred  to  German  species  are  different  from  the  type  \Sphenolepis  kurriana  (Dunker), 
Sphenolepis  sternbergiana  (Dunker),  Pseudocycas  dunkeriana  (Schenk)]  and  these 
were  described  as  new  species  (Watson  1964).  But  for  the  vast  majority  of  species 
no  such  comparison  is  possible.  It  is  also  now  known  that  the  German  and  English 


FIG.  i.     Inferred  present  extent  of  Wealden  strata.     (After  Allen  1967) 


Wealden  are  not  wholly  contemporaneous  (see  Table  I).  For  these  reasons  I  feel 
that  it  is  of  little  value  to  produce  comparative  lists  of  species  for  the  various  locali- 
ties. This  has  been  done  frequently  in  the  past  and  perhaps  most  recently  and 
comprehensively  by  Delcourt  &  Sprumont  (1955).  Such  lists,  however,  are  probably 
highly  inaccurate  and  will  remain  so  until  there  is  a  wholesale  revision  of  all  Lower 
Cretaceous  fossil  floras. 

Techniques 

Techniques  used  were  of  the  simplest.  With  few  exceptions  the  usual  method  for 
bulk  macerations  was  used  (see  Harris  1926). 

Stratigraphical  background 

The  stratigraphy  of  the  Wealden  has  been  described  by  White  (1928),  Allen  (1954, 
1955,  1959,  1960,  1965,  1967),  Hughes  (1958)  and  Howitt  (1964).  Owing  to  its 
alleged  relict  fauna  and  flora  of  Jurassic  affinity  the  English  Wealden  has  on 
occasions  been  assigned  to  the  Jurassic  System.  It  is  now,  however,  generally 


ENGLISH   WEALDEN    FLORA    I 


213 


TETHYAN 

BOREAL 

South-  east 
France 

North  -  west 
Germany 

Southern 
England 

CRETACEOUS  (part) 

Barremian 

Barremian 

Weald 
Clay 

CRETACEOUS  (part) 

Hauterivian 

Hauterivian 

Valanginian 

'Upper  &  Middle 
Valdensis 

Hastings 
Beds 

Berriasian 

Wealden 

Durlston 

^            Beds 
U 

UJ 

Upper 
Serpulite 

Lower     Serpulite 

~  O£ 

13          Lulworth 
Beds 

JURASSIC  (part) 

Munder 
Marls 

JURASSIC  (part) 

Tithonian 

Portland 
Beds 

Eimbeckhausen 
Plaltenkalk 

TABLE,  i — Correlation  of  N.  W.  European  "  Wealden  "  (from  table  4  in  Dodson,  Rex,  Casey  & 

Allen  1964). 


accepted  as  part  of  the  Cretaceous.  Part,  or  even  all,  of  the  Purbeck  might  also  be 
Cretaceous.  Table  I  is  a  correlation  table  for  the  Wealden  of  north-west  Europe 
recently  proposed  by  Dodson,  Rex,  Casey  &  Allen  (1964).  On  the  basis  of  glau- 
conite  studies  the  radiometric  age  of  the  Jurassic /Cretaceous  boundary  is  estimated 
to  be  about  135  million  years.  Text-fig,  i  shows  the  present  extent  of  the  Wealden. 
The  Fairlight  Clays  have  yielded  most  of  the  fossil  plants,  particularly  in  the  region 
of  Ecclesbourne  Glen.  The  large  number  of  specimens  collected  by  Mr.  P.  Rufford 
and  others  has  led  to  the  false  impression,  which  has  been  perpetuated  (Gallois  1965), 
that  plants  are  abundant  in  the  Fairlight  Clays.  This  is  not  so:  even  small  frag- 


214  A   REVISION   OF   THE 

ments  are  quite  rare.  It  has  not  been  possible  to  ascertain  the  exact  localities  and 
horizons  at  which  Rufford  collected;  nor  is  it  known  over  what  period  of  time  the 
collection  was  assembled.  In  a  brief  description  of  the  Hastings  Beds  by  Rufford 
(Seward  1894  :  xvii)  he  refers  to  the  Fairlight  Clays  "  yielding,  very  locally,  Ferns, 
Cycads  and  Conifers  "  and  also  to  "  the  very  limited  extent  of  outcropping  fossili- 
ferous  strata  ".  In  the  early  years  of  this  century  (around  1910)  a  further  important 
collection  of  Wealden  plants  was  made  by  P.  Teilhard  de  Chardin  and  Felix  Pelletier, 
two  Jesuit  priests  who  stayed  in  the  area  for  four  years.  Their  specimens  (Seward 
1913)  were  collected  mainly  in  the  neighbourhood  of  Fairlight  which  is  about  a  mile 
to  the  east  of  Ecclesbourne  Glen.  The  collection  was  subsequently  divided  between 
the  British  Museum  (Natural  History)  and  the  Hastings  Museum.  The  Hastings 
Museum  also  has  a  large  number  of  Rufford's  specimens,  many  of  which  are  the 
counterparts  of  the  specimens  in  the  British  Museum  (Natural  History).  Attempts 
at  collecting  in  recent  years  have  yielded  nothing  to  compare  with  the  magnificent 
specimens  (fronds  three  feet  long)  obtained  by  these  earlier  collectors. 

ACKNOWLEDGEMENTS 

This  work  forms  part  of  a  Ph.D.  thesis  of  the  University  of  Reading,  supervised 
by  Professor  T.  M.  Harris,  F.R.S.  and  I  thank  him  for  his  unfailing  patience,  kind- 
ness and  encouragement.  I  wish  also  to  thank  Professor  P.  Allen  for  much  help  and 
for  permission  to  reproduce  his  text-figures. 

Some  of  the  work  was  carried  out  in  the  Department  of  Palaeontology,  British 
Museum  (Natural  History),  some  in  the  Botany  Department,  University  of  St. 
Andrews  and  some  in  the  Herbarium,  Manchester  Museum  where  I  was  given  research 
facilities  for  which  I  am  most  grateful.  Thanks  are  due  to  the  Keeper  of  Palaeon- 
tology, British  Museum  (Natural  History)  for  permission  to  work  on  the  collections. 

Tenure  of  a  D.S.I. R.  Studentship  is  gratefully  acknowledged. 

II.  SYSTEMATIC  DESCRIPTIONS 

Unless  otherwise  indicated  in  the  text  all  specimens  are  in  the  collections  of  the 
British  Museum  (Natural  History)  and  are  prefixed  by  V. 

THALLOPHYTA 

CHLOROPHYCEAE.     CHARALES 
Organ  Genus  CIRCONITELLA  nov. 

DIAGNOSIS.  Gyrogonite  uncalcified,  cutinised;  surrounded  by  five  sinistrally 
spiralled  cells;  shape  ellipsoidal,  circular  in  transverse  section.  Apical  pole  formed 
by  all  five  cells;  basally  ending  against  a  basal  plate;  without  basal  cage. 

TYPE  SPECIES.     Chara  knowltoni  Seward  1894;  13,  Text-fig.  I. 

DISCUSSION.  This  genus  is  recognized  for  the  type  species  which  was  formerly 
included  in  Nitellites  Horn  af  Rantzien  in  my  Ph.D.  thesis  of  1964.  The  diagnosis  of 
Nitellites,  however,  requires  that  size  and  shape  should  be  very  close  to  that  of  the 


ENGLISH   WEALDEN   FLORA    I  215 

Recent  genus  Nitella  but  C.  knowltoni  differs  in  being  somewhat  larger  and  un- 
flattened. 

Dijkstra  (1959)  describes  an  uncalcified  gyrogonite  which  he  refers  to  as  "  Prae- 
chara  symmetrica  Peck  cf.  ".  I  am  satisfied,  from  his  diagnosis  and  photographs 
that  this  fossil  is  identical  with  C.  knowltoni  (Seward).  Whilst  this  species  fits  into 
Praechara  Horn  af  Rantzien  (1954)  on  its  general  shape  and  apical  structure, 
Praechara  has  hitherto  been  used  only  for  members  of  the  Characeae,  i.e.  calcified 
gyrogonites  which  cannot  be  placed  in  other  well-defined  genera. 

Circonitella  knowltoni  (Seward)  comb.  nov. 
PI.  i,  figs.  1-5 ;  Text-figs.  2-6. 

1894     Char  a  knowltoni  Seward  :  13,  text-fig,  i. 

X959     Praechara  symmetrica  Peck  cf. ;  Dijkstra  :  15,  pi.  2,  figs.  20-23. 

EMENDED  DIAGNOSIS.  Gyrogonite  uncalcified,  ellipsoidal  in  side  view,  apical  and 
basal  halves  similar,  4007*  to  665  JLL  long  and  380  ju  to  550  ju,  wide.  Spiral  cells  five, 
sinistrally  spiralled,  about  45  /*  wide  near  equator  and  showing  about  twelve  con- 
volutions laterally.  Cells  usually  slightly  concave  with  distinct  intercellular  ridges 
3  /*  to  15  ft  wide,  sometimes  flat  and  occasionaUy  convex.  Outer  cutinised  membrane 
(ectosporostine  of  Horn  af  Rantzien)  marked  with  coarse  but  very  low  and  ill- 
defined  tubercles  up  to  5  fi  diameter.  Inner  membrane  (endosporostine  of  Horn 
af  Rantzien)  present,  delicate  and  without  a  patterned  surface.  Apical  pole  formed 
by  ends  of  spiral  cells  meeting  in  a  slightly  eccentric  pattern ;  basal  pole  with  rounded 
plate  100  /i  wide;  no  cage  or  basal  claws  present. 

NEOTYPE.    ¥.51555. 

DESCRIPTION.  Numerous  black  cutinized  gyrogonites  are  embedded  in  an  argil- 
laceous matrix,  the  majority  being  flattened  and  presenting  a  lateral  view  but  a 
few  being  orientated  so  as  to  show  polar  views.  A  few  gyrogonites  preserved  by 
pyritization  are  completely  unsquashed  and  were  obtained  by  macerating  small 
pieces  of  the  rock  in  water.  These  specimens  show  that  the  gyrogonite  was  originally 
completely  circular  in  section  and  that  flattening  occurred  during  preservation. 
Whereas  most  of  the  flattened  specimens  have  concave  spiral  cells  with  intercellular 
ridges  the  solid  ones  have  convex  spirals  with  depressions  between  and  tend  to  be 
somewhat  larger.  However,  one  of  them  (¥.51555)  has  part  of  the  convex  surface 
broken  away  revealing  the  familiar  concave  cells  with  ridges  below.  These  two 
surfaces  are  interpreted  as  the  outer  and  inner  walls  of  the  spiral  cells. 

Apical  poles  are  not  seen  nearly  so  frequently  as  basal  poles  but  a  few  undistorted 
specimens  were  obtained.  It  may  be  noted  that  in  the  material  from  the  Nether- 
lands, Dijkstra  (1959)  also  found  the  apical  poles  of  the  gyrogonites  less  well  pre- 
served. The  tips  of  the  spiral  cells  tend  to  be  swollen  and  meet  in  a  slightly  irregular 
pattern.  Many  more  basal  poles  are  seen  with  the  easily  identifiable  circular  plate 
against  which  the  five  spiral  cells  end  in  a  regular  manner.  Their  width  near  the 
pole  is  the  same  as  at  the  equator. 

Laterally  up  to  twelve  convolutions  of  spiral  cells  are  seen  each  making  more  than 
two  turns.  Thus  the  spiral  is  so  flat  that  the  furrows  and  ridges  appear  almost 


2l6 


A   REVISION    OF   THE 


horizontal.  As  mentioned  above  the  spiral  cells  are  concave  in  most  gyrogonites 
but  convex  in  a  few  specimens.  There  are  also  many  in  which  they  are  flat  and 
their  margins  are  then  nearly,  or  quite,  invisible  in  the  untreated  fossil.  For 
variety  in  intercellular  ridges  compare  PL  i,  figs.  2,  4. 

The  tubercles  on  the  ectosporostine  are  irregularly  arranged  about  8  /*  apart, 
usually  about  four  tubercles  across  the  width  of  a  spiral  cell.     Many  gyrogonites 


FIGS.  2-6.  Circonitella  knowltoni  (Seward).  2,  3.  Gyrogonites  showing  apical  pole. 
Fig.  2,  specimen  accidentally  destroyed.  Fig.  3,  ¥.51549.  x  100.  4.  Side  view  of  a 
gyrogonite  with  concave  cells,  showing  the  very  flat  spiral.  V. 5 1550.  X  75.  5. 
Basal  pole  of  a  gyrogonite.  ¥.3114.  x  100.  6.  Cutinized  wall  showing  part  of  three 
spiral  cells  with  tubercles  on  the  outer  layer.  The  inner  layer  of  the  wall  is  seen  on  the 
right  of  the  middle  cell.  Slide  V.  5 1551.  x  350. 

gave  no  satisfactory  cuticles  at  all  and  some  that  did  yield  cuticles  were  less  clearly 
marked.  The  cuticle  is  usually  reddish-brown  in  colour  but  in  parts  the  inner  mem- 
brane of  the  wall  is  seen  alone  as  a  straw-coloured  homogeneous  layer  without 
markings  of  any  kind.  The  coronula  is  not  known,  nor  is  the  oospore. 

Dijkstra  suggested  that  his  fossils  were  originally  calcified  but  became  decalcified 
during  preservation.  It  is  much  more  likely,  however,  that  the  present  specimens 
were  never  calcified.  There  is  evidence  that  both  the  concave  inner  wall  and  the 


ENGLISH   WEALDEN   FLORA   I  217 

convex  outer  wall  of  the  gyrogonite  cells  are  composed  of  organic  matter  and  if  my 
interpretation  of  these  layers  is  correct  then  it  seems  conclusive.  It  is  clear  that 
in  the  specimens  with  only  the  inner  organic  layer  present  the  fine  clay  matrix  is 
everywhere  pressing  against  this  layer;  there  is  no  gap  at  all  such  as  would  be  left 
if  a  lime-shell  had  been  dissolved.  We  would  have  to  suppose  that  decalcification 
occured  first,  then  preservation.  This  species  is  preserved  in  a  unique  way  in  speci- 
mens from  two  widely  separated  localities  and  it  seems  unlikely  to  have  been  caused 
by  the  same  unusual  event  at  two  distant  places. 

The  absence  of  a  calcareous  shell  on  the  outside  of  the  gyrogonite  immediately 
suggests  Nitellaceous  affinities  but  further  study  shows  that  in  other  features  it  is 
more  like  the  Characeae.  I  make  no  assumption  about  the  correct  classification  of 
the  plant  producing  this  gyrogonite. 

In  all  the  Nitellaceae  the  gyrogonites  are  laterally  compressed  whereas  Circonitella 
knowltoni  is  quite  round  in  transverse  section  as  are  members  of  the  Characeae.  In 
studying  a  number  of  living  Nitellaceae,  Horn  af  Rantzien  (1959)  found  Nitella 
hyalina  to  be  the  least  flattened,  a  typical  specimen  being  340  /£  long,  260  fi  wide 
and  180  ju,  thick.  He  also  gives  details  concerning  the  average  size  of  Nitella  gyrogo- 
nites :  '  .  .  .  generally  speaking  the  oosporangia  (gyrogonites)  of  Nitella  are  small,  in 
most  species  the  length  does  not  exceed  350  fi '.  C.  knowltoni  is  much  larger, 
attaining  almost  twice  this  length,  although  Nitella  gyrogonites  with  larger  dimen- 
sions do  occur,  e.g.  N.  flexilis  which  reaches  575  fi  long  (Groves  &  Bullock- Webster 
1917). 

Twelve  convolutions  of  spiral  cells  seen  in  lateral  view  is  common  in  the  Characeae 
but  in  the  genus  Nitella  the  largest  number  known  is  eleven,  in  N.  polygyra.  A 
more  usual  number  in  this  group  is  five  or  six  and  hence  the  angle  of  the  spiral  is 
steeper  than  in  C.  knowltoni. 

Amongst  British  Charales  the  most  similar  ectosporostine  pattern  is  seen  in  N. 
opaca,  although  the  tubercles  are  more  numerous  and  larger.  Most  of  the  Characeae 
have  a  much  smoother  granulate  surface.  The  absence  of  basal  claws  accords  with 
Recent  species  of  Nitella. 

Apart  from  Quaternary  fossils  the  only  other  uncalcified  gyrogonites  are  Nitel- 
lites  sahnii  Horn  af  Rantzien  and  Nitellites  sp.  (Horn  af  Rantzien  1957)  from  the 
Middle  to  Upper  Jurassic  Rajmahal  series  of  India.  Both  differ  from  C.  knowltoni 
in  having  more  in  common  with  the  Recent  genus  Nitella.  They  are  much  smaller, 
being  only  160  /*  at  the  longest,  and  they  are  laterally  compressed.  The  spiral 
cells  show  only  six  to  eight  convolutions  and  the  ectosporostine  has  a  reticulate 
surface. 

Genus  GYROGONA  Lamarck 
Gyrogona  medicaginula  Lamarck 

Selected  references- 

1804     Gyrogona  medicagimtla  Lamarck  :  355. 

1829     Chara  medicaginula  (Lamarck);  Lyell  :  91,  pi.  13,  figs.  1-8. 

I933     Gyrogona  medicaginula  Lamarck;  Groves  :  24,  25,  39. 

1965     Gyrogona  medicaginula  Lamarck;  Grambast  &  Paul  :  239,  pi.  i,  figs.  1-4. 


218  A   REVISION   OF   THE 

REMARKS.  Several  gyrogonites  of  this  species  are  present  in  a  piece  of  pure 
siliceous  rock  (V .  2928)  which  is  included  in  the  Wealden  Collections  (British  Museum, 
Natural  History).  The  specimen  has  the  appearance  of  a  large  pebble  from  the 
beach  and  it  seems  probable  that  it  is  not  Wealden  in  age.  G.  medicaginula  is 
otherwise  only  known  from  the  Tertiary  (Lyell  1829,  Grambast  &  Paul  1965)  and  is 
found  in  the  Oligocene  of  the  Isle  of  Wight.  The  specimen  is  accompanied  by  a 
note  written  by  Groves  and  is  probably  the  specimen  to  which  he  refers  (Groves 
T933  :  25)  when  saying  "  Fruits  collected  from  a  Wealden  deposit,  Fairlight  Glen, 
Hastings  (Sussex),  England  .  .  .  are  considered  by  the  present  writer  to  be  referable 
to  this  species  ". 

The  matrix  cannot  be  matched  to  anything  in  the  Hastings  Beds  and  the  gyro- 
gonites of  G.  medicaginula  are  much  larger  than  is  usual  in  pre-Tertiary  species. 
It  seems  probable,  therefore,  that  the  specimen  was  mistakenly  identified  as  Wealden ; 
perhaps  washed-up  on  the  beach  near  Hastings. 

BRYOPHYTA 

MUSCI 

?  Bryophyte  leaf. 
Text-figs.  7-9 

Leaf  elliptical,  nearly  2  mm.  long,  just  over  i  mm.  wide.  Margin  entire,  one  end 
(possibly  both)  rounded.  No  midrib  present.  Lamina  flat,  one  cell  thick,  cells 
forming  about  eight  longitudinal  rows,  marginal  cells  smaller,  about  as  long  as  broad. 
Cell  walls  thin,  not  locally  thickened  at  corners;  no  surface  thickenings  observed. 

MATERIAL.     V .  2857, v  •  28570, v  •  28576. 

This  small  fossil  is  known  from  three  isolated  specimens  which  are  very  similar; 
one  has  a  counterpart.  The  above  description  assumes  it  is  like  a  Bryophyte  leaf 
but  I  have  been  unable  to  find  a  Bryophyte  leaf  with  similarly  arranged  cells.  In 
the  leafy  liverworts  the  cells  tend  to  be  isodiametric  or  rounded  and  do  not  form 
longitudinal  rows.  In  Naiadita  lanceolata  Buckman  (Harris  1938)  the  cell  rows 
are  markedly  transverse.  In  the  mosses  they  are  isodiametric  to  elongated  but 
they  may  form  longitudinal  rows.  Transversely  elongated  cells  are  very  rare  in 
mosses  but  they  are  seen  in  a  few  Tortula  species.  However  they  have  leaves  which 
are  otherwise  different. 

Ventral  scales  and  rounded  appendages  of  the  scales  of  the  Marchantiales  have 
more  or  less  elongated  cells  and  where  rows  are  recognizable  they  are  transverse, 
never  longitudinal.  In  fern  ramenta  the  cells  are  often  in  longitudinal  rows  but  are 
usually  very  narrow. 

HEPATICAE 

The  fossils  described  here  are  all  thalloid  and  branch  dichotomously  and  some  of 
them  form  a  rosette-like  growth.  Some  show  definite  Hepatic  features  rather  than 
Algal,  but  others  do  not,  although  they  look  like  Hepatics.  Various  generic  names 
have  been  used  for  such  fossils  by  earlier  authors  (e.g.  Marchantites  Brongniart) 
suggesting  rather  precise  affinity.  Walton  (1925)  in  giving  fine  details  of  such  fossils 


ENGLISH   WEALDEN   FLORA   I  219 

for  the  first  time  revised  their  classification.  A  fossil  with  definite  Hepatic  features 
he  called  Hepaticites.  This  feature  in  practice  is  usually  the  presence  of  rhizoids. 
Fossils  in  which  no  rhizoids  were  demonstrated  he  called  Thallites.  According  to 
this  scheme  the  genus  Marchantites  is  now  restricted  to  forms  showing  definite 
Marchantialean  characters.  Thus  Marchantites  zeilleri  Seward  was  referred  to  the 
genus  Thallites  Walton  (Harris  1942).  Further  study  of  this  plant  has  revealed  the 
presence  of  rhizoids  and  it  is  here  included  in  Hepaticites  Walton. 

Lundblad  (1954)  separated  from  Hepaticites  those  species  showing  Marchantialean 
ventral  scales  as  Marchantites  and  those  with  Marchantialean  air-chambers  as 
Marchantiolites.  In  the  present  work  the  two  genera  of  Walton  suffice,  but  one 
species  which  almost  certainly  has  Marchantialean  ventral  scales  would  have  been 
placed  in  Marchantites  had  it  been  well  enough  preserved  for  the  facts  to  be  secure. 

Genus  HEPATICITES  Walton  1925  :  565 

Hepaticites  zeilleri  (Seward)  comb.  nov. 

PL  2,  figs.  1-3 

1894     Marchantites  Zeilleri  Seward  :  18,  pi.  i,  fig.  3. 
1942     Thallites  zeilleri  (Seward)  Harris  :  397. 
1954     Thallites  zeilleri  (Seward);  Lundblad  :  408. 

EMENDED  DIAGNOSIS.  Plant  thalloid,  dichotomously  branched,  distance  between 
dichotomies  not  constant.  Thallus  about  3  mm.  broad,  composed  of  dark,  well 
defined  midrib  I  mm.  wide  and  delicate  lamina  probably  one  cell  thick.  Margins 
entire.  Cells  of  thallus  in  rows,  longitudinal  in  midrib  but  inclined  at  an  angle  of 
about  45°  in  lamina;  cells  about  30  fi  wide.  Near  apex  midrib  tending  to  divide 
into  number  of  divergent  dark  bands  separated  by  thinner  tissue.  Numerous 
unicellular  thin  walled,  non-tuberculate  rhizoids  arising  on  midrib,  up  to  5  mm. 
long  and  typically  40  p  wide. 

HOLOTYPE  .     V .  2330 . 

DESCRIPTION.  The  holotype  suggests  a  rosette  shaped  thallus  in  the  position  of 
growth  (PL  2,  fig.  i  shows  the  counterpart).  A  second  specimen  shows  smaller 
pieces  of  thallus  mixed  with  fragments  of  Onychiopsis  and  is  more  likely  to  have 
been  deposited  by  water. 

The  thallus  branches  are  delicate  towards  the  tip  and  the  apex  is  nowhere  visible. 
Here  the  midrib  is  more  delicate  and  seems  to  divide.  In  this  delicate  part  longi- 
tudinal rows  of  cells  can  be  seen  but  elsewhere  the  midrib  is  very  dark  and  opaque 
and  presumably  several  cells  thick.  The  rows  of  cells  in  the  lamina  are  clearly 
seen  only  in  a  few  places  and  then  only  the  longitudinal  walls  are  preserved. 

There  is  nothing  suggesting  Marchantialean  air  chambers  or  ventral  scales.  The 
rhizoids  although  clear  are  only  moderately  well  preserved,  unbranched  and  without 
septa;  their  walls  appear  to  be  plain  rather  than  tuberculate.  They  form  groups 
arising  from  the  midrib  and  usually  point  forwards.  A  number  of  detached  rhizoid 
bunches  are  present  beside  the  thallus. 

COMPARISON.  H.  zeilleri  is  the  only  British  Wealden  liverwort  with  a  clearly 
defined  midrib  and  lamina.  Among  other  fossils  it  bears  a  most  striking  resemblance 


220  A   REVISION   OF  THE 

to  H.  laevis  Harris  (1931)  from  the  Lower  Lias  of  East  Greenland.  The  rows  of 
cells  in  that  species  are  more  clearly  defined  and  the  midrib  is  0-5  mm.  wider  than  in 
H.  zeilleri  but  otherwise  they  look  the  same. 

Other  similar  species  are:  H.  arcuatus  (Lindley  &  Hutton)  Harris  (1942  :  394, 
1961  :  7)  which  is  of  similar  dimensions  and  has  a  well  defined  midrib,  but  the  lamina 
is  three  cells  thick,  and  there  are  fewer  rhizoids.  Thallites  yabei  (Kryshtofovich 
1932,  1938,  Oishi  1940,  Harris  1942)  is  a  larger  plant  of  which  no  microscopic  details 
are  known. 

CLASSIFICATION.  H.  zeilleri  looks  very  like  Pellia  and  agrees  with  it  in  the  cell- 
rows  of  the  lamina  and  in  the  rhizoids.  Plants  of  the  Pellia  form  are  familiar  in  the 
Jungermanniales  anacrogynae  but  scarcely  occur  in  other  groups  and  accordingly 
H.  zeilleri  is  placed  here. 

Hepaticites  ruffordi  sp.  nov. 
PL  2,  figs.  4,  5 

DIAGNOSIS.  Thallus  branched  dichotomously  at  intervals  of  about  i  cm.,  width 
about  3-4  mm.  Margins  entire,  apex  probably  notched.  Greater  part  of  thallus 
marked  with  grooves  (considered  to  be  ventral  scales)  at  intervals  of  0-5  mm. 
Grooves  strongly  arched,  longitudinal  in  middle  region  but  curving  outwards  and 
then  slightly  backwards  at  their  ends.  Middle  region  of  older  part  of  thallus,  about 
2  mm.  wide,  marked  by  two  lines  appearing  thicker  and  beyond  which  grooves  do 
not  extend. 

HOLOTYPE  .     V .  2343 . 

DESCRIPTION.  The  only  specimen  known  is  preserved  as  an  impression  in  fine 
sandstone ;  a  little  of  the  plant  substance  remaining  as  a  dark  stain  but  this  has 
cracked  and  no  cells  are  seen  satisfactorily.  Faint  marks  crossing  the  grooves  may 
be  the  impression  of  cells  but  there  is  nothing  more.  No  rhizoids  were  seen  nor  any 
structure  in  the  lamina.  H.  ruffordi  is  tentatively  placed  in  Hepaticites  because  it  is 
thought  to  show  a  distinctive  Hepatic  character — ventral  scales — even  though 
rhizoids  are  not  preserved.  These  ribs,  or  ventral  scales,  are  of  interest  since  they 
are  confined  to  the  Marchantiales.  Unfortunately  the  specimen  is  too  poorly  pre- 
served to  be  sure  that  they  are  indeed  scales.  I  have,  therefore,  refrained  from 
including  it  in  Marchantites  in  Lundblad's  sense. 

COMPARISON.  Marchantites  hallei  Lundblad  (1955)  is  a  liverwort  from  the  Lower 
Cretaceous  of  Patagonia.  It  is  similar  in  dimensions  to  H.  ruffordi  but  is  known  in 
much  more  detail,  having  two  sorts  of  rhizoids  and  well  preserved  ventral  scales. 
Hepaticites  haiburnensis  Harris  (1961  :  9)  is  another  similar  form  in  which  the  thallus 
extends  further  beyond  the  scales. 

Genus  THALLITES  Walton  1925  :  564. 

For  discussion  of  Thallites  see  p.  219.  The  two  species  described  here  are  associa- 
ted in  considerable  numbers  on  certain  bedding  planes.  Seward  placed  them  in 


ENGLISH  WEALDEN   FLORA   I  221 

Algites  thinking  them  to  be  marine  red  algae.  Closer  study  shows  that  they  are 
more  likely  to  be  terrestrial  liverworts  though  proof  is  lacking.  They  are  certainly 
not  marine  for  the  rocks  in  which  they  occur  are  of  purely  freshwater  origin  and 
indeed  the  specimens  are  crowded  with  the  small  freshwater  lamellibranch,  Neo- 
miodon.  Despite  careful  study  the  two  forms  described  here  show  so  few  details  that 
they  can  hardly  be  considered  to  be  properly  characterized  species.  However, 
they  have  been  described  as  such  and  as  their  names  are  fairly  well  known  they  are 
here  retained. 

Thallites  valdensis  (Seward)  comb.  nov. 

Text-figs.  12-15 
1894     Algites  valdensis  Seward  :  4,  pi.  i,  figs,  i,  2. 

EMENDED  DIAGNOSIS.  Plant  thalloid;  thallus  forming  rosette  of  segments  branch- 
ing dichotomously  three  or  four  times.  Diameter  of  whole  thallus  up  to  5-5  cm.; 
width  of  branches  typically  4-5  mm.,  branches  ending  bluntly.  Midrib  represented 
by  median  dark  line  TOO  /*  wide  and  dichotomizing  5  mm.  behind  dichotomy  of 
thallus.  Cells  in  midrib  region  not  differing  from  rest  of  lamina.  Dark  ribs  passing 
obliquely  across  lamina  at  intervals  of  about  300  /i.  Lamina  apparently  composed 
of  one  layer  of  thin  walled  cells  ranging  in  size  from  about  15-75  /£. 

HOLOTYPE  .     V .  2857 . 

DESCRIPTION.  Seven  specimens  were  examined  on  three  blocks  of  fine-grained 
calcareous  sandstone.  The  preservation  is  poor  and  splitting  of  the  rock  has 
resulted  also  in  splitting  of  the  fossil  into  two  parts.  The  rosette  of  branches  strongly 
suggests  that  the  plant  has  been  preserved  in  position  of  growth.  It  accords  with  a 
suggestion  made  by  Walton  (1949)  for  a  Carboniferous  thallus  which  he  called  cf. 
Hepaticites.  Like  Walton's  specimen  there  are  several  very  perfect  rosettes  and  it 
looks  as  though  the  central  and  oldest  part  of  the  thallus  had  decayed  away  before 
the  plants  were  buried  and  preserved.  Fragments  would  be  unlikely  to  drift  to- 
gether into  a  neat  rosette. 

The  general  appearance  of  the  cells  is  constant  throughout  the  whole  thallus 
though  they  vary  considerably  in  size  and  shape.  They  are  on  the  whole  thin  walled 
but  additional  organic  material  is  present  which  tends  to  conceal  their  walls.  The 
positions  of  the  median  line  of  the  thallus  and  the  ribs  are  merely  marked  by  a  greater 
concentration  of  this  dark  structureless  matter.  The  central  strand  is  at  best  faint 
and  is  clear  in  only  two  thallus  branches.  The  rest  show  no  central  strand  at  all. 
The  oblique  ribs  though  not  always  seen  are  more  widespread.  I  think  that  the 
pattern  of  cell-like  polygons  represents  true  cells  and  is  not  some  effect  of  poor 
preservation,  for  instance  the  imprint  of  sand  grains  on  a  thin  coaly  layer.  At 
best  they  certainly  look  like  cells  and  associated  with  T.  valdensis  are  small  frag- 
ments of  equally  delicate  plant  tissue,  also  one  layer  thick  but  with  quite  differently 
shaped  cells.  One  of  these  fragments  is  the  small  leaf  described  here  as  PBryophyte 
leaf;  another  is  more  like  a  fern  ramentum  with  very  narrow  cells.  In  neither  is 
there  any  possibility  that  the  cellular  pattern  has  been  imposed  by  sand  grains. 


222 


A   REVISION   OF  THE 


10 


kr:*^£^'  , -••""' 


14 


15 


FIGS.  7-9.  (?)  Bryophyte  leaf.  7,  8,  9.  Leaf  showing  a  single  layer  of  cells.  Figs.  8, 
9  are  part  and  counterpart.  Fig.  7,  V. 28576.  Figs.  8,  9,  ¥".2857.  All  x  20. 

FIGS,    10,    ii.     Thallites    catenelloides   (Seward).     10.     Part   of   specimen  showing  cells. 
V.2857.      x  100.     n.     Part  of  thallus.     ¥.2857.      x  4. 

FIGS.  12-15.  Thallites  valdensis  (Seward).  12.  Part  of  specimen  showing  cells.  ¥.2857. 
X  100.  13.  Rosette  shaped  thallus  suggesting  preservation  in  situ.  V. 28576.  x  i. 
14.  A  branch  of  the  thallus  showing  thinner  peripheral  region  and  ribs  of  the  thicker 
central  region.  V. 28576  x  4.  15.  Apex  of  branch  showing  the  pattern  of  diver- 
gent ribs.  ¥.28576.  x  15. 


ENGLISH   WEALDEN   FLORA   I  223 

No  Marchantialean  air  chambers  are  visible,  nor  is  there  any  clear  suggestion  of 
two  superimposed  layers  of  cells,  though  presumably  further  layers  do  occur,  at 
least  on  the  midrib  and  lateral  ribs.  No  rhizoids  are  present. 

Transfers  were  made  but  these  yielded  no  additional  information.  Chondms 
crispus,  the  red  alga  with  which  Seward  compared  T.  valdensis  was  found  to  be  quite 
unlike  it  in  having  a  much  thicker  thallus  and  cells  of  very  different  shape. 

COMPARISON.  T.  catenelloides  has  a  thallus  of  similar  texture  but  is  a  much  smaller 
plant.  Of  the  other  British  Wealden  forms  Hepaticites  zeilleri  differs  in  its  clearly 
marked  midrib  and  strong  rhizoids.  H.  ruffordi  differs  in  its  strong  pattern  of 
arcuate  ribs. 

Most  thalloid  liverwort-like  plants  have  a  broad  and  strongly  marked  midrib. 
The  following  which  are  of  similar  dimensions  to  T.  valdensis  have  only  a  faint 
midrib. 

T.  umlensis  Kryshtofovich  &  Prynada  (1933)  from  the  Rhaeto-Liassic  of  U.S.S.R. 
is  rather  narrower  and  has  more  pointed  lobes.  No  microscopic  details  are  known. 

Hepaticites  rosenkrantzi  Harris  (1931, 1937)  from  the  Lower  Lias  of  East  Greenland 
has  no  midrib  but  is  two  cells  thick,  the  cells  tending  to  be  elongated  and  in  rows. 

H.  amauros  Harris  (1937),  also  from  the  Lower  Lias  of  East  Greenland,  has  a 
midrib  which  is  a  dark,  narrow  strip  but  the  thallus  is  many  cells  thick. 


Thallites  catenelloides  (Seward)  comb.  nov. 
Text-figs.  10,  ii 

1894     Algites  catenelloides  Seward  :  7,  pi.  i,  figs,  i,  2. 

EMENDED  DIAGNOSIS.  Plant  thalloid;  thallus  dichotomously  branched,  branches 
about  0-5  mm.  wide;  margins  probably  entire.  Thallus  composed  of  uniform  tissue 
one  cell  thick;  cells  thin  walled,  diameter  about  50/4. 

HOLOTYPE.     ¥.2857. 

DESCRIPTION.  Numerous  specimens  occur  on  the  same  blocks  as  T.  valdensis 
and  in  some  cases  they  are  closely  associated.  The  specimens  of  T.  catenelloides 
vary  from  small  fragments  to  what  are  probably  whole  thalli  measuring  i  cm.  across. 
They  do  not  form  rosettes.  The  narrow  ribbon-like  thallus  branches  frequently 
and  in  parts  appears  to  have  an  undulating  though  entire  margin.  The  specimen  in 
Text-fig,  ii  is  the  same  as  that  figured  by  Seward.  His  suggestion  that  the  thallus 
has  constrictions  similar  to  the  red  alga  Catenella  seems  to  be  based  only  on  the 
appearance  of  part  of  this  particular  thallus.  It  is  not  seen  in  the  other  specimens 
and  is  probably  due  to  the  undulating  margin. 

The  cells  of  the  thallus  are  identical  with  those  of  T.  valdensis  and  the  description 
for  that  species  will  serve  for  both.  Transfers  were  made  but  these  showed  only  the 
single  layer  of  cells  composing  the  thallus. 

COMPARISON.  In  Catenella  opuntia  the  thallus  is  several  cells  thick  and  has  a 
surface  layer  of  cells  much  smaller  than  those  of  T.  catenelloides,  whilst  the  central 
cells  are  larger  and  elongated. 

GEOL.   17,  5  19 


224  A  REVISION   OF  THE 

All  other  Mesozoic  hepatics,  including  the  three  other  English  Wealden  species, 
are  much  larger  than  T.  catenelloides.  The  only  plants  of  similar  dimensions  are 
from  the  Carboniferous.  Two  rather  similar  fossils  are  Thallites  willsi  Walton  (1925, 
1928,  1949)  and  Hepaticites  langi  Walton  (1928).  Neither  has  a  definite  midrib, 
both  differ  in  being  more  than  one  cell  thick  and  H.  langi  has  well  marked  rhizoids. 
T.  willsi  which  is  more  comparable  has  a  rather  broader  thallus. 

PTERIDOPHYTA 

LYGOPODIALES 

SELAGINELLACEAE 

Genus  SELAGINELLA  Spring 

Only  one  other  Selaginella  from  the  Mesozoic,  S.  hallei  Lundblad  (19500),  is 
known  in  as  much  detail  as  the  Wealden  species  described  below  and  referred  here 
to  the  living  genus. 

The  genus  Selaginellites  Zeiller  (1906)  was  originally  used  for  heterosporous  fossils 
having  more  than  four  megaspores  in  each  megasporangium  and  was  thought  to 
differ  in  this  respect  from  the  Recent  genus  Selaginella.  Subsequently  it  was  used 
by  Halle  (1907)  for  all  heterosporous  fossils  regardless  of  the  number  of  megaspores. 
Later  work  proved  that  living  species  may  produce  more  than  four  megaspores  in 
each  megasporangium  (Duerden  1929) .  Miner  (1932)  used  Selaginellites  for  isolated 
megaspores  which  would  ordinarily  have  been  called  Triletes  but  Harris  (1935) 
recommended  that  its  use  should  be  restricted  to  fossil  shoots  which  showed  evidence 
of  being  closer  to  Selaginella  than  any  other  genus.  Dijkstra  (1949)  was  the  first 
to  place  the  English  Wealden  species  in  Selaginella  but  this  may  have  been  inadver- 
tent as  he  later  returned  to  the  use  of  Selaginellites  without  comment.  I  consider 
that  the  Wealden  species  in  no  way  differs  from  a  typical  anisophyllous  Selaginella 
with  the  normal  number  of  four  megaspores  in  each  megasporangium  and  it  has, 
therefore,  been  referred  to  the  living  genus. 

Selaginella  dawsoni  (Seward) 
Text-figs.  16-24 

1894     Plantae  incertae  sedis  Seward  :  20,  pi.  i,  figs.  8,  9. 

1913     Selaginellites  Dawsoni  Seward  :  87,  text-fig,  i. 

I9i3«  Selaginellites  Dawsoni  Seward;  Seward  :  86,  pi.  4,  figs.  1-7. 

1949     Selaginella  Dawsoni  (Seward)  Dijkstra  :  19.  (name  in  list). 

DIAGNOSIS.  Fertile,  dichotomously  branched  shoots;  sterile  portion  of  shoot  in 
part  known  about  i  mm.  wide,  dorsiventral,  covered  with  small  overlapping  leaves. 
Leaves  anisophyllous,  in  four  ranks;  leaves  of  upper  two  rows  alternating  and 
appressed  to  stem;  rhomboid  in  shape,  about  0-5  mm.  long;  leaves  of  lower  two 
rows  longer,  probably  about  I  mm.  long,  slightly  spreading  laterally  but  pointing 
towards  apex.  Strobilus  terminal,  unbranched,  2  mm.  wide,  at  least  3  cm.  long. 
Overlapping  sporophylls  decussately  arranged,  apparently  all  alike  and  longer  than 
leaves,  about  1-5  mm.  Sporangia  more  or  less  rhomboid  in  outline  about  0-5  mm. 


ENGLISH  WEALDEN   FLORA  I  225 

across.  Megasporangia  fewer  but  distributed  throughout  length  of  strobilus;  each 
containing  four  megaspores.  Megaspore  with  rounded  dorsal  surface,  ventral 
surface  formed  by  three  flattened  contact  facets ;  diameter  285  ^-340  /*  (mean  306  /*) ; 
height  about  15  %  less ;  tri-radiate  ridges  very  broad,  about  30  ju,  high,  edge  rounded, 
not  divided;  length  equal  to  radius  of  spore;  arcuate  ridges  absent.  Ventral  surface 
crowded  with  tubercles,  circular  or  oblong  and  5-12  fi  across ;  tubercles  also  present 
on  surface  of  tri-radiate  ridges  but  less  prominent.  Dorsal  surface  of  spore  having 
a  coarser  pattern  of  widely  spaced  ribs,  irregular  in  shape,  about  4  ju  wide  and  up  to 
60  [i  long;  spore  surface  smooth  between  ribs.  Spore  wall  easily  separable  into 
two  distinct  layers,  outer  20-30  fi  thick,  inner  about  2  fi  thick.  Inner  membrane 
straw-coloured  and  indistinctly  marked  with  small  pits.  Substance  of  wall,  in- 
cluding tubercles  and  ribs,  light  brown  in  colour  and  composed  of  minute  granules. 

Microsporangia  contain  numerous  microspores  which  are  almost  spherical  but 
with  flattened  contact  facets;  average  diameter  45  /i  (range  38-52  fi}\  tri-radiate 
mark  fine  but  distinct,  equal  to  about  two  thirds  of  spore  radius.  Contact  facets 
seen  as  three  lobes  separated  by  tri-radiate  marks  not  quite  reaching  notches  between 
lobes;  surface  of  facets  more  or  less  covered  with  indistinct  tubercles,  1-2  ^  in  dia- 
meter. Spore  wall  distinctly  two  layered,  together  only  about  2  ju,  thick  at  facets 
but  5-6  fi  thick  elsewhere.  Outside  facets  both  layers  uneven  in  thickness;  inner 
layer  becoming  thicker  opposite  a  thin  place  in  the  outer.  Maximum  thickness  of 
inner  layer  about  3  /*,  outer  about  5  fi. 

HOLOTYPE  .     V .  2328 . 

DESCRIPTION.  The  only  material  available  consists  of  two  fragments  of  shoot 
and  a  few  lengths  of  cone  which  are  present  on  two  blocks  of  ironstone.  Some  of 
the  cone  specimens  have  been  much  damaged.  The  two  shoots  are  each  about  3  cm. 
long  and  frequently  branch  dichotomously.  For  the  most  part  the  sterile  portion 
of  the  plant  is  seen  merely  as  a  faint  outline  on  the  rock,  the  stem  and  some  of  the 
leaves  being  replaced  by  a  white  mineral  and  scarcely  any  carbonaceous  plant  material 
remains.  On  only  one  or  two  branches  can  the  phyllotaxis  be  worked  out.  Here 
the  mineralized  outlines  of  the  dorsal  leaves  are  seen  in  surface  view  and  the  ventral 
leaves  in  side  view.  This  is  the  portion  of  shoot  seen  in  Text-fig.  17.  Although 
the  margins  of  the  leaves  are  shown  as  entire  it  is  impossible  to  tell  whether  they 
are  correctly  represented  since  the  precise  margins  are  unrecognizable.  All  that  is 
known  of  the  shape  of  the  ventral  leaves  is  from  what  they  show  in  side  view.  They 
are  seen  to  have  thicker  tissue  in  the  centre.  The  exact  arrangement  of  the  dorsal 
leaves  just  behind  a  dichotomy  is  uncertain. 

The  most  complete  strobilus  is  3  cm.  long  but  is  probably  incomplete,  ending  as 
it  does,  very  sharply  at  the  edge  of  the  rock.  Although  this  strobilus  does  not  have 
any  attached  sterile  part,  each  of  the  two  shoots  have  bases  of  similar  strobili 
attached  to  one  of  their  branches.  Three  portions  of  cones  lying  close  to  one  another 
and  parallel  indicate  that  a  number  of  them  were  borne  terminally  on  the  plant. 

The  sporangial  walls  are  not  preserved  though  the  outlines  of  the  microspore 
masses  are  clearly  seen.  Only  one  tetrad  of  megaspores  is  present  in  the  megas- 
porangium. 


226 


A   REVISION   OF  THE 


The  sporophylls  are  seen  only  at  the  periphery  of  the  strobilus  in  side  view,  their 
general  shape  being  unknown. 

Slides  of  detached  spores  prepared  by  Dr.  W.  Chaloner  and  kindly  lent  by  him 
show  in  most  of  the  megaspores  the  two  very  distinctive  ornamentations  of  the 


16 


17 


\ 


20 


24 


FIGS.  16-24.  Selaginella  dawsoni  (Seward).  16.  Part  of  a  strobilus  with  microsporangia, 
megaspores  in  outline  and  sporophylls  in  side  view.  V.3I5I.  x  10.  17.  Part  of  a 
sterile  shoot  showing  anisophylly.  Holotype,  ¥.2328.  x  10.  18.  Ventral  surface 
of  microspore  showing  contact  facets,  tri-radiate  marks  and  two-layered  wall  in  optical 
section.  V.  31510.  X  500.  19.  Dorsal  surface  of  the  same  spore.  V.  31510.  x  500. 

20.  Detail  from  the  same  spore  showing  tubercles  on  contact  facets.    V. 31510.    x  2500. 

21.  Ventral  face  of  megaspore.     Specimen  lost,      x  100.     22.     Detail  of  megaspore 
showing   broad  tri-radiate   ridge   and  facet  with  tubercles.     V.3i5iy.      x  250.       23. 
Single  rib  from  dorsal  surface  of  megaspore.     V.3i5ry.      x  250.     24.     Megaspore  in 
side  view  showing  dorsal  surface,  two  facets  and  a  tri-radiate  ridge.     V.3i5ry.      X  100. 


ENGLISH   WEALDEN   FLORA   I  227 

different  surfaces  and  a  number  of  broken  specimens  show  the  two  separate  layers 
of  the  spore  wall.  The  thin  inner  layer  is  easily  removed  and  mounted  in  glycerine 
jelly  without  further  treatment.  Very  brief  maceration  of  the  the  thick  outer 
coat  renders  it  transparent  before  mounting.  Some  of  the  megaspores  do  not  show 
the  two  distinctive  sculptures  quite  so  clearly  as  the  figured  specimens.  In  these 
the  dorsal  ribs  are  not  so  coarse  and  well  defined.  Lack  of  abundant  material 
prevents  more  detailed  study  of  this  variation. 

Some  of  the  mounted  microspores  remain  in  tetrads  but  most  of  them  occur  singly. 
The  layers  of  the  wall  are  clearly  seen  in  optical  section,  their  irregular  adjacent 
surfaces  closely  interlocking,  the  one  being  at  its  thickest  where  the  other  is  thinnest. 
The  pits  in  the  outer  wall  while  most  numerous  on  the  dorsal  side  also  occur  on  the 
ventral  side  but  not  on  the  facets. 

By  squashing  the  spores  the  thin  walled  facets  may  be  detached  and  their  surface 
more  easily  studied.  In  some  specimens  the  tuberculate  pattern  covers  the  whole 
surface  of  the  facet  but  is  only  marginal  in  others. 

COMPARISON.  Only  one  other  Mesozoic  Selaginella  shoot  with  cone  has  been 
described,  namely  5.  hallei  Lundblad  (1950^)  from  the  Rhaetic  of  Sweden.  The 
sterile  shoot  is  similar  in  size  but  with  slightly  larger  and  more  spreading  leaves. 
Like  S.  dawsoni  it  is  anisophyllous,  but  the  leaves  are  known  in  more  detail.  The 
strobilus  of  5.  hallei  is  much  shorter  and  has  sporophylls  which  are  described  as 
spirally  arranged  but  may  equally  well  be  described  as  in  alternating  whorls  of 
about  four.  There  are  four  megaspores  in  each  megasporangium.  Selaginellites 
polaris  Lundblad  (1948)  is  a  Selaginelloid  cone  from  the  Trias  of  Greenland.  It  is 
much  stouter  than  5.  dawsoni  (7  mm.  wide)  and  has  numerous  megaspores  in  each 
megasporangium. 

Selaginella  dichotoma  Velenovsky  (1931)  is  a  sterile  shoot  from  the  Cretaceous  of 
Bohemia  with  more  spreading  leaves. 

Selaginellites  nosikovii  Kryshtofovich  &  Prynada  (1932)  is  a  sterile  shoot  from  the 
Jurassic  of  Siberia  which  also  has  larger,  more  spreading  leaves  than  5.  dawsoni. 

Among  living  species  5.  sanguinolenta  is  very  similar  in  size  and  leaf  arrangement, 
but  the  spores  are  different  the  megaspores  being  of  similar  size  but  with  much  more 
prominent  tubercles  on  the  dorsal  surface. 

5.  emmeliana  possesses  strikingly  similar  megaspores  to  those  of  5.  dawsoni  with 
the  same  shape  and  sculpturing  both  on  the  dorsal  and  ventral  surfaces.  S.  emmel- 
iana, however,  possesses  other  megaspores  which  show  a  great  deal  of  variation 
both  in  size  and  surface  ornament.  Not  all  have  different  dorsal  and  ventral  pat- 
terns, some  having  the  coarse  ribbed  pattern  on  both  surfaces  whilst  others  have  the 
fine  tuberculate  pattern  on  both.  The  foliage  of  5.  emmeliana  is  quite  different  from 
that  of  5.  dawsoni. 

EQUISETALES 

Genus  EQUISETITES  Sternberg 

REMARKS.  The  external  morphology  of  E.  lyelli  has  been  fully  described  by 
Allen  (1941).  Additional  information  about  this  species  (Watson  1964)  was  obtained 


228  A   REVISION   OF   THE 

from  some  petrified  fragments  found  by  Mr.  P.  Whybrow  (British  Museum,  Natural 
History).  Very  recently  Mr.  Whybrow  has  found  much  more  extensive  petrified 
material  and  in  addition  new  compression  material  is  available  with  easily  prepared 
cuticle.  It  will,  therefore,  be  necessary  to  make  a  complete  revision  of  the  species 
using  all  available  material,  including  that  described  by  Allen  (1941). 

Descriptions  of  all  three  species  of  Wealden  Equisetites  (Seward  1894)  will  be 
included  in  a  later  volume. 


FILICALES 
Family  MATONIACEAE 

Genus  MATONID1UM  Schenk  1871  :  219 
Matonidium  goepperti  (Ettingshausen) 

1852  A  lethopteris  goepperti  Ettingshausen  :  8,  pi.  7,  fig.  7. 

1894  Matonidium  Gopperti  (Ett.);  Seward  :  63,  text-figs.  7,  8. 

1913  Matonidium  goepperti  (Ett.);  Seward  :  89,  text-fig.  zC,  pi.  14,  fig.  3a. 

1961  Matonidium  goepperti  (Ett.) ;  Harris  :  112,  text-figs.  37,  38.  (see  also  for  earlier  references). 

Matonidium  goepperti  (Ett.)  has  been  fully  revised  by  Harris  (1961)  using  abundant, 
well  preserved  material  from  the  Yorkshire  Jurassic.  The  English  Wealden  material 
is  scanty  and  ill-preserved  but  agrees  in  all  available  characters  with  the  Yorkshire 
material  and  I  regard  the  two  as  specifically  identical. 


Family  DIPTERIDACEAE 

Genus  HAUSMANNIA  Dunker  1846  :  12 

Hausmannia  dichotoma  Dunker 

PI.  3,  figs,  i,  2 

1846  Hausmannia  dichotoma  Dunker  :  12,  pi.  5,  fig.  i ;  pi.  6,  fig.  12. 

1894  Dictyophyllum  Roemeri  Schenk;  Seward  :  140,  text-figs.  16,  17. 

1906  Hausmannia  dichotoma  Dunker;  Richter  :  18,  pis.  3-6. 

1913  Hausmannia  pelletieri  Seward  :  89,  pi.  14,  figs.  1-3. 

1961  Hausmannia  dichotoma  Dunker;  Harris  :  126,  text-fig.  43. 

This  species  is  widespread  in  the  Lower  Cretaceous  of  Europe  and  less  common  in 
the  Jurassic.  It  has  been  revised  by  Harris  (1961)  who  gives  an  emended  diagnosis 
based  on  Richter's  material  (Lower  Cretaceous ;  Germany)  and  on  Yorkshire  Jurassic 
specimens.  The  leaves  of  H.  dichotoma,  both  sterile  and  fertile,  are  quite  well  known 
but  nothing  is  yet  known  of  the  sorus.  The  English  Wealden  material  consists  of  a 
few  imperfect  sterile  leaves  which  add  nothing  to  our  knowledge  of  the  species. 
The  best  examples  are  figured  in  PI.  3,  figs,  i,  2. 


ENGLISH   WEALDEN    FLORA    I  229 

Family  SCHIZAEACEAE 

Genus  RUFFORDIA  Seward  1894  :  75 
Ruffordia  goepperti  (Bunker) 

PL  4,  figs.  1-7;  PL  5,  figs.  1-4;  Text-figs.  25-31. 

Selected  references : 

1843     Cheilanthites  Gopperti  T>\mk.&c  :  6. 

1894     Ruffordia  Gopperti  (Dunker)  Seward  :  76,  pi.  3,  figs.  5,  6;  pi.  4;  pi.  5,  figs.  1-5. 

1894     Ruffordia  Gopperti  (Dunker)  var.  latifolia  Seward  :  85,  pi.  6,  figs,  i,  ia. 

1910     Ruffordia  Goepperti  (Dunker) ;  Seward  :  350,  text-fig.  260  A-C.   (figures  repeated  from 

Seward  1894). 

1913     Ruffordia  goepperti  (Dunker);  Seward  :  91,  text-fig.  2A.  (drawing  of  a  spore). 
1921     Ruffordia  Goepperti  (Dunker);  Halle  :  10,  pi.  2,  figs.  1-8. 

1958     Ruffordia  goepperti  (Dunker);  Couper  :  109,  pi.  17,  figs.  4-6.  (photographs  of  spores). 
1966     Ruffordia  goepperti  (Dunker);  Hughes  &  Moody-Stuart  :  274,  pi.  43,  figs.  1-12.  (spores). 

EMENDED  DIAGNOSIS.  Frond  triangular,  tripinnate  to  quadripinnate ;  rachis 
slender.  Pinnae  triangular  or  ovate;  primary  pinnae  nearly  opposite;  secondary 
pinnae  increasingly  alternate.  Sterile  and  fertile  parts  distinct.  Sterile  pinnules 
variable  in  width,  decurrent;  narrow  pinnules  linear,  apices  mucronate  or  acute, 
venation  Coenopterid;  wider  pinnules  ovate,  Sphenopterid,  with  small  marginal 
tooth  at  each  vein  ending.  Fertile  pinnules  reduced  in  length  but  not  breadth; 
more  widely  outspread,  not  decurrent,  margins  undulating,  apices  bluntly  rounded. 
Sporangia  borne  singly  on  underside  of  pinnules  without  indusium ;  typically  twelve 
on  a  pinnule,  arranged  in  two  longitudinal  rows.  Annulus  transverse,  apical,com- 
posed  of  about  fifteen  to  twenty-five  long  narrow  cells  radiating  from  a  central  plate ; 
extending  at  least  to  middle  of  sporangium.  (Remaining  parts  of  sporangium 
unknown,  spore  number  uncertain).  Spores  rounded-triangular  with  convex  sides, 
typically  45  /£  equatorial  diameter  (range  35  fi  to  60  //.),  tri-radiate  cracks  almost 
reaching  corners.  Apical  parts  of  facets  smooth,  rest  of  facets  and  whole  of  distal 
surface  covered  with  ribs  separated  by  grooves.  Ribs  forming  three  sets,  each  set 
starting  on  contact  face  and  extending  on  to  distal  surface ;  in  each  set  ridges  nearly 
parallel.  Ridges  of  a  set  joining  near  tri-radiate  cracks  and  also  joining,  though 
less  regularly,  along  continuation  of  tri-radiate  cracks  onto  back  of  spore.  Smooth 
apical  parts  of  facets  as  thick  as  ribs. 

DESCRIPTION  AND  DISCUSSION.  Ruffordia  goepperti  (Dunker)  is  represented  by  a 
large  number  of  sterile  specimens  which  show  a  wide  range  in  the  form  of  the  pin- 
nules, the  extremes  of  which  are  shown  in  PL  4,  figs.  I,  4.  The  specimens  with 
broadest  pinnules  were  designated  R.  goepperti  var.  latifolia  (Seward  1894)  but  all 
are  now  included  in  R.  goepperti  (Dunker)  as  there  seems  to  be  intergradation, 
though  the  possibility  of  the  widest  forms  belonging  to  a  different  species  is  not 
excluded.  There  are  three  fertile  specimens  amongst  the  English  material  (V.2i6o 
V .  2295,  V .  2192) .  Two  are  completely  fertile  (V .  2160  and  V .  2192)  whilst  the  other 
(¥.2295)  has  basal  fertile  pinnae  and  a  sterile  top  (PL  5,  fig.  i).  Specimens  ¥.2192 
and  ¥.2295  are  preserved  in  the  characteristic  red  ironstone  and  only  the  spore 
masses,  and  sometimes  the  sporangium  annulus,  are  well  preserved,  the  rest  of  the 


230 


A   REVISION   OF   THE 


31 


FIGS.  25-31.  Ruffordia  goepperti  (Dunker).  25.  Broad  form  of  pinnules  showing  vena- 
tion. V.2357.  x  8.  26.  Fertile  pinnules  showing  undulating  margins.  Broken 
lines  indicate  positions  of  sporangia.  V.2i6o.  x  15.  27.  Sporangium  showing  thick- 
ened walls  of  annulus.  V.22Q5.  x  100.  28.  Spore  in  equatorial  view  showing 
widely  spaced  ribs  and  shapes  of  ribs  in  section,  i.e.  rounded,  flattened,  grooved.  Slide 
from  V.2i6o,  x  750.  29,  30.  Proximal  and  distal  surfaces  of  the  same  spore  showing 
wide  ribs  closely  spaced.  Slide  from  V.  21 60,  X  750.  31.  Spore  with  narrow  ribs  and 
wide  grooves.  Slide  from  V.  2 1926,  x  750. 


ENGLISH   WEALDEN   FLORA   I  231 

frond  being  only  a  faint  impression  on  the  rock.  This  is  a  typical  condition  of  the 
plants  in  this  matrix  but  it  is  sometimes  useful  for  details  of  venation  (PL  4,  fig.  7). 
Specimen  V .  2160  in  a  soft,  fine,  grey  siltstone  matrix  is  much  better  preserved  though 
the  frond  is  heavily  varnished  and  badly  cracked.  Text-figure  26  shows  the  rounded 
lines  of  the  fertile  pinnules  with  a  lobed  appearance.  The  specimen  presents  the 
upper  surface  of  the  frond  and  the  sporangia  below  are  seen  only  in  faint  outline. 
A  balsam  transfer  revealed  no  details  of  the  sporangia. 

The  spores  of  R.  goepperti  (Dunker)  were  inadequately  figured  by  Seward  (1913, 
text-fig.  2A)  using  ¥.2192  and  by  Couper  (1958,  pi.  17,  figs.  4-6)  using  Seward's 
slide  preparation.  Recently  the  spores  have  been  studied  in  detail  by  Hughes  & 
Moody-Stuart  (1966)  using  ¥.2192  and  Seward's  slide  mentioned  above.  I  have 
found  that  V .  2192  yields  spores  which  are  somewhat  different  from  those  of  V .  2295 
and  V.2i6o.  The  spores  agree  in  size  range  and  general  form  but  differ  in  details 
of  sculpturing.  The  spores  from  V .  2295  and  V .  2160  have  ribs  (muri)  from  2  /*  to  5  /* 
wide  separated  by  furrows  (lumina)  0-5  [i  to  2  fi  wide  but  the  ribs  of  any  particular 
spore  are  always  wider  than  the  furrows  and  the  majority  of  spores  have  ribs  about 
3  ju,  wide  and  furrows  I  /£  wide  (Text-figs.  28-30).  The  ribs  vary  in  profile  and  may 
be  rounded,  flattened  or  indented;  this  variation  is  seen  in  Text-fig.  28.  Spores 
from  ¥.2192  have  the  sculpture  in  almost  exactly  the  reverse  proportions  (Text-fig. 
31),  the  ribs  being  i  ju,  to  2  /i  wide  and  spaced  2  fi  to  5  /£  apart.  Hughes  &  Moody- 
Stuart  (1966)  give  a  detailed  diagnosis  of  these  spores  and  numerous  figures.  I 
have  prepared  spores  from  various  parts  of  ¥.2192  but  I  have  been  unable  to  find 
any  with  broad  ribs.  Neither  ¥.2295  nor  V.2i6o  yielded  spores  with  narrow  ribs 
and  I  could  find  no  suggestion  of  gradation  between  the  two  types  of  spores. 

The  identity  of  ¥.2192  is  not  in  doubt,  however,  because  in  all  other  respects  it 
agrees  perfectly  with  the  other  two  specimens.  It  is  closely  similar  to  the  basal 
pinnae  of  ¥.2295  and  also  has  undulating  pinnules  as  in  V.2i6o. 

Fertile  specimens  of  R.  goepperti  (Dunker)  from  Manchuria  were  figured  by  Halle 
(1921)  including  photographs  of  the  sporangia,  each  represented  by  a  well  preserved 
annulus.  This  annulus  is  certainly  an  apical  ring  of  the  Schizaeaceous  type.  Halle's 
photographs  show  the  long  annulus  cells  radiating  from  a  small  circular  space  or 
"  plate  ".  This  suggests  that  the  annulus  cells  in  Text-fig.  27  do  not  show  their 
full  length,  although  he  does  report  cases  where  the  "  central  area  seems  to  be  much 
wider  than  others  ".  Halle  removed  what  he  considered  to  be  a  whole  sporangium 
and  gives  three  countings  in  the  region  of  400.  I  have  removed  what  appeared  to  be 
whole  sporangia  from  the  balsam  transfers  and  obtained  various  counts  under  100. 
I  regard  my  counts  as  highly  suspect  as  it  was  difficult  to  be  sure  that  the  spore 
masses  were  whole  sporangia.  One  cannot  know  how  reliable  Halle's  count  is  but 
he  reports  that  "  in  some  cases  it  was  possible  to  remove  the  whole  contents  of  a 
sporangium  more  or  less  intact  ".  In  living  members  of  the  Schizaeaceae  the  spore 
output  is  128  or  256.  Hughes  &  Moody-Stuart  (1966)  regard  the  spores  of  Halle's 
material  as  of  a  different  type  from  those  found  in  R.  goepperti  (Dunker)  from  the 
English  Wealden.  However,  his  photographs  of  sterile  and  fertile  fronds  appear  to 
be  very  similar  to  the  English  specimens.  One  of  them  (pi.  2,  fig.  2)  shows  short, 
undulating,  fertile  pinnules.  Halle's  photograph  of  the  spores  (pi.  2,  fig.  8)  is  a 


232  A   REVISION   OF   THE 

poor  one  and  considering  the  spore  differences  in  the  English  specimens  of  R. 
goepperti  (Dunker)  I  have  retained  the  Manchurian  material  within  the  species 
until  it  can  be  proved  otherwise.  Certain  details  of  the  sporangium  in  the  diagnosis 
above  are  based  on  Halle's  account. 

Among  living  genera  of  the  Schizaeaceae,  R.  goepperti  (Dunker)  most  resembles 
Anemia  with  its  distinct  and  reduced  fertile  pinnae.  The  sterile  frond  of  Anemia 
adiantifolia  is  very  similar  in  form  to  the  broader  specimens  of  R.  goepperti  whilst 
the  narrow  pinnules  closely  resemble  Anemia  cuneata  (Bower  1926,  text-fig.  440). 

The  sculptured  spores  of  the  genus  Mohria  are  of  exactly  the  same  type  as  R. 
goepperti  (Dunker).  This  type  of  spore  also  occurs  in  the  other  Wealden  Schizaea- 
ceous  fern,  Pelletieria  valdensis  Seward  and  a  comparison  is  included  in  the  descri- 
tion  of  that  species. 


Genus  PELLETIERIA  Seward  1913  :  91 

Pelletieria  valdensis  Seward 
PI.  6,  figs,  i,  2;  Text-figs.  32-40 

1913     Pelletieria  valdensis  Seward  :  91,  text-figs.  2B,  3,  4;  pi.  12,  figs.  I2a,  I2&;  pi.  14,  fig.  5. 

1958     Pelletieria  valdensis  Seward ;  Couper  :  109. 

1966     Pelletieria  valdensis  Seward;  Hughes  &  Moody-Stuart,  pi.  44,  figs.  1-8;  pi.  45,  figs.  1-7. 

EMENDED  DIAGNOSIS.  Known  only  from  fertile  specimens.  Frond  at  least 
quadripinnate ;  consisting  of  slender  axis  branched  repeatedly,  with  no  lamina  except 
at  tips  of  ultimate  branches.  Tips  bearing  groups  of  fertile  pinnules;  each  pinnule 
tightly  rolled,  completely  enclosing  the  sporangia.  Each  rolled  pinnule  more  or 
less  spherical,  2  mm.  across,  enclosing  at  least  ten  sporangia;  sporangium  elongated, 
sessile  or  shortly  stalked,  wall  composed  of  small  elongated  cells,  longitudinally 
orientated.  (Annulus  unknown).  Spores  rounded-triangular  with  tri-radiate  cracks 
of  medium  length ;  surface  sculptured  with  a  pattern  of  flat  ribs  3-7  /*  wide  separated 
by  narrow  grooves  0-5-2  ju,  wide. 

LECTOTYPE.     ¥.2329.     Selected  Hughes  &  Moody-Stuart  1966. 

DESCRIPTION.  P.  valdensis  Seward  is  represented  by  several  specimens  all  of 
which  are  fertile.  Fragments  of  pinnae  and  detached  fertile  "  capsules  "  are 
abundant  but  there  are  only  two  assembled  groups  of  pinnae,  specimens  ¥.2368 
and  ¥.2329.  Both  of  these  syntypes  were  figured  by  Seward  (1913,  text-figs.  36,  B'; 
pi.  12,  figs.  I2«,  b.)  and  ¥.2329  (Text-fig.  32)  has  recently  been  selected  as  Lectotype 
by  Hughes  &  Moody-Stuart  (1966).  After  I  picked  away  the  matrix  ¥.2368  is 
now  a  better  specimen,  more  extensive  and  better  preserved  than  ¥.2329  (PI.  6, 
figs,  i,  2,  Text-fig.  39). 

The  exact  form  of  the  fertile  "  capsules  "  is  far  from  fully  understood.  A  number 
of  isolated  "  capsules  "  were  dissolved  out  of  the  rock  in  HF  but  revealed  nothing 
more  about  their  structure  than  is  shown  in  the  Lectotype  which  clearly  shows  the 
folded  nature  of  the  pinnules  (Text-fig.  32).  A  number  of  specimens  show  hairs  in 
the  matrix  where  "  capsules  "  have  been  removed  (Text-fig.  32)  but  such  hairs  could 


ENGLISH   WEALDEN    FLORA   I 


233 


FIGS.  32-36.  Pelletieria  valdensis  Seward.  32.  Lectotype  showing  folded  "  capsules  ", 
hairs  and  cell  directions  (indicated  by  dotted  lines).  V.232Q.  x  6.  33.  Small 
specimen  with  three  ribs  on  the  "  capsules  ".  V. 51846.  x  6.  34.  Largest  assem- 
bled shoot  showing  dissected  form  of  frond.  ¥.2368.  x  3.  35.  Spore  showing 
thickness  of  wall  (indicated  by  dotted  line)  V. 51841.  x  750.  36.  Eroded  "cap- 
sule "  with  one  sporangium  showing  cells  of  the  wall.  Others  showing  sporer  V. 51847. 
X  35- 


234 


A   REVISION   OF   THE 


not  be  found  on  any  of  the  specimens  dissolved  out.  Certain  specimens  show  the 
axis  continued  as  a  rib  over  the  back  of  the  rolled  fertile  pinnules  and  this  rib  is 
branched  in  ¥.51846  (Text-fig.  33).  The  pinnules  of  ¥.51846  appear  less  tightly 
rolled  than  others  and  might  well  prove  worth  sacrificing  to  a  balsam  transfer. 
That  this  specimen  is  fertile  is  not  in  doubt  as  spores  can  be  seen  through  a  broken 
part  at  the  top. 

Because  the  sporangia  are  completely  enclosed  they  could  only  be  studied  in 
certain  eroded  "  capsules  "  which  show  sporangia  in  various  attitudes  (Text-fig.  40) . 


38 


FIGS.  37—40.  Pelletieria  valdensis  Seward.  37,  38.  Proximal  and  distal  surfaces  of  the 
same  spore.  Slide  from  V. 2329,  x  750.  39.  Equatorial  view  of  spore  showing  widely 
spaced  ribs.  Slide  from  V. 51841,  x  750.  40.  Eroded  "  capsule  "  with  folded 
pinnules  in  section  and  spores  in  sporangia.  ¥.51840.  x  35. 


ENGLISH  WEALDEN   FLORA   I  235 

Attempts  to  isolate  a  sporangium  failed  because  the  plant  material  crumbles  very 
easily  into  small  fragments:  thus  the  spore  number  is  not  known.  However,  on 
such  a  sporangium  as  in  the  centre  of  Text-figure  40  around  fifty  spores  can  be  counted 
on  the  surface  but  because  only  one  layer  can  be  seen  it  seems  probable  that  the 
contents  are  at  least  twice  this  figure.  One  sporangium  seen  in  surface  view  (Text- 
fig.  36)  shows  the  cells  of  the  wall,  small,  elongated  and  longitudinally  orientated; 
although  there  is  no  indication  of  the  annulus. 

The  spores  of  P.  valdensis  Seward  have  recently  been  redescribed  by  Hughes  & 
Moody-Stuart  (1966)  using  Seward's  figured  slide  preparation  from  ¥.2329  (1913) 
and  new  preparations  from  ¥.2329.  They  found  the  size  range  to  be  52-98  p  with 
80  %  of  specimens  between  63  jn  and  84  ju,  maximum  diameter.  My  preparations 
from  V .  2329  and  V .  21841  agree  well  with  these  figures  but  V .  2368  yielded  somewhat 
larger  spores,  56-115  /*  (100  measured)  with  75  %  between  75  ju  and  105  fi.  Hughes 
&  Moody-Stuart  (1966)  have  diagnosed,  described  and  figured  the  spores  in  detail 
and  there  is  nothing  more  to  add. 

COMPARISON.  The  ribbed  spores  of  P.  valdensis  Seward  are  typically  Schizaea- 
ceous  and  very  similar  to  those  of  various  living  and  fossil  species.  However,  the 
form  of  the  fertile  fronds  cannot  be  matched ;  the  living  members  of  the  Schizaeaceae 
merely  having  curled  margins  protecting  the  sporangia.  The  only  comparable 
structure  I  know  is  in  the  species  Onoclea  sensibilis  Linn,  where  the  fertile  pinnules 
are  narrow  and  so  tightly  inrolled  as  to  completely  enclose  the  sporangia. 

The  spores  of  P.  valdensis  are  similar  in  form  to  those  of  Ruffordia  goepperti, 
particularly  the  form  with  the  broad  ribs  and  narrow  grooves,  but  they  are  quite 
easily  distinguishable  on  size  difference  alone.  The  upper  limit  found  for  the  dia- 
meter of  R.  goepperti  was  60  /*  and  the  lower  limit  for  P.  valdensis  63  /*.  A  most 
distinct  difference  is  the  thickness  of  the  exine;  in  R.  goepperti  about  4  /*  (including 
the  thickness  of  the  ribs)  whilst  in  P.  valdensis  (Text-fig.  35)  it  varies  from  about 
5  /£  to  15  fji.  This  variation  depends  upon  the  spaces  between  ribs  being  open  or 
closed  and  is  dealt  with  in  some  detail  by  Hughes  &  Moody-Stuart  (1966  :  280-282) . 
A  further  difference  is  that  the  ribbed  pattern  in  P.  valdensis  extends  on  to  the 
contact  facets  whilst  in  R.  goepperti  the  facets  are  smooth. 


Family  POLYPODIACEAE 

Genus  ASPIDISTES  Harris,  1964  :  181 

The  type  species  of  this  genus,  Aspidistes  thomasi  Harris,  is  the  only  other  Mesozoic 
fern  I  know  with  the  indusium  superum  and  other  aspidioid  characters.  Although 
various  diagnostic  characters  of  Aspidistes  are  not  known  for  Aspidistes  sewardi 
sp.  nov.  the  form  of  its  indusium  and  spores  give  it  a  good  measure  of  agreement. 
The  slight  difference  in  spore  number  (A.  thomasi  has  forty-eight  )is  not  important. 
However,  further  details  of  the  sorus  and  sporangium  may  well  show  it  to  be  generi- 
cally  distinct.  Various  other  genera  exist  whose  names  suggest  close  affinity  with 
Aspidium  or  Polystichum  but  are  based  only  on  sterile  specimens. 


236  A   REVISION   OF  THE 

Aspidistes  sewardi  sp.  nov. 
PL  6,  figs.  3,  4;  Text-figs.  41-43 
1894     Weichseliamantelli  (Brongniart)  Seward  :  121. 

DIAGNOSIS.  (Based  on  single  fertile  pinna  V.2i73«).  Pinnules  arising  from 
upper  surface  of  rachis  at  an  angle  of  50-60°;  pinnules  2  mm.  long,  1-5  mm.  broad, 
apex  rounded;  joined  to  each  other  at  base  by  a  narrow  web;  margin  of  pinnules  and 
web  rolled.  Groups  of  three  sori  borne  on  under  surface  of  each  pinnule ;  two  larger 
adjacent  to  the  rachis.  Each  sorus  with  a  thick  peltate  indusium  covering  about 
twelve  sporangia  arranged  in  a  circle.  Sporangium  containing  thirty-two  spores 
(sporangial  wall  unknown).  Spores  trilete,  smooth  walled,  thickened  at  corners; 
narrow  bands  of  thickening  flanking  triradiate  crack.  Arms  of  triradiate  crack 
nearly  reaching  corners,  sometimes  bifid  at  ends ;  diameter  60-75  /£. 

HOLOTYPE.     V .  21730:,  the  only  specimen. 

DESCRIPTION.  The  single  specimen  (¥.2173^)  is  6-5  cm.  long  and  was  formerly 
referred  to  Weichselia  mantelli  (=  W.  reticulata  Stokes  &  Webb)  by  Seward  (1894) 
its  form  being  very  similar  to  the  smaller  pinnules  of  that  species.  However  its 
abaxial  surface  is  uppermost  and  on  moistening  with  paraffin  the  sori  are  clearly 
visible  to  the  naked  eye  (PI.  6,  fig.  4).  It  is  certainly  not  Weichselia  reticulata  for 
fertile  material  of  that  species  is  included  in  the  Bommer  Collection  and  it  is  quite 
different  (correspondence  with  Dr.  K.  L.  Alvin). 

Unfortunately  the  specimen  is  heavily  impregnated  with  glue  and  the  substance 
of  the  fossil  is  badly  crumbling,  making  it  very  difficult  to  examine  the  details  of 
the  sorus.  The  water  soluble  gum  swells  rather  violently  when  wet  and  lifts  the 
fossil  fragments  off  the  surface  of  the  rock.  Careful  picking  with  a  fine  needle  re- 
vealed a  certain  amount  of  information.  The  circular  indusium  appears  to  be  quite 
thick  and  is  probably  attached  by  a  central  stalk;  details  of  its  cellular  structure 
could  not  be  seen.  The  sporangia  are  distinct  pear-shaped  masses,  about  twelve 
in  number,  in  a  ring  under  the  indusium  and  sometimes  a  few  under  the  rolled  margin 
of  the  pinnule.  They  were  easily  removed  intact  to  count  the  spores.  Of  the  twelve 
sporangia  macerated  and  counted  nine  contained  thirty-two  spores  and  three  others 
contained  twenty-nine,  thirty,  and  thirty-one.  The  sporangial  wall  was  not  seen 
either  in  the  untreated  sporangium  or  during  maceration. 

Among  the  Wealden  ferns  there  are  few  species  of  sterile  fronds  to  which  this 
specimen  could  belong  but  it  is  similar  to  one  described  as  Dichopteris  delicatula 
Seward  (1913  pi.  n,  figs.  6a,  b).  Possibly  further  specimens,  sterile  or  fertile,  are 
among  the  Weichselia  reticulata  material  but  nothing  is  known  of  its  venation  and 
it  is  not  always  clear  in  Weichselia.  Furthermore  any  specimen  with  the  adaxial 
side  upwards  would  easily  be  missed. 

COMPARISON.  Amongst  living  genera  A.  sewardi  is  most  similar  to  Polystichum 
agreeing  in  the  form  of  the  indusium  and  the  spore  output.  However  all  living 
species  of  the  Aspidieae  have  wedge-shaped  spores  whereas  both  A.  sewardi  and 
A .  thomasi  Harris  have  trilete  spores. 


ENGLISH  WEALDEN   FLORA   I 


237 


42 


44 


43 


FIGS.  41-43.  Aspidistes  sewardi  sp.  nov.  41.  Three  pinnules  showing  shape,  rolled  mar- 
gins and  sori.  V.2i73a,  x  15.  42,  43.  Two  spores.  Triradiate  crack  in  Fig.  43  with 
bifid  ends.  Fig.  42,  slide  ii  from  V.  2173(3,  x  750.  Fig.  43,  slide  i  from  V.  21730,  x  500. 

FIG.  44.  Nilssonia  schaumburgensis  Dunker.  Base  of  leaf .  Specimen  L. 5532  Manchester 

Museum,  x  5. 


A .  sewardi  is  named  after  Sir  Albert  Charles  Seward  who  first  described  the  English 
Wealden  flora. 

UNCLASSIFIED  FERNS 

The  following  are  ferns,  described  by  Seward  (1894),  to  which  I  can  add  little  or 
nothing. 

Cladophlebis  longipennis  Seward 

1894     Cladophlebis  longipennis  Seward  :  89,  pi.  9,  figs,  i,  la. 

Cladophlebis  albertsii  (Dunker) 

1894     Cladophlebis  Albertsii  (Dunker)  Seward  :  91,  text-fig.  9;  pi.  8. 


238  A   REVISION   OF  THE 

Cladophlebis  browniana  (Dunker) 

1894     Cladophlebis  Browniana  (Dunker)  Seward  :  99,  pi.  7,  fig.  4. 
1913     Cladophlebis  Browniana  (Dunker)  Seward  :  95,  pi.  8,  figs,  i,  2. 

Cladophlebis  dunkeri  (Schimper) 

1894     Cladophlebis  Dunkeri  (Schimper)  Seward  :  100,  pi.  7,  fig.  3. 

1950     Cladophlebis  (Klukia)  dunkeri  (Schimper);  Radforth  &  Woods  :  780,  pi.  i. 

The  various  English  Wealden  specimens,  referred  by  Seward  to  the  four  species 
above,  appear  to  be  indistinguishable  from  each  other  and  Cladophlebis  longipennis 
Seward  is  particularly  ill-defined.  Indeed  Seward  himself  later  wrote  (1913)  of  the 
"  difficulty — or  indeed,  impossibility — of  separating  Cladophlebis  browniana  and 
Cladophlebis  dunkeri  ".  Modern  treatment  of  Canadian  material  referred  to  C, 
dunkeri  (Schimper)  by  Radforth  &  Wood  (1950)  shows  the  fern  to  have  Schizaeaceous 
sporangia  and  spores,  though  the  latter  are  scarcely  distinguishable  in  the  photo- 
graph. 

Sphenopteris  ruffordi  (Seward)  comb.  nov. 
1894     Acrostichopteris  Ruffordi  Seward  :  61,  pi.  6,  fig.  3. 

Sphenopteris  fittoni  Seward 
1894     Sphenopteris  Fittoni  Seward  :  107,  pi.  6,  fig.  2;  pi.  7,  fig.  i. 

Sphenopteris  fontainei  Seward 

1894     Sphenopteris  Fontainei  Seward  :  106,  pi.  7,  fig.  2. 

Leckenbya  valdensis  (Seward) 

1894  Nathorstia  valdensis  Seward  :  145,  pi.  7,  fig.  5;  pi.  9,  figs.  2,  2«. 

1894  Leckenbya  valdensis  (Seward)  Seward  :  384. 

1895  Leckenbya  valdensis  (Seward);  Seward  :  225. 
1911  Gleichenites  cycadina  (Schenk);  Seward  :  664. 

Teilhardia  valdensis  Seward 

1913     Teilhardia  valdensis  Seward  :  96,  pi.  n,  figs,  ja-gb. 

Genus  ONYCHIOPSIS  Yokoyama  1889  :  26 
Onychiopsis  psilotoides  (Stokes  &  Webb) 

1824     Hymenopteris  psilotoides  Stokes  &  Webb  :  423,  pi.  46,  fig.  7. 

1827  Hymenopteris  psilotoides  Stokes  &  Webb;  Mantell  :  55,  pi.  i,  figs.  3a,  3b;  pi.  3,  figs.  6,  7; 
pi.  3*,  fig.  2. 

1828^  Sphenopteris  Mantelli  Brongniart  :  50. 

1828  Sphenopteris  Mantelli  Brongniart;  Brongniart  :  170,  pi.  45,  figs.  3-7. 
1833     Sphenopteris  Mantelli  Brongniart ;  Mantell  :  241. 

1894     Onychiopsis  Mantelli  (Brongniart)  Seward  :  41,  pi.  2,  fig.  i;  pi.  3,  figs.  1-4. 
1894     Onychiopsis  elongata  (Geyler) ;  Seward  :  55. 

1905     Onychiopsis  psilotoides  (Stokes  &  Webb)  Ward  :  155,  506,  518,  pi.  39,  figs.  3-6;  pi.  in, 
fig.  4;  pi.  113,  fig.  i. 


ENGLISH   WEALDEN    FLORA    I  239 

1910     Onychiopsis  Mantelli  (Brongniart) ;  Seward  :  378,  fig.  280,  A,  B. 

1913     Onychiopsis mantelli  (Brongniart);  Seward  :  96. 

1961     Onychiopsis  psilotoides  (Stokes  &  Webb);  Tattersall  :  349,  pi.  12,  figs.  1-3. 

DISCUSSION.  This  species  was  of  world  wide  distribution  in  the  Lower  Cretaceous 
and  its  remains  are  known  from  England,  Germany,  France,  Belgium,  Portugal, 
Russia,  Canada,  U.S.A.,  Egypt,  South  Africa,  India  and  Japan.  Although  the 
species  has  been  described  numerous  times,  including  fertile  parts  (Tattersall  1961), 
it  is  still  imperfectly  known.  Spores  were  readily  obtained  but  resemble  those  of  a 
number  of  families  and  do  not  help  in  classification.  Details  of  sporangia  are  still 
unknown ;  though  some  fertile  pinnules  have  a  lumpy  surface  suggestive  of  sporangia. 
However  the  whole  surface  is  covered  by  an  indusium-like  sheet  of  elongated  cells 
and  maceration  leaves  no  trace  of  sporangial  walls. 

I  have  been  unable  to  add  usefully  to  our  knowledge  of  this  species  and  consider 
that  the  figures  of  Seward  (1894)  and  Tattersall  (1961)  cannot  be  improved  upon. 
In  the  light  of  a  recent  review  of  this  species  by  Sukh  Dev  (1965)  with  an  exhaustive 
synonomy  I  have  given  only  those  references  which  include  the  important  name 
changes  or  refer  to  the  English  material. 


Genus  WEICHSELIA  Stiehler 

Weichselia  reticulata  (Stokes  &  Webb) 

PI.  6,  fig.  5 

Selected  references : 

1824  Pecopteris  reticulata  Stokes  &  Webb  :  423,  pi.  46,  fig.  5;  pi.  47,  fig.  3. 

1827  Pecopteris  reticulata  Stokes  &  Webb;  Mantell  :  56,  pi.  3,  fig.  5;  pi.  3*,  fig.  3. 
18280  Lonchopter is  Mantelli  Brongniart;  60. 

1828  Lonchopteris  Mantelli  Brongniart  :  369,  pi.  131,  figs.  9,  10. 
1894  Weichselia  Mantelli  (Brongniart)  Seward  :  114,  text-figs.  12,  13. 
1899  Weichselia  reticulata  (Stokes  &  Webb)  Ward  :  651,  pi.  160,  figs.  2-4. 
1919  Weichselia  reticulata  (Stokes  &  Webb);  Florin  :  305,  figs,  i,  2-5. 

1965^  Weichselia  reticulata  (Stokes  &  Webb);  Reyman6wna  :  16,  text-fig,  i;  pi.  i,  figs.  7-11. 
1965     Weichselia  reticulata  (Stokes  &  Webb) ;  Sukh  Dev  :  73,  figs.  7,  8. 

DISCUSSION.  This  widespread  Lower  Cretaceous  species  has  recently  been  re- 
vised by  Reymanowna  (19650)  and  Sukh  Dev  (1965).  Most  of  the  English  specimens 
are  mere  fragments  of  pinnae,  many  of  them  fusainized,  but  there  is  one  large, 
assembled  frond  in  the  collection.  This  is  figured  by  Seward  (1894,  pi.  10,  fig.  3.) 
and  shows  the  characteristic  habit  of  the  plant  very  well. 

The  lower  epidermis  of  W.  reticulata  (Stokes  &  Webb)  was  first  figured  by  Florin 
(1919)  using  Belgian  material  and  Sukh  Dev  (1965)  has  figured  the  epidermis  from 
English  Wealden  specimens.  Reymanowna  (1965)  gives  figures  of  stomata  together 
with  photographs  of  indusia,  but  the  fertile  material  of  W.  reticulata  (Stokes  & 
Webb)  has  yet  to  be  fully  described.  None  of  the  English  material  is  fertile. 

Sukh  Dev  (1965)  gives  extensive  references  to  older  work  so  that  only  selected 
references  have  been  cited  above. 

GEOL.   17,  5  2O 


240  A   REVISION   OP  THE 

SPERMATOPHYTA 
PTERIDOSPERMAE 
Genus  PACHYPTERIS  Brongniart 

1828  Pachypteris  Brongniart  :  49  (Name  only) . 

1829  Pachypteris  Brongniart  :  166. 

1964     Pachypteris  Brongniart ;  Harris  :  121  (Emended  diagnosis). 

Pachypteris  lanceolata  Brongniart 
PL  i,  fig.  6;  Text-figs.  45-47 

1828  Pachypteris  lanceolata  Brongniart  :  50  (Name  only). 

1829  Pachypteris  lanceolata  Brongniart  :  167,  pi.  45,  fig.  i. 

1895     Dichopteris  sp.  cf.  D.  laevigata  (Phillips)  Seward  :  184,  pi.  12,  fig.  6. 
1964     Pachypteris  lanceolata  Brongniart;  Harris  :  137,  text-figs.  55-58. 

MATERIAL.  V.  3245 

DESCRIPTION.  The  following  description  is  based  on  the  only  English  Wealden 
specimen  known.  It  is  a  single  pinna,  8  cm.  long,  i  cm.  wide  at  the  widest  part, 
with  two  rows  of  pinnules,  leathery  in  texture.  The  rachis,  up  to  700  /*  wide,  is 
very  prominent  on  the  underside  of  the  pinna.  The  pinnules  arise  alternately, 
usually  slightly  overlapping  but  not  completely  divided  and  are  up  to  6  mm.  long 
and  3  mm.  wide,  narrowing  slightly  but  attached  to  the  rachis  by  a  broad  and 
decurrent  base.  The  pinnules  are  typically  oval  and  blunt  ended.  Pinnules  at  the 
apex  are  smaller  and  less  divided;  the  rachis  is  not  so  prominent  here.  The  venation 
is  visible  on  the  lower  side  of  some  pinnules  projecting  slightly  above  the  surface, 
but  concealed  in  most,  the  veins  consisting  of  a  midrib  with  a  few  laterals  given  off 
at  a  fairly  low  angle.  The  midrib  is  broad  and  situated  above  the  middle  of  the 
pinnule  where  it  joins  the  rachis.  The  margins  of  the  pinnules  are  not  thickened. 

The  upper  epidermis  lacks  stomata  and  has  a  thick  cuticle  showing  isodiametric 
cells  with  smooth  walls  [and  without  pits.  These  cells  are  irregularly  arranged 
except  at  the  margin  where  they  are  elongated  and  tend  to  be  in  longitudinal  rows, 
the  cells  here  having  thicker  anticlinal  walls.  The  lower  cuticle  is  thinner,  the  cells 
of  the  lower  epidermis  being  thin  walled,  isodiametric  and  irregularly  arranged, 
except  at  the  margin  and  in  the  midrib  region.  The  position  of  the  midrib  is  indi- 
cated by  a  region  of  elongated,  thicker  walled  cells  in  longitudinal  rows.  The  margin 
is  like  that  of  the  upper  epidermis.  Stomata  are  numerous  on  the  lower  side  but 
absent  from  the  midrib  and  marginal  regions.  They  are  slightly  sunken,  irregu- 
larly orientated  and  distributed  and  are  almost  always  separated  by  ordinary 
epidermal  cells.  The  guard  cells  are  thinly  cutinized  except  along  the  edges  of  the 
stomatal  aperture  where  there  is  a  thicker  rim;  the  aperture  is  elongated.  Subsidiary 
cells,  usually  six  in  number,  form  a  ring  and  are  generally  smaller  than  the  other 
epidermal  cells.  They  have  thicker  cuticle  on  the  periclinal  walls  which  forms  a 
ring  round  the  stoma  but  are  not  thus  thickened  adjacent  to  the  stomatal  pit. 
Encircling  cells  are  occasional  and  unspecialized.  Trichomes  are  absent. 

DISCUSSION.  The  preservation  is  good  though  the  surface  is  somewhat  cracked 
and  because  of  the  shortage  of  material  available  only  a  little  could  be  macerated. 


ENGLISH  WEALDEN   FLORA   I 


241 


However,  it  was  established  that  the  specimen  is  attached  to  the  rock  by  its  upper 
surface  and  that  whilst  the  lower  cuticle  is  complete  most  of  the  upper  is  missing, 
only  a  narrow  strip  remaining  around  the  edge  of  the  pinnules.  Thus  very  little 
upper  cuticle  has  been  examined  and  it  is  not  known  whether  it  had  any  stomata 
nearer  the  middle  of  the  pinnule.  Only  a  few  of  the  pinnules  show  the  positions  of 
the  midrib  and  the  laterals  on  the  underside,  and  at  best  they  are  indistinct.  An 
attempt  was  made  to  see  the  venation  by  transmitted  light  during  maceration  but 
this  failed.  The  prepared  cuticle,  whilst  showing  the  position  of  the  midrib  by  the 
elongated  cells  and  lack  of  stomata,  gives  no  indication  of  the  position  of  the  lateral 
veins. 


47 


FIGS.  45-47.  Pachypteris  lanceolata  Brongniart.  45.  Part  of  pinna  showing  venation. 
¥.3145.  x  4.  46.  Reconstructed  transverse  section  of  pinna  showing  prominent 
rachis  on  lower  side.  X4-  47.  A  single  stoma.  The  darker  stippling  indicates  the  ring  of 
thicker  cuticle.  Slide  V.  31456.  x  700. 


The  surface  of  the  specimen  viewed  in  reflected  light  shows  very  clearly  the  cells 
of  the  lower  epidermis  with  a  divergent  trend  in  relation  to  veins.  In  the  frag- 
mentary cuticle  preparations  this  trend  is  not  apparent  and  the  cells  appear  hap- 
hazard in  arrangement.  One  very  prominent  cell  was  seen,  the  nature  of  which  is 
obscure  though  it  is  certainly  not  stomatal.  It  is  possibly  a  trichome  base  though 
no  similar  cells  could  be  identified  with  certainty. 

COMPARISON.  A  careful  comparison  was  made  with  specimens  of  Pachypteris 
lanceolata  Brongn.  from  the  Yorkshire  Jurassic  (Harris  1964).  The  Yorkshire 
material  is  known  in  much  more  detail,  numerous  specimens  being  available  some 
of  which  are  complete  leaves.  A  notable  feature  of  P.  lanceolata  is  its  wide  range 
in  form  with  intergradation  between  the  extreme  types.  These  include  forms  with 
small  oval  pinnules,  others  with  small  narrow  pinnules  and  much  larger  forms  with 


242  A   REVISION   OF  THE 

lanceolate  pinnules.  Forms  just  like  the  present  specimen  occur.  The  cuticles  of  the 
various  forms  although  basically  the  same  show  a  certain  variety  in  details  such  as 
number  and  size  of  stomata.  Some  have  a  large  number  of  stomata  close  together 
while  in  others  they  are  more  widely  spaced.  The  cuticle  of  the  present  fossil  shows 
more  widely  spaced  stomata  than  most  of  the  specimens  attributed  by  Harris  to 
P.  lanceolata  but  some  of  the  latter  approach  the  Wealden  form  very  closely.  These 
include  specimens  from  Boulby  Alum  Quarry,  Shale  2B ;  Whorlton,  Scugdale,  Rank 
Crag  Soft  Jet  and  Eston  Moor,  Beacon  Quarry.  It  should  be  pointed  out  that  these 
are  not  ordinary  specimens  but  merely  cuticle  fragments  obtained  by  maceration 
and  identified  on  their  microscopic  characters. 

As  in  the  Wealden  fossil,  the  stomata  of  the  Jurassic  P.  lanceolata  avoid  the  midrib 
and  margins  and  are  indistinguishable  in  structure.  There  is  also  very  close  simil- 
larity  in  size  and  shape  of  the  ordinary  epidermal  cells,  indeed  some  of  the  cuticle 
fragments  of  the  two  fossils  could  scarcely  be  told  apart. 

What  little  is  known  of  the  venation  of  the  Wealden  form  simply  affords  more 
evidence  of  the  similarity  between  these  two  fossils  and  consequently  I  have  placed 
the  Wealden  specimen  in  the  same  species. 


GYMNOSPERMAE 

GYGADALES 

Genus  NILS  S  ONI  A  Brongniart 
Nilssonia  schaumburgensis  (Dunker) 

Text-figs.  44,  48-52 

Selected  references : 

1846    Pterophyllum  schaumburgense  Dunker  :  15,  pi.  i,  fig.  7;  pi.  2,  fig.  i;  pi.  6,  figs.  5-10. 

1871     Anomozamites  schaumburgense  (Dunker)  Schenk  :  231,  pi.  33,  figs.  1-9. 

1890     Nilssonia  cf.  schaumburgensis  (Dunker)  Nathorst  :  5,  9,  13,  pi.  i,  figs.  6-ga. 

1895     Nilssonia  schaumburgensis  (Dunker) ;  Seward  :  53,  text-fig.  3. 

1905     Nilssonia  schaumburgensis  (Dunker);  Fontaine  in  Ward  :  303,  pi.  72,  figs.  17-21. 

EMENDED  DIAGNOSIS.  Leaf  linear;  width  usually  3-5  mm.,  rarely  up  to  12  mm.; 
length  unknown  but  at  least  9  cm.  Apex  rounded  and  notched;  base  tapering. 
Midrib  1-2  mm.  wide,  depressed  on  upper  surface,  depression  not  narrowing  towards 
apex.  Lamina  entire  or  more  or  less  divided.  Surface  of  lamina  flat  or  slightly 
convex  upwards.  Veins  fairly  prominent,  projecting  above  and  below;  arising  from 
midrib  at  an  angle  between  70°  and  80°;  concentration  of  veins  typically  three  to 
four  per  mm. 

Cuticle  of  moderate  thickness  but  anticlinal  walls  often  obscure.  Marginal  cells 
of  leaf  and  pinnae  elongated  parallel  to  edges  and  having  thicker  and  more  distinct 
anticlinal  walls.  Cells  in  main  part  of  lamina  tending  to  be  elongated  almost  at 
right  angles  to  edge,  parallel  with  veins.  Upper  cuticle  without  stomata  or  tri- 
chomes,  positions  of  veins  scarcely  distinguishable.  Lower  cuticle  not  showing 
positions  of  veins  clearly  but  cells  with  slight  tendency  to  be  more  elongated  over 
veins.  Stomata  scattered  between  veins;  small,  inconspicuous  and  scarcely  sunken. 


ENGLISH   WEALDEN   FLORA   I 

50 


243 


52 


FIGS.  48-52.  Nilssonia  schaumburgensis  (Dunker).  48.  Leaf,  slightly  divided,  showing 
form  of  apex.  Position  of  rachis  indicated  by  dotted  line  or  solid  line  where  lamina  is 
broken  and  rachis  is  exposed.  V.3762.  x  2.  49.  More  divided  leaf  showing  the 
veins.  Position  of  rachis  shown  by  dotted  lines.  V.2I7I.  x  5.  50.  Cuticle  showing 
elongated  cells  of  leaf  edge  in  centre,  lower  surface  to  the  left,  upper  surface  to  the  right. 
Slide  from  ¥.1436.  x  300.  51.  Lower  cuticle  showing  a  resin  body,  a  stoma  and 
two  single-celled  trichome  bases.  Slide  from  V.2I7I**.  x  300.  52.  Single  stoma 
showing  exposed  guard  cells.  Slide  from  ¥.3762.  X  700. 


244  A   REVISION   OF   THE 

Guard  cells  exposed;  30-50/4  long  and  about  15  {i  wide,  surrounded  by  several 
unspecialized  and  ill-defined  subsidiary  cells.  Unicellular  trichome  bases  scattered 
over  whole  lower  surface;  thickly  cutinized  except  for  central  circle.  Resin  bodies 
occasionally  seen,  about  50  fi  across.  Reproductive  organs  unknown. 

DESCRIPTION.  This  small  Nilssonia  is  represented  by  more  than  a  dozen  speci- 
mens from  the  English  Wealden.  They  show  a  good  deal  of  variety  in  the  form  of 
the  lamina,  some  being  entire,  others  slightly  and  irregularly  divided,  others  re- 
gularly and  completely  divided  almost  to  the  midrib.  Only  two  specimens  are 
more  than  6  mm.  wide,  V  .2ijia  which  is  12  mm.  wide  (Seward  1895  :  53,  text-fig. 
30)  and  V.373I  which  Seward  labelled  as  Taeniopteris  beyrichii  var  superba.  Both 
agree  with  the  others  in  their  cuticle.  All  but  one  specimen  have  the  adaxial  surface 
exposed  and  in  these  the  midrib  is  seen  only  when  the  substance  of  the  lamina  is 
broken  away  (Text-fig.  48)  but  in  all  specimens  its  position  is  indicated  by  a  depres- 
sion of  the  lamina.  This  depression  retains  its  width  to  the  apex  of  the  leaf.  V .  3731 
is  the  only  specimen  with  the  lower  surface  uppermost  and  is  the  largest  specimen, 
9  cm.  long  (incomplete)  and  u  mm.  wide,  tapering  to  4  mm.  near  the  base.  The 
rachis  is  2-5  mm.  wide  whilst  in  most  specimens  it  is  little  more  than  I  mm. 

Resin  bodies  are  rare;  they  are  not  seen  on  the  surface  of  the  fossils  and  only  three 
were  seen  in  macerated  cuticle  fragments. 

The  stomata  are  particularly  difficult  to  distinguish  and  were  identified  with 
certainty  on  only  a  very  few  pieces  of  lower  cuticle.  The  stoma  in  Text-fig.  52 
is  the  best  example  seen  and  shows  a  slight  surface  thickening  in  the  middle  region 
of  the  guard  cells. 

DISCUSSION.  Nilssonia  schaumburgensis  has  been  identified  by  numerous  authors 
from  several  countries.  In  all  cases  there  is  very  little  detail  on  which  to  compare  the 
various  accounts.  From  the  few  details  available,  i.e.  size,  shape  and  vein  concentra- 
tion the  English  material  agrees  well  with  the  type-material  described  by  Dunker 
(1846)  and  the  specimens  figured  by  Schenk  (1871),  Nathorst  (1890)  and  Fontaine 
in  Ward  (1905) . 

COMPARISON.  N.  revoluta  Harris  (1964)  from  the  Jurassic  (Upper  Deltaic)  of 
Yorkshire  is  the  only  other  species  I  know  which  is  as  small  as  N.  schaumburgensis. 
The  leaves  are  similar  in  size  and  are  entire;  also  the  vein  concentration  is  similar, 
up  to  thirty  per  cm.  Another  similarity  is  that  the  veins  are  scarcely  shown  in  the 
upper  cuticle,  however  they  are  strongly  indicated  in  the  lower  cuticle.  Rather  differ- 
erent  characters  shown  by  N.  revoluta  include  conspicuous  cell  outlines  with  nodular 
thickenings  and  bulging  subsidiary  cells. 

Genus  BECKLESIA  Seward  1895  :  179 
Becklesia  anomala  Seward 

Text-figs.  53-56 
1895     Becklesia  anomala  Seward  :  179,  pi.  14,  figs.  2,  3. 

EMENDED  DIAGNOSIS.  Leaf  simply  pinnate,  probably  much  longer  than  broad 
(whole  leaf  unknown),  consisting  of  a  slender  rachis  bearing  two  lateral  rows  of 


ENGLISH   WEALDEN   FLORA   I 


245 


long  straight  pinnae.  Rachis  6  mm.  wide,  pinnae  arising  at  intervals  of  approxi- 
mately i  cm.,  at  an  angle  of  about  40°.  Pinnae  at  least  up  to  10  cm.  long,  3  mm. 
wide,  narrowing  to  about  2  mm.  near  rachis  (apex  unknown),  rounded  in  transverse 
section;  margins  of  pinnae  straight.  Lower  side  of  pinna  having  broad,  flat  midrib 


53 


54 


55 


FIGS.  53-56.  Becklesia  anomala  Seward.  53.  A  single  stoma.  One  hypodermal  cell 
is  seen  on  the  right  indicated  by  a  single  line;  Slide  from  V. 2361  a.  x  700.  54.  A 
piece  of  cuticle  from  a  non-stomatal  region  of  the  leaf,  showing  a  four-celled  trichome 
base.  Slide  from  V.  23610,.  x  200.  55.  An  unsquashed  portion  of  the  pinna  showing 
the  two  narrow  stomatal  grooves  on  the  lower  side.  V.236ia.  x  10.  56.  A  recon- 
structed section  of  a  pinna,  x  10. 


246  A    REVISION    OF   THE 

flanked  by  two  very  narrow,  shallow  stomatal  grooves  with  lateral  non-stomatal 
regions  about  as  broad  as  the  midrib;  upper  surface  flat  but  some  pinnae  showing 
numerous  raised  trichome  bases. 

Cuticle  about  2  IJL  thick  in  non-stomatal  regions,  very  delicate  in  stomatal  grooves. 
Ordinary  epidermal  cells  of  upper  and  lower  surface  large,  thin  walled  and  arranged 
in  longitudinal  rows ;  very  varied  in  shape,  either  square,  rectangular  or  quite  irreg- 
ular; typically  50  fi  broad.  Cells  over  midrib  and  at  edges  of  pinna  tending  towards 
diamond  shape.  Anticlinal  walls  cutinized,  usually  straight  but  sometimes  slightly 
sinuous.  Presence  of  a  hypodermis  occasionally  indicated,  particularly  near  trichome 
bases.  Trichome  bases  mostly  occurring  on  upper  surface  but  sometimes  also  in 
non-stomatal  regions  of  lower  surface ;  roughly  circular  in  surface  view,  about  60-70  fi 
diameter,  composed  of  up  to  four  small,  thick-walled  cells  tending  to  overlap  other 
epidermal  cells.  Trichome  bases  not  present  on  all  pinnae.  Stomata  longitudinally 
orientated  and  scattered  in  grooves  about  45  fi  wide.  Guard  cells  thinly  cutinized, 
about  45  /*  long  with  six  to  seven  subsidiary  cells  surrounding  and  slightly  overlapp- 
ing them;  guard  cells  and  subsidiary  cells  slightly  sunken  in  shallow  stomatal  pit, 
surrounded  by  thicker  walled  cells,  inner  walls  of  which  form  thickly  cutinized  pit 
rim.  Ordinary  epidermal  cells  between  stomata  much  smaller  than  those  in  non- 
stomatal  regions. 

HOLOTYPE.     V.  23610. 

DESCRIPTION.  This  species  is  described  from  two  specimens  both  of  which  are 
incomplete  leaves  with  no  complete  pinnae.  Whilst  most  of  the  pinnae  have  been 
squashed  during  preservation,  two  fragments  are  so  preserved  that  they  retain  their 
original  shape  and  thickness.  One  is  about  i  mm.  thick  and  shows  particularly 
well  that  the  stomatal  grooves  are  shallow,  that  the  midrib  is  flat,  and  that  the  margin 
is  rounded  in  section. 

Much  of  the  plant  substance  is  missing  from  the  two  specimens  and  what  remains 
is  very  cracked,  so  that  all  cuticle  preparations  are  small  pieces.  Cuticle  from  the 
non-stomatal  regions  was  easily  obtained  but  it  was  only  with  considerable  difficulty 
that  preparations  of  the  stomatal  grooves  were  made.  All  grooves  are  filled  with 
matrix  and  have  extremely  delicate  cuticle  which  disintegrates  even  when  most 
carefully  macerated.  The  stoma  shown  in  Text-fig.  53  is  from  the  best  preparation 
obtained  in  which,  unfortunately,  all  the  stomatal  pits  contain  debris  which  obscures 
some  of  the  details. 

DISCUSSION.  While  the  characters  of  this  species  are  distinctly  Cycadalean  it 
is  not  closely  comparable  to  any  other  genus,  living  or  fossil.  It  is  thus  convenient 
to  retain  the  original  generic  name.  The  stomatal  grooves  are  much  narrower 
than  in  any  other  Cycad.  In  Cycas  stomata  are  present  over  the  whole  of  the  lower 
surface  except  the  midrib  and  margins.  The  trichome  bases  of  B.  anomala  are 
similar  to  those  in  Cycas  revoluta  (Pant  &  Mehra  1962,  text-fig.  330)  but  in  that 
species  they  are  two-celled. 

Amongst  fossil  Cycads  Paracycas  cteis  Harris  (1964)  from  the  Jurassic  (Lower 
Deltaic)  of  Yorkshire  is  similar  in  size  but  differs  in  such  features  as  wider  stomatal 
bands  and  irregularly  arranged  stomata. 


ENGLISH   WEALDEN   FLORA   I 

Becklesia  sulcata  sp.  nov. 


247 


PI.  2,  fig.  6;  Text-figs.  57-58 

DIAGNOSIS.  Pinnae  up  to  2  mm.  wide  (length  unknown),  with  two  narrow  longi- 
tudinal grooves  on  lower  side;  stomata  confined  to  these  grooves.  Grooves  so 
placed  that  the  central  non-stomatal  region  is  twice  as  wide  as  lateral  non-stomatal 
regions.  Margins  smooth  and  entire.  Cuticle  moderately  thick  but  much  thinner 
in  stomatal  grooves.  Non-stomatal  region  between  grooves  composed  of  cells  in 
longitudinal  rows,  transverse  anticlinal  walls  usually  oblique,  sometimes  slightly 


FIGS.  57,  58.  Becklesia  sulcata  sp.  nov.  57.  Cuticle  from  a  stomatal  groove  showing 
stomata  and  papillae.  Holotype,  V. 51524.  x  300.  58.  Cuticle  of  upper  side. 
V. 51524.  x  300. 


sinuous.  Cells  of  other  non-stomatal  regions  not  arranged  in  rows,  very  varied  in 
shape  and  irregular  in  arrangement;  walls  often  slightly  sinuous.  Guard  cells 
extremely  thinly  cutinized,  except  for  slight  thickening  around  stomatal  aperture. 
Guard  cells  about  50  /*  long,  not  at  all  sunken,  irregularly  orientated  but  majority 
more  or  less  longitudinal.  Subsidiary  cells  very  variable,  not  distinguished  from 
other  ordinary  epidermal  cells  in  groove,  all  of  which  are  irregular  in  shape  and  size, 
smaller  than  epidermal  cells  of  non-stomatal  regions.  Subsidiary  cells  often  shared 
by  adjacent  stomata.  Some  of  these  cells  bearing  large,  hollow,  globular  or  elongated 
papillae;  usually  one  such  papilla  near  each  stoma. 

HOLOTYPE.    ¥.51524. 

DESCRIPTION.  This  species  is  known  only  from  cuticle  fragments  which  are 
assumed  to  be  pieces  of  pinnae  although  there  is  no  direct  evidence  that  they  are 


248  A   REVISION   OF   THE 

from  a  pinnate  leaf.  However,  it  is  so  similar  to  Becklesia  anomala  Seward  that  I 
have  placed  it  in  the  same  genus.  The  main  similarities  are  in  such  features  as 
width  of  pinnae,  position  and  width  of  stomatal  grooves  and  form  of  ordinary  epider- 
mal cells. 

COMPARISON.  Becklesia  sulcata  lacks  the  multicellular  trichome  bases  of  B. 
anomala,  has  thinner  more  exposed  guard  cells  and  has  large  papillae. 

If  the  form  of  ordinary  epidermal  cells  is  any  criterion  in  considering  the  affinities 
of  these  little  known  plants,  these  irregular  yet  characteristic  cells  are  almost  exactly 
the  same  as  in  many  living  Cycads  e.g.  Cycas  revoluta. 

GINKGOALES 
Genus  PSEUDOTORELLIA  Florin  1933  :  142 

EMENDED  DIAGNOSIS.  Leaves  entire,  linear  to  elliptic;  apex  rounded,  narrowing 
at  base.  Two  or  more  longitudinal  veins  formed  by  dichotomies  chiefly  in  basal 
part,  ending  blindly.  Stomata  confined  to  lower  side,  always  longitudinally  orien- 
tated; haplocheilic,  monocyclic  or  incompletely  amphicyclic;  guard  cells  sunken, 
strongly  cutinized.  Subsidiary  cells  four  or  more.  Epidermal  cells  with  straight 
or  faintly  sinuous  outlines. 

The  new  fossil  described  here  is  at  least  the  ninth  species  of  this  genus  to  be  des- 
cribed and  it  now  seems  necessary  to  emend  the  generic  diagnosis  as  the  original 
diagnosis  is  too  narrowly  defined  for  present  purposes,  based  as  it  was  on  the  single 
species  P.  nordenskioldi  Florin.  The  simplified,  broader  diagnosis  above  will  better 
serve  to  admit  such  a  fossil  as  the  new  English  Wealden  species  with  its  wide  range 
of  leaf  form  and  scattered  stomata. 

TYPE  SPECIES.     Pseudotorellia  nordenskioldi  Florin  1933. 

Pseudotorellia  heterophylla  sp.  nov. 
PI.  6,  figs.  6,  7;  Text-figs.  59-64. 

DIAGNOSIS.  Leaves  entire,  showing  wide  range  of  size  and  shape  from  narrow  and 
needle-like  to  broad  and  elliptic.  Dimensions  of  holotype  12  mm.  long  and  3  mm. 
wide ;  longest  leaf  known  3  cm.  long  and  i  mm.  wide ;  other  leaves  ranging  between 
these  two.  Apex  unknown.  Petiole  unknown. 

Veins  varying  in  number  from  two  in  narrowest  leaf  to  eight  in  broadest,  formed  by 
dichotomies  near  base  of  leaf.  Veins  may  converge  near  apex  but  end  blindly. 
Interstitial  ducts  sometimes  present  between  veins. 

Cuticle  thick  and  leathery.  Stomata  present  on  one  side  of  leaf  only,  scattered 
over  whole  surface  (presumably  lower),  not  avoiding  veins;  longitudinally  orientated, 
typically  80  [i  long.  Ordinary  epidermal  cells  elongated,  arranged  in  longitudinal 
rows,  sometimes  with  elongated,  thickened  ridge  along  surface.  Surface  of  cuticle 
frequently  sculptured. 

Guard  cells  sunken  beneath  subsidiary  cells,  strongly  thickened  at  common 
junction  wall  with  overlying  subsidiary  cells.  Subsidiary  cells  six,  usually  papillose, 


ENGLISH   WEALDEN   FLORA   I  249 

papillae  projecting  over  stomatal  aperture.  Stomata  usually  not  upsetting  se- 
quence of  cell  rows,  subsidiary  cells  continuous  in  their  lines  of  ordinary  epidermal 
cells.  Sometimes  small  subsidiary  cells  coinciding  with  outer  edge  of  guard  cells, 
forming  ring  or  partial  ring  of  more  specialized  subsidiary  cells.  Encircling  cells 
rare.  Guard  cells  sometimes  having  long  polar  appendages. 

Upper  epidermis  without  stomata,  composed  of  longitudinal  rows  of  elongated 
cells  with  fairly  thick  walls,  sometimes  in  "  packets  ",  often  having  finely  sculp- 
tured cuticle.  Positions  of  veins  not  indicated  on  upper  surface.  Edges  of  leaf 
smooth  with  fairly  thick  rim  of  cuticle.  Cuticle  of  all  forms  of  leaf  closely  agreeing; 
only  minor  differences  in  size  of  stomata,  degree  of  thickening  and  form  of  polar 
appendages  of  guard  cells. 

HOLOTYPE.     ¥.51525. 

DESCRIPTION.  A  considerable  number  of  whole  and  broken  leaves  were  obtained 
by  bulk  maceration  of  the  same  coaly  material  which  yielded  Becklesia  sulcata. 
Besides  these  a  few  of  the  larger  needles  were  picked  off  specimen  V.2222  in  the 
British  Museum  (Natural  History)  collections.  Several  fragments  were  also  iden- 
tified in  washings  from  the  core  of  a  borehole  drilled  near  Winchester.  Most  of 
the  linear  leaves  were  found  only  in  short  lengths  whilst  the  large  elliptical  leaves 
were  almost  whole.  However,  on  none  of  these  is  an  apex  or  petiole  present. 
The  petiole  presumably  had  one  vein  which  dichotomized  at  the  base  of  the  leaf  to 
give  the  two  main  veins  which  may  further  give  off  branches  on  their  inner  sides  in 
the  lower  part  of  the  leaf.  The  veins  are  then  parallel  along  the  length  of  the  leaf 
but  may  converge  slightly  near  the  apex  (Text-figs.  61,  62.)  although  they  all  appear 
to  end  blindly.  The  interstitial  ducts  which  are  often  present  between  veins  are 
of  unknown  nature  but  are  possibly  resin  ducts  or  even  subsidiary  veins. 

A  number  of  leaves  have  what  appear  to  be  abortive  stomata.  One  of  these  is 
seen  in  Text-fig.  63.  No  guard  cells  are  developed  but  a  ring  of  papillae  surround 
the  "  guard  cell  mother-cell  ".  A  few  other  "  freak  "  stomata  were  seen  where 
the  guard  cells  were  misplaced  under  the  stomatal  pit.  Only  one  transversely 
orientated  stoma  was  seen. 

The  differences  in  polar  appendages  of  guard  cells  may  be  compared  in  Text-figs. 
63,  64.  Text-fig.  64  shows  a  stoma  with  the  longest  and  most  complex  appendages 
projecting  well  beyond  the  ends  of  the  guard  cells.  In  Text-fig.  63  the  appendages 
are  quite  short  and  pointed  or  scarcely  developed  at  all.  Also  shown  in  Text-fig.  63 
are  some  stomata  with  the  subsidiary  cells  following  the  line  of  the  guard  cells  as 
mentioned  in  the  diagnosis. 

COMPARISON.  P.  heterophylla  shows  a  much  wider  range  of  leaf  form  than  other 
species.  P.  nordenskioldi  (Nathorst)  described  by  Florin  (1933),  P.  minuta  (Nathorst) 
by  Lundblad  (1957),  P.  ensiformis  (Heer)  and  P.  crassifolia  (Prynada)  by  Doludenko 
(Vachrameev  &  Doludenko  1961)  all  have  the  greatest  width  of  their  leaf  situated 
towards  the  apex.  P.  grojecensis  Reymanowna  (1963),  P.  ephela  (Harris  1935)  and 
the  elliptic  leaves  of  P.  heterophylla  have  the  greatest  width  of  the  leaf  situated 
in  the  middle  region.  P.  angustifolia  Doludenko  and  P.  longifolia  Doludenko 


250 


A   REVISION   OF   THE 


(Vachrameev  &  Doludenko  1961)  both  have  long  linear  leaves  of  similar  dimen- 
sions to  some  forms  of  P.  heterophylla. 

A  common  feature  of  the  Pseudotorellia  cuticle  is  the  presence  of  median  ridges  of 
cuticle  on  the  epidermal  cells.  P.  heterophylla,  P.  nordenskioldi,  P.  minuta,  P. 
ephela  and  P.  ensiformis  all  have  these  ridges.  In  all,  the  epidermal  cells  are  similar 
in  form,  longitudinally  elongated.  The  packets  of  cells  seen  in  the  upper  epidermis 


59 


61     r? 


60    ; 


FIGS.  59-64.  Pseudotorellia  heterophylla  sp.  nov.  59-62.  Various  forms  of  leaf  showing 
veins.  Figs.  60-62  showing  interstitial  ducts.  Fig.  59,  ¥.51526,  Fig.  60,  V. 51527, 
Fig.  61,  Holotype,  V.  51525,  Fig.  62,  V.  51528.  All  x  10.  63.  Cuticle  showing stomata, 
abortive  stoma  and  ridges  on  the  epidermal  cells.  V.  51529.  x  150.  64.  Stoma  with 
long  polar  appendages  Slide  from  V.  2222.  x  300. 


of  P.  heterophylla  (PI.  6,  fig.  6)  are  surpassed  in  development  by  those  seen  in  P. 
grojecensis.  In  that  species  the  packets  consist  of  up  to  five  cells  and  form  a  con- 
spicuous pattern  at  right  angles  to  each  other  (Reymanowna  1963,  text-fig.  8A.) 

The  stomata  of  P.  heterophylla  appear  to  be  more  heavily  cutinized  than  in  other 
species.  For  instance  in  P.  ephela  and  P.  grojecensis  the  guard  cells  are  heavily 
cutinized  at  the  outer  edges  and  along  the  stomatal  aperture  but  in  P.  heterophylla 
the  whole  upper  surface  of  the  guard  cells  is  heavily  cutinized,  but  sometimes  slightly 
thinner  around  the  aperture. 

The  polar  appendages  of  P.  heterophylla  are  unique  among  the  species  under  con- 
sideration for  the  rest  have  thin  polar  areas. 

Papillate  subsidiary  cells  like  those  in  P.  heterophylla  are  also  present  in  P.  ephela, 
P.  minuta  and  P.  ensiformis  while  P.  grojecensis  has  a  cuticular  diaphragm  on  the 
surface  of  the  guard  cells  around  the  aperture. 

III.  REFERENCES 

ALLEN,  P.  1941.  A  Wealden  Fossil  Soil  Bed  with  Equisetum  lyelli  (Mantell).  Proc.  Geol. 
Ass.,  Lond.,  52  :  362-374. 

—  1954.     Geology  and  Geography  of  the  London-North  Sea  Uplands  in  Wealden  Times. 
Geol.  Mag.,  Lond.,  91  :  498-508. 

• I955-     Age  of  the  Wealden  in  North- Western  Europe.     Geol.  Mag.,  Lond.,  92  :  265-281. 

1959-     The   Wealden    Environment:    Anglo-Paris    Basin.     Phil.    Trans.    R.   Soc.   Lond., 

242B  :  283-346. 

—  1960.     Geology  of  the  Central  Weald:  The  Hastings  Beds.     Geologist's  Ass.  Guide,  London, 
24  :  1-28,  figs.  1-5. 

1965.     L'age  du  Purbecko-Wealdien  D'Angleterre.     Compte  rendu  du  colloque  sur  le  Cretace 

inferieur,  Lyon  1963.     Mem.  Bur.  Recherches  geol.  min.,  Paris,  34  :  321-326. 

—  1967.     Origin  of  the  Hastings  Facies  in  North- Western  Europe.     Proc.  Geol.  Ass.  Lond., 
78  :  27-105. 

ANDREWS,  H.  N.     1955.     Index  of  Generic  Names  of  Fossil  Plants,  1820-1950.     Bull.  U.S. 

geol.  Surv.,  Washington,  1013  :  1-262. 
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252  A   REVISION   OF  THE 

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EXPLANATION  OF  PLATES 


Most  of  the  figured  specimen,  are  in  the  British  Museum  (Natural  History)  and  bear  the 
prefix  V. 


GEOL.  17,  5 


PLATE    i 
Circonitella  knowltoni  (Seward) 

FIGS,  i,  2.     Gyrogonites  showing  five  spiral  cells  meeting  at  the  apical  pole.     Fig.  i,  V. 5 1552, 
Fig  2,  specimen  lost.      X  80. 

FIG.   3.     Basal  pole  of  gyrogonite  showing  circular  basal  plate.     V. 51553.      x  80. 
FIG.  4.     Pyritised,  unsquashed  gyrogonite  with  convex  spiral  cells,  showing  basal  plate. 
V. 51554.      x  80. 

FIG.  5.     Neotype  with  convex  spiral  partly  broken  away.     V. 5 1555.      x  80. 
FIGS.  1-5,  individual  specimens  all  from  one  block  numbered  V. 5 1548  (re-registered  from 

V.i  0700). 

Pachypteris  lanceolate  Brongniart 

FIG.  6.     Apical  part  of  pinna,  lower  side  uppermost,  showing  prominent  midrib.     "^3245. 

X  2. 


Bull.  Br.  Mus.  mat.  Hist.  (Geol.)  17,  5 


PLATE  i 


GEOL.  17,  5 


PLATE  2 
Hepaticites  zeilleri  (Seward) 

FIG.  i.     General  form  of  thallus  showing  dichotomous  branching.     Holotype,  V. 233005      x  i. 

FIG.  2.     Thallus  showing  thick  midrib  region  and  delicate  lamina  also  rhizoids  including 
detached  bunches.     V.233oa.      x  3. 

FIG.  3.     Enlargement  showing  rhizoids  pointing  forwards  between  a  dichotomy  of  the  thallus. 
V.233oa.      x  10. 

Hepaticites  ruff  or  di  sp.  nov. 

FIGS.  4,  5.     Thallus  showing  arcuate  grooves,  Fig.  4  immersed  in  paraffin,  Fig.  5  coated  with 
ammonium  chloride.     Holotype,  V.2343.      x  3. 

Becklesia  sulcata  sp.  nov 

FIG.  6.     Part  of  a  pinna  showing  stomatal  grooves.     V. 5 1540.      x  20. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  5 


PLATE  2 


I 


PLATE  3 
Hausmannia  dichotorna  Dunker 

FIG.   i.     Small  fan-shaped  leaf  with  broken  apices.     Sterile.     ¥.12317.      x  3. 

FIG.   2.     Larger  more  divided  leaf  with  strap-shaped  segments.     Sterile.     ¥.12349.      X  3. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  5 


PLATE  3 


PLATE  4 
Ruffordia  goepperti  (Dunker) 

FIG.  i.  Frond  with  narrowest  form  of  segments.     V.2I57.      x  i. 

FIG.  2.  Frond  with  wider  sterile  pinnules  at  the  top.  Fertile  pinnae  at  the  base  scarcely 
visible  (photographed  dry) .     V.2295.      x  I- 

FIGS.  3,  4.     Fronds  with  widest  pinnules.     Fig.  3,  V.2357.     xi.     Fig.  4,       ¥.12331.     xi. 

FIG.  5.  Partof  V.2I57.      x  3. 

FIG.  6.  Sterilepart  of  V.2295.      x  3- 

FIG.  7.  Pinnaof  V.2357  showing  ventation.      x  3- 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  5 


PLATE  4 


PLATE  5 
Ruffordia  goepperti  (Bunker) 

FIG.  i.  Specimen  with  sterile  top  and  fertile  base,  immersed  in  paraffin.     V.2295.      x  I 

FIG.  2.  Fertile  frond.     V.2i6o.      x  i. 

FIG.  3.  Fertile  frond.     V.2i92«.      x  i. 

FIG.  4.  Frond  in  Fig.  3  enlarged  to  show  the  undulating  margins  of  the  fertile  pinnules. 

V. 21920.  x  3. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol).  17,  5 


PLATE  5 


PLATE  6 
Pelletieria  valdensis  Seward 

FIGS,  i,  2.  Largest  fertile  specimen  picked  out  of  the  matrix.  V.23&8.  Fig.  i,  x  i,  Fig. 
2,  X  3. 

Aspidistes  sewardi  sp.  nov. 

FIG.  3.     Specimen  immersed  in  paraffin,  sori  visible  on  most  of  pinnules.     Holotype,  V.  21730 
x  i. 
FIG.  4.     Middle  part  of  same  specimen  enlarged  to  show  rachis,  sori  and  rolled  edges,      x  3. 

Weichselia  reticulata  (Stokes  &  Webb) 

FIG.  5.  Part  of  pinna  showing  typical  form  with  fleshy  pinnules.  Indication  of  reticulate 
venation  just  visible  in  two  bottom-left  pinnules.  V.  21740.  X  3. 

Pseudotorellia  heterophylla  sp.  nov. 

FIG.  6.  Cuticle  of  upper  surface  showing  cells  in  "  packets  ",  Geological  Survey  &  Museum 
PF2758,  x  200. 

FIG.  7.  Cuticle  of  upper  surface  from  same  specimen  showing  sculptured  surface  of  the 
cells,  x  500. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  5 


PLATE  6 


PRINTED  IN  GREAT  BRITAIN 
BY  ADLARD  &  SON  LIMITED 
BARTHOLOMEW  PRESS,  DORKING 


TWO  NEW  DICYNODONTS  FROM 

THE  TRIASSIC  NTAWERE 

FORMATION,  ZAMBIA 


C.  B.  COX 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
GEOLOGY  Vol.  17  No.  6 

LONDON:   1969 


TWO  NEW  DICYNODONTS  FROM  THE 
TRIASSIC  NTAWERE  FORMATION,  ZAMBIA 


BY 

C.  B.  COX 


Department  of  Zoology,  King's  College,  London 


Pp.  255-294;  23  Text-figures 


BULLETIN  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

GEOLOGY  Vol.  17  No.  6 

LONDON:  1969 


THE     BULLETIN     OF    THE    BRITISH    MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

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ready.  Volumes  will  contain  about  three  or  four 
hundred  pages,  and  will  not  necessarily  be  completed 
within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
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This  paper  is  Vol.  17,  No.  7  of  the  Geological 
Palaeontological  series.  The  abbreviated  titles  of 
periodicals  cited  follow  those  of  the  World  List  of 
Scientific  Periodicals. 


World  List  abbreviation : 
Bull.  Br.  Mus.  nat.  Hist.  (Geol.). 


Trustees  of  the  British  Museum  (Natural  History)  1969 


TRUSTEES    OF 
THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 

Issued  7  March,  1969  Price  Seventeen  Shillings 

I 


TWO  NEW  DICYNODONTS  FROM  THE 
TRIASSIC  NTAWERE  FORMATION,  ZAMBIA 

By  CHRISTOPHER  BARRY  COX 

CONTENTS 

Page 
I.     HISTORICAL  INTRODUCTION     .......         257 

II.     STRATIGRAPHY       .........          258 

III.  SYSTEMATIC  DESCRIPTIONS      .......         260 

Genus  Zambiasaurus  submersus  nov.  ....  262 

Genus  Sangusaiirus  edentatus  nov.       .....  286 

IV.  AGE  OF  THE  NTAWERE  FAUNAS         ......  290 

V.     ACKNOWLEDGMENTS       ........  293 

VI.     REFERENCES          .........          293 

SYNOPSIS 

Two  new  types  of  dicynodont  from  the  upper  fossiliferous  horizon  of  the  Ntawere  Formation 
of  Zambia  are  described.  The  first,  Zambiasaurus  submersus,  is  represented  by  the  fragmentary 
remains  of  at  least  eighteen  juveniles  and  one  adult,  all  of  which  were  apparently  drowned  in  a 
sudden  flood.  Zambiasaurus  is  a  stahleckeriid  closely  related  to,  and  probably  directly  ancestral 
to,  Stahleckeria  of  the  Middle  Triassic  of  Brazil.  It  is  the  earliest  known  stahleckeriid,  and  the 
first  known  outside  South  America.  The  earlier  history  of  the  group  is  unknown. 

The  other  new  dicynodont,  Sangusaurus  edentatus,  is  known  only  from  a  few  fragments  of  the 
skull.  It  is  a  kannemeyeriid,  and  shows  some  similarities  to  1 schigualastia  of  the  Middle  or 
Upper  Triassic  of  Argentina. 

Both  the  faunas  of  the  Ntawere  Formation  appear  to  be  intermediate  in  age  between  the 
Cynognathus  Zone  fauna  of  South  Africa  and  the  Manda  fauna  of  East  Africa.  Its  age,  in  terms 
of  the  standard  Triassic  sequences,  is  provisionally  estimated  as  lowermost  middle  Triassic 
(Lower  Anisian). 

I.  HISTORICAL  INTRODUCTION 

THE  Luangwa  River  is  a  major  tributary  of  the  Zambezi  River,  and  is  the  main 
river  of  the  eastern  half  of  Zambia.  Fossil  reptiles  were  first  found  in  the  upper 
Luangwa  River  valley  by  Prentice  in  1925,  and  the  area  was  reconnoitred  and  later 
investigated  by  Dixey  in  1928  and  1935  (see  Dixey  1936,  1937).  He  recorded  fossil 
reptiles  from  six  different  horizons  within  the  Karroo  succession,  and  also  from  a 
higher  horizon  which  he  named  the  "  Dinosaur  Beds  "  and  which  he  regarded  as 
Cretaceous  in  age. 

The  upper  Luangwa  River  valley  is  remote,  and  is  inaccessible  to  vehicles  unless 
these  can  negotiate  rough  dirt  tracks.  As  a  result,  the  area  was  not  re-examined 
until  1960  and  1961,  when  Dr.  A.  R.  Drysdall  of  the  Geological  Survey  of  Northern 
Rhodesia,  and  James  Kitching,  Field  Officer  of  the  Bernard  Price  Institute  for 
Palaeontological  Research  of  Witwatersrand  University,  spent  a  total  of  four  months 
in  the  area.  They  examined  the  area  north-west  of  the  small  village  of  Sit  we,  on 
the  River  Luwumbu,  125  miles  north  of  Lundazi,  the  administrative  centre  of  this 
north-eastern  portion  of  the  Eastern  Province  of  Zambia.  They  confirmed  that  the 
area  was  rich  in  fossil  reptiles,  and  collected  nearly  500  specimens. 

GEOL.   iy,   6.  23 


258  TWO   NEW   TRIASSIC   DICYNODONTS    FROM   ZAMBIA 

Karroo  fossils  were,  of  course,  originally  found  in  South  Africa,  and  many  of  the 
early  specimens  were  sent  to  the  British  Museum  (Natural  History)  in  London  for 
appraisal  and  description.  The  resulting  collection  is  of  historic  interest  and  contains 
many  type  specimens,  but  many  are  imperfect  and  most  of  them  are  poorly  docu- 
mented and  lack  post-cranial  elements.  It  was  therefore  decided  to  mount  a  major 
expedition  to  Africa,  with  the  aim  of  making  a  large  and  representative  collection 
of  fossil  Permian  and  Triassic  vertebrates,  using  up-to-date  methods  of  collecting 
and  recording  full  details  as  to  their  localities  and  stratigraphical  origin.  In  view 
of  the  known  richness  of  the  area,  it  was  decided  to  rely  upon  the  upper  Luangwa 
River  valley  for  the  collection  of  the  bulk  of  the  Permian  fossils,  and  afterwards  to 
visit  the  south-western  region  of  Tanganyika,  where  fossil  vertebrates  of  both 
Permian  and  Triassic  age  were  known  to  occur. 

The  resulting  expedition,  the  British  Museum  (Natural  History)  and  University 
of  London  Joint  Palaeontological  Expedition  to  Northern  Rhodesia  and  Tanganyika, 
spent  six  weeks  collecting  in  the  Sit  we  area  in  the  summer  of  1963,  and  a  preliminary 
account  of  the  results  has  been  published  (Attridge,  Ball,  Charig  &  Cox  1964). 
The  expedition  was  greatly  helped  by  the  presence  of  James  Kitching,  whose  services 
had  been  very  kindly  loaned  by  the  Bernard  Price  Institute.  As  a  result  there  was 
no  difficulty  in  finding  the  fossil  localities,  once  the  necessary  tracks  for  vehicle 
access  had  been  constructed,  and  a  total  of  220  specimens,  weighing  some  2\  tons, 
was  collected. 

II.  STRATIGRAPHY 

As  a  result  of  their  examination  in  1960-61,  Drysdall  &  Kitching  were  able  to 
give  a  detailed  account  of  the  stratigraphy  and  geology  of  the  area  (Drysdall  & 
Kitching  1963),  from  which  the  details  in  this  section  are  taken. 

Some  alterations  to  Dixey's  earlier  interpretation  of  the  stratigraphy  of  the  area 
became  necessary.  Drysdall  &  Kitching  found  that  all  of  Dixey's  six  Karroo 
horizons  were  part  of  a  single  richly  fossiliferous  formation,  which  they  named  the 
Madumabisa  Mudstone.  The  fauna  of  this  is  identical  with  that  of  the  Lower 
Beaufort  Endothiodon  and  Kistecephalus  zones  of  South  Africa. 

Above  the  Madumabisa  Mudstone,  and  separated  from  it  by  an  hiatus  of  probably 
minor  nature,  Drysdall  &  Kitching  defined  a  major  lithological  unit  which  they 
have  subdivided  to  give  three  formations  of  more  convenient  thickness.  The  lowest, 
the  Escarpment  Grit,  is  of  varying  thickness  (230-  over  700  feet),  whilst  the  upper- 
most, the  Red  Marl,  is  230-400  feet  thick.  Between  the  two  lies  the  Ntawere 
Formation,  again  of  varying  thickness  (400-  over  3,500  feet)  and  consisting  of  a 
succession  of  thin,  alternating  arenaceous  and  argillaceous  beds.  The  whole  unit 
is  subdivided  into  formations  on  the  basis  of  the  different  proportions  of  mudstone 
and  grit. 

Above  the  Red  Marl  lies  the  non-fossiliferous  Upper  Grit,  over  3,000  feet  thick; 
in  some  places  the  two  are  separated  by  a  thin  transition  zone  of  intercalated  grits 
and  mudstones,  while  in  others  there  is  a  sharp  transition. 

Drysdall  &  Kitching  found  two  fossiliferous  horizons  in  the  Ntawere  Formation 
and  Red  Marl.  The  lower  horizon  lies  within  the  Ntawere  Formation;  its  fauna 


TWO   NEW   TRIASSIC    DICYNODONTS    FROM    ZAMBIA  259 

consisted  predominantly  of  large  dicynodonts,  with  fairly  common  large  amphibians, 
and  also  yielded  an  excellent  Diademodon  skull  which  has  already  been  described 
(Brink,  1963).  The  upper  horizon  comprised  the  uppermost  beds  of  the  Ntawere 
Formation  and  the  lower  part  of  the  Red  Marl;  its  fauna  contained  the  remains  of 
both  vertebrates  and  molluscs. 

Drysdall  &  Kitching  consider  that  Dixey's  "  Dinosaur  Beds  "  can  only  be  identi- 
fied, both  lithologically  and  geographically,  with  the  fossiliferous  Ntawere  Forma- 
tion. Dixey  based  his  assessment  of  the  "  Dinosaur  Beds  "  upon  some  large,  frag- 
mentary, rolled  bones  which  he  found  in  situ  and  which  Swinton  tentatively 
identified  as  possibly  dinosaurian.  However,  Drysdall  &  Kitching  found  no  bones 
in  situ  at  the  localities  indicated  on  Dixey's  map,  but  they  were  able  to  identify  a 
thin  superficial  pebble  deposit  which  Dixey  regarded  as  the  last  debris  of  the  almost 
completely  eroded  Dinosaur  Beds.  This  pebble  deposit  contained  large  fragmentary 
bones,  mostly  identifiable  as  dicynodont.  Bones  of  comparable  size  and  appearance 
are  common  in  the  nearby  Madumabisa  Mudstone,  so  that  the  bones  found  in  the 
pebble  deposit  are  probably  derived  fossils  originating  in  the  Madumabisa  Mudstone. 
The  bones  observed  in  situ  by  Dixey  have  not  been  relocated  and  may  also  have 
been  derived  fossils.  Alternatively  they  may  have  come  from  as  yet  unrecognized 
fossiliferous  beds  of  the  Ntawere  Formation,  which  is  now  known  to  contain  pseudo- 
suchians — whose  remains  are  not  readily  distinguishable  from  those  of  dinosaurs. 

The  fossiliferous  beds  themselves  are  dark  red,  soft  mudstones,  with  feldspathic 
grit  bands.  Bones  almost  always  occur  individually;  no  articulated  remains  have 
been  found.  The  bones  are  usually  cracked  and  weathered,  suggesting  pro- 
longed exposure  before  burial;  they  are  often  covered  with  a  purple-red  layer  of 
haematite,  or  with  a  thick  layer  of  calcite.  Fragments  of  bone  are  common  in 
some  of  the  grit  bands,  and  may  be  the  remains  of  skeletons  disarticulated  by 
shortlived  floods  which  were  responsible  for  the  coarser  sediment  forming  the  grit 
bands. 

Drysdall  &  Kitching  state  (1963  :  22)  that  amphibians  and  molluscs  (Unio 
karooensis]  are  present  towards  the  base  of  the  upper  fossiliferous  horizon,  where  the 
commonest  reptiles  are  the  cynodonts.  The  cynodonts  persist  into  the  higher 
levels,  where  the  predominant  reptile  is  now  a  large  dicynodont,  where  the  reptiles 
outnumber  the  amphibians,  and  where  the  molluscs  are  now  rarer.  One  of  the 
cynodonts  has  already  been  described  (Brink,  1963)  as  Luangwa  drysdalli. 

The  fossils  are  found  in  the  systems  of  small  gulleys  and  eroding  flats  at  the  upper 
ends  of  the  small  tributaries  of  the  River  Sangu.  Each  of  the  dicynodont  genera 
described  in  this  paper  was  found  in  an  individual  collecting  area  of  this  kind,  within 
the  general  area  described  as  "  locality  15  "  by  Drysdall  &  Kitching,  and  hence  within 
the  upper  fossiliferous  horizon.  During  the  1963  expedition  we  were  not  able  to 
define  the  stratigraphic  inter-relationships  of  these  individual  areas,  so  that  the 
positions,  within  this  horizon,  of  the  two  dicynodont  genera  described  here  are  not 
known. 

Fossils  from  the  Ntawere  Formation  are  usually  fairly  scarce  and  covered  with 
layers  of  haematite  or  calcite.  The  Zambiasaurus  material  makes  an  outstanding 
exception  to  this  rule:  a  portion,  about  ten  yards  long  and  seven  yards  wide,  of  the 

ENTOM.    17,   7.  23§ 


260  TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 

side  of  a  small  ridge  was  found  to  be  strewn  with  fragments  of  bone,  nearly  all  of  which 
were  completely  free  from  matrix.  The  whole  area  was  stripped  of  vegetation  and 
swept  clean,  with  the  result  that  500-600  fragments  of  bone  were  collected.  Though  no 
bones  could  be  found  in  situ,  some  of  the  fragments  were  embedded  in  pieces  of 
coarse,  unsorted,  highly  feldspathic  grit.  The  proportion  of  the  alkali  feldspar,  quartz 
and  mafic  minerals  suggests  that  the  parent  rock  was  of  granite  composition.  The 
poor  sorting  and  lack  of  rounding  of  the  grains,  and  unaltered  condition  of  the 
feldspar,  suggests  rapid  deposition.  The  whole  collection  was  therefore  deposited 
by  one  of  the  floods  mentioned  above. 

III.    SYSTEMATIC    DESCRIPTIONS 

Genus  ZAMBIASAURUS  nov. 

The  new  genus  has  been  named  Zambiasaurus  after  the  new  name  of  its  country 
of  origin. 

GENERIC  DIAGNOSIS  :  Large  dicynodont :  the  composite  restored  immature  skull 
is  23  cm.  long  and  21  cm.  broad,  while  the  adult  might  have  had  a  skull  about  45  cm. 
long  and  40  cm.  broad.  No  teeth  in  upper  or  lower  jaws.  Greatest  width  of  skull 
is  across  occiput;  skull  tapers  anteriorly.  Wide  inter-orbital  region,  narrow  inter- 
temporal  region.  Blunt  snout.  Short  median  suture  between  nasals.  Preparietal 
bone  absent;  pineal  foramen  completely  surrounded  by  parietals.  Parietals  are 
slightly  concave  antero-posteriorly  and  form  bulk  of  intertemporal  bar.  No  sharp 
median  intertemporal  ridge.  Interparietal  bone  does  not  extend  far  forwards. 
Sharp  transition  between  dorsal  and  occipital  surfaces.  Occipital  wings  of  squa- 
mosal  extend  laterally  and  somewhat  posteriorly.  Palatal  surface  of  premaxilla 
bears  pair  of  anterior  ridges. 

At  least  four  sacral  ribs.  Scapular  blade  tall  and  narrow,  with  low  ridge  running 
up  antero-external  edge;  acromion  process  probably  poorly  developed.  Coracoid 
foramen  wholly  within  precoracoid  bone.  Small  pubis. 

TYPE  SPECIES:  Zambiasaurus  submersus  sp.  nov. 

MATERIAL:  Unless  otherwise  stated,  all  specimen  numbers  refer  to  the  collection 
deposited  in  the  British  Museum  (Natural  History). 

The  bone  fragments  were  mainly  parts  of  the  post-cranial  skeleton.  Apart  from 
unidentified  fragments  and  portions  of  ribs,  the  collection  included : 

Vertebral  centra           .          .  58  Ulna           ....  29 

Sacral  ribs            ...  7  Ilium           ....  40 

Scapula      ....  39  Ischium  16 

Coracoid     ....  13  Pubis          ....  9 

Precoracoid          ...  4  Femur         .          .          .          .  51 

Clavicle       ....  6  Tibia           .          .          .          .24 

Sternum     ....  3  Fibula         ....  13 

Interclavicle        .          .          .  Metapodials,  podials  etc.       .  15 

Humerus    ....  68  Premaxillae         ...  4 

Radius        ....  19  Maxillae      ....  9 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA  261 

Nasals         ....  2  Quadrates  ....  2 

Frontals     ....  6  Squamosals          .          .          .  27 

Postorbital           .          .          .  I  Dentaries    ....  2 

Intertemporal  region    .          .  8  Articular  region  of  lower  jaw  10 

Braincase  elements       .          .  8  Other  fragments  of  lower  jaw  6 

As  found,  none  of  the  limb  bones  was  complete,  but  it  was  subsequently  found  that 
the  fragments  included  the  whole  of  an  ulna,  two  femora  and  two  tibiae.  Apart 
from  a  few  cynodont  bones,  and  one  other  exception  discussed  below,  there  is  no 
variation  in  the  morphology  of  the  bones,  which  therefore  appear  all  to  belong  to  a 
single  species  of  dicynodont.  This  species  is  represented  by  the  remains  of  at  least 
eighteen  individuals,  as  there  are  eighteen  right  distal  ends  of  humeri;  however,  it 
is  almost  certain  that  there  was  really  a  somewhat  greater  number  of  individuals, 
all  of  which  are  only  incompletely  represented.  These  individuals  come  from  a 
fairly  restricted  size  range;  in  the  humeri,  for  example,  the  proximal  ends  are  from 
8-5  to  io-o  cm.  across  and  the  distal  ends  are  from  7-8  to  n-o  cm.  across. 

In  addition  to  these  individuals,  there  are  the  remains  of  a  single  individual  of 
considerably  larger  size,  the  distal  end  of  whose  humerus  is  17  cm.  across.  This 
specimen  could  be  regarded  either  as  a  larger  member  of  the  same  species  as  the 
numerous  smaller  individuals,  or  as  belonging  to  a  different,  larger  species. 

There  are  several  reasons  for  believing  that  the  many  smaller  individuals  are 
immature  forms,  and  that  the  single  large  individual  represents  the  adult  of  the 
same  species.  The  immaturity  of  the  small  specimens  is  indicated  by  the  lack  of 
co-ossification  and  sutural  union  of  the  skull  bones  (even  the  bones  of  the  braincase 
have  not  become  firmly  united)  and  by  the  poorly  defined  nature  of  the  articular 
surfaces  of  the  limb  bones.  On  the  humerus,  for  example,  the  articular  areas  for 
the  glenoid,  radius  and  ulna  are  hardly  defined  at  all,  while  the  articular  surfaces 
of  the  radius,  tibia  and  fibula  are  almost  featureless  and  give  the  impression  of 
having  been  covered  in  life  with  a  thick  capping  of  cartilage.  Such  surfaces  on  this 
and  other  bones  have  a  characteristic  appearance:  the  surface  is  smooth  but  inter- 
rupted by  a  large  number  of  tiny  round  holes,  the  edges  of  which  are  often  slightly 
raised,  giving  the  appearance  of  tiny  volcanic  craters.  In  living  reptiles  this  appear- 
ance is  characteristic  of  surfaces  covered  by  cartilage,  and  it  will  be  referred  to  as 
"  cartilage  ornament  "  in  the  descriptive  sections  below. 

The  bones  of  the  larger  individual,  on  the  other  hand,  have  very  well  developed 
articular  areas,  but  are  otherwise  identical  with  those  of  the  smaller  individuals. 
It  is  therefore  regarded  as  the  adult  of  the  species,  but  is  described  separately,  after 
the  smaller  individuals  have  been  described,  so  as  to  facilitate  any  taxonomic  change 
in  its  status,  should  this  subsequently  be  found  necessary. 

A  death-assemblage  composed  of  a  single  adult  and  many  juveniles  is  rather 
unusual.  The  flood  which  deposited  the  accompanying  coarse  grit  may  provide  a 
possible  explanation ;  it  is  conceivable  that  such  a  flood  might  cause  a  heavy  mort- 
ality in  the  young  dicynodonts,  but  that  only  a  few  of  the  larger  adults  would  be 
caught  and  overwhelmed.  The  adult  might  even  be  a  carcass  of  an  individual 
which  had  died  recently  and  which  was  caught  up  by  the  flood  waters. 


262  TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 

Zambiasaurus  submersus  sp.  nov. 
Text-figs.  1-20 

The  specific  name  submersus  refers  to  the  likelihood  that  the  specimens  were 
drowned. 

HOLOTYPE  OF  Z.  submersus:  Livingstone  Museum  (Zambia)  specimen  No. 
LM/NH  9/2,  consisting  of  interparietal,  left  parietal  and  part  of  left  squamosal  of 
presumed  immature  individual. 

PARATYPES:  British  Museum  (Natural  History)  catalogue  Nos.  R.  9001^.9140; 
Livingstone  Museum  (Zambia)  specimen  Nos.  LM/NH  9/3-9/35. 

HORIZON  AND  LOCALITY  :  Upper  fossilif erous  horizon  of  Triassic  Ntawere  Formation ; 
from  locality  15  of  Drysdall  &  Kitching  (1963),  about  3^  miles  west  of  Sitwe,  in  the 
upper  Luangwa  Valley,  Eastern  Province,  Republic  of  Zambia  (Field  No.  15  A/I). 

DESCRIPTION.  As  explained  below  the  original  skull  morphology  has  had  to  be 
reconstructed  and  deduced  from  the  eighty-five  fragments  listed  above. 

The  most  extensive  fragment,  which  has  been  designated  as  the  type  specimen 
(Livingstone  Museum  specimen  no.  LM/NH  9/2;  cast  in  B.M.(N.H.)  is  R.gooo) 
comprises  an  interparietal,  left  parietal  and  part  of  the  left  squamosal  (Text-fig, 
i,  b-e).  The  right  parietal  has  become  detached  and  lost,  so  that  the  sutural 
surfaces  by  which  it  was  attached  to  the  interparietal  and  left  parietal  are  visible. 
It  can  therefore  been  seen  (Text-fig,  id)  that  the  interparietal  does  not  extend  far 
forwards  between  the  two  parietals.  The  interparietal  also  does  not  extend  far 
laterally  before  it  joins  the  squamosal,  neither  does  it  overlap  the  occipital  surface 
of  the  squamosal  to  any  great  extent.  Though  there  is  a  sharp  transition  between 
the  dorsal  surface  of  the  intertemporal  bar  and  the  posterior  surface  of  the  inter- 
parietal, there  is  also  a  recessed  area  in  the  dorso-median  region  of  the  interparietal, 
where  the  nuchal  ligaments  were  presumably  attached.  Below  this  recess  lies  the 
occipital  surface  of  the  interparietal,  in  which  there  is  a  pair  of  nutrient  foramina. 

The  dorsal  surface  of  the  left  parietal  is  smooth  and  slightly  concave  antero- 
posteriorly.  Though  it  is  also  slightly  concave  transversely,  so  that  there  would 
have  been  a  very  slight  median  ridge,  the  intertemporal  bar  as  a  whole  is  flattened, 
rather  than  forming  an  inverted  V  shape.  The  dorsal  and  lateral  surfaces  of  the 
parietal  meet  at  an  acute  angle  (about  60°)  and  the  lateral  surface  is  shallowly 
concave  dorso-ventrally.  The  posterior  end  of  the  lateral  surface  of  the  parietal 
is  slightly  damaged,  but  extended  back  to  overlap  the  squamosal.  The  upper  part  of 
the  lateral  surface  of  the  parietal  is  slightly  recessed  and  bears  the  pitted  and  grooved 
ornament  characteristic  of  a  surface  overlapped  by  another  bone.  There  can  be 
no  doubt  that  this  surface  was  covered  by  the  posterior  extension  of  the  post- 
orbital.  Another  specimen  (R .  9020)  shows  this  surface  even  more  plainly,  and  it 
is  clear  that  the  postorbital  extended  back  as  far  as  the  junction  between  the  parietal 
and  the  interparietal. 


TWO    NEW   TRIASSIC   DICYNODONTS    FROM   ZAMBIA 


263 


The  parietal  in  the  holotype  forms  the  whole  of  the  border  of  the  left  half 
of  the  pineal  foramen  (Text-fig,  ib).  Another  specimen  (R.9020)  in  which  this 
region  is  slightly  better  preserved,  shows  that  the  parietal  forms  a  thin  strip  of  the 
dorsal  surface  both  lateral  to,  and  anterior  to,  the  pineal  foramen  (Text-fig.  la). 
In  neither  specimen  is  there  any  trace  of  a  suture  delimiting  a  separate  preparietal 
bone,  and  this  bone  must  have  been  absent. 


D 


VSUf. 


FIG.  i.  Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile,  x  f .  A,  dorsal  view  of  anterior 
end  of  left  parietal,  R.go2o.  B-E,  holotype  (Livingstone  Museum  No.  LM/NH  9/2) 
in  B,  dorsal  view;  C,  lateral  view;  D,  medial  view;  E,  anterior  view.  Abbreviations: 
d.su.f.,  dorsal  sutural  area  for  frontal;  IP,  interparietal ;  P,  parietal;  pin.,  pineal  opening; 
rec.pf?,  recess  for  Ppostfrontal;  SQ,  squamosal;  su.p.,  sutural  area  for  right  parietal; 
su.po.,  sutural  area  for  postorbital;  v.su.f.,  ventral  sutural  area  for  frontal. 


264 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM   ZAMBIA 


Much  of  the  anterior  end  of  the  dorsal  surface  of  the  parietal  is  covered  by  ridged, 
grooved  and  pitted  areas  of  overlap  (Text-fig.  ia,  b).  The  most  medial  of  these 
areas  must  have  received  the  posterior  end  of  the  frontal  (Text-fig,  ia,  b)  which 
therefore  reached  as  far  posteriorly  as  the  level  of  the  posterior  border  of  the  pineal 
foramen  and  also  approached  very  close  to  its  lateral  and  anterior  borders  (Text- 
figs.  ia,  2b,  2ob).  The  most  lateral  area  of  sutural  overlap  on  the  antero-dorsal 
region  of  the  parietal  is  that  for  the  postorbital.  Between  this  and  the  area  for  the 
frontal  there  is  a  deep  posteriorly-directed  recess  (Text-fig.  la-c:  rec.pf  ?).  This 


N 


FIG.  2.  Zambiasaurus  submersus  gen.  et  sp.  nov.  Drawings  showing  relationships  of 
preserved  fragments  of  skull  to  reconstruction  of  complete  juvenile  skull,  x  J.  A,  lateral 
view;  B,  dorsal  view;  C,  occipital  view.  Abbreviations:  F,  frontal;  N,  nasal;  MX, 
maxilla;  PMX,  premaxilla;  Q,  quadrate;  SQ,  squamosal. 


TWO    NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA  265 

recess  may  have  received  a  small  separate  postfrontal  bone.  If  it  was  present,  this 
bone  may  have  been  concealed  by  a  superficial  meeting  between  the  postorbital  and 
the  frontal. 

There  is  a  considerable  angle  between  the  more  posterior  part  of  the  parietal  and 
those  areas  to  which  the  frontal  attached.  This  fact,  and  the  concave  outline  of  the 
dorsal  edge  of  the  parietal  in  lateral  view  (Text-fig,  ic),  suggests  that  the  intertemporal 
bar  projected  postero-dorsally  above  the  level  of  the  rest  of  the  dorsal  surface  of 
the  skull  (Text-fig.  2a).  This  is  confirmed  by  fragment  R.goi4  (Text-fig.  3a),  which 
comprises  the  region  around  the  front  end  of  the  pineal  foramen.  The  line  of  the 
dorsal  edge  of  the  intertemporal  bar  apparently  continued  forwards  for  a  short 
distance  on  to  that  part  of  the  frontal  which  lies  antero-lateral  to  the  pineal  foramen, 
for  this  part  of  the  bone  is  slightly  thickened  above  the  level  of  the  rest  of  the  frontal. 
This  fragment  also  confirms  that  there  is  no  preparietal  bone  and  that  the  frontal 
overlaps  the  anterior  end  of  the  parietal.  The  frontal  also  extends  posteriorly  for  a 
short  distance  under  the  parietal,  where  there  is  a  wide  sutural  union  between  these 
two  bones  (Text-fig,  id,  e:  v.su.f.). 

The  morphology  of  the  area  immediately  in  front  of  the  pineal  foramen,  including 
the  interorbital  width,  is  shown  by  specimen  R.goiS.  This  is  a  large  piece  of  a  right 
frontal,  including  both  the  midline  suture  and  part  of  the  edge  of  the  orbit.  Speci- 
men R  .9016  is  an  almost  identical  fragment  of  a  left  frontal,  on  the  postero-medial 
region  of  which  can  be  seen  the  suture  for  attachment  to  the  parietal.  This  fact 
is  important,  for  it  establishes  the  relationship  between  these  frontal  fragments,  inclu- 
ding the  upper  margin  of  the  orbit,  and  the  intertemporal  bar.  The  resulting  position 
of  the  orbit  is  relatively  far  back,  so  that  the  posterior  edge  of  the  post-orbital  bar 
is  at  the  level  of  the  pineal  foramen. 

It  is  possible,  but  not  certain,  that  the  most  antero-lateral  corner  of  the  frontal 
fragment  R .  9015  bears  a  short  stretch  of  the  surface  to  which  the  prefrontal  bone  was 
attached.  Even  if  it  does  not,  the  suture  cannot  have  been  far  anterior  to  this  point, 
since  that  bone  normally  forms  the  antero-dorsal  corner  of  the  orbit. 

Specimen  R.90I2  is  part  of  a  right  nasal  bone  (Text-fig.  3b,  c).  The  bone  was 
overlapped  postero-medially  by  the  anterior  end  of  the  frontal,  and  antero-medially 
by  the  posterior  end  of  the  premaxilla.  These  surfaces  are  separated  by  only  a 
short  median  sutural  surface  for  the  left  nasal.  In  another,  slightly  larger,  right 
nasal  fragment  (specimen  R.90I3)  the  overlap  surfaces  for  the  premaxilla  and  frontal 
meet,  completely  covering  the  median  suture  between  the  two  nasals.  The  antero- 
lateral  corner  of  these  nasals  bears  a  number  of  foramina. 

The  dorsal  surface  of  the  nasal  R.90I2  consists  of  two  planes,  which  meet  along  a 
line  which  runs  antero-laterally.  The  more  postero-lateral  plane  faces  dorso- 
laterally  and  represents  the  lateral  surface  of  the  snout.  The  anterior  plane  faces 
antero-dorsally  and  represents  the  beginning  of  the  down-turning  of  the  snout. 
This  plane  provides  a  clue  as  to  the  relative  position  of  the  nasal  and  of  the  anterior 
end  of  the  snout  since,  together  with  the  anterior  surface  of  the  premaxilla,  it  must 
form  a  smooth  curve  when  seen  in  lateral  view  (Text-fig.  2a).  Some  hint  as  to  the 
distance  between  the  nasal  and  the  anterior  end  of  the  premaxilla  is  also  provided  by 
the  angle  of  the  suture  between  these  two  bones,  as  seen  along  the  front  surface  of 


266 


TWO   NEW   TRIASSIC    DICYNODONTS    FROM   ZAMBIA 


the  nasal.     In  most  dicynodonts  this  suture  runs  to  the  antero-dorsal  corner  of  the 
nostril  in  an  almost  straight  line. 

The  front  edge  of  the  premaxilla  is  blunt,  running  transverse  to  the  main 
axis  of  the  skull.  The  grooves  which  lie  lateral  to  the  paired  anterior  palatal 
ridges  therefore  run  on  to  the  anterior  edge  of  the  premaxilla  (specimen  R.QOOI; 
Text-fig.  3d). 


pin. 


su.n. 


su.m. 


FIG.  3.  Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile,  x  f .  Dorsal  view  of  fragment 
R.QOI4  comprising  area  around  anterior  end  of  pineal  foramen.  B,  C,  fragment  of  right 
nasal  (R.goia)  in  dorsal  (B)  and  medial  (C)  views;  D,  ventral  view  of  premaxilla  (R.gooi). 
Abbreviations:  F,  frontal;  P,  parietal;  pin.,  pineal  foramen;  su.f.,  sutural  area  for  frontal; 
su.m.,  sutural  area  for  maxilla;  su.n.,  sutural  area  for  left  nasal;  su.pmx.,  sutural  area  for 
premaxilla. 


There  is  little  difficulty  in  fitting  a  maxilla  (e.g.  specimen  R .  9004)  to  the  premaxilla. 
All  nine  maxillae  are  tuskless.  Like  the  premaxilla  and  the  anterior  part  of  the 
nasal,  the  maxilla  bears  a  number  of  nutrient  foramina.  There  is  a  slight  posteriorly 
directed  flange  down  its  postero-lateral  edge. 

At  no  point  is  there  any  junction  between  the  preserved  parts  of  the  nasal- 
premaxilla-maxilla  section  and  the  more  posterior  section  of  the  skull.  It  was 
therefore  only  possible  to  attempt  to  estimate  the  original  relationship  between 
these  two  sections  by  assembling  each  independently  in  a  plasticene  matrix  and  then 
matching  them  together,  keeping  the  palatal  surface  of  the  premaxilla  horizontal. 
The  reconstruction  shown  appears  quite  plausible  and  does  not  violate  any  known 
normal  feature  of  dicynodont  morphology.  Nevertheless,  the  exact  distance  between 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM   ZAMBIA  267 

the  two  sections  (and  therefore  the  exact  length  of  the  frontal)  must  remain  in 
doubt. 

At  the  other  end  of  the  skull,  there  is  similarly  no  certain  relationship  between  the 
intertemporal-interparietal  region  and  the  remainder  of  the  squamosal  and  braincase 
(Text-fig.  2).  Most  of  the  fragments  of  squamosal  comprise  the  region  where  the 
zygomatic  arch  arises  from  the  front  surface  of  the  occipital  wing.  The  orientation 
of  the  base  of  the  zygomatic  arch,  and  also  the  orientation  of  the  piece  of  squamosal 
attached  to  the  interparietal  of  the  type  specimen  (Text-fig,  ib)  show  that  the 
squamosals  extended  laterally  and  somewhat  posteriorly  from  the  midline.  The 
approximate  width  across  the  occiput  follows  from  the  interorbital  width,  since  the 
zygomatic  arches  must  have  continued  anteriorly  and  somewhat  medially  into  the 
suborbital  bar. 

The  fragments  of  braincase  include  a  good  basioccipital-basisphenoid  (specimen 
R .  9027)  and  a  good  exoccipital-opisthotic-prootic  (specimen  R .  9028) .  The  occipital 
condyle  is  of  the  normal  tripartite  dicynodont  pattern.  The  tuber  around  the 
fenestra  ovalis  is  formed  partly  by  the  basisphenoid  and  partly  by  the  opisthotic. 
The  opisthotic  also  forms  the  whole  of  the  distal  end  of  the  paroccipital  process; 
its  anterior  surface  is  covered  by  the  prootic.  Dorso-laterally,  both  the  prootic 
and  the  opisthotic  end  abruptly  in  a  thick  surface,  the  ornament  of  which  shows  that 
it  was  capped  by  cartilage,  and  which  presumably  met  the  squamosal.  The  dorsal 
edge  of  the  opisthotic,  which  slopes  upwards  and  outwards,  bears  a  similar  ornament 
and  must  have  joined  the  supraoccipital.  The  medial  surface  of  the  prootic  and 
opisthotic  bears  the  excavation  for  the  inner  ear. 

The  quadrate  is  of  usual  dicynodont  type.  Once  again,  its  exact  relationship  to 
the  remainder  of  the  skull  is  uncertain,  and  it  has  merely  been  placed  in  the  position 
normal  in  the  group,  at  about  the  same  level  as  the  ventral  margin  of  the  premaxilla 
and  ventro-lateral  to  the  distal  end  of  the  paroccipital  process. 

Most  of  the  fragments  of  lower  jaw  are  of  the  articular  region,  which  is  of  normal 
dicynodont  type.  Two  fragments  of  the  front  end  of  the  dentary  (specimens 
R.9039,  9040)  show  that  this  was  wide  and  blunt,  as  might  have  been  expected  from 
the  shape  of  the  premaxilla. 

Postcranial  morphology.  To  facilitate  comparison,  all  bones  are  illustrated  as 
viewed  from  the  left  side.  Where  necessary,  bones  from  the  right  side  have  been 
reversed  in  the  drawings  so  as  to  appear  as  left  side  bones. 

There  is  a  considerable  number  of  fragmentary  vertebrae.  The  prezygapophyses 
are  usually  concave,  the  postzygapophyses  convex.  The  antero-dorsal  corner  of 
the  lateral  surface  of  the  centrum  bears  a  small  facet  for  the  lower  end  of  the  rib 
head.  The  remainder  of  the  facet  for  the  rib  head  extends  postero-dorsally  up  the 
side  of  the  neural  arch.  This  facet  extends  to  a  varying  degree  up  the  ventral 
surface  of  the  transverse  process,  which  is  dorso-laterally  directed. 

It  is  not  possible  to  obtain  any  useful  information  from  the  fragments  of  ribs. 

The  scapula  illustrated  (Text-fig.  4)  is  reconstructed  from  two  complementary 
fragments:  a  lower  fragment  (R.goGS)  and  an  upper  fragment  ^.9069).  The  blade 
is  very  long  and  narrow.  There  is  a  rather  low  scapular  spine  which  runs  for  some 
distance  up  the  antero-lateral  edge  of  the  bone  (Text-fig.  4b)  but  which  does  not 

GEOL.   17,  7. 


268 


TWO   NEW   TRIASSIC    DICYNODONTS    FROM   ZAMBIA 


FIG.  4.  Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.  Composite  restoration.  A,  lateral 
view  of  scapula,  precoracoid  and  coracoid  (scapula  and  precoracoid  are  composite  recon- 
structions) ;  B,  section  through  scapula  at  level  a-b ;  C,  view  of  ventral  end  of  scapula,  lateral 
surface  uppermost,  x  f.  Abbreviations:  cor.f.,  coracoid  foramen;  rug.  trie.,  rugosity 
for  ligament  of  triceps  muscle. 


TWO   NEW   TRIASSIC    DICYNODONTS    FROM   ZAMBIA 


269 


reach  its  upper  end.  The  acromion  process  is  damaged,  but  does  not  appear  to 
have  been  very  large.  There  is  a  pronounced  rugose  area  on  the  postero-dorsal 
edge  of  the  scapula  a  short  distance  above  the  glenoid;  this  is  probably  the  area  of 


FIG.  5.     Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.     Coracoid  in  A,  dorsal  view 

and  B,  medial  view,   x  f 


FIG.  6.     Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.     Sternum  in  dorsal  view,  x  f. 


origin  of  the  scapular  ligament  of  the  triceps  muscle.  The  glenoid  surface  is  sub- 
circular,  but  the  surface  for  attachment  of  the  precoracoid  is  quite  thin  (Text-fig.  40) . 
The  coracoid  illustrated  (specimen  1^.9073;  Text-figs.  4,  5)  is  probably  from  a 
somewhat  smaller  individual  than  the  scapula  illustrated,  as  the  scapular  facet 
on  the  coracoid  is  narrower  than  the  coracoid  facet  on  the  scapula.  The  outlines 
of  these  two  surfaces  do  not,  in  any  case,  match;  this  is  probably  because  there  was 


270 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 


still  much  cartilage  between  the  elements  of  the  girdles  of  these  presumably  juvenile 
individuals. 

The  precoracoid  (Text-fig.  4)  is  incomplete;  it  is  restored  from  three  incomplete 
specimens  (R. 9078-80).  Part  of  the  edge  of  the  coracoid  foramen  is  preserved; 
since  there  is  no  notch  in  the  lower  edge  of  the  scapula,  this  foramen  must  have 
lain  wholly  within  the  precoracoid. 

Though  a  few  fragments  of  clavicle  are  preserved,  these  give  no  useful  information 
about  the  bone.  No  fragments  of  interclavicle  have  been  identified. 

The  sternum  (Text-fig.  6)  is  roughly  hexagonal  in  outline.  Its  ventral  surface  is 
slightly  concave.  On  its  dorsal  surface  lie  a  pair  of  postero-lateral  bosses,  on  the 
ends  of  which  can  be  seen  cartilage-ornament.  These  bosses  probably  gave  insertion 
to  the  ventral  ends  of  the  anterior  thoracic  ribs. 

Since  no  complete  humerus  is  known,  it  has  reconstructed  mainly  from  two  comple- 
mentary fragments,  a  proximal  fragment  R.goSS  and  a  distal  fragment 


A  B 

FIG.  7.  Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.  Composite  restoration  of 
humerus  in  A,  ventral  view  and  B,  posterior  view,  x  f .  Abbreviations :  dp.crest,  delto- 
pectoral  crest;  ent.f.,  entepicondylar foramen. 


TWO    NEW   TRIASSIC    DICYNODONTS    FROM    ZAMBIA  271 

(Text-figs.  7,  8).  It  is  strongly  twisted.  As  in  all  the  limb-bones,  the  areas  of 
muscular  insertion  and  of  articulation  are  poorly  defined,  as  might  be  expected  in 
juvenile  animals.  There  is  thus  little  trace  of  the  proximal  condyle,  nor  of  the 
condyles  for  the  radius  and  ulna.  Traces  of  cartilage-ornament  can  be  seen  over 
many  of  these  surfaces,  proving  that  the  absence  of  the  condyles  is  not  due  simply 
to  erosion . 

No  complete  radius  is  known,  and  the  bone  has  been  reconstructed  (Text-fig.  9) 
from  two  proximal  fragments  (specimens  1^.9093,  9094)  and  two  distal  fragments 
(specimens  R.gogs,  9096).  It  is  a  slender  bone  and  has  been  reconstructed  to  be, 
as  is  normal,  slightly  longer  than  the  ulna.  The  proximal  end  is  slightly  convex, 
the  distal  end  slightly  concave. 

A  single  complete  right  ulna  is  known  (specimen  R.gog8,  Text-fig.  9).  The  bone 
is  fairly  slender.  Only  the  horizontal  surface  of  the  glenoid  notch  is  represented. 
Since  the  whole  of  the  proximal  surface  of  the  ulna  is  preserved  (as  proved  by  its 
covering  of  cartilage-ornament) ,  the  remainder  of  the  sigmoid  notch  must  have  been 


B 

FIG.  8.  Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.  Composite  restoration  of 
humerus  in  A,  dorsal  view  and  B,  anterior  view,  x  f.  Abbreviations:  dp.crest, 
delto-pectoral  crest. 


272 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 


borne  on  a  separately-ossified  olecranon  process.  The  distal  end  of  the  ulna  is 
strongly  convex. 

Like  those  of  the  pectoral  girdle,  the  bones  of  the  pelvic  girdle  (Text-figs.  10-12) 
cannot  be  fitted  together  accurately,  and  much  of  the  regions  between  the  bones 
must  still  have  been  cartilaginous  at  this  stage  of  growth. 

The  outline  of  the  ilium  has  been  based  mainly  on  that  of  specimen R.  9103,  except 
for  the  extreme  posterior  preserved  portion  of  the  blade  which  is  taken  from  speci- 
men R.9io6.  The  sacral  facets  are  reconstructed  from  specimens  R. 9103-05. 

The  outer  surface  of  the  blade  is  slightly  concave  both  dorso-ventrally  and  antero- 
posteriorly,  while  the  inner  surface  is  slightly  convex  in  both  these  planes.  The 


A  B 

FIG.  9.  Zambiasaurus  submersus  gen.,  et  sp.  nov.,  juvenile.  A,  radius  (composite  restoration) 
and  ulna  in  posterior  and  distal  views.  B,  medial  view  of  ulna;  C,  radius  (composite 
restoration)  and  ulna  in  anterior  and  proximal  views,  x  f.  Abbreviations:  RD,  radius; 
UL,  ulna. 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA  273 

acetabular  surface  of  the  ilium  faces  ventro-laterally  and  slightly  anteriorly.  The 
facets  for  at  least  four  sacral  ribs  can  be  distinguished  on  the  inner  surface  of  the 
blade  (Text-fig.  12). 

Most  of  the  outline  of  the  ischium  (Text-figs.  10,  n)  is  based  upon  specimen  R .  9108 ; 
most  of  the  upper  edge  and  a  little  of  the  posterior  edge  is  complete.     Specimen 


pu.tub. 


FIG.  10.     Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.     Lateral  view  of  pelvis.   X 
Abbreviations:  obt.l,  obturator  foramen;  pu.tub.,  pubic  tubercle. 


274 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 


IL 


IS 


FIG.  ii.  Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.  Views  of  acetabular-articu- 
latory  surfaces  of  the  ilium  (IL),  ischium  (IS)  and  pubis  (PU).  x  f .  Lateral  surfaces  of 
the  bones  are  peripheral,  medial  surfaces  are  central. 


FIG.  12.     Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.     Medial  view  of  composite 

restoration  of  ilium,    x    f . 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 


275 


R .  9109  includes  a  little  more  of  the  ventral  portion  of  the  bone.  The  anterior  edge 
of  the  ischium  is  notched  for  the  obturator  foramen.  The  postero-dorsal  region 
of  the  bone  slants  somewhat  inwards,  so  that  there  is  a  stout  ridge  between  this 
surface  and  the  more  antero-ventral  region. 

All  edges  of  the pubis  (specimen  R. 9113,  Text-figs.  10,  n)  bear  cartilage-ornament, 
so  that  the  figures  show  the  bone  complete  at  this  stage  of  ossification.  It  bears  a 
large  convex  face  which  must  include  the  surfaces  which  articulated  with  the  ilium 
and  ischium,  and  also  the  pubic  contribution  to  the  acetabulum.  Below  this  region 
the  pubis  projects  antero-laterally  to  form  a  rather  elongate  pubic  tubercle.  The 
posterior  edge  of  the  bone  is  notched  for  the  obturator  foramen. 

The  femur  illustrated  (Text-fig.  13)  is  specimen  R.girS,  which  is  complete  and 
from  the  right  side;  it  is  i8'5  cm.  long.  The  articular  facets  are  poorly  developed. 
In  side  view  it  can  be  seen  that  the  distal  condyles  were  probably  directed  some- 
what posterior  to  the  main  axis  of  the  bone. 


FIG.  13.     Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.    Femur  in  A,  anterior  view ; 
B,  lateral  view;  C,  posterior  view,   x  f. 


276 


TWO   NEW   TRIASSIC    DICYNODONTS    FROM    ZAMBIA 


The  tibia  illustrated  (Text-fig.  14)  is  specimen  R.QI23,  which  is  also  complete  and 
from  the  right  side.  The  proximal  surface  bears  a  pair  of  concavities  for  the  femoral 
condyles.  The  lateral  surface  of  this  end  of  the  bone  is  also  slightly  notched  where  it 
adjoined  the  proximal  end  of  the  fibula.  The  cnemial  crest  is  represented  by  a 
thickening  of  the  antero-lateral  surface,  which  ends  proximally  in  a  surface  which 
faces  slightly  anteriorly.  The  distal  surface  of  the  tibia  can  be  divided  into  a 
moderately  flat  lateral  region,  and  a  markedly  convex  medial  region. 

The  fibula  is  a  slender,  slightly  curved  bone  (Text-fig.  14).  The  proximal  end 
figured  is  specimen  R .  9128,  while  the  distal  end  is  specimen  R.  9129.  The  proximal 
end  is  convex;  it  is  crescentic  in  outline,  so  that  its  medial  edge  curves  around  the 
lateral  surface  of  the  tibia.  The  distal  articular  surface  is  also  convex,  but  is  oval 
in  outline. 

Though  a  number  of  elements  from  the  hand  and  foot  are  preserved,  no  useful 
information  can  be  derived  from  them. 


FIG.  14.  Zambiasaurus  submersus  gen.  et  sp.  nov.,  juvenile.  A,  tibia  and  fibula  in  posterior 
and  proximal  views;  B,  medial  view  of  tibia;  C,  tibia  and  fibula  in  anterior  and  distal 
views,  x  f.  Abbreviations:  FIB,  fibula;  TIB,  tibia. 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 


277 


ADULT  SPECIMEN.  Together  with  the  remains  of  the  immature  specimens,  there 
was  preserved  also  a  number  of  much  larger  dicynodont  bones.  Since  no  individual 
bone  was  duplicated,  and  since  all  of  these  bones  were  of  commensurate  size,  it 
seems  likely  that  all  are  derived  from  a  single  individual.  They  have  therefore  all 
been  allocated  a  single  specimen  number,  1^.9140.  They  include  a  few  uninformative 
fragments  of  skull  and  ribs,  a  few  neural  arches,  two  vertebral  centra,  a  complete 
left  coracoid  and  fragments  of  a  pair  of  scapulae,  a  left  humerus,  a  right  radius, 
ulna  and  tibia,  and  a  single  phalanx. 

One  of  the  neural  arches  is  almost  complete,  and  is  shown  (Text-fig.  15)  mounted 
upon  one  of  the  centra.  The  rib  facet  extends  from  the  corner  of  the  centrum  up  the 


FIG.  15.    Zambiasaurus  submersus  gen.  et  sp.  nov.,  adult  (R.gi4o).  Neural  arch  and  centrum 
(not  originally  associated)  in  A,  anterior  view;  B,  lateral  view;  C,  posterior  view,  x  \. 

side  of  the  dorso-laterally  directed  transverse  process,  but  does  not  reach  its  distal 
end.  The  outline  of  the  dorsal  end  of  the  neural  spine  is  restored  from  another  speci- 
men from  this  large  individual.  It  is  rather  narrow  antero-posteriorly,  and  slopes 
backwards  at  a  considerable  angle. 

The  outline  and  morphology  of  the  scapula  (Text-fig.  16)  and  coracoid  are  identical 
to  those  of  the  immature  specimen  (cf.  Text-fig.  4).  In  anterior  or  posterior  view 
it  can  be  seen  that  the  blade  of  the  scapula  is  curved  to  conform  to  the  outline  of  the 
rib  cage. 

A  fragment  of  the  postero-dorsal  corner  of  the  proximal  end  of  the  humerus  is 
preserved  and  shows  a  well  developed  articular  condyle  (Text-fig.  I7a).  The  distal 
half  of  the  bone  (Text-fig,  ryb,  c)  shows  well  developed  condyles  for  the  radius  and 
ulna. 


278 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 


\ 


\ 


TWO   NEW   TRIASSIC   DICYNODONTS   FROM     ZAMBIA 


279 


The  proximal  ends  of  the  radius  and  ulna,  and  the  distal  end  of  the  ulna,  are 
preserved.  This  fragment  of  radius  shows  no  features  of  interest.  The  olecranon 
region  of  the  ulna  is  lacking,  and  the  proximal  surface  of  the  remainder  of  this  end 
of  the  bone  is  unfortunately  too  poorly  preserved  for  it  to  be  possible  to  ascertain 
whether  there  was  a  separate  olecranon  ossification,  as  in  other  Triassic  dicynodonts. 

The  proximal  end  of  the  tibia  is  well  preserved  (Text-figs.  18,  19).  The  pair  of 
depressions  for  the  femoral  condyles  are  clearly  visible.  The  cnemial  ridge  running 
up  the  antero-lateral  face  of  the  bone  is  better  developed  than  in  the  immature 
specimen  (cf.  Text-fig.  14),  and  terminates  in  a  rounded  surface  which  is  directed 
dorsally  and  anteriorly.  A  rugose  area  (Text-fig.  19,  rug.)  on  the  postero-lateral 


FIG.  17.  Zambiasaurus  submersus  gen.  et  sp.  nov.,  adult  (1^.9140).  Humerus,  x  f. 
A,  B,  dorsal  views  of  fragments  (A)  of  proximal  end  and  (B)  of  distal  end ;  C,  ventral  view 
of  fragment  of  distal  end.  Abbreviation:  prox.  cond.,  proximal  condyle. 


a8o 


TWO    NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 


corner  of  the  bone  may  mark  the  position  where  the  proximal  end  of  the  fibula 
contacted  the  tibia. 

If  measurements  (in  cm.)  of  the  bones  of  the  adult  are  compared  with  those  of  the 
immature  specimens  figured,  the  following  results  are  obtained. 


Length  of  scapula 

Length  of  coracoid 

Width  of  distal  end  of  humerus 

Width  of  proximal  end  of  radius 

Width  of  proximal  end  of  tibia 


Juvenile 
24 
6-3 

9 
4-6 


Adult 

43 
10-5 

17 

TO 

10-5 


Adult/ 
Juvenile 

1-8 
1-7 
1-9 

2-2 
2-1 


cnem. 
crest 


B 


FIG.  18.     Zambiasaurus  submersus  gen.  etsp.  nov.,  adult.  (R. 914°)     Tibia,  x  £.    A,  medial 
view;  B,  anterior  and  proximal  views.  Abbreviation:  cnem.  crest,  cnemial  crest. 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA  281 

It  can  be  seen  that  the  adult  bones  are  about  twice  the  size  of  the  juvenile  bones. 
If  the  skull  of  the  adult  was,  similarly,  about  twice  the  size  of  the  restored  juvenile 
skull  and  retained  a  similar  length  :  width  ratio,  it  would  have  been  about  45  cm. 
long  and  40  cm.  wide  across  the  occiput. 

DISCUSSION.  Taxonomy  and  relationships  of  Zambiasaurus.  There  can,  first  of 
all,  be  little  doubt  that  Zambiasaurus  is  a  stahleckeriid,  rather  than  a  kannemeyeriid. 
Its  blunt  snout,  wide  but  low  occiput,  short  temporal  opening  and  lack  of  a 


cnem. 
crest 


rug.-f* 


FIG.  19.     Zambiasaurus  submersus  gen.  et  sp.  nov.,  adult  (R. 9140).     Tibia,  x  J.     A,  lateral 
view;  B,  posterior  view.    Abreviation:  cnem.  crest,  cnemial  crest;  rug.,  rugosity. 


high  intertemporal  crest,  are  all  similar  to  these  diagnostic  features  of  the  family 
Stahleckeriidae  as  originally  defined  (Cox,  1965). 

Zambiasaurus  is  the  first  stahleckeriid  known  outside  South  America.  The 
Stahleckeriidae  at  present  includes  three  genera:  Stahleckeria  of  the  Santa  Maria 
Formation  of  Brazil,  described  by  von  Huene  (1935-42) ;  Dinodontosaurus  from  the 
above  Brazilian  Formation  (Cox,  1965)  and  also  from  the  Chanares  Formation  of 
Argentina  (Cox,  1968)  and  Chanaria,  also  from  the  latter  Formation  (Cox,  1968). 
Both  Dinodontosaurus  and  Chanaria  have  retained  tusks  and  a  preparietal  bone, 
in  contrast  to  Stahleckeria  in  which  both  these  features  are  absent.  Their  absence 
in  Zambiasaurus  suggests  that  this  new  genus  is  more  closely  related  to  Stahleckeria 
than  to  the  other  two  stahleckeriid  genera. 


282 


TWO   NEW  TRIASSIC   DICYNODONTS   FROM     ZAMBIA 


Though  tusks  are  frequently  lost  in  dicynodonts,  the  absence  of  the  preparietal 
is  more  rare.  Comparison  of  the  skulls  of  Zambiasaurus  and  Stahleckeria  (Text-fig. 
20)  reveals  several  other  similarities.  Despite  some  superficial  differences,  the 
morphology  of  the  whole  intertemporal  bar  is  very  similar  in  the  two  genera.  The 
parietals  in  Zambiasaurus  can  still  be  seen  anterior  to  the  pineal  foramen,  where  they 
have  occupied  the  space  normally  filled  by  the  preparietal.  In  Stahleckeria  the 


B 


FIG.  20.  Zambiasaurus  submersus  gen.  et  sp.  nov.,  reconstructed  juvenile  skull  x  J  (left) 
and  Stahleckeria  potens,  adult  skull  x  c.  1/12  (right).  A,  lateral  views:  B,  dorsal  views; 
C,  occipital  views.  (Stahleckeria  after  Camp,  1956). 


TWO   NEW   TRIASSIC    DICYNODONTS    FROM    ZAMBIA  283 

frontals  have  extended  slightly  further  back,  enclosing  the  anterior  half  of  the  pineal 
foramen.  In  both  genera  the  frontal  meets  the  parietal  at  a  level  half  way  along  the 
pineal  foramen,  the  parietal  forms  a  rather  wide  intertemporal  bar  which  is  almost 
flat  in  transverse  section,  and  the  interparietal  contributes  little  to  the  intertemporal 
bar. 

Another  similarity  between  Zambiasaurus  and  Stahleckeria  is  that  in  both  the 
nasals  either  meet  for  only  a  very  short  distance  in  the  midline  or  are  separated  by 
a  junction  between  the  frontals  and  the  premaxilla.  In  Stahleckeria  this  is  probably 
a  result  of  the  very  wide,  massive  snout ;  if  the  nasals  lie  far  apart,  the  processes  of 
ossification  outwards  from  the  centre  of  each  bone  might  not  reach  the  midline 
before  this  area  had  been  ossified  by  the  frontal.  As  far  as  one  can  tell,  the  snout 
of  Zambiasaurus  also  was  rather  wide ;  this  follows  from  the  shape  of  the  incomplete 
nasals,  and  the  centre  of  ossification  of  the  bone  certainly  lies  more  lateral  than  these 
fragments. 

As  far  as  the  post-cranial  skeleton  is  concerned,  it  is  in  any  case  difficult  to  find 
many  systematic  differences  between  the  Triassic  dicynodonts  (see  Cox,  1965  for  a 
general  survey).  This  difficulty  is  compounded  in  Zambiasaurus  by  the  fact  that 
most  of  the  bones  are  known  only  in  their  juvenile  state,  with  poorly  developed 
processes  for  muscular  insertion  and  articular  condyles.  A  few  similarities  between 
Zambiasaurus  and  Stahleckeria  can  nevertheless  be  observed.  The  narrow,  pos- 
teriorly-directed neural  spine  of  Zambiasaurus  (Text-fig.  15)  is  very  like  that  of  the 
anterior  vertebrae  of  Stahleckeria.  The  coracoid,  sternum  and  pelvis  are  in  general 
similar  in  the  two  genera,  there  being  a  striking  similarity  between  the  ischium  and 
pubis  of  Zambiasaurus  and  those  of  a  juvenile  Stahleckeria  figured  by  von  Huene 
(1935-42,  PI.  9,  fig.  4).  The  difficulties  mentioned  above  make  it  impossible  to 
make  useful  comparisons  between  the  limb  bones  of  the  two  genera. 

The  absence  of  tusks,  and  of  the  preparietal  bone,  the  morphology  of  the  inter- 
temporal bar  and  of  the  snout,  and  some  post-cranial  features  thus  all  suggest  that 
Zambiasaurus  may  be  quite  closely  related  to  Stahleckeria.  The  Ntawere  Formation, 
in  which  Zambiasaurus  is  found,  is  certainly  older  than  the  Santa  Maria  Formation, 
which  contains  Stahleckeria.  It  is  therefore  possible  that  Zambiasaurus  may  be 
actually  ancestral  to  Stahleckeria,  and  the  known  differences  between  the  two  genera 
may  now  be  examined  with  this  possibility  in  mind. 

As  far  as  one  can  tell,  the  occipital  wings  of  the  squamosal  are  directed  more 
posteriorly  in  Zambiasaurus  than  in  Stahleckeria,  so  that  its  occiput  is  not  as  flat 
as  that  of  Stahleckeria,  and  its  temporal  opening  is  slightly  longer.  The  occiput 
itself,  as  restored,  is  not  as  wide  in  Zambiasaurus  as  in  Stahleckeria.  However,  this 
feature  could  not,  from  the  material  preserved,  be  restored  with  any  great  degree  of 
accuracy.  It  is  also  worth  noting  that  the  restored  skull  is  that  of  a  juvenile  and 
that  the  occiput  apparently  does  increase  in  relative  size  as  the  animal  grows. 
This  is  suggested  by  the  relatively  larger  occiput  of  the  large  specimen  of 
Dinodontosaurus  turpior  which  was  described  as  a  possible  "  old  bull  "  (Cox,  1965, 

pp.  489-494)- 

Another  difference  between  the  occiputs  of  the  two  genera  is  that  the  interparietal 
of  Stahleckeria  may  extend  further  laterally,  as  shown  in  Camp's  (1956)  reconstruction 


284  TWO   NEW   TRIASSIC   DICYNODONTS    FROM   ZAMBIA 

based  on  further  study  of  von  Huene's  material.  However,  the  outline  of  this  bone 
is  shown  dotted  in  his  figure  (1956,  Fig.  45)  and  this  point  is  therefore  uncertain. 
In  any  case,  there  is  no  obvious  reason  why  the  interparietal  should  not  become 
wider  during  the  evolution  of  the  very  wide  occiput  of  Stahleckeria. 

The  snout  of  the  "  old  bull  "  Dinodontosaurus  mentioned  above  is  also  more  massive 
than  that  of  the  smaller  specimens,  and  this  should  be  borne  in  mind  when  comparing 
the  immature  skull  of  Zambiasaurus  with  the  adult  skull  of  Stahleckeria,  as  these 
show  a  similar  difference.  The  snout  of  Stahleckeria  is  extremely  heavy  and  short, 
so  that  the  centre  of  the  maxilla  lies  under  the  anterior  edge  of  the  orbit.  As 
reconstructed,  the  maxilla  of  Zambiasaurus  lies  further  forwards.  However,  the 
exact  inter-relationship  between  its  maxilla  and  orbit  cannot  be  determined  from  the 
fragments  available  (cf.  Text-fig.  2)  and,  in  any  case,  the  condition  shown  in  Zambia- 
saurus could  well  be  ancestral  to  that  found  in  Stahleckeria. 

The  interorbital  region  of  Stahleckeria  also  differs  somewhat  from  that  of  Zambia- 
saurus in  that  its  nasal  and  prefrontal  extend  further  posteriorly,  so  that  the  anterior 
half  of  the  upper  margin  of  the  orbit  is  formed  by  the  prefrontal  instead  of  by  the 
frontal  as  in  Zambiasaurus.  This,  too,  may  be  a  result  of  the  shorter  snout  of 
Stahleckeria,  since  the  centres  of  ossification  of  the  bones  of  this  region  must  lie 
further  posteriorly,  so  that  they  will  encroach  on  the  area  formerly  occupied  by  the 
frontal. 

The  only  obvious  difference  between  the  post-cranial  skeletons  of  Zambiasaurus 
and  Stahleckeria  is  the  shape  of  the  scapula.  In  Stahleckeria  (von  Huene,  1935-42, 
PI.  7,  figs.  1-3)  the  scapula  narrows  above  the  glenoid  region,  but  further  dorsally 
it  becomes  progressively  wider ;  there  is  a  strong,  antero-laterally  directed  acromion 
process,  from  which  a  stout  spine  runs  up  the  outer  surface  of  the  blade  near  its 
anterior  edge.  In  Zambiasaurus,  on  the  other  hand,  the  scapula  does  not  expand 
dorsally,  probably  had  a  small  anteriorly  directed  acromion  process,  and  has  a  very 
low  spine.  However,  there  is  no  reason  to  suppose  that  the  above  features  of  the 
scapula  of  Stahleckeria  could  not  have  evolved  from  the  condition  found  in 
Zambiasaurus. 

To  summarize,  none  of  the  known  differences  between  the  two  genera  precludes 
the  possibility  that  Zambiasaurus  is  not  only  closely  related  to  Stahleckeria,  but  is 
also  directly  ancestral  to  it.  It  is  impossible,  on  the  basis  of  present  material,  to 
test  this  hypothesis  further ;  such  tests  will  depend  on  the  discovery  of  stahleckeriids 
of  an  age  intermediate  between  that  of  the  Ntawere  Formation  and  that  of  the  Santa 
Maria  Formation. 

Origin  of  the  stahleckeriids.  As  will  be  shown  later  in  this  paper,  the  age  of 
the  Ntawere  Formation  may  provisionally  be  estimated  as  lower  Anisian.  Zambia- 
saurus is  therefore  the  oldest  known  stahleckeriid.  As  has  already  been  shown,  it 
is  also  probably  ancestral  to  Stahleckeria  and  less  closely  related  to  the  tusked  genera 
Dinodontosaurus  and  Chanaria.  The  line  leading  to  these  tusked  stahleckeriids 
must,  then,  have  diverged  from  the  tuskless  Zambiasaurus-Stahleckeria  line  in  or 
before  the  Lower  Triassic  (Scythian). 

I  have  earlier  (1965)  suggested  that  the  stahleckeriids  (including  all  the  above 
genera)  may  be  separated  from  the  kannemeyeriids  on  the  basis  of  the  form  of  the 


TWO    NEW   TRIASSIC    DICYNODONTS    FROM    ZAMBIA  285 

palate  and  occiput.  The  morphology  of  these  areas  is,  of  course,  determined  by  the 
orientation  and  volume  of  the  jaw  muscles,  and  by  the  way  in  which  the  jaws  are 
used  during  feeding.  The  modifications  in  the  kannemeyeriid  line  (seen  most  clearly 
in  Kannemeyeria  and  Ischigualastid)  seem  mainly  to  be  directed  towards  increasing 
the  antero-posterior  length  of  the  jaw  muscles.  This  has  been  achieved  by  some 
exaggeration  of  that  posterior  extension  of  the  squamosal  which  is  common  in 
Permian  dicynodonts,  and  also  by  the  postero-dorsal  extension  of  the  posterior 
region  of  the  inter-temporal  bar.  These  features  seem  to  accompany  a  rather 
pointed  premaxilla  (e.g.  Sangusaurus,  see  Text-fig.  22b)  in  which  the  anterior  ends 
of  the  paired  anterior  palatal  ridges  of  the  premaxilla  meet  the  converging  lateral 
walls  of  the  mouth  cavity.  As  a  result  of  this,  the  grooves  which  lie  lateral  to  these 
ridges  meet  the  antero-lateral  margins  of  the  snout,  not  its  anterior  edge. 

The  modifications  of  the  stahleckeriids,  instead,  seem  to  have  been  directed 
towards  an  increase  in  the  width  of  the  skull  and  anterior  end  of  the  jaws.  The  pos- 
terior extension  of  the  squamosal  is  lost  altogether  and  the  temporal  opening  is  very 
short,  but  that  part  of  the  squamosal  which  lies  lateral  to  the  root  of  the  zygomatic 
arch  is  greatly  developed.  This  lateral  development  presumably  provided  attach- 
ment for  muscles  which  would  have  allowed  more  accurate  control  of  the  lateral 
movements  of  the  jaw  than  would  have  been  possible  in  the  kannemeyeriids.  This 
may  be  correlated  with  the  transversely  widened  anterior  surface  of  both  the  pre- 
maxilla and  the  dentary.  As  a  result  of  the  width  of  this  region,  the  paired  palatal 
ridges  of  the  premaxilla,  and  the  grooves  lateral  to  these  ridges,  run  on  to  the  anterior 
edge  of  the  snout  (Text-fig.  3d). 

I  have  earlier  (1965)  suggested  that  these  differences  may  be  explained  by  their 
feeding  habits,  the  kannemeyeriids  having  been  browsers  feeding  on  fronds  and  leafy 
branches,  while  the  stahleckeriids  were  grazers,  feeding  on  any  low  herb-cover  similar 
to  that  provided  by  grasses  today.  Further  understanding  of  the  functional  mech- 
anical differences  between  these  two  groups  will  be  possible  only  when  well  preserved 
undistorted  skulls  and  jaws  are  available.  These  would  permit  the  study  of  the 
sequences  of  possible  jaw  movements,  in  the  same  way  as  Crompton  &  Hotton's 
(1967)  investigation  of  the  functional  morphology  of  two  Permian  dicynodonts, 
Emydops  and  Lystrosaurus. 

It  is  at  present  impossible  to  suggest  confidently  any  links  between  the  Triassic 
dicynodont  groups  and  the  varied  dicynodonts  of  the  Upper  Permian  Kistecephalus 
Zone.  Immediately  above  this  zone  lies  the  Lystrosaurus  Zone,  in  which  tiny 
Myosaums  and  the  aberrant  lystrosaurids  are  the  only  dicynodonts  known.  Above 
this  is  the  Cynognathus  Zone  from  which  Kannemeyeria  is  the  only  dicynodont 
described.  Though  the  Sinokannemeyeria  fauna  of  China  is  about  the  same  age 
as  the  Cynognathus  Zone  fauna  (though  Bonaparte  (1966)  had  tentatively  suggested 
that  the  Sinokannemeyeria  fauna  might  be  younger  than  this,  he  no  longer  considers 
that  this  is  likely  [Bonaparte,  personal  communication]),  the  possible  kannemeyeriids 
Sinokannemeyeria  and  Parakannemeyeria  are  very  specialized  in  their  long  snouts, 
as  Bonaparte  (1966)  has  pointed  out,  a  modification  quite  unlike  the  general  trend 
in  dicynodont  evolution. 

As  a  result  of  these  facts,  little  can  be  said  of  the  history  of  the  stahleckeriids  and 


286  TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 

the  kannemeyeriids  during  the  Scythian  or  (if  their  differentiation  had  taken  place 
even  earlier)  during  the  late  Permian.  Dicynodonts  with  both  the  high,  narrow 
type  of  skull  (e.g.  Daptocephalus,  Dinanomodon)  and  the  low,  wide  type  of  skull 
(e.g.  Aulacephalodori)  are  found  in  the  Upper  Permian  Kistecephalus  Zone  fauna  of 
South  Africa.  There  may  or  may  not  be  phyletic  connections  between  these  Permian 
forms  and  the  corresponding  Triassic  groups;  it  is,  at  present,  impossible  to  decide. 


Genus  SANGUSAURUS  nov. 

DIAGNOSIS.  Large  dicynodont :  complete  skull  would  probably  have  been  35-40 
cm.  long.  No  teeth  in  upper  or  lower  jaws.  Premaxilla  tapers  anteriorly  to  blunt 
point.  Median  region  of  intertemporal  bar  markedly  concave  in  transverse  section. 
Low  boss  immediately  posterior  to  pineal  foramen.  Interparietal  forms  posterior 
end  of  intertemporal  bar. 

TYPE  SPECIES.     Sangusaurus  edentatus  sp.  nov. 


Sangusaurus  edentatus  sp.  nov. 
Text-figs.  21-23 

HOLOTYPE.  Livingstone  Museum  (Zambia)  specimen  No.  LM/NH  9/1  (Field  No. 
15  H/4)  consisting  of  the  following  damaged  bones:  premaxilla,  right  and  left 
maxillae,  right  and  left  quadrates;  postero-dorsal  corners  of  both  squamosals ;  right 
prefrontal;  intertemporal  bar,  consisting  of  interparietal  and  right  and  left  parietals. 

HORIZON  AND  LOCALITY.  Upper  fossiliferous  horizon  of  Triassic  Ntawere  Forma- 
tion; from  locality  15  of  Drysdall  &  Kitching  (1963),  about  3^  miles  west  of  Sitwe, 
in  the  upper  Luangwa  Valley,  Eastern  Province,  Republic  of  Zambia. 

DESCRIPTION.  The  material  was  collected  as  isolated  fragments  over  an  area  of 
about  50  square  yards  but,  since  all  the  bones  are  of  commensurate  size  and  no 
individual  element  is  duplicated,  it  seems  very  likely  that  all  belong  to  a  single  skull. 
Though  much  of  this  skull  was  not  recovered,  the  fragments  preserved  are  sufficient 
to  demonstrate  that  it  belongs  to  a  new  genus  of  dicynodont.  The  generic  name 
Sangusaurus  is  derived  from  that  of  a  nearby  stream,  the  Sangu,  while  the  specific 
name  edentatus  refers  to  the  lack  of  tusks. 

The  most  characteristic  feature  of  the  new  genus  is  the  intertemporal  bar  (Text- 
fig.  21).  The  median  region  of  the  parietals  is  quite  deeply  concave  in  section,  as 
can  be  seen  in  posterior  view  (Text-fig.  2ic).  The  sides  of  this  trough  form  ridges, 
parts  of  the  lateral  surfaces  of  which  may  have  been  originally  covered  by  the  post- 
orbitals,  though  the  bone  is  too  badly  damaged  for  the  areas  of  overlap  to  be 
distinguishable. 

There  is  a  low,  median  rugose  boss  immediately  behind  the  pineal  foramen. 
The  anterior  ends  of  the  parietals  are  damaged,  but  what  remains  of  these  surfaces 


TWO    NEW   TRIASSIC    DICYNODONTS    FROM    ZAMBIA 


287 


bears  a  pattern  of  vertical  grooves  similar  to  those  found  at  the  anterior  end  of  the 
parietal  of  Zambiasaurus  (cf.  Text-fig.  le).  This  suggests  that  these  surfaces  are 
those  to  which  the  frontals  and  preparietal  were  sutured. 

The  interparietal  forms  a  wide  contribution  to  the  posterior  end  of  the  inter- 
temporal  bar  and  has  a  W-shaped  suture  with  the  parietals.  There  is  an  abrupt 
transition  between  this  region  of  the  interparietal  and  its  occipital  region  (Text-fig. 
2ib,  occ.  surf,  ip),  since  these  two  regions  lie  at  a  very  acute  angle  (c.  25°)  to  one 
another.  As  preserved,  the  occipital  surface  of  the  interparietal  therefore  slopes 
very  sharply  antero-ventrally,  and  it  seems  unlikely  that  this  is  due  to  crushing, 
since  this  region  of  the  skull  is  extremely  solid.  If  it  is  undistorted,  this  orientation 
of  the  occipital  surface  of  the  interparietal  implies  that  the  whole  intertemporal 


pin.f. 


occ.  surf.  ip. 


FIG.  21.  Sangusaurus  edentatus  gen.  et  sp.  nov.,  intertemporal  bar  of  type  specimen,  x  ^. 
A,  dorsal  view;  B,  lateral  view;  C,  posterior  view.  Abbreviations:  IP,  interparietal; 
occ.surf.ip.,  occipital  surface  of  interparietal;  P,  parietal;  pin.f.,  pineal  foramen. 


288 


TWO    NEW   TRIASSIC    DICYNODONTS    FROM   ZAMBIA 


bar  was  directed  postero-dorsally,  so  that  its  posterior  end  projected  backwards 
above  the  occiput. 

The  palatal  surface  of  the  premaxilla  (Text-fig.  22b)  bears  a  pair  of  anterior  palatal 
ridges.  The  premaxilla  tapers  anteriorly,  so  that  the  anterior  ends  of  these  ridges 
meet  the  converging  lateral  surfaces  of  the  bone.  The  grooves  lateral  to  these 
ridges  therefore  do  not  emerge  along  the  anterior  edge  of  the  premaxilla,  but  along 
its  antero-lateral  edges,  and  they  are  more  shallow  than  the  median  groove  which 
lies  between  the  palatal  ridges.  The  outer  surface  of  the  anterior  end  of  the  pre- 
maxilla bears  a  median  ridge  and  a  pair  of  lateral  ridges. 

The  right  maxilla  (Text-fig.  3)  is  more  complete  than  the  left.  There  is  no  sign 
of  a  tusk  either  in  the  caniniform  processes  or  in  either  maxillary  antrum.  A 
fairly  well  developed  flange  runs  down  the  postero-lateral  edge  of  the  bone.  The 


B 


FIG.  22. 


saurus  edentatus  gen.  et.  sp.  nov.,  premaxilla  of  holotype.    x 
A.  lateral  view;  B,  ventral  view. 


TWO   NEW   TRIASSIC    DICYNODONTS    FROM    ZAMBIA 


289 


palatal  surface  of  the  maxilla  is  slightly  recessed  anteriorly  where  it  would  have  been 
overlapped  by  the  premaxilla.  If  the  maxilla  is  orientated  so  that  its  caniniform 
process  is  directed  vertically,  the  ventral  edge  of  this  recess  runs  antero-ventrally 
at  a  considerable  angle  to  the  horizontal.  However,  the  premaxilla-maxilla  suture 
in  this  region  in  most  dicynodonts  runs  approximately  horizontally,  parallel  to  the 
palatal  surface.  Such  a  result  can  be  achieved  only  if  the  caniniform  process  is 
swung  forwards  so  as  to  have  a  very  marked  antero-ventral  orientation. 

The  isolated  right  pref rental  and  a  few  other  damaged  and  unidentifiable  fragments 
show  no  features  of  significance. 


B 


FIG.  23.  Sangusaurus  edentatus  gen.  etsp.  nov.,  maxilla  of  holotype.  x  |.  A,  lateral  view; 
B,  section  through  maxilla  at  a-b.  Abbreviation:  gr.pmx.,  groove  marking  lower  limit 
of  area  of  maxilla  originally  covered  by  premaxilla. 


DISCUSSION.  Taxonomy  and  relationships  of  Sangusaurus.  There  can,  first  ot  all, 
be  no  doubt  that  Sangusaurus  is  generically  distinct  from  all  other  dicynodonts, 
since  the  features  mentioned  in  the  diagnosis  are  not  found  together  in  any  previously 
known  genus. 

Several  features  indicate  that  Sangusaurus  is  a  kannemeyeriid  rather  than  a 
stahleckeriid.  These  features  include  the  postero-dorsally  directed  intertemporal 
bar  (if  the  posture  of  this  region  has  been  interpreted  correctly  above)  and  anteriorly 
taping  premaxilla,  the  palatal  ridges  of  which  meet  the  lateral  edges  of  the  bone, 


2Qo  TWO   NEW   TRIASSIC   DICYNODONTS    FROM   ZAMBIA 

not  the  anterior  edge.  The  well  developed  flange  down  the  posterior  surface  of  the 
maxilla  is  also  commonly  found  in  kannemeyeriids  (though  a  small  flange  is  present 
in  the  stahleckeriid  Zambiasaurus) . 

The  Kannemeyeriidae  at  present  includes  six  genera:  Kannemeyeria,  from  the 
Lower  Triassic  Cynognathus  Zone  of  South  Africa,  the  Middle  Triassic  Manda 
Formation  of  East  Africa,  and  the  Middle(?)  Triassic  Puesto  Vie  jo  Formation  of 
Argentina  (Bonaparte,  1966,  1967) ;  Sinokannemeyeria  and  Parakannemeyeria,  both 
from  the  Lower  Triassic  Er-ma-ying  Formation  of  Shansi,  China;  Barysoma,  from 
the  Middle  Triassic  Santa  Maria  Formation  of  Brazil;  Ischigualastia,  from  the  Middle 
or  Upper  Triassic  Ischigualasto  Formation  of  Argentina,  and  Placerias  from  the 
Upper  Triassic  Chinle  Formation  of  Arizona,  U.S.A.  The  morphology  of  Ischigua- 
lastia and  Placerias,  and  the  inter-relationships  of  all  the  above  genera  have  been 
discussed  in  an  earlier  paper  (Cox,  1965). 

Compared  with  the  above  kannemeyeriids  (except  Barysoma,  whose  skull  is  known 
only  from  a  partial  occiput),  the  structure  of  the  intertemporal  bar  of  Sangusaurus 
is  closest  to  that  of  Ischigualastia.  In  both  genera  the  median  region  of  the  parietals 
is  concave  in  transverse  section,  there  is  a  W-shaped  suture  between  the  parietals 
and  the  interparietal,  and  the  posterior  end  of  the  intertemporal  bar  (again,  if 
correctly  interpreted  in  Sangusaurus)  projects  posteriorly  above  the  occiput.  The 
intertemporal  bars  of  the  other  genera  differ.  That  of  Placerias  is  much  wider  than 
that  of  Sangusaurus  and  is  flatter  dorsally  in  cross-section.  That  of  Kannemeyeria 
is  very  narrow,  forming  a  sharp  median  crest.  Those  of  both  Sinokannemeyeria 
and  Parakannemeyeria  are  moderately  wide  and  are  concave  dorsally,  but  are  not 
postero-dorsally  directed,  and  there  is  therefore  a  more  gradual  transition  from  the 
dorsal  surface  of  the  intertemporal  bar  on  to  the  occiput. 

Though  similar  to  one  another  in  general,  the  intertemporal  bars  of  Sangusaurus 
and  Ischigualastia  differ  in  that  the  interparietal  is  much  longer  in  Ischigualastia 
and  forms  the  whole  width  of  the  posterior  end  of  the  intertemporal  bar,  and  the 
dorsal  surface  of  its  interparietal  is  convex,  not  concave  as  in  Sangusaurus. 

Another  similarity  between  Sangusaurus  and  Ischigualastia  is  the  lack  of  tusks 
in  both  genera.  All  the  remaining  kannemeyeriids  are  tusked.  (This  region  is 
unknown  in  Barysoma,  which  appears  to  be  closely  related  to  Ischigualastia.)  How- 
ever, this  similarity  is  less  significant  than  the  similarities  noted  above,  as  many 
dicynodont  genera  have  lost  the  tusks.  If  the  antero-ventral  orientation  of  the 
caniniform  process  of  Sangusaurus  suggested  above  is  correct,  this  is  another  feature 
known  only  in  this  genus  and  in  Ischigualastia. 

The  above  comparisons  suggest  that  Sangusaurus  is  more  closely  related  to  Ischi- 
gualastia than  to  the  other  known  Triassic  kannemeyeriids.  However,  Sangusaurus 
is  so  incompletely  known  that  it  would  be  unwise  to  place  too  much  reliance  upon 
these  resemblances,  or  to  regard  a  relationship  between  the  two  genera  as  definitely 
established. 

IV.  AGE  OF  THE  NTAWERE  FAUNAS 

Before  the  significance  of  the  faunas  of  the  Ntawere  Formation  can  be  appreciated, 
it  is  first  necessary  to  establish  the  Triassic  faunal  sequence  into  which  they  must  be 


TWO   NEW   TRIASSIC   DICYNODONTS    FROM   ZAMBIA  291 

inserted.  I  shall  therefore  first  discuss  the  faunas  and  ages  of  three  other  units: 
the  Cynognathus  Zone  of  South  Africa,  the  Molteno  Beds  of  South  Africa  and  the 
Manda  Formation  of  Tanganyika  (Tanzania) . 

1.  Cynognathus  Zone.     This  is  the  uppermost  unit  of  the  Beaufort  Series.     The 
fauna  includes  cynognathid,  diademodont  and  trirachodont  cynodonts,  the  very 
primitive  rhynchosaurs  Howesia  and  Mesosuchus,  and  the  primitive  pseudosuchians 
Euparkeria  and  Erythrosuchus.     The  only  dicynodont  which  has  been  described  is 
Kannemeyeria ;  however,  two  other  undescribed  genera  are  known,  one  in  the  collec- 
tion of  the  Bloemfontein  Museum,  South  Africa,  and  the  other  in  that  of  the  Smith- 
sonian Institution,  Washington.     Though  the  Cynognathus  Zone  is  up  to  2,000  feet 
thick,  this  fauna  is  found  only  in  the  lowest  500-600  feet  (Hotton  &  Kitching,  1963). 

I  have  recently  (Cox,  1967)  suggested  that  the  Cynognathus  Zone  may  be  best 
regarded  as  of  Upper  Scythian  age.  This  estimate  was  based  upon  the  presence  in 
the  underlying  Lystrosaurus  Zone  (itself  regarded  as  Lower  Scythian  in  age)  of  the 
capitosaurid  labyrinthodont  Wetlugosaurus,  which  had  also  been  identified  from  a 
bed  in  Greenland  above  a  zone  containing  a  dateable  ammonite  fauna.  Though 
Welles  &  Cosgriff  (1965)  have  pointed  out  the  unreliability  of  the  Greenland  specimens 
in  particular,  and  of  this  "  genus  "  in  general,  Cosgriff  has  also  provided  a  better 
basis  for  the  dating  of  the  Cynognathus  Zone.  He  has  described  (Cosgriff,  1965) 
a  new  Australian  temnospondyl  which  is  associated  with  microfloras  and  marine 
fossils,  both  of  which  indicate  a  Lower  Scythian  age.  It  is  closely  related  to 
Peltostega  of  the  Upper  Scythian  Poisdonomya  Beds  of  Spitzbergen,  and  to  Rhytido- 
steus  of  the  Cynognathus  Zone.  Cosgriff  (1965)  also  points  out  that  comparison  of 
the  temnospondyl  amphibians  of  the  Cynognathus  Zone  with  those  of  the  Lystro- 
saurus Zone  suggests  that  there  was  probably  an  appreciable  time  lapse  between  these 
two  faunas.  Relevant  here  is  Hotton  &  Kitching's  (1963)  comment,  based  on 
geological  interpretation,  that  the  Cynognathus  Zone  may  have  been  deposited  after 
a  considerable  interval  of  erosion.  All  these  facts  may  be  reconciled  and  accommo- 
dated if  the  Lystrosaurus  Zone  is  regarded  as  of  Upper  Permian  age  (as  Cosgriff, 
1965,  suggests),  and  is  followed  by  a  gap  in  the  record,  equivalent  to  the  Lower 
Scythian,  which  is  in  turn  followed  by  the  Upper  Scythian  fauna  from  the  lower  part 
of  the  Cynognathus  Zone. 

2.  Molteno  Beds.     These  beds  were  originally  regarded  as  barren.     Boonstra 
(1947)  described  a  cynognathid  cynodont,  Cynidiognathus  longiceps,  collected  by 
Stockley  in  what  he  thought  were  lower  Molteno  Beds  of  Basutoland  (now  Lesotho) . 
This  species  is  known  from  the  Cynognathus  Zone,  and  it  is  now  thought  possible 
that  this  specimen  may  have  come  from  nearby  exposures  of  that  Zone.     More 
recently,    Crompton    &    Ellenberger    (1957)    described    a    traversodont    cynodont, 
Scalenodontoides,  from  the  upper  Molteno  Beds ;  however,  Crompton  (1968,  personal 
communication)  now  considers  that  the  level  from  which  this  specimen  originated 
lies  in  the  strata  which  are  transitional  between  the  Molteno  Beds  and  the  overlying 
Red  Beds,  not  in  the  typical  Molten  Beds.     As  a  result,  once  again,  vertebrate 
fossils  are  unknown  from  the  Molteno  Beds.     These  beds,  up  to  1,900  feet  thick, 
may  therefore  cover  all  or  part  of  the  time  between  the  underlying  Upper  Scythian 


292  TWO   NEW   TRIASSIC    DICYNODONTS    FROM      ZAMBIA 

Cynognathus  Zone  and  the  overlying  Red  Beds.  These  latter  contain  a  varied 
dinosaur  fauna,  including  melanorosaurs,  thecodontosaurs  and  sauropod-like  foot- 
prints, and  are  probably  of  Upper  Triassic  age  (Charig,  Attridge  &  Crompton,  1965). 
No  more  accurate  assessment  can  at  present  be  made  than  that  the  Molteno  Beds 
therefore  represent  part  or  all  of  the  Middle  Triassic  (Ladinian  and  Anisian). 

3.  Manda  Formation.     This  fauna  lacks  the  more  primitive  cynognathid  and 
diademodont   cynodonts  found  in   the   Cynognathus   Zone,   but   instead  includes 
traversodont  cynodonts.     Both  the  rhynchosaur  and  the  diverse  pseudosuchians  are 
far  more  advanced  and  common  than  are  those  of  the  Cynognathus  Zone.     The 
dicynodonts  include  a  greater  variety  of  forms  than  the  Cynognathus  Zone,  but  the 
genus  Kannemeyeria  is  still  present.     The  Manda  fauna  has  provisionally  been 
regarded  as  of  Anisian  (possibly  Upper  Anisian)  age  (Cox,  1965,  1967). 

4.  N  taw  ere  Formation.     As   already  mentioned,    this   formation   contains   two 
fossiliferous  horizons,  which  will  be  discussed  separately.     That  from  the  lower  part 
of  the  formation  contains  two  undescribed  new  tusked  dicynodonts  (in  the  collection 
of  the  Bernard  Price  Institute  for  Palaeontological  Research,  Johannesburg),  the 
cynodont  Diademodon  rhodesiensis ,  and  fragments  of  large  labyrinthodonts.    The  age 
of  this  fauna  must  at  present  depend  upon  the  characters  of  the  cynodont.     Brink 
(1963  :  79)  states  that  this  is  "  very  typically  a  Diademodon  and  it  is  very  difficult 
to  find  a  particular  feature  about  the  skull  on  the  strength  of  which  it  can  emphatically 
be  differentiated  from  a  known  species  like  D.  polyphagus  "  (of  the  Cynognathus 
Zone).     Brink  notes,   however,   that   the  post-dentary  bones   are   more  reduced, 
and  the  coronoid  process  of  the  dentary  better  developed,  than  in  the  Cynognathus 
Zone  species.     These  somewhat  advanced  features  may  indicate  that  this  Ntawere 
fauna  is  slightly  younger  than  that  of  the  Cynognathus  Zone,  and  it  might  conveni- 
ently be  regarded  as  equivalent  to  the  upper,   non-fossiliferous,   portion  of  the 
Cynognathus  Zone. 

The  upper  fossiliferous  horizon  extends  over  the  topmost  beds  of  the  Ntawere 
Formation  and  the  lower  part  of  the  Red  Marl.  Its  fauna  includes  the  stahleckeriid 
dicynodont  Zambiasaurus,  the  kannemeyeriid  dicynodont  Sangusaurus,  two 
traversodont  cynodonts  (Luangwa  drysdallensis  Brink  1963  and  another  specimen 
which  will  be  described  by  Crompton),  fragments  of  pseudosuchians,  and  the  remains 
of  the  largest  known  labyrinthodont  amphibian,  approximately  twice  the  size  of 
Paracydotosaurus  (which  will  be  described  by  Panchen). 

The  absence  of  diademodonts  (as  far  as  is  known),  the  presence  of  traversodonts 
and  its  stratigraphical  position  above  the  lower  fossiliferous  horizon  (itself  of  possible 
Cynognathus  Zone  affinities)  all  indicate  that  this  upper  Ntawere  fauna  is  younger 
than  the  Cynognathus  Zone.  The  above  characters  of  its  cynodont  fauna  similarly 
indicate  a  closer  relationship  to  the  Manda  fauna.  The  only  remaining  problem  is 
whether  this  Ntawere  Fauna  should  be  regarded  as  older  than,  or  the  same  age  as, 
the  Manda  Formation.  The  only  fact  relevant  to  this  decision  is  the  absence  from 
the  Ntawere  fauna  of  the  rhynchosaurs  which  form  such  a  large  element  in  the  Manda 
fauna.  Though,  as  has  been  pointed  out  previously  (Cox,  1967),  this  group  is  notably 
unreliable  as  a  faunal  indicator,  it  must  be  accepted  here  in  the  absence  of  any  more 


TWO   NEW   TRIASSIC    DICYNODONTS    FROM    ZAMBIA  293 

reliable  evidence.  The  upper  fossiliferous  horizon  of  the  Ntawere  Formation  is 
therefore  provisionally  to  be  regarded  as  slightly  older  than  the  Manda  Formation. 
If  this  is  so,  the  suggestion  of  Dixey  (1936)  and  of  Drysdall  &  Kitching  (1964)  that 
the  Escarpment  Grit  (which  underlies  the  Ntawere  Formation)  may  be  equivalent 
to  the  Kingori  Sandstone  (which  underlies  the  Manda  Formation)  must  be  incorrect 
unless  the  Kingori  Sandstone  is  equivalent  to  both  the  Ntawere  Formation  and  the 
Escarpment  Grit.  It  is,  perhaps,  more  likely  that  the  Kingori  Sandstone  represents 
a  later  phase  in  the  sedimentary  cycle  which  began  with  the  Escarpment  Grit. 

TABLE  i 

South  Africa  Zambia  Tanzania 

Rhaetian  Cave  Sandstones 

Norian  Red  Beds 

Carnian  "1 

Ladinian  >     Molteno  Beds 

U.  Anisian         J  Manda  Formation 

L.  Anisian  Unfossiliferous  Ntawere  Formation  Kingori  Sandstone 

U.  Cynognathus  Zone 
U.  Scythian  Cynognathus  Zone  Escarpment  Grit 

fauna 

L.  Scythian 
U.  Permian  Lystrosaurus  Zone 

Kistecephalus  Zone  Madumabisa  Kawinga  Formation 

Mudstone 


V.  ACKNOWLEDGMENTS 

Most  of  the  expenses  of  the  1963  expedition,  during  which  these  specimens  were 
collected,  were  met  by  grants  from  the  Department  of  Scientific  and  Industrial 
Research  and  the  Royal  Society ;  grants  were  also  kindly  made  by  the  Percy  Sladen 
Memorial  Fund,  the  Godman  Exploration  Fund  and  Shell  Research  Ltd.  All  the 
members  of  the  Expedition  are  grateful  to  these  bodies  for  their  support,  which  made 
the  Expedition  possible. 

The  drawings  in  this  paper  are  by  Miss  J.  Joffe  and  Mr.  P.  Hutchinson,  who  were  in 
turn  employed  as  my  research  assistant.  The  salary  of  this  post  was  met  from  a 
special  grant  of  the  Natural  Environment  Research  Council,  for  whose  support  I  am 
greatly  indebted. 

VI.  REFERENCES 

ATTRIDGE,  J.,  BALL,  H.  W.,  CHARIG,  A.  J.  &  Cox,  C.  B.     1963.     The  British  Museum  (Natural 

History) — University  of  London  Joint  Palaeontological  Expedition  to  Northern  Rhodesia 

and  Tanganyika,  1963.     Nature,  Lond.  201  :  445-449,  4  figs. 
BONAPARTE,   J.   F.     1966.     Una  nueva  "  fauna  "  Triasica  de  Argentina  (Therapsida:   Cyno- 

dontia  Dicynodontia) .     Consideraciones  filogeneticas  y  paleobiogeograficas.     Ameghiniana, 

Buenos  Aires,  4  :  243-296,  29  figs.,  2  pis. 
—  1967.     New  vertebrate  evidence  for  a  southern  transatlantic  connexion  during  the  Lower 

or  Middle  Triassic.     Palaeontology,  London,  10  :  554-563,  7  figs. 


294  TWO   NEW   TRIASSIC   DICYNODONTS    FROM    ZAMBIA 

BOONSTRA,  L.  D.  1947.  Notes  on  some  Stormberg  fossil  bones  from  Basutoland.  In  G.  M. 
Stockley's  Report  on  the  geology  of  Basutolani,  pp.  94-95,  i  pi.  Maseru. 

BRINK,  A.  S.  1963.  Two  cynodonts  from  the  Ntawere  Formation  in  the  Luangwa  Valley  of 
Northern  Rhodesia.  Palaeont.  afr.,  Johannesburg,  8  :  77-96,  figs.  12-15. 

CAMP,  C.  L.  1956.  Triassic  dicynodont  reptiles.  Part  II.  Triassic  dicynodonts  compared. 
Mem.  Univ.  Calif.,  Berkeley,  13  :  305-341,  figs.  42-62. 

CHARIG,  A.  J.,  ATTRIDGE,  J.  &  CROMPTON,  A.  W.  1965.  On  the  origin  of  the  sauropods 
and  the  classification  of  the  Saurischia.  Proc.  Linn.  Soc.  Lond.  176  :  197-221. 

COSGRIFF,  J.  W.  1965.  A  new  genus  of  Temnospondyli  from  the  Triassic  of  Western  Australia. 
/.  Proc.  R.  Soc.  West.  Aust.,  Perth,  48  :  65-90,  13  figs. 

Cox,  C.  B.  1965.  New  Triassic  dicynodonts  from  South  America,  their  origins  and  relation- 
ships. Phil.  Trans.  R.  Soc.,  London,  248B  :  457-516,  30  figs. 

—  1967.     Changes  in  terrestrial  vertebrate  faunas  during  the  Mesozoic.     In:  Harland,  W.  B. 
et  al.  eds.  The  Fossil  Record,  London  (Geological  Society),  pp.  77-89,  i  fig. 

—  1968.     The  Chanares   (Argentina)   Triassic  reptile  fauna.     IV.     The  dicynodont  fauna. 
Breviora,  Cambridge,  Mass.  No.  205:  1-27,  12  figs. 

CROMPTON,  A.  W.  &  ELLENBERGER,  F.  1957.  On  a  new  cynodont  from  the  Molteno  Beds  and 
the  origin  of  the  tritylodontids.  Ann.  S.  Afr.  Mus.,  Cape  Town,  44  :  1-14,  5  figs.,  i  pi. 

CROMPTON,  A.  W.  &  HOTTON,  N.  1967.  Functional  morphology  of  the  masticatory  apparatus 
of  two  dicynodonts  (Reptilia,  Therapsida).  Postilla,  New  Haven,  No.  109  :  1-51,  7  figs. 

DIXEY,  F.  1936.  The  Karroo  of  the  upper  Luangwa  valley  of  north-east  Rhodesia.  Rep. 
geol.  Surv.  Dep.  Nyasald.,  Livingstone,  24  : 

—  1937.     The  geology  of  part  of  the  upper  Luangwa  valley,  north-eastern  Rhodesia.      Q.  Jl 
geol.  Soc.  Lond.  93  :  52-74,  i  fig.,  pi.  iv. 

DRYSDALL,  A.  R.  &  XITCHING,  J.  W.     1963.     A  re-examination  of  the  Karroo  succession  and 

fossil  localities  of  part  of  the  upper  Luangwa  Valley.     Mem.  geol.  Surv.  Dep.  N.  Rhod., 

Lusaka,  1  :  1-62,  4  figs.  7  pis. 
HOTTON,  N.  &  KITCHING,  J.  W.     1963.     Speculations  on  Upper  Beaufort  deposition.     S.  Afr. 

J.  Sci.,  Cape  Town,  59  :  254-258. 
HUENE,  F.  VON     1935-42.     Die  fossilen  Reptilien  des  sudamerikanischen  Gondwanalandes  an  der 

Zeitenwende.     332  pp.,  66  figs.,  38  pis.  Tubingen. 
WELLES,  S.  P.  &  COSGRIFF,  J.  W.     1965.     A  revision  of  the  labyrinthodont  family  Capito- 

sauridae.     Univ.  Calif.  Publs.  Bull.  Dep.  Geol.,  Berkeley,  54  :  1-148,  48  figs.,  i  pi. 


PRINTED  IN  GREAT  BRITAIN 
BY  ADLARD  &  SON  LIMITED 
BARTHOLOMEW  PRESS,  DORKING 


LOWER  CAMBRIAN  L2S 

ARCHAEOCYATHA  FROM  THE 

AJAX  MINE, 
BELTANA,  SOUTH  AUSTRALIA 


FRANCOISE  DEBRENNE 


BULLETIN  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

GEOLOGY  Vol.  17  No.  7 

LONDON:  1969 


LOWER  CAMBRIAN  ARCHAEOCYATHA 

FROM  THE  AJAX  MINE, 
BELTANA,  SOUTH  AUSTRALIA 


BY 

FRANCOISE  DEBRENNE 

Museum  National  d'Histoire  Naturelle,  Paris 


Pp.  295-376;  18  Plates,  15  Text-figures. 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
GEOLOGY  Vol.  17  No.  7 

LONDON:  1969 


THE     BULLETIN    OF    THE    BRITISH    MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they  become 
ready.  Volumes  will  contain  about  three  or  four 
hundred  pages,  and  will  not  necessarily  be  completed 
within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  17,  No.  7  of  the  Geological 
(Palaeontological)  series.  The  abbreviated  titles  of 
periodicals  cited  follow  those  of  the  World  List  of 
Scientific  Periodicals. 


World  List  abbreviation  : 
Bull.  Br.  Mus.  nat.  Hist.  (Geol.). 


Trustees  of  the  British  Museum  (Natural  History)  1969 


TRUSTEES    OF 
THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 

Issued  15  April,  1969  Price  £4  155. 


LOWER  CAMBRIAN  ARCHAEOCYATHA 

FROM  THE  AJAX  MINE, 
BELTANA,  SOUTH  AUSTRALIA 

By  FRANCOISE  DEBRENNE 


CONTENTS 

Page 

I.     INTRODUCTION          .........  299 

II.     SYSTEMATIC  DESCRIPTIONS         .......  302 

Phylum  ARCHAEOCYATHA       .          .          .  .          .          .302 

Class  REGULARIA  Vologdin,  1937          •          •          •          •          •  3°2 

Order  MONOCYATHIDA  Okulitch,  1935         ....  302 

Family  MONOCYATHIDAE  R.  &  W.  R.  Bedford,  1934         •  3°2 

Genus  MONOCYATHUS  R.  &  W.  R.  Bedford,  1934  302 

M.  porosus  R.  &  W.  R.  Bedford    .          .          .  303 

M.  mellifer  R.  &  W.  R.  Bedford    ...  303 

Genus  TUMULIOLYNTHUS  Zhumvlevz,  1963       .  304 

T.  irregularis  (R.  &  W.  R.  Bedford)        .          .  304 

Order  AJACICYATHIDA  R.  &  J.  Bedford,  1939      .          .          .  305 

Family  DOKIDOCYATHIDAE  R.  &  W.  R.  Bedford,  1936      .  305 

Genus  ALPHACYATHUS  R.  &  J.  Bedford,  1939    .  305 

A.  cf.  annularis  (R.  &  W.  R.  Bedford)    .          .  305 

Family  ACANTHINOCYATHIDAE  R.  &  W.  R.  Bedford,  1936  306 

Genus  ACANTHINOCYATHUS  R.  &  W.  R.  Bedford,  1936  306 

A.  apertus  R.  &  J.  Bedford  ....  307 

Family  AJACICYATHIDAE  R.  &  J.  Bedford,  1939       .          .  308 

Genus  ARCHAEOCYATHELLUS  Ford,  1873          .  308 

Subgenus  STAPICYATHUS  Debrenne,  1964         .  309 

A.  (S.)  stapipom  (Taylor)       .          .          .  309 

Genus  LOCULICYATHUS  Vologdin,  1931      .          .  310 

Subgenus  LOG  ULICYATHELLUS  nov.       .          .  310 

L.  (L.)  floreus  (R.  &  W.  R.  Bedford)       .  310 

Family  ROBUSTOCYATHIDAE  Debrenne,  1964   .          .          .  311 

Genus  ROBUSTOCYATHUS  Zhuravleva,  1960         .  311 

R.  magnipora  (R.  &  W.  R.  Bedford)       .          .  311 

R.  subacutus  (R.  &  W.  R.  Bedford)         .          .  312 

Genus  ZONACYATHUS  R.  &  J.  Bedford,  1937        .  314 

Z.  retevallum  (R.  &  W.  R.  Bedford)        .          .  315 

Z.  retezona  (Taylor)      .          .          .          .          .  316 

Family  BRONCHOCYATHIDAE  R.  &  J.  Bedford,  1936  .  317 

Genus  CYATHOCRICUS  nov 318 

C.  tracheodentatus  (R.  &  W.  R.  Bedford)          .  319 
C.  dentatus  (Taylor)      .          .          .          .          .321 

Family  ETHMOCYATHIDAE  nov.       .....  322 

Genus  ETHMOCYATHUS  R.  &  W.  R.  Bedford,  1934  323 

E.  lineatus  R.  &  W.  R.  Bedford     ...  323 

GEOL.    17,  7  24 


298  LOWER   CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA 

Family  ERISMACOSCINIDAE  Debrenne,  1964      .          .          .  325 

Genus  ERISMA  COSCIN  US  Debrenne,  1958   .          .  325 

E.  rugosus  (R.  &  W.  R.  Bedford)  .  .  .      .          .  325 

E.  equivallum  (Taylor)  .          .          .          .  326 

E.  quadratus  (R.  &  W.  R.  Bedford)         .          .  328 

E.  textilis  (R.  &  W.  R.  Bedford)    ...  329 

E.  cellularis  (R.  &  W.  R.  Bedford)          .          .  330 

E.  peter  si  (R.  &  W.  R.  Bedford)    .          .          .  331 

E.  retifer  (R.  &  W.  R.  Bedford)     ...  332 

Family  POLYCOSCINIDAE  Debrenne,  1964         .          .          .  334 

Genus  TOMOCYATHUS  Rozanov,  1960  .          .  334 

Subgenus  ERUGATOCYATHUS  nov.  .          .  334 

T.  (E.)  papillatus  (R.  &  W.  R.  Bedford)  334 

Family  SALAIROCYATHIDAE  Zhuravleva,  1955  .          .  336 

Genus  SALAIROCYATHUS  Vologdin,  1940   .          .  336 

Subgenus  S.  (SALAIROCYATHUS)  Vologdin,  1940  336 

?  S.  (S.)  annulatus  (R.  &  W.  R.  Bedford)  337 

Family  ALATAUCYATHIDAE  Zhuravleva,  1955  .          .  338 

Subfamily  TUMULOCOSCININAE  Zhuravleva,  1960  .  338 

Genus  ETHMOCOSCINUS  Simon,  1939  .          .  338 

E.  papillipom  (R.  &  W.  R.  Bedford)      .          .  339 

Subfamily  ALATAUCYATHINAE  Zhuravleva,  1960  .          .  340 

Genus  ALATAUCYATHUS  Zhuravleva,  1955          .  340 

Subgenus  A NAPTYCTOCYATHUS  nov.     .          .  340 

A .  (A.)  cribripora  R.  &  W.  R.  Bedford  .  340 

A.  (A.)  flabellus  sp.  nov.        .          .          .  341 

Class  IRREGULARIA  Vologdin,  1937       .....  342 

Order  ARCHAEOCYATHIDA  Okulitch,  1935  .      .  .          .  342 

Family  DICTYOCYATHIDAE  Taylor,  1910  .          .          .  342 

Genus  PIN  A  COC  YA  TH  US  R.  &  W.  R.  Bedford,  1 934  342 

P.  spicularis  R.  &  W.  R.  Bedford.          .          .  343 

Family  FLINDERSICYATHIDAE  R.  &  J.  Bedford,  1939        .  344 

Genus  FLINDERSICYATHUS  R.  &  J.  Bedford,  1937  344 

Subgenus  FLINDERSICYATHUS      R.       &      J. 

Bedford,  1937      .          •          •          •          •  34^ 

F.  (F.)  graphicus  (R.  &  W.  R.  Bedford) .  346 

F.  (F.)  irregularis  (Taylor)     .          .          .  348 

F.  (F.)  major  (R.  &  W.  R.  Bedford)        .  349 

F.  (F.)  speciosus  (R.  &  W.  R.  Bedford)  .  350 

Subgenus  PYCNOIDOCYATHUS  Taylor,  1910    .  350 

F.  (P.)  synapticulosus  (Taylor)        .          .  351 

F.  (P.)  simplex  (Taylor)         .          .          .  352 

F.  (P.)  maximipora  (R.  &  W.  R.  Bedford)  352 

F.  (P.)  vicinisepta  (R.  &  W.  R.  Bedford)  .  353 

Family  METACYATHIDAE  R.  &  W.  R.  Bedford,  1934          •  354 

Genus  METALDETES  Taylor,  1910        ...  355 

M.  dissepimentalis  (Taylor)  ....  358 

M.  irregularis  (R.  &  W.  R.  Bedford)       .          .  359 

M.  taylori  (R.  &  W.  R.  Bedford)   ...  360 

Genus  METAFUNGIA  R.  &  W.  R.  Bedford,  1934  .  361 

M.  reticulata  R.  &  W.  R.  Bedford  .          .  362 

Family  METACOSCINIDAE  R.  &  W.  R.  Bedford,  1936         .  363 

Genus  METACOSCINUS  R.  &  W.  R.  Bedford,  1934  3^3 

M.  reteseptatus  R.  &  W.  R.  Bedford       .          .  364 


LOWER   CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA  299 

?  Family  METACOSCINIDAE   ......  367 

Genus  PYCNOIDOCOSCINUS  R.  &  W.  R.  Bedford, 

1936.          .  ....  367 

P.  pycnoideum  R.  &  W.  R.  Bedford        .          .  367 

III.  STRATIGRAPHICAL  CONCLUSIONS  .          ......         368 

IV.  SUMMARY         ..........          371 

V.     ACKNOWLEDGMENTS  ........         372 

VI.     REFERENCES   ..........         373 

APPENDIX     GLOSSARY  .......         376 

SYNOPSIS 

Specimens  from  the  Ajax  Mine,  first  described  by  R.  &  W.  R.  Bedford  (1934,  J936)  and  now  in 
the  British  Museum  (Natural  History),  have  been  re-examined  and  are  revised.  Fifteen  families, 
21  genera,  4  subgenera  and  41  species  are  discussed  and  arranged  according  to  current  classifica- 
tion. One  family  (Ethmocyathidae),  one  genus  (Cyathocricus) ,  three  subgenera  (Loculicyathellus, 
Erugatocyathus,  Anaptyctocyathus)  and  one  species  (Anaptyctocyathus  flabellus)  are  considered 
new.  The  two  genera  Metacyathus  and  Bedfordcyathus  are  shown  to  be  synonyms  of  Metaldetes. 
The  excellent  preservation  of  this  silicified  material  has  enabled  the  internal  structures  of  some 
species  to  be  determined  accurately  and  resulted  in  the  discovery  of  new  wall  and  intervallum 
types. 

Stratigraphical  correlation  shows  that  there  are  clear  affinities  between  the  Ajax,  Kameshki 
and  Sanashtygkol  faunas.  The  presence  of  both  advanced  and  simple  forms  in  the  Ajax  fauna 
indicates  that  it  is  probably  of  Upper  Kameshki-Lower  Sanashtygkol  age,  i.e.  the  middle  of  the 
lower  division  of  the  Lower  Cambrian. 

Ce  travail  a  pour  but  de  reviser  les  Archeocyathes  qui  firent  1'objet  du  premier  m6moire  de 
R.  et  W.  R.  Bedford  (1934)  et  qui  sont  actuellement  conserv6s  dans  les  collections  du  B.M.(N.H.). 
15  families,  21  genres,  4  sous-genres  et  41  especes  sont  decrits  et  classes  selon  les  criteres  actuels 
de  classification.  Une  nouvelle  famille  (Ethmocyathidae),  un  nouveau  genre  (Cyathocricus), 
trois  sous-genres  (Loculicyathellus,  Erugatocyathus,  Anaptyctocyathus}  et  une  nouvelle  espece 
ont  ete  etablies.  Bedfordcyathus  et  Metacyathus  tombent  en  synomymie  avec  Metaldetes.  La 
fossilisation  exceptionnelle  du  materiel  silicifi6  a  permis  la  definition  de  nouvelles  structures 
murales  et  intervallaires. 

Les  faunes  d'Ajax  ont  des  affinites  avec  les  faunes  des  horizons  de  Kameshki  et  Sanashtygkol, 
c'est  a  dire  qu'elles  datent  du  milieu  de  la  partie  inferieure  du  Cambrien  inferieur. 

I.  INTRODUCTION 

THE  present  work  is  a  revision  of  material  in  the  British  Museum  (Natural  History) 
collection,  collected  and  originally  described  by  R.  &  W.  R.  Bedford  in  1934.  The 
specimens  come  from  a  Lower  Cambrian  exposure  in  a  limestone  hill  behind  the  Ajax 
copper  mine,  ten  miles  north-east  of  Beltana,  in  the  Flinders  Range,  South  Australia. 
T.  G.  Taylor  was  the  first  to  visit  this  locality  in  February,  1906  and  his  subsequent 
monograph  on  Australian  Archaeocyatha  (1910)  was  the  first  work  to  give  detailed 
descriptions  of  such  fossils. 

In  a  series  of  papers  from  1934-1939,  the  Bedfords,  R.,  W.R.  &  J.,  described 
material  collected  in  South  Australia.  The  specimens  mentioned  in  their  papers  now 
belong  to  the  following  museums,  according  to  correspondence  in  the  British  Museum 
(Natural  History)  from  Dr.  Dorothy  Hill:  Memoir  I — B.M.  (N.H.) ;  Memoir  2 — South 
Australian  Museum ;  Memoirs  3,  4  and  6 — Princeton  University,  U.S.A.  The  remain- 


300  LOWER   CAMBRIAN    ARCH  AEOC  Y  ATH  A   FROM   AUSTRALIA 


Unmefarrtorpriosec/     racers, 
sands/ones,     sh»/rs. 


CAMBRIAN        OUTCROPS 

OF 
SOUTH         AUSTRALIA 


»vi»h      sliqht       modifications     from     the 

Geological      Map      of     South     Australia 

D«p».     of      Min«s,     Adelaide.    1953 


30° 


FIG.  i.     The  Cambrian  outcrops  of  South  Australia.     After  B.  Daily,  1956. 


LOWER   CAMBRIAN    ARCH  AEOC  Y  ATH  A   FROM    AUSTRALIA  301 

der  of  the  Bedford  collection  was  acquired  by  Prof.  D.  A.  Brown  of  the  Geology  Dept, 
Australian  National  Museum,  Canberra. 

The  silicified  Australian  material  enabled  both  Taylor  and  the  Bedfords  to  discover 
details  of  the  finer  internal  structure  by  etching  their  specimens.  Some  credit  is  due 
to  the  Bedfords  for  the  classification  of  the  Archaeocyatha  and  their  recognition  of  the 
systematic  importance  of  ontogeny  in  such  a  classification. 


FIG.  2.     Section  from  a  generalized  geological  map  of  the  Flinders  Ranges,  produced  by  the 
Geol.  Survey  of  South  Australia  and  published  in  Glaessner,  M.  F.  &  Parkin,  L.  W.  1958. 

The  study  of  Archaeocyatha  has  figured  prominently  in  assigning  a  Cambrian  age  to 
these  rocks.  Daily  (1956)  recognized  5  faunal  assemblages  in  the  Ajax  limestones,  but 
considered  that  the  material  collected  by  both  Taylor  and  the  Bedfords  could  only 
come  from  his  faunal  assemblage  No.  I,  which  is  given  a  Lower  Cambrian  age.  Yet 
Walter  (1967)  states  that  the  Ajax  fauna  cannot  be  placed  in  Daily's  scheme  of  faunal 
units. 

Recently  Dorothy  Hill  (1965),  in  her  work  on  Antarctic  Archaeocyatha,  revised  the 
phylum  and  provided  good  illustrations  of  type  material  in  Adelaide  University,  the 
South  Australian  Museum,  Princeton  University  and  the  British  Museum  (Nat.  Hist.). 
At  her  suggestion,  I  undertook  the  study  of  the  Bedford  material  in  the  latter  collec- 
tion, with  the  intention  of  providing  further  description,  re-classification  and,  above 
all,  better  figures  of  the  specimens.  It  has  often  been  difficult  in  the  past  to  assess  the 
characters  of  genera  and  species  mentioned  by  the  Bedfords,  as  their  figures  are  either 
composite,  or  cannot  be  located  on  the  actual  specimen.  The  genus  Syringocnema  is 
not  dealt  with  in  this  paper,  although  well  represented  in  that  part  of  the  Bedford 
collection  in  the  B.M.  (N.H.),  because  of  its  thorough  description  by  both  Taylor 
(1910  :  153)  and  Gordon  (1920  :  699). 


302  LOWER   CAMBRIAN   ARCH  AEOCY  ATM  A   FROM   AUSTRALIA 

"  Archaeocyatha  are  known  from  all  the  continents  except  South  America  and  are 
characteristic  of  the  calcareous  facies  of  the  Lower  Cambrian  "  (Hill  1965  :  30).  This 
widespread  distribution  makes  them  particularly  useful  for  stratigraphical  correla- 
tion; yet,  as  Walter  (1967)  points  out,  their  use  is  limited  for,  being  benthonic, 
dispersal  is  restricted.  "  Recognition  of  the  extinct  Archaeocyatha  as  a  separate 
phylum  near  the  Porifera  and  Coelenterata,  is  now  general  "  (Hill  1965  :  45).  In  the 
past,  their  systematic  position  has  been  the  subject  of  considerable  debate,  with 
opinion  ranging  from  calcareous  algae  to  sponges.  However,  they  are  regarded  by 
Hill  (1964  :  253)  "  as  single  multicellular  organisms,  with  organization  higher  than 
that  of  the  Protozoa,  but  with  less  differentiation  than  the  Porifera  ". 

A  short  glossary  of  the  more  important  descriptive,  morphological  terms  is  included 
as  an  Appendix. 

II.  SYSTEMATIC  DESCRIPTIONS 

Phylum  ARCHAEOCYATHA  Vologdin,  1937 

Class  REGULARIA  Vologdin,  1937 
Order  MONOCYATHIDA  Okulitch,  1935 
Family  MONOCYATHIDAE  R.  &  W.  R.  Bedford,  1934 
DIAGNOSIS.     Single-walled    cup.     Pore    system    simple    to    slightly    complex. 

COMPOSITION  OF  THE  FAMILY.  Monocyathus  Bedford,  R.  &  W.R.  1934,  Rhabdolyn- 
thus  Zhuravleva  1960,  Tumuliolynthus  Zhuravleva  1963,  ?  Tunkia  Bedford,  R.  &  J. 
1936. 

Genus  MONOCYATHUS  R.  &  W.  R.  Bedford,  1934 

1899  Rhabdocyathus  von  Toll  (non  Brooks  1893) :  45,  pi.  8,  figs.  2c,  6  and  7,  t.-figs.  4-7. 

1910  Archaeolynthus  Taylor  :  157,  pi.  5. 

1934  Monocyathus  R.  &  W.  R.  Bedford  :  2,  pi.  i,  fig.  i. 

1936  Monocyathus  R.  &  W.  R.  Bedford  :    12,  pi.  10,  fig.  46. 

1937  Rhabdocnema  Okulitch  :  251 

1939     Monocyathus  R.  &  W.  R.  Bedford;  R.  &  J  Bedford  :  69,  pi.  42,  fig.  161. 
1939     Archaeolynthus  Taylor;  Simon  :  21. 
1949     Archaeolynthus  Taylor;  Zhuravleva  :  549. 

TYPE  SPECIES.  Monocyathus  porosus  R.  &  W.  R.  Bedford,  1934,  selected  by  R.  & 
W.  R.  Bedford,  1936. 

DISCUSSION.  Okulitch  (1950),  Debrenne  (1964)  and  Hill  (1965)  consider  Archaeo- 
lynthus Taylor  1910  an  invalid  name,  as  the  type  specimen  was  not  designated  by  the 
author  and  the  reference  material  was  destroyed  by  serial  sectioning. 

DIAGNOSIS.  Small  conical  cups  with  a  simple  porous  single  wall  and  the  vertical 
rows  of  pores  in  a  quincunx  pattern. 

COMPOSITION  OF  THE  GENUS.  Monocyathus  absolutus  (Vologdin  1940),  M.  bilateralis 
(Vologdin  1962),  M.  contractus  Hill  1965,  M.  copulatus  (Vologdin  1940),  M.  kuzneskii 
(Vologdin  1931),  M.  lebedevae  (Vologdin  1937),  M.  macrospinosus  (Zhuravleva  1963), 
M.  mellifer  R.  &  W.  R.  Bedford,  1936,  M.  nalivkini  (Vologdin  1939),  M.  operculatus 


LOWER   CAMBRIAN   ARCH  AEOC  Y  ATM  A    FROM   AUSTRALIA  303 

Maslov  1960,  M.  partibus  (Vologdin  1963),  M.  polaris  (Vologdin  1937),  M.  porosus 
R.  &  W.  R.  Bedford,  1934,  M.  robustus  R.  &  W.  R.  Bedford,  1936,  M.  sibiricus  (von 
Toll  1899),  M.  simplex  (Vologdin  1940),  M.  sparsipom  R.  &  W.  R.  Bedford,  1936,  M. 
spinosus  R.  &  W.  R.  Bedford,  1936,  M.  tenuimurus  (Vologdin  1940),  M.  tolli  (Krasno- 
peeva  1937),  M.  unimurus  (Vologdin  1940). 

Monocyathus  porosus  R.  &  W.  R.  Bedford 

(PL  i,  ng.  3) 

1934  Monocyathus  porosus  R.  &  W.  R.  Bedford  :  2,  pi.  i,  fig.  i. 

1939  Monocyathus  porosus  R.  &  J.  Bedford  :  69,  fig.  161. 

1963  Archaeolynthus  porosus  (Bedford  &  Bedford) ;  Zhuravleva  :  88-89,  fig.  39. 

1965  Monocyathus  porosus  R.  &  W.  R.  Bedford;  Hill  :  52,  pi.  2,  fig.  i. 

LECTOTYPE.     B.M.  (N.H.)  S  4140  selected  by  Hill  (1965). 
OTHER  MATERIAL.     Syntypes  B.M.  (N.H.)  S  7630-31,  S  4783-8. 

DESCRIPTION.  Conical  cup,  slightly  waved,  with  a  single  wall  of  constant  thickness 
and  pores  in  quincunx.  The  upper  edge,  in  the  material  studied,  is  not  turned  back 
into  the  central  cavity  as  an  incipient  pelta.  The  pores  are  proportionately  smaller 
and  more  numerous  towards  the  upper  part  of  the  cup.  The  bigger  specimens  have 
smaller  and  more  regular  pores. 

DIMENSIONS 

Lectotype  S  7630-7631 

(mm.)  (mm.) 

Height  (pars)                                                                  15  31 

Diameter                                                                            6  9-5 
Wall: 

Diameter  of  the  pores                                        o  •  27-0  -20  o  •  10 

Distance  between  the  vertical  rows                0-33-0-27  °'33 

Distance  between  the  horizontal  rows              o  •  54-0  -67  o  •  35 

DISCUSSION.  This  species  is  very  similar  to  the  Siberian  species  M.  nalivkini 
(Vologdin).  Zhuravleva  (1963  : 80)  distinguishes  the  two,  mainly  on  the  supposed 
presence  of  a  pelta  and  inner  rugosity  in  porosus,  which  was  described  and  figured  by 
R  &  W.  R.  Bedford  (1936).  However,  these  peculiar  structures  are  neither  seen  in 
the  lectotype,  nor  on  the  other  specimens  in  the  British  Museum  collection.  It  is 
therefore  possible  that  the  two  species  are  synonymous. 

Monocyathus  mellifer  R.  &  W.  R.  Bedford 

(PI.  i,  fig.  4) 

1936     Monocyathus  mellifer  R.  &  W.  R.  Bedford  :  12,  pi.  10,  fig.  49. 
MATERIAL.     B.M.  (N.H.)  S  4821.   The  whereabouts  of  the  holotype  is  not  known. 
REMARKS.     The  specimen  is  a  cylindrical  fragment  of  a  single-walled  cup.     The 
wall  is  like  a  honeycomb,  formed  by  short  hexagonal  pipes  with  a  hexagonal  section 
on  the  outer  side  and  a  circular  section  on  the  inside. 


304         LOWER  CAMBRIAN  ARCH AEOC Y ATH A  FROM  AUSTRALIA 
DIMENSIONS 

(mm.) 

Height  (pars)  16 

Diameter  3  •  72 
Wall: 

Inner  diameter  of  pores  o  •  40 

Pipe  wall  thickness  0-27 

Wall  thickness  0-67 

DISCUSSION.  This  fragmentary  piece  is  similar  to  the  type-species  briefly  described 
by  R.  &  W.  R.  Bedford  1936. 

Genus  TUMULIOLYNTHUS  Zhuravleva,  1963 

1932     Rhabdocyathus  von  Toll  :  Vologdin  (pars)  :  65. 
1963     Tumuliolynthus  Zhuravleva  :  101. 

TYPE  SPECIES.  Rhabdocyathus  tubexternus  Vologdin  1932,  by  original  designation 
Zhuravleva  (1963  :  101). 

DIAGNOSIS.  One-walled  Archaeocyatha,  with  the  pores  protected  externally  by 
simple  tumuli. 

COMPOSITION  OF  THE  GENUS.  (After  Zhuravleva  1963) :  T.  irregularis  (R.  &  W.  R. 
Bedford,  1934),  T.  karakolensis  Zhuravleva  1963,  T.  musatovi  (Zhuravleva  1961), 
T.  tubexternus  (Vologdin  1932),  T.  vologdini  (Yakovlev  1956). 

Tumuliolynthus  irregularis  (R.  &  W.  R.  Bedford) 
(PI.  i,  fig.  i) 

1934     Monocyathus  irregularis  R.  &  W.  R.  Bedford  :  2,  pi.  i,  fig.  2. 

1939     Monocyathus  irregularis  R.  &  W.  R.  Bedford;  R.  &  J.  Bedford  :  68,  fig.  160. 

1963     Tumuliolynthus  irregularis  (R.  &  W.  R.  Bedford)  Zhuravleva  :  no,  fig.  58. 

HOLOTYPE.     B.M.  (N.H.)  S  4141. 

OTHER  MATERIAL.     B.M.  (N.H.)  S  7643-6,  S  4771,  S  4774-7,  S  4764-5. 

REMARKS.  The  holotype  is  a  small  cylindrical  fragment  with  several  large  aper- 
tures, each  of  which  is  probably  the  basal  trace  of  a  pore.  The  scattered  tumuli 
occur  on  the  lower  parts  and  have  a  large  opening  at  the  top.  The  poor  preservation 
and  small  size  of  this  specimen  prevent  a  more  detailed  description.  The  other 
specimens  are  more  complete,  generally  larger,  and  have  a  thicker  wall  with  irregular 
apertures,  each  with  a  tumulus  perforated  at  the  top. 

DIMENSIONS 

S  4141  S  4774-4777          S  4817-4820 

(mm.)  (mm.)  (mm.) 
Cup: 

Height                                                                6-1  18 

Diameter                                                            1-69  0-33  0-27 
Wall: 

Diameter  of  pores                                       o •  60-0 -23  0-20  0-27 

Average  distance  between  pores                      . .  0-67 

Thickness                                                             o-io  0-33  0-27 


LOWER   CAMBRIAN    ARCH  AEOC  Y  ATH  A   FROM   AUSTRALIA  305 

DISCUSSION.  R.  &  W.  R.  Bedford,  (1934)  and  R.  &  J.  Bedford,  (1939)  noted  the 
presence  of  "  papillae  "  and  only  placed  this  species  in  the  genus  Monocyathus 
provisionally.  Zhuravleva  (1963)  uses  the  term  "  tumulus  "  when  referring  to  the 
papillae  and  after  examining  the  material,  I  consider  her  change  of  terminology 
justified. 

Tumuliolynthus  irregularis  differs  from  other  species  of  the  genus  by  the  irregular 
size  and  pattern  of  its  pores. 

Order  AJACICYATHIDA  R.  &  J.  Bedford,  1939 
Family  DOKIDOCYATHIDAE  R.  &  W.  R.  Bedford,  1936 

DIAGNOSIS.  Two-walled  cup,  walls  simply  porous,  connected  by  radial  horizontal 
rods. 

COMPOSITION  OF  THE  FAMILY,  Dokidocyathus  Taylor  1910,  Alphacyathus  R.  &  J. 
Bedford,  1939. 

Genus  ALPHACYATHUS  R.  &  J.  Bedford 
!Q39     Alphacyathus  R.  &  J.  Bedford  :  72. 

TYPE  SPECIES.  Dictyocyathus  annularis  R.  &  W.  R.  Bedford,  1936,  by  original 
designation  R.  &  J.  Bedford,  (1939  :  72,  fig.  55). 

DIAGNOSIS.  Cup  with  two  simply  porous  walls.  In  the  intervallum,  radial 
cylindrical  bars  are  arranged  in  regularly  spaced  horizontal  planes  and  are  connected 
by  synapticulae  that  are  opposite  from  loculus  to  loculus,  forming  a  continuous 
ring  in  the  centre  of  the  intervallum. 

COMPOSITION  OF  THE  GENUS.     A  single  species  Alphacyathus  annularis  (R.  &  W.  R. 
Bedford,  1936). 

Alphacyathus  cf.  annularis  (R.  &  W.  R.  Bedford) 

(PI.  I,  fig.  2) 

cf.  1936     Dictyocyathus  annularis  R.  &  W.  R.  Bedford  :  13,  pi.  u,  fig.  55. 
1939     Alphacyathus  annularis  R.  &  W.  R.  Bedford)  R.  &  J.  Bedford  :  72. 

HOLOTYPE.  P  942  in  the  South  Australian  Museum,  Adelaide  according  to  Hill 
(1965  :  55)- 

OTHER  MATERIAL.     B.M.  (N.H.)     S  4822,  S  4766. 

DESCRIPTION.  Small  cylindrical  cups.  Outer  wall  is  pierced  by  circular  pores  in 
quincunx,  the  skeletal  tissue  between  the  pores  ("  linteaux  ")  is  of  constant  thickness 
and  equal  in  width  to  the  pore  diameter.  Inner  wall  is  built  on  the  same  pattern  with 
regular  pores  and  considerable  skeletal  tissue.  One  inner  wall  pore  at  each  inter- 
radial  space  and  at  each  horizontal  level. 


306         LOWER  CAMBRIAN  ARCH  AEOCY  ATM  A  FROM  AUSTRALIA 
DIMENSIONS 

S  4822  S  4766 

(mm.)  (mm.) 

Cup: 

Height        .......          5  unknown 

Diameter    .......          2  2  •  45 

Intervallum         .          .          .          .          .          .         0-70  0-50 

Central  cavity     .          .          .          .          .          .         0-56  0-90 

Outer  wall: 

No.  of  pores  between  2  bars           .          .          .           2-3  2-3 

Diameter    .          .          .          .          .          .          .         0-07  0-07 

Vertical  partitions        .          .          .          .          .         0-07  0-07 

Horizontal  partitions   .          .          .          .          .         0-07  0-07 

Thickness  .          .          .          .          .          .          .         0-07  0-07 

Inner  wall : 

No.  of  pores  between  2  bars                                         i  i 

Diameter    .          .          .          .          .          .          .          0-15  0-13 

Vertical  partitions        .          .          .          .          .         0-07  0-15 

Horizontal  partitions   .          .          .          .          .         0-07  0-15 

Thickness  .......         0-07  0-15 

Rods: 

Interradial  space           .          .          .          .          .            . .  0-2 

Vertical  space      .          .          .          .          .          .0-2 

Diameter    .          .          .          .          .          .          .         0-15  0-15 

DISCUSSION.     Neither  specimen  provides  a  sufficiently  complete  transverse  section 
for  showing  the  synapticulae  and  they  cannot  be  assigned  with  certainty  to  annular  is. 


Family  ACANTHINOCYATHIDAE  R.  &  W.  R.  Bedford,  1936 

DIAGNOSIS.  Two-walled  cups  with  intervallar  horizontal  rods.  Inner  wall  simple, 
but  outer  wall  has  protected  pores,  the  lower  skeletal  part  of  each  pore  bearing  a  long 
spinous  process,  projecting  upwards  and  outwards  from  the  wall. 

COMPOSITION  OF  THE  FAMILY.     Acanthinocyathus  R.  &  W.  R.  Bedford,  1934. 


Genus  ACANTHINOCYATHUS  R.  &  W.  R.  Bedford,  1936 

1934     Acanthocyathus  R.  &  W.  R.  Bedford  (non  Edwards  &  Haime  1848,  which  is  a  hexacoral)  :  4. 
1936     Acanthinocyathus  R.  &  W.  R.  Bedford  :  n. 

TYPE  SPECIES.  Acanthocyathus  apertus  R.  &  W.  R.  Bedford  (1934  :  4,  fig.  20),  by 
monotypy. 

DIAGNOSIS.  Two-walled  cups  with  radial,  horizontal,  or  upwardly  oblique  rods. 
The  outer  wall  pores  are  partly  obscured  by  scales.  The  inner  wall  consists  of  a  net  of 
polygonal  to  circular  pores. 

COMPOSITION  OF  THE  GENUS.  A.  apertus  (R.  &  W.  R.  Bedford),  ?  A.  transiens 
R.  &  J.  Bedford  1939. 


LOWER   CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA  307 

Acanthinocyathus  apertus  R.  &  W.  R.  Bedford 
(PI.  2,  Text-fig.  3) 

1934     Acanthocyathus  apertus  R.  &  W.  R.  Bedford  :  4,  fig.  20. 

1936     Acanthinocyathus  apertus  (R.  &  W.  R.  Bedford);  R.  &  W.  R.  Bedford  :  n,  fig.  45. 

LECTOTYPE.     B.M.  (N.H.)  S  4166  chosen  here. 

OTHER  MATERIAL.     Para-lectotypes  B.M.  (N.H.)  S  4167-8. 

DESCRIPTION.     Cylindrical  rods  cross  the  intervallum  and  are  orientated  horizont- 
ally, or  sometimes  obliquely  upwards  and  outwards  from  the  inner  to  the  outer  wall. 


FIG.  3.     Acanthinocyathus  apertus  R.  &  J.  Bedford 

The  inner  wall  is  well  shown  in  S  4168 ;  it  consists  of  a  large  pored  net,  with  a  mesh  of 
irregular  size  and  shape  formed  by  skeletal  threads  of  constant  thickness.  The  outer 
wall  is  the  most  characteristic  feature  of  this  species ;  the  large  pores  are  in  quincunx 
and  of  various  shapes ;  the  skeletal  tissue  occupies  a  smaller  area  than  the  pores  and 
carries  long  spines  that  are  directed  upwards  and  outwards.  Unfortunately,  these 
long  protective  spines  are  often  broken,  but  when  complete  they  may  reach  the 
centre  of  the  overlying  pore,  that  is  to  say,  since  the  rows  alternate,  the  centre  of  the 
pore  two  rows  above.  The  base  of  the  spine  is  level  with  the  lower  third  of  the  pore. 


308         LOWER  CAMBRIAN  ARCH AEOCY ATH A  FROM  AUSTRALIA 
DIMENSIONS 

Holotype 

S  4166                       S  4168  S  4167 

(mm.)                        (mm.)  (mm.) 

Cup: 

Height  (pars)  34  32                              25 

Upper  diameter  1 7  •  5  13                              1 1 

Lower  diameter  15  12-5                            8-5 

Upper  intervallum  3-5  3-4                            3 

Lower  inter  vallum  3-4  3-4                            2-37 

Outer  wall: 

Diameter  of  pores  i  -7-2  . .                  i  -oi-i  -35 

scales  4  •  75-6  . .                              3-4 

Vertical  partitions  i  •  35  . .                            0-67 

Horizontal  partitions  1-35  . .                            1-35 

Thickness  0-67  . .                            0-6 

Inner  wall: 

Diameter  of  pores  0-75                   0-2  7-2  -03  0-6-1-15 

Partitions  +  0-61                            0-4 

Thickness  0-40  0-47                           0-4 

Rods  0-4 

DISCUSSION.  R.  &  W.  R.  Bedford,  thought  the  inner  wall  was  a  scaffolding  of  tri- 
radiate  spicules,  the  side  rays  surrounding  the  pores  and  the  median  ray  being  pro- 
longed into  a  spinous  process  directed  upwards  and  outwards.  They  considered  that 
the  spicular  elements  of  Acanthinocyathus  could  indicate  a  phylogenetic  link  between 
Sponges  and  Archaeocyatha.  This  suggestion  is  not  borne  out  by  observation.  The 
wall  of  Acanthinocyathus  is  a  porous  sheet  with  the  elongation  of  its  horizontal  skeletal 
parts  into  scales  of  an  unusual  size. 

Family  AJACICYATHIDAE  R.  &  J.  Bedford,  1939 

Genus  ARCHAEOYCATHELLUS  Ford,  1873 

Subgenus  STAPICYATHUS  Debrenne,  1964 

1873     Archaeocyathellus  Ford  (Pars). 

1964     Archaeocyathellus  (Stapicyathus)  Debrenne  :  127. 

TYPE  SPECIES.  Archaeocyathus  stapipora  Taylor  (1910  :  118),  by  original  designa- 
tion of  Debrenne  (1964  :  127). 

DIAGNOSIS.  Conical  cups;  outer  and  inner  walls  with  simple  pores  as  in  true 
Ajacicyathidae.  Intervallum  crossed  by  imperf orate  radial  septa.  The  neighbour- 
ing loculi  only  open  into  one  another  and  into  the  central  cavity,  by  the  stirrup-pores 
of  the  inner  wall. 

REMARKS.  Debrenne  (1964  :  127)  considered  this  form  a  non-corrugated  subgenus 
of  Archaeocyathellus  Ford. 

COMPOSITION  OF  THE  SUBGENUS.  A.  (S.)  stapipora  (Taylor  1910)  and  ?  A.  (S.) 
yukonensis  (Okulitch  1957). 


LOWER   CAMBRIAN    ARCH AEOC Y ATM A   FROM   AUSTRALIA 


309 


Archaeocyathellus  (Stapicyathus)  stapipora  (Taylor) 

(PI-  i,  %.  5) 

1910  Archaeocyathus  stapipora  Taylor  :  118,  pi.  7,  figs.  37  and  38,  pi.  3,  fig.  10,  t-fig.  14. 

1939  Archaeocyathus  stapipora  Taylor;  R.  &  J.  Bedford  :  75. 

IQ39  Archaeocyathus  (Protocyathus)  stapipora  (Taylor)  Simon  :  54. 

1964  Archaeocyathellus  (Stapicyathus)  stapipora  (Taylor)  Debrenne  :  127. 

1965  Robustocyathus  stapipora  (Taylor)  Hill  :  61  and  68. 

LECTOTYPE.  Taylor  1910,  PI.  7,  fig.  38  G,  chosen  here.  The  specimen  should  be 
in  the  University  of  Adelaide. 

OTHER  MATERIAL.  B.M.  (N.H.)  S  4733,  S  4351-2,  S  4138,  S  4528,  S  4817-20,  S 
7621. 

DESCRIPTION.  Narrow  intervallum  of  constant  width  from  the  basal  conical  apex 
to  the  upper  bowl-shaped  cup.  Some  specimens  are  more  cylindrical.  Outer  wall 
thin,  pierced  by  round  pores  that  are  arranged  quincunxially.  Inner  wall  thicker, 
with  one  vertical  row  of  pores  in  front  of  each  septa ;  each  pore  is  excavated  into  its 
septum  so  that  stirrup-pores  are  formed.  These  inner  wall  pores  are  also  arranged 
into  horizontal  lines. 


DIMENSIONS 


Cup: 

Height  (pars) 

Diameter 

Intervallum  coeff. 

Interseptum 

Loculi 
Outer  wall: 

No  of  pore  rows  per 
interseptum 

Diameter  of  pores 

Vert,  partitions 

Thickness 
Inner  wall : 

No.  of  pore  rows  per 
interseptum 

Diameter  of  pores 

Vertical  part. 

Horizontal  part 

Thickness 
Septa 

Thickness 


§4733 
(mm.) 

30 

about  40 
0-02 
o-339 
1/3-4 


3 

0-07 
0-068 
0-150 


two   1/2 
0-27 
0-27 

0-16 
o-i 

non  porous 
0-06 


S  4351 
(mm.) 

40 

about  20 
0-45-0-66 

0-61 
from  i/i  -8  to  1/2 


3 

0-07 
0-068 
0-23 


two   1/2 
O'27 

o-33 
0-27 
o-i 

non  porous 
0-06 


84352 

(mm.) 

S  4817-4820 
(mm.) 

22 

about  12 

15 
about  35 

o-i 
0-61 

1/2 

0-042 
0-74 

1/2 

3 
0-06 
0-07 

3  to  4 
0-075 
0-068 

0-15 

o-i 

tWO  1/2 

tWO  1/2 

O-2O 

O-27 

o-33 

0-40 

O-2O 

O-27 

O-I 

0-  I 

non  porous 

non  porous 

0-07 

0-07 

S  7621 
(mm.) 

20 

about  10 
o-i 
o-37 
i/3'3 


3 

0-07 
o-i 
o- 1 


tWO  1/2 
0'27 
O-27 
O-2O 
O-I 

non  porous 
0-07 


DISCUSSION.  Only  one  species  known.  The  different  shapes  i.e.  cylindrical,  or 
bowl-shaped  could  conceivably  represent  different  species,  but  their  coefficients  are 
not  sufficiently  different. 


310  LOWER  CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA 

Genus  LOCULICYATHUS  Vologdin,  1931 
Subgenus  LOCULICYATHELLUS  nov. 

TYPE  SPECIES.     Archaeocyathus  floreus  R.  &  W.  R.  Bedford  1934. 

DIAGNOSIS.  Cup  has  the  generic  characters  of  Loculicyathus :  thin  porous  walls  and 
septa,  vesicular  tissue  crossing  intervallum  and  central  cavity.  The  external  longitu- 
dinal corrugations  on  the  outer  wall,  like  those  of  Ajacicyathellus,  subgenus  of 
Ajacicyathm  (see  Debrenne  1964  :  127),  or  Archaeocyathellus  Ford  1873,  distinguish  it 
as  a  subgenus. 

DISCUSSION.  R.  &  W.  R.  Bedford  (1937)  compared  A.  floreus  with  Archaeo- 
cyathellus. They  singled  out  forms  with  radial  imperforate  septa  and  stirrup-pores 
at  the  inner  wall,  from  others  with  regularly  porous  septa.  The  holotype  of  floreus 
(S  4144),  the  paratype  and  other  material  in  the  B.M.  (N.H.)  collection,  show  sparse 
pores  but  no  stirrup-pores. 

The  only  species  recognized  at  present  is  Loculicyathus  (Loculicyathellus}  floreus 

(R.  &  W.  R.  Bedford  1934). 

Loculicyathus  (Loculicyathellus}  floreus  (R.  &  W.  R.  Bedford) 

(PI.  3,  figs,  i,  2,  4) 

1934     Archaeocyathus  floreus  R.  &  W.  R.  Bedford  :  2,  fig.  4. 

1937     Archaeocyathus  floreus  R.  &  W.  R.  Bedford;  R.  &  J.  Bedford  :  35,  figs.  I44A  &  B. 

HOLOTYPE.    B.M.  (N.H.)  S  4144. 

OTHER  MATERIAL.  Paratype  B.M.  (N.H.)  S  4145.  Also  B.M.  (N.H.)  S  4730, 
S  4739  and  S  7635. 

DIAGNOSIS.  Small  cylindrical  cup  with  vertical  corrugation  between  two  neigh- 
bouring septa,  so  that  transverse  sections  recall  scleractinian  corolla.  Outer  wall  with 
horizontal  and  vertical  rows  of  non-alternating  pores.  The  pores  of  the  inner  wall  and 
septa  are  in  quincunx.  A  few  dissepiments  occur  across  the  intervallum  and  central 
cavity. 

DIMENSIONS 

S  4144  S  4145        S  4739 

(mm.)  (mm.)         (mm.) 
Cup: 

Height  (pars)                      9-5  10 

flow    5  •  o  flow    5-0  6-77 

Diameter  <  <  " 

\upp.  6-0  VUPP-  6'5 

Interseptum  o  •  75-2  •  o  o  •  75-2  •  o  0-95 

Interv.  coeff.  0-4  ..  1-6 

Parietal  coeff.  i  -o  . .  i  -6 

Outer  Wall: 

No.  of  pore  rows  per  intersept  6 

Diameter  of  pores  0-13 

Vertical  skel.  part.  0-13 

Horizontal  skel.  part.  °'33 


LOWER  CAMBRIAN  ARCH AEOC Y ATH A  FROM  AUSTRALIA 


DIMENSIONS — continued 


Inner  Wall : 

No.  of  pore  rows  per  intersept 

Diameter  of  pores 

Vertical  skel.  part. 

Horizontal  skel.  part. 
Septa : 

Diameter  of  pores 

Vertical  skel.  part. 

Horizontal  skel.  part. 


S  4144 
(mm.) 

2 
0-23 

o-33 

0-23 


0-23 

o-33 
0-23 


s  4145 

(mm.) 

0-26-0 -37 
o-35 

O-22 
0-30 
0-26 


S  4739 
(mm.) 


DISCUSSION.  The  specimens  from  Ardrossan  figured  by  R.  &  J.  Bedford,  (1939, 
figs.  144  A  &  B)  only  have  one  pore  per  intersept  at  the  inner  wall  and  a  non- 
corrugated  outer  wall.  They  would  therefore,  seem  to  be  a  quite  different  form. 


Family  ROBUSTOCYATHIDAE  Debrenne,  1964 
Genus  ROBUSTOCYATHUS  Zhuravleva,  1960 

TYPE  SPECIES.  Archaeocyathus  robustus  Vologdin  1937,  by  original  designation  of 
Zhuravleva  (1960  :  133). 

DIAGNOSIS.  Solitary  cups  with  simply  porous  outer  wall  and  single  vertical  row  of 
apertures  per  intersept  at  the  inner  wall,  so  that  each  loculus  opens  into  the  central 
cavity. 

COMPOSITION  OF  THE  GENUS.  R.  argentus  (Okulitch  1935),  R.  annulatus  Zhuravleva 
1960,  R.  artecaveatus  (Vologdin  19400),  R.  biohermicus  Zhuravleva  1960,  R.  densus 
Debrenne  1964,  R.  hupei  Debrenne  1964,  R.  levigatus  (Vologdin  19400),  R.  magnipora 
(R.  &  W.  R.  Bedford  1934),  R.  moori  (Vologdin  1937),  R.  novus  Zhuravleva  1960,  R. 
polyseptatus  (Vologdin  19400),  R.  proskurjakowi  (von  Toll  1889),  R.  pseudotichus 
(Vologdin  19400),  R.  salebrosus  (Vologdin  1931),  R.  spinosus  Zhuravleva  1960, 
R.  subacutus  (R.  &  W.  R.  Bedford,  1934),  R.  sucharichensis  Zhuravleva  1960,  R.  ? 
tennis  (Vologdin  1940). 


Robustocyathus  magnipora  (R.  &  W.  R.  Bedford) 
(PI-  3,  fig-  5) 

1934     Archaeocyathus  magnipora  R.  &  W.  R.  Bedford  :  2,  pi.  i,  fig.  7. 

1937     Paranacyathus  magnipora  (R.  &  W.  R.  Bedford)  R.  &  J.  Bedford  :  34. 

HOLOTYPE.     B.M.  (N.H.)  S  4146. 

DESCRIPTION.     The  holotype,  a  partially  destroyed  cup,  was  the  only  specimen 
examined.     The  radial  septa  are  few,  straight  and  pierced  by  three  alternate  vertical 

GEOL.   17,   7  25 


312  LOWER   CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA 

rows  of  circular  pores,  which  are  not  connected  to  the  wall  pores.  The  outer  wall  is  a 
thin  plate  with  circular  pores  in  quincunx.  The  inner  wall  has  a  single  vertical  row  of 
large  pores  per  intersept  and  pores  of  neighbouring  rows  alternate.  The  septa,  which 
spring  out  of  the  inner  wall,  form  boundaries  to  new  pores  by  splitting  the  previous 
one. 

DIMENSIONS 

Cup:  (mm.) 

Height  (pars)  24 

Diameter  8  •  8 

Intervallum  i  •  83 

Interseptum  o  •  74 
Loculus  (trapezoid)                                              °'47  and  0-6  for 

Height  =  0-9 
Parietal  coefficient  +  unknown 

Outer  wall : 

Vertical  rows  of  pores  per  intersept  4 

Diameter  of  pores  o  •  20 

Skeletal  partitions  o  •  13 

Thickness  0-13 

Inner  wall :  i 

Vertical  diameter  o  •  40 

Horizontal  diameter  o  •  60 

Skeletal  partitions  0-33 

Thickness  o  •  33 

Septa : 

Alternating  vertical  rows  of  pores  per  septum  3 

Diameter  0-61 

Skeletal  partitions  0-61 

DISCUSSION.  The  specimens  figured  by  R.  &  J.  Bedford  (1937,  figs  I42A-E)  and 
described  as  magnipora  are  unlike  the  holotype  mentioned  above.  They  differ  by 
having  various  forms  and  patterns  of  outer  wall  and  septal  pores.  These  characters, 
together  with  the  presence  of  an  Archaeopharetra-type  apex,  caused  R.  &  J.  Bedford 
to  place  magnipora  in  the  genus  Paranacyathus.  Unfortunately,  it  was  not  possible 
to  compare  the  holotype  with  these  specimens.  The  holotype  (S  4146)  shows  the 
true  characters  of  Robustocyathus :  regular  walls  and  septal  pores,  which  are  sufficient 
to  place  it  in  the  Class  Regularia,  although  the  initial  stages  are  not  present  in  the 
specimen.  At  the  moment,  it  is  not  possible  to  decide  the  true  systematic  position 
of  the  specimens  figured  by  the  Bedfords  in  1937. 


Robustocyathus  subacutus  (R.  &  W.  R.  Bedford) 
(PL  3,  figs.  6,  7,  Text-fig.  4) 

1934     Archaeocyathus  subacutus  R.  &  W.  R.  Bedford  :  2,  pi.  i,  figs.  3a-c. 
1937     Archaeocyathus  acutus  Bornemann;  Ting  :  358,  pi.  9,  figs.  1-2. 
1961     Ajacicyathus  walliseri  F.  &  M.  Debrenne  :  696,  pi.  19,  fig.  3. 


LOWER   CAMBRIAN    ARCH  AEOC  Y  ATM  A    FROM    AUSTRALIA  313 

HOLOTYPE.     B.M.  (N.H.)  S  4142. 

OTHER  MATERIAL.     B.M.  (N.H.)  S  4143,  S  4747,  S  4792,  S  7620. 

DESCRIPTION.  Small  cylindrico-conical  cup  with  thin  skeletal  plates  and  few  pores. 
The  outer  wall  only  has  one  pore  in  the  middle  of  each  intersept;  these  pores  are 
arranged  in  widely  separated  vertical  rows  and  more  closely  set  horizontal  ones.  The 
inner  wall  is  thin  and  perforated  by  a  single  row  of  pores  per  intersept.  The  aperture 


FIG.  4.     Robustocyathus  subacutus  (R.  &  W.  R.  Bedford) 


of  each  pore  is  larger  than  the  width  of  the  interseptum.  The  inner  edges  of  the  septa, 
which  laterally  delineate  the  pore,  have  vertical  undulations  (see  Text-fig.  4). 
Numerous  radial  septa,  having  a  few  small  isolated  pores,  are  found  near  the  outer 
wall,  but  they  have  only  been  seen  in  specimen  S  4747. 

The  undulations  on  the  inner  edges  of  the  septa,  close  to  the  inner  wall,  are  different 
from  those  seen  in  Ethmophyllum  Meek.  The  amplitude  of  the  waves  is  smaller,  while 
the  sinuous  edges  do  not  join  together  to  close  up  the  interseptum  and  form  another 
wall  that  connects  the  septa,  but  simply  limit  the  inner  wall  pores.  The  inner  wall, 
itself,  remains  as  a  completely  separate  sheet. 


3H     LOWER  CAMBRIAN  ARCH AEOCY ATH A  FROM  AUSTRALIA 

DIMENSIONS 

S  4142  S  4143  S  4747  S  7620  walliseri 

(mm.)  (mm.)  (mm.)  (mm.)  (mm.) 

Cup: 

Height  (pars)  30  20  25  +  + 

Diameter  10-12  +  8-10  7  13 

Intervallum  coeff.                                    0-3  0-39  0-5  .  .  0-23 

Parietal  coeff.                                         5-4  +  5-5  5-5  5-2 

Interseptum                                            0-30  °'33  0-20  0-25 

Loculus                                                     1/6  1/8  1/7  . .  1/4 

Outer  wall: 

No.  of  pore  rows  per  intersept            i  +  +  i  i 

Diameter                                                 0-13  ..  ..  0-7  0-13 

Skeletal  vert.  part.                                0-40  ..  ..  0-40  + 

Skeletal  horiz.  part.                               0-13  ..  ..  0-13  o-n 

Thickness                                                 0-06  . .  0-06  0-07  0-07 

Inner  wall : 

No.  of  pore  rows  per  intersept            i  i  i  i  i 

Diameter                                                 0-33  0-40  0-27  0-27  0-26 

Skeletal  vert.  part.                                0-13  0-13  +  0-13  0-13 

Skeletal  horiz.  part.                               0-13  0-13  +  0-13  + 

Thickness                                                 0-33  0-40  0-33  0-27  0-26 

Septa 

Thickness                                                 o-io  o-io  +  o-io  0-06 

DISCUSSION.  R.  walliseri  Debrenne,  in  spite  of  its  very  large  size,  seems  to  fall 
between  the  limits  of  specimens  of  subacutus.  Another  species,  R.  pseudotichus 
(Vologdin)  has  similar  undulations  on  the  inner  parts  of  the  septa  that  border  the 
large  inner  wall  pores,  but  in  addition,  has  long  spines  on  the  vertical  edges  of  the 
pores. 

Genus  ZONACYATHUS  R.  &  J.  Bedford,  1937 

1937     Zonacyathus  R.  &  J.  Bedford  :  36. 

I94oa  Ethmophyllum  Meek;  Vologdin  (pars)  :  66-68. 

TYPE  SPECIES.  Archaeocyathus  retevallum  R.  &  W.  R.  Bedford  (1934  :  2,  fig.  6),  by 
monotypy. 

DIAGNOSIS.  Porous  two-walled  cups  with  no,  or  few,  porous  radial  septa.  The 
outer  wall  has  regular  pores.  The  inner  wall  has  branching  pore-tubes;  the  initial 
tube  is  located  in  the  middle  of  each  interseptum  and  then  branches  so  that  the 
secondary  tubes  open  in  front  of  the  septa.  The  tubes  may  lengthen  and  curve  into 
the  central  cavity.  The  pore-tubes  are  never  formed  by  septal  fluting. 

DISCUSSION.  The  type  species  retevallum  is  known  from  only  the  three  specimens 
in  the  B.M.  (N.H.)  collection.  The  skeletal  tissue  is  extremely  thin  and  breakable, 
with  the  result  that  structures  are  either  crushed  or  destroyed,  making  it  difficult  to 
distinguish  the  pore  systems.  The  validity  of  a  genus  based  on  such  fragments  may 
be  questionable.  Nevertheless,  from  studying  a  very  close  but  simpler  species,  Z. 
retezona  Taylor,  which  has  an  inner  wall  built  of  shorter  pore-tubes  that  branch  in  the 
same  way,  it  is  possible  to  understand  the  more  complex  pattern  present  in  retevallum. 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A   FROM   AUSTRALIA  315 

COMPOSITION  OF  THE  GENUS.  Zonacyathus  retevallum  (R.  &  W.  R.  Bedford  1934), 
Z.  retezona  (Taylor  1910).  The  species  Ethmophyllum  poletevae  Vologdin  19400;,  E. 
vermiculatum  Vologdin  1938  and  E.flexum  Vologdin  1940^,  are  placed  in  Zonacyathus 
with  reservation. 

Zonacyathus  retevallum  (R.  &  W.  R.  Bedford) 
(PL  4,  figs.  4,  5) 

1934     Archaeocyathus  retevallum  R.  &  W.  R.  Bedford  :  2,  fig.  6. 

1937     Zonacyathus  retevallum  (R.  &  W.  R.  Bedford,)  R.  &  J.  Bedford  :  36,  figs.  I53A-E. 

1965     Zonacyathus  retevallum  (R.  &  W.  R.  Bedford):  Hill  :  76,  pi.  4,  figs.  3a-b. 

HOLOTYPE.     B.M.  (N.H.)  S  4147. 

OTHER  MATERIAL.     B.M.  (N.H.)  S  4726-4727. 

DESCRIPTION.  Bowl-shaped  cup  with  narrow  waved  intervallum,  easily  distorted 
but  without  giving  any  obvious  folds.  Owing  to  the  bad  preservation  of  the  material, 
all  measurements  are  approximate  and  observations  incomplete. 

Outer  wall  simple  with  two  alternating  rows  of  oval  pores,  arranged  in  quincunx,  to 
each  interseptum.  The  septa  are  difficult  to  see  in  longitudinal  section  in  the  Bed- 
ford specimens.  The  septa  appear  to  be  imperf orate,  but  may  have  small  scattered 
pores.  The  inner  wall  has  branching  pore-tubes  of  Y-form,  which  lengthen  and  curve 
into  the  central  cavity.  It  is  suspected  that  some  lateral  communication  occurs 
between  the  pore-tubes  that  penetrate  into  the  central  cavity. 

DIMENSIONS 

S  4147  :  i  S  4147  :  2  S  4726-4727 

(mm.)  (mm.)                     (mm.) 

Cup: 

Height  25  40 

Diameter  20  12-30                     10 

Intervallum  coefficient  o  •  1 1  0-12                       0-2 

Parietal  coefficient                                                 .  .  6-6 

Interseptum  0-27  0-27                       0-27 

Loculus  J/7'2  J/4'8                      I/5'5 

Outer  wall: 

No.  of  rows  of  pores  per  intersept  2  2 

Diameter  of  pores                                                ..  0-13X0-06 

Vertical  partitions                                               ..  o-io 

Horizontal  partitions                                          ..  o-io 

Thickness  o-io  0-13                       o-io 

Inner  wall : 

No.  of  rows  of  pores  per  intersept  i  i                             i 

Diameter  of  pores  0-20  0-27                        0-20 

Vertical  partitions  o-io  0-13                        0-13 

Horizontal  partitions  o-io  0-13                       0-13 

Thickness                                                      at  least  0-6  0-8  0-6 

Septa : 

Thickness  o  •  i  o  o  •  i  o 


3i6 


LOWER   CAMBRIAN    A RCH AEOC Y ATM A    FROM   AUSTRALIA 


DISCUSSION.  Ethmophyllum  flexum  Vologdin  (19400  :  66,  pi.  15,  fig.  4)  is  very 
similar  to  Z.  retevallum  R.  &  W.  R.,  Bedford,  having  the  same  irregular  shape  and 
size. 

Zonacyathus  retezona  (Taylor) 

(PI.  4,  figs,  i,  2,  3,  Text-fig.  5) 
1910     Archaeocyathus  retezona  (Taylor)  :  121,  pi.  7,  fig.  38A,  pi.  6,  fig.  3iE. 

MATERIAL.  B.M.  (N.H.)  S  4353,  S  4764,  S  4774,  S  4778,  S  4803-4808,  S  4811-4816, 
S  7623,  S  7634,  S  4731. 

DESCRIPTION.  Conical,  externally  smooth  cups.  The  outer  wall  is  a  porous  sheet, 
with  vertical  alternating  rows  of  slightly  elliptical  pores  surrounded  by  skeletal  tissue 


FIG.  5.     Zonacyathus  retezona  (Taylor) 

of  constant  width.  Radial  septa  are,  probably,  imperforate.  Inner  wall  independent 
from  septa  and  has  two  vertical  rows  of  pores,  one  row  opposite  each  septa  and  the 
other  in  the  middle  of  each  interseptum.  Each  pore  opposite  a  septum,  is  the  opening 
for  two  oblique  pore-tubes,  one  coming  from  each  loculus.  On  the  inner  face  of  the 
intervallum,  a  central,  radial  pore-tube  is  joined  by  a  branch  from  each  of  its  neigh- 
bours to  form  one  elliptical  pore.  All  of  these  pore-tubes  are  horizontal.  Z.  retezona 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A    FROM   AUSTRALIA 


317 


differs  from  Z.  retevallum  because  its  tubes  are  short  and  do  not  lengthen,  or  curve  into 
the  central  cavity. 

DIMENSIONS 

S  4811-  S  4730- 


S  4774- 


Cup: 

Height 

Diameter 

Intervallum  coeff. 

Loculus 

Parietal  coeff. 
Outer  wall : 

No.  or  rows  of  pores 
per  intersept 

Diameter  of  pores 

Vertical  partitions 

Horizontal  partitions 

Thickness 
Inner  wall : 

No.  of  rows  of  pores 
per  intersept 

Diameter  of  pores      ( 

Vertical  partitions 

Horizontal  partitions 

Thickness 
Septa : 

No.  of  pore  rows 

Diameter  of  pores 

Partitions 

Thickness 


S4353 

(mm.) 

20 
10 
O-2I 

3'9 


4 

0-06 

0-07 

0-06 

0-13 


87623 
(mm.) 


8 

O-22 

4-2 


0-13 


4816 

(mm.) 

45 
15 

O-I2 


3 

o-io 

0-07 

0-06 

0-13 


4732 

(mm.) 

42 
27 
0-9 


4 

0-06 

0-06 

0-07 

0-13 


S  4764 
(mm.) 

4777 
(mm.) 

S7634 
(mm.) 

84778 
(mm.) 

20 

pars  15 

.  . 

13 

5-8 

9 

10 

10 

1/2 

4-8  to  3*9 

0-25 

O-2I 

3'4 

O-2I 

4-6 

0-07 

0-06 

0-06 

o-io 


I 

i 

i 

i 

I 

20-0-27 

0-27 

0-13-0  -2 

0-27 

0-27 

0-16 

0-13 

O-IO 

0-13 

0-13 

0-13 

0-13 

O-2O 

O-20 

0-40 

0-40 

o-35 

0-40 

0-40 

4-6         4-6 


0-13 


I 

0-27 

0-16 


0-54 


4 

0-07 

0-05 


I 

O-2O 

0-13 
0-40 


0-07 


O-IO 


2 

0-13 
0-13 
O-IO 


0-13 


O-IO 


0-06       0-07 


DISCUSSION.  Taylor  (1910  :  122)  gave  the  following  interpretation  of  the  inner 
wall  of  Z.  retezona:  "  a  simply  porous  inner  wall,  with  one  or  two  pores  between  each 
pair  of  adjacent  septa  ".  In  fact,  if  one  refers  to  his  fig.  27,  it  is  seen  that  one  pore 
occurs  in  each  septum  and  the  other  in  the  middle  of  the  interseptum.  I  consider 
that  the  regular  inner  wall  is  composed  of  pore  tubes,  that  join  together  and  open 
into  the  central  cavity.  Taylor  thought  the  inner  parts  of  the  septa  were  modified 
into  curved  rods  which  supported  the  simple  inner  wall.  However,  the  excellent 
silicified  material  in  the  Bedford  collection  enables  the  path  of  the  canals  to  be 
followed  throughout  their  length. 

Family  BRONCHOCYATHIDAE  R.  &  J.  Bedford,  1936 

1936  Bronchocyathidae  R.  &  J.  Bedford  :  25 

1937  Stillicidocyathidae  Ting  :  367. 
1951  Thalamocyathidae  Zhuravleva  :  98. 
1955  Ethmophyllidae  Okulitch  :  E  12. 
J959  Cyclocyathellidae  Zhuravleva  :  426. 

1965     Bronchocyathidae  R.  &  J.  Bedford;  Hill  :  93. 


TYPE  GENUS. 
1936). 


Thalamocyathus  Gordon  1920  (=Bronchocyathus  R.  &  J.  Bedford, 


3i8  LOWER   CAMBRIAN    ARCH AEOCY ATH A   FROM   AUSTRALIA 

COMPOSITION  of  GENUS.  Thalamocyathus  Gordon  1920,  Stillicidocyathus  Ting  1937. 
Polystillicidocyathus  Debrenne  1959,  Cyathocricus  gen.  nov. 

REMARKS.  Thalamocyathidae  Zhuravleva  1951  has  not  won  general  acceptance 
under  Article  400  of  the  International  Code  of  Zoological  Nomenclature,  1961  and 
while  this  rule  stands  in  its  present  form,  this  family  name  must  be  rejected. 

Genus  CYATHOCRICUS  nov. 

TYPE  SPECIES.     Archaeocyathus  tracheodentatus  R.  &  W.  R.  Bedford,  1934. 
DERIVATION.     From  KPIKOS,  cricos  =  a  ring. 

DIAGNOSIS.  Cup  with  a  simply  porous  outer  wall;  straight,  sparsely  perforated 
radial  septa  and  an  annulate  inner  wall.  Annuli  consist  of  undulating,  horizontal,  or 
slightly  inclined  plates  that  are  neither  S-,  nor  V-shaped ;  their  axial  rim  is  cogged. 

COMPOSITION  OF  THE  GENUS.  C.  tracheodentatus  (R.  &  W.  R.  Bedford  1934),  C. 
dentatus  (Taylor  1910)  and  C.  annulispinosus  (Vologdin  1931). 

DISCUSSION.  Cyathocricus  differs  from  other  annulate  genera  in  that  the  rings 
appear  to  be  straight  in  vertical  section.  The  ft  component,  closest  to  the  septa,  is 
generally  horizontal  and  the  a  component,  projecting  into  the  central  cavity,  is 
complicated  by  cogs  that  are  more  or  less  fused  together. 

This  new  genus  is  established  for  those  species  incorrectly  referred  to  the  genus 
Bronchocyathus  R.  &  J.  Bedford  1936.  Hill  (1965  :  94)  gives  an  account  of  this 
situation,  involving  Thalamocyathus  and  Bronchocyathus. 

Gordon  (1920  :  687)  created  the  genus  Thalamocyathus  and  included  the  species 
Archaeocyathus  tubavallum  Taylor,  A.  trachealis  Taylor,  A.  infundibulum  Bornemann, 
A .  ichnusae  Meneghini  and  T.  flexuosus  Gordon,  but  did  not  designate  a  type  species. 

R.  &  J.  Bedford  (1936  :  25)  erected  Bronchocyathus,  designating  B.  trachealis 
(Taylor)  as  the  type  species  and  including  Ethmophyllum  dentatum  Taylor  and 
Archaeocyathus  tracheodentatus  R.  &  W.  R.  Bedford. 

The  type  species  of  Thalamocyathus  Gordon  was  subsequently  designated  by  Ting 
(J937  :  368)  as  T.  trachealis  (Taylor),  after  elimination  of  the  other  syntype  species 
because  they  possessed  cribrose  inner  walls.  Hill  (1965  :  94)  states  that  she  con- 
siders this  to  be  a  valid  designation.  Simon  (30th  Dec.  1939  :  40)  later  named  A. 
trachealis  as  the  type  species  of  the  genus,  in  case  of  any  doubts  that  Ting  had  done  so. 

Thus  Bronchocyathus  R.  &  J.  Bedford,  is  a  junior  objective  synonym  of  Thalamo- 
cyathus Gordon  since  both  have  the  same  type  species  (Int.  Code  Nomen.,  Article  6ib). 

R.  &  J.  Bedford  (1939  :  75)  pointed  out  that  A.  trachealis  Taylor  belonged  to 
Cyclocyathus  Vologdin  1931,  a  genus  of  which  they  were  unaware  in  1936.  As 
Vologdin  had  not  designated  a  type  species,  they  cited  trachealis,  but  this  is  not 
acceptable  since  the  species  was  not  listed  in  the  original  description  of  Cyclocyathus, 
although  the  Bedfords  considered  that  Vologdin's  use  of  tubavallum  Taylor  was  a 
lapsus  calami  for  trachealis  Taylor.  In  referring  to  tubavallum,  the  Bedfords 
mentioned  that  it  was  the  first  species  listed  by  Gordon  in  his  description  of  Thalamo- 
cyathus and  was  founded  on  a  single  fragment  of  which  conflicting  figures  were  pro- 
vided, and  that  the  whereabouts  of  the  specimen  is  uncertain.  Finally,  R.  &  J. 


LOWER   CAMBRIAN   ARCH AEOCY ATH A    FROM   AUSTRALIA  319 

Bedford  (1939  :  75)  decided  to  apply  the  name  Bronchocyathus  to  those  forms  having 
a  more  complex  inner  wall  than  Cyclocyathus  and  gave  Ethmophyllum  dentatum 
Taylor  as  the  type  species.  This  does  not  conform  to  the  rules  of  zoological  nomen- 
clature, for  Article  61  states  "the  type  of  any  taxon,  once  fixed  in  conformity  with  the 
provisions  of  the  Code,  is  not  subject  to  any  change  except  by  exercise  of  the  plenary 
powers  of  the  Commission  ".  It  is,  therefore,  necessary  to  provide  another  name  for 
the  genus  containing  dentatum  Taylor.  Archaeocyathus  tracheodentatus  R.  &  J. 
Bedford,  is  selected  as  the  type  species  of  the  new  genus  Cyathocricus,  as  it  is  more 
thoroughly  known  than  dentatum. 

Hill  (1965  :  94)  mentions  other  nomenclatural  problems  of  Thalamocyathus  and  the 
species  trachealis  Taylor.  Zhuravleva  (1960  :  220)  cites  Bronchocyathus  as  the  genus 
of  the  family  Bronchocyathidae,  distinguishing  this  group  from  the  Stillicidocyathidae 
by  the  presence  of  inter-communicating  pore-tubes.  However,  this  was  based  on  the 
use  of  E.  dentatum  Taylor  as  type-species  of  Bronchocyathus. 

Cyathocricus  tracheodentatus  (R.  &  W.  R.  Bedford) 
(PL  5,  figs.  4,  5,  PL  6,  fig.  4,  Text-fig.  6) 

1934     Archaeocyathus  tracheodentatus  R.  &  W.  R.  Bedford  :  2,  fig.  5. 

1936     Bronchocyathus  tracheodentatus  (R.  &  W.  R.  Bedford)  R.  &  J.  Bedford  :  25,  fig.  104. 

HOLOTYPE.     B.M.  (N.H.)  S  4148. 
OTHER  MATERIAL.     B.M.  (N.H.)  S  4754. 

DESCRIPTION.  The  type  specimen  is  a  conical  cup  with  a  broken  apex.  Corruga- 
tions on  the  upper  part  provide  good  tangential  and  longitudinal  sections  of  the  inner 
wall  (see  Text-fig.  6) .  The  numerous  radial  septa  are  straight  and  are  perforated  by  a 
few  vertical  rows  of  fine  pores.  Part  of  the  outer  wall  is  preserved,  showing  the  some- 
what irregularly  placed  polygonal  pores.  The  inner  wall  consists  of  a  vertical  series 
of  thick,  horizontal,  ring-shaped  plates,  which  are  triangular  in  cross-section  and  are 
joined  to  the  inner  edges  of  the  septa  by  the  /?  component.  The  annuli  thin  toward 
the  central  cavity,  are  slightly  wavy  on  their  upper  face  and  their  free  rim  (a  com- 
ponent) is  regularly  cogged. 

DIMENSIONS 

S  4148  (Upper)         S  4148  (Lower)         S  4754-4755 

(mm.)  (mm.)  (mm.) 

Cup: 

Height  (pars).  31  31  5 

Upper  Diameter  1 1  6-5  7 

Lower  Diameter  0-2  0-3  0-3 

Parietal  coefficient  . .  8-3 

Interseptum  0-27  0-13  0-13 

Loculus  0-27/1-6  0-13/1-29 

Outer  wall : 

No.  of  pore  rows  per  intersept  3-4  .  .  2 

Diameter  0-06  0-06  0-05 

Vertical  partitions  0-05  0-05  0-05 

Horizontal  partitions  0-05  0-05  0-05 

Thickness  o-io  o-io  0-13 


32° 


LOWER   CAMBRIAN    ARCH AEOC Y ATM A    FROM   AUSTRALIA 


DIMENSIONS — continued 


Inner  wall: 

Space  between  rings 

Thickness  of  ring 

Length  of  cog 

Width  of  ring 

Space  between  cogs 
Septa : 

No.  of  pores 

Thickness 


S  4148  (Upper)         S  4148  (Lower) 


(mm.) 


0-27 

0-13-0-20 
0-67 
o-33 


Imperforate 


0-06 


4754-4755 
(mm.) 

O-2O 
O-IO 
0-67 
O-27 


Imperforate 
0-05 


DISCUSSION.  C.  annulispinosus  (Vologdin  1931)  is  similar  morphologically,  but  its 
specific  coefficients  are  not  well  known.  According  to  Vologdin  (1931  :  fig.  42)  the 
diameter  is  approximately  5-6  mm,  the  intervallum  coefficient  would  be  0-4  and  the 
parietal  coefficient  12 — figures  that  are  quite  close  to  those  of  specimen  S  4148.  The 
essential  difference  is  in  the  downward  orientation  of  the  annuli  into  the  central 
cavity  present  in  annulispinosus. 


FIG.  6.     Cyathocricus  tracheodentatus  (R.  &  W.  R.  Bedford) 


LOWER   CAMBRIAN    ARCH  AEOC  Y  ATH  A    FROM   AUSTRALIA  321 

Cyathocricus  dentatus  (Taylor) 
(PI.  6,  figs.  1-3,  Text-fig.  7) 

1910     Ethmophyllum  dentatum  Taylor  :  129,  pi.  10,  fig.  59;  pi.  13,  fig.  76;  pi.  16,  fig.  89;  t.-figs. 

23  and  31. 

1936     Bronchocyathus  dentatus  (Taylor)  R.  &  J.  Bedford  :  25,  fig.  103. 
1960     Bronchocyathus  dentatus  (Taylor);  F.  &  M.  Debrenne  :  703,  pi.  20,  fig.  8. 

HOLOTYPE.     Not  designated. 

OTHER  MATERIAL.    B.M.  (N.H.)  S  4752-4753,  S  4756,  S  4355. 

DESCRIPTION.  Cylindrical-conical  cup  that  may  have  a  large  diameter  and  a 
wide  central  cavity.  The  intervallum  is  full  of  straight  radial  septa,  that  have  about 
2-3  vertical  rows  of  small  pores  near  the  outer  wall.  The  outer  wall  is  perforated  by 
pores  of  the  same  size  as  those  of  the  septa,  with  2-3  alternate  vertical  rows  per 
interseptum.  There  are  no  stirrup-pores  connecting  the  two  systems.  The  inner  wall 
is  formed  by  a  series  of  vertical  annular  plates,  which  are  as  thick  as  the  septa.  These 
plates  extend  into  the  inner  part  of  the  intervallum,  as  well  as  into  the  central  cavity. 


FIG.  7.     Cyathocricus  dentatus  (Taylor) 


322  LOWER   CAMBRIAN    ARCH  A  EOC  Y  ATM  A   FROM   AUSTRALIA 

Their  free  edge  is  serrated  into  toothlike  projections,  that  are  2-3  loculi  wide  at  the 
base,  and  which  are  situated  either  in  front  of  a  septum,  or  in  the  centre  of  an  inter- 
septum  (PI.  6,  fig.  2).  The  tips  of  these  projections  are  flat  on  top  and  bend  slightly 
upwards.  When  examining  the  material,  layers  of  the  teeth  were  prised  apart  without 
discovering  any  order  of  appearance.  On  the  lower  part  of  each  tooth  there  is  a  curved 
support  which  joins  it  to  the  underlying  ring  plate. 

DIMENSIONS 


S4355 

(mm.) 


S4753 

(mm.) 


12 

o-3 

0-13 

2 

O-O6 

0-05 

0-05 

O-O6 


30 
18 
0-15 

O-2O 

2-3 

O-O6 
0-05 
O-O5 
O-O6 


S4756 

(mm.) 

53 
15-16 


S  4752 

(mm.) 


9 
0-13 


not  seen 


0-06 


0-13 


Cup: 

Height 

Diameter 

Intervallum  coefficient 

Interseptum 
Outer  wall: 

No.  of  pore  rows  per  intersept 

Diameter 

Vertical  partitions 

Horizontal  partitions 

Thickness 
Inner  wall: 

Distance  between  annuli 

Thickness  of  annuli 

Width  of  annuli 

Projections 
Septa : 

Thickness 
Diameter 
Vertical  partitions 
Horizontal  partitions 

DISCUSSION.  The  interpretation  of  the  inner  wall  structure  of  dentatus  is  difficult 
owing  to  the  limited  number  of  known  specimens.  In  common  with  other  forms,  the 
term  "  vesicular  "  has  been  used  in  the  past  to  describe  the  various  poor  oblique 
sections  of  specimens,  which  have  a  complex  inner  wall  structure.  It  was  not 
possible  to  examine  the  holotype,  but  there  are  specimens  of  dentatus  in  the  Bedford 
collection,  wrongly  called  tracheodentatus,  which  provided  various  views  of  this 
structure.  From  these  specimens  (S  4355,  S  4753  and  S  4752)  it  was  possible  for  me 
to  reconstruct  the  inner  wall  of  dentatus  with  some  certainty.  This  species  differs 
from  tracheodentatus  and  annulispinosus  by  having  thinner  annuli,  vertical  supports 
between  the  annuli,  and  the  triangular  shape  and  different  distribution  of  the  teeth. 


0-20 
O-O6 

0-27 
0-06 

0-20 
0-06 

O-2O 

o-  10 

0-40 
0-40 

Imperforate 

o-33 
0-67 
Perforate 
on  outer  side 

0-27 
0-27 
Imperforate 

o-33 
0-67 

0-06 

0-06 
0-06 

0-05 

0-06 

0-27 

Family  ETHMOCYATHIDAE  nov. 

DIAGNOSIS.  Simple  outer  wall;  uniserially  pored  inner  wall  covered  over  on  the 
innermost  (central  cavity)  side  with  a  secondary  wall  of  annular  plates.  This  feature 
is  characteristic  of  the  new  family.  Ethmocyathus  R.  &  W.  R.  Bedford  is  the  type 
genus. 


LOWER  CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA  323 

DISCUSSION.  It  is  necessary  to  create  a  separate  family  within  the  Ajacicyathacea 
to  place  the  genus  Ethmocyathus.  There  is  no  justification  for  placing  Ethmocyathus 
in  the  family  Ethmophyllidae  (Hill  1965  :  76;  Zhuravleva  1960  :  162),  for  its  inner 
wall  is  not  formed  by  horizontal  fluting  of  the  inner  edges  of  the  septa,  as  they 
suggested.  Neither  is  the  genus  a  doubtful  member  of  the  Tumulocyathidae 
(Debrenne  1964  :  113),  since  the  outer  wall  has  simple  pores  and  its  annuli  have  a 
peculiar  form. 


Genus  ETHMOCYATHUS  R.  &  W.  R.  Bedford,  1934 
1934     Ethmocyathus  R.  &  W.  R.  Bedford  :  2,  fig.  8. 

TYPE  SPECIES.     Ethmocyathus  lineatus  R.  &  W.  R.  Bedford,  by  monotypy. 

DESCRIPTION.  Cup  with  straight  radial,  sparsely  porous  septa.  The  outer  wall 
has  close,  simple  pores.  The  inner  wall  is  composed  of  a  thin  sheet  of  hexagonal  pores 
screened  from  the  central  cavity  by  thin,  narrow  horizontal  annuli. 

DISCUSSION.  Sometimes,  both  of  the  horizontal  sides  of  the  inner  wall  hexagonal 
pores  are  reduced  so  that  the  openings  become  rhombic.  The  inner  edges  of  the 
septa  are  sinuous,  following  the  outlines  of  the  inner  wall  pores,  but  do  not  contribute 
to  their  formation  by  horizontal  fluting. 

COMPOSITION  OF  THE  GENUS.     Ethmocyathus  lineatus  R.  &  W.  R.  Bedford. 


Ethmocyathus  lineatus  R.  &  W.  R.  Bedford 
(PI.  5,  figs.,  1-3,  Text-fig.  8) 

1934     Ethmocyathus  lineatus  R.  &  W  .R.  Bedford  :  2,  fig.  8. 

1965     Ethmocyathus  lineatus  R.  &  W.  R.  Bedford;  Hill  :  76,  pi.  4,  figs.  2a-b,  t.-fig.  16.  2. 

HOLOTYPE.     B.M.  (N.H.)  S  4149. 

DESCRIPTION.  This  is  based  on  a  single  specimen  consisting  of  a  fragment  from  a, 
probably,  cylindrical  cup.  Radial  septa  with  sparse  pores  on  the  outer  two-thirds  of 
each  septum.  The  annular  plates  join  the  inner  wall  net  of  hexagonal  pores,  without 
any  visible  thickening.  The  outer  wall  is  simple  with  regular  oval  pores. 

DIMENSIONS 

Cup:  (mm.) 

Height  (pars).  15 

Diameter  (approx.)  13 

Interseptum  o  •  1 8 

Intervallum  coefficient  0-13 

Outer  wall :  (only  seen  between  2  neighbouring  septa) 

3  pores  lengthened  horizontally  and  arranged  in  quincunx 

Diameter  o  •  05-0  •  03 

Skeletal  partitions  o  •  03 

Thickness  o  •  07 


LOWER  CAMBRIAN  ARCHAEOCYATHA  FROM  AUSTRALIA 


DIMENSIONS  —  continued 


Inner  wall: 

Vertical  diameter 

Horizontal  diameter 

Skeletal  partitions 

Thickness 

Thickness  of  annuli 

Distance  between  annuli 

5  annular  plates  cover  one  pore  (vertically) 
Septa: 

Imperf orate  for  0-37  mm.  from  the  inner  wall 

Diameter  of  pores 

Vertical  partitions 

Horizontal  partitions 

DISCUSSION.     Only  this  one  species  known. 


(mm.) 
0-27 

0-2 
0-075 

o-37 
0-03 
0-03 


0-03 

0-15 
0-18 


FIG.  8.     Ethmocyathus  lineatus  R.  &  W.  R.  Bedford 


LOWER   CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA  325 

Family  ERISMACOSCINIDAE  Debrenne,  1964 

DIAGNOSIS.  Cup  with  radial  septa  and  porous  tabulae.  Thin,  simply  porous 
outer  wall.  Inner  wall  with  several  vertical  series  of  pores  to  each  interseptum, 
sometimes  with  thickening  of  the  pore  framework  ("  linteaux  ")  and  ornamental 
spines. 

COMPOSITION  OF  THE  FAMILY.  Erismacoscinus  (Pluralicoscinus)  Debrenne  1963, 
Erismacoscinus  Debrenne  1958,  Asterocyathus  Vologdin  1940,  Retecoscinus  Zhurav- 
leva  1960,  Tuvacyathus  Vologdin  1940,  Geniculicyathus  Debrenne  1960,  Coscinoteichus 
Debrenne  1964,  Coscinoptycta  Broili  1915. 


Genus  ERISMACOSCINUS  Debrenne,  1958 

TYPE  SPECIES.  Erismacoscinus  marocanus  Debrenne  1958  :  65,  pi.  3,  figs.  11-16, 
by  monotypy. 

DIAGNOSIS.  Intervallum  with  regularly  porous  septa  and  tabulae.  Simple  outer 
wall,  generally  perforated  by  numerous  openings  in  quincunx.  Thick  inner  wall, 
always  has  several  vertical  series  of  pores  to  each  interseptum.  The  inner  wall  pores 
frequently  carry  spines,  that  vary  in  shape  and  distribution. 

COMPOSITION  OF  THE  GENUS.  Erismacoscinus  contains  all  those  species,  having 
several  vertical  series  of  pores  to  each  interseptum,  that  were  previously  included  in 
the  genus  Coscinocyathus  Bornemann  (see  list  of  Debrenne  1964  :  166-167). 

DISCUSSION.  Hill  (1965  :  108-109)  suggests  this  genus  might  be  synonymous  with 
Tannuolacyathus  Vologdin  1957,  but  the  vesicular  tissue  and  double  porous  wall  of 
the  latter,  have  led  me  to  consider  that  genus  as  belonging  to  a  distinct  family,  the 
Tannuolacyathidae  (Debrenne  1964  :  188) . 


Erismacoscinus  rugosus  (R.  &  W.  R.  Bedford) 
(PI.  8,  figs.  2,  3) 

1934     Coscinocyathus  rugosus  R.  &  W.  R.  Bedford  :  3,  fig.  n. 

1964     Erismacoscinus  rugosus  (R.  &  W.  R.  Bedford)  Debrenne  :  167. 

HOLOTYPE.     B.M.  (N.H.)  S  4152. 

DESCRIPTION.  A  single  specimen  of  a  funnel-shaped  cup  with  a  wide  intervallum 
and  rather  narrow  central  cavity.  Septa  simply  porous  and  not  always  reaching 
the  inner  wall.  Tabulae  form  a  regular  network  of  pores.  The  inner  wall  is  simple 
with  some  spines.  The  pores  of  the  outer  wall  are  in  vertical  rows,  between  which  the 
"  linteaux  "  project.  These  keels  continue  towards  the  exterior,  giving  the  fossil  a 
rough  appearance  from  which  its  specific  name  is  derived.  The  pores  of  neighbouring 
rows  alternate  slightly. 


326  LOWER  CAMBRIAN   ARCH AEOCYATH A   FROM   AUSTRALIA 

DIMENSIONS 

(mm.) 

Cup: 

Height  (pars).  6 
Diameter                                                                   from  4  •  75-5  •  5 

Interseptum  (outer)  i-oi 

(inner)  0-33 

Intertabulum  I  •  35 

Loculi  1/2,  8/2,  8 

Intervallum  coefficient  0-8 

Parietal  coefficient  3  •  6 

Outer  wall : 

2-3  rows  of  pores  per  interseptum 

Vertical  diameter  of  pores  o  •  20 

Horizontal  diameter  of  pores  o  •  30 

Vertical  thickness  0-13 

Horizontal  thickness  o  •  05 

Inner  wall: 

2  rows  of  alternating  pores  per  interseptum 

Diameter  of  pores  o  •  10 

Linteaux  0-16 

Thickness  of  wall  o-io 

Septa : 

10  rows  of  pores  per  septum 

Diameter  of  pores  o  •  05 

Partitions  0-15 

Tabulae : 

7  pores  per  interseptum 

Diameter  of  pores  o  •  05 

Skeletal  partitions  0-06 

DISCUSSION.     The  presence  of  outer  keels  is  a  rare  modification.     It  seems  that 
E.  equivallum  (Taylor)  has  outer  keels  in  front  of  each  septum  (see  below). 


Erismacoscinus  equivallum  (Taylor) 
(PI.  7,  figs.  1-3  and  5) 

1908     "turbinate  Archaeocyathinoid"  Taylor  :  426,  pi.  i,  fig.  i. 

1910     Coscinocyathus  equivallum  Taylor  :  138-139,  pi.  15,  fig.  85,  t.-fig.  12. 

1964     Erismacoscinus  equivallum  (Taylor)  Debrenne  :  167 

HOLOTYPE.     In  the  University  of  Adelaide. 

OTHER  MATERIAL.  B.M.  (N.H.)  S  200  is  the  counterpart  (i.e.  the  other  face  of  the 
section)  of  the  holotype.  S  198,  naturally  etched  specimen  figured  by  Taylor  (1908, 
pi.  i,  fig.  2  :  423). 

DESCRIPTION.  Outer  wall  has  projections  of  skeletal  tissue  corresponding  to  the 
septa.  As  I  did  not  see  a  specimen  entirely  free  of  matrix,  it  is  only  possible  to 
assume  that  this  feature  is  similar  to  the  keels  present  in  E.  rugosus.  These  projec- 


LOWER   CAMBRIAN    ARCH  AEOCY  ATM  A   FROM   AUSTRALIA  327 

tions  only  occur  at  the  junction  of  the  septa  and  the  outer  wall,  and  not  between  the 
rows  of  pores  in  the  interseptum.  The  pores  are  elliptical  and  in  quincunx. 

The  inner  wall  has  round,  alternate  pores.  The  linteaux  are  smooth  on  the  inter- 
vallum  side  but  small  projections  of  spines  and  "  bristles  "  occur  on  the  central  cavity 
face. 

Thick,  straight  septa  have  round  regularly  scattered  pores  in  elongated  quincunx. 
The  skeletal  tissue  occupies  a  greater  surface  area  than  the  pores.  Slightly  convex 
tabulae  have  a  fine  network  of  small,  more  or  less  regular  polygonal  pores. 

DIMENSIONS 

S  200  S  198 

(mm.)  (mm.) 

Cup: 

Height  22 

Diameter  10  30-18 

Intervallum  coefficient  0-46  °'43 

Parietal  coefficient                                      Not  measurable  66 

Interseptum  o  •  3  7  0-41 

Intertabulum  o  •  44 

Outer  wall: 

No.  of  pore  rows  per  intersept  2  2 

Diameter  of  pores  0-15  0-22-0-15 

Vertical  partitions  o-n  0-11-0-15 

Horizontal  partitions  0-15  o  •  1 1 

Thickness  o  •  1 1  o  •  1 1 

Inner  wall : 

No.  of  pore  rows  per  intersept  2  2 

Diameter  of  pores  0-07  0-15 

Vertical  partitions  0-07  o  •  1 1 

Horizontal  partitions  0-07  o  •  1 1 

Thickness  o  •  1 1  o  •  1 1 

Septa : 

No.  of  pore  rows  per  intersept  9-10  15 

Diameter  of  pores  o  •  1 1  0-07 

Vertical  partitions  0-15  0-22 

Horizontal  partitions  0-15  0-22 

Thickness  0-07  0-15 

Tabulae: 

No.  of  pore  rows  per  intersept  14  4-22 

Diameter  of  pores  0-05  o  •  05-0  •  15 

Vertical  partitions  0-05  o  •  03-0  •  07 

Horizontal  partitions  0-05 

Thickness  0-07  0-07 

DISCUSSION.  Specimen  S  198  is  somewhat  corrugated.  At  the  level  of  the  folds 
the  septa  are  not  radial  but  parallel,  as  seen  in  Polystillicidocyathus  Debrenne  and 
other  colonial  forms. 

The  similarity  in  structure  of  the  two  walls  is  not  quite  so  evident  as  Taylor  stated 
(1910  :  138).  The  specimen  S  200  shows  that  the  inner  wall  is  a  little  thinner  and  has 
ragged  "  linteaux  "  and  spines. 

GEOL.  17,  7  26 


328  LOWER   CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA 

Erismacoscinus  quadratus  (R.  &  W.  R.  Bedford) 
(PI.  7,  fig-  4>  PL  8,  fig-  5) 

1934     Coscinocyathus  quadratus  R.  &  W.  R.  Bedford  :  3,  pi.  2,  fig.  10. 
1964     Erismacoscinus  quadratus  (R.  &  W.  R.  Bedford)  Debrenne  :  167. 

HOLOTYPE.     B.M.  (N.H.)  S  4157. 

DESCRIPTION.  Only  one  incomplete  specimen  known;  a  little  deformed  and  there- 
fore, difficult  to  measure  accurately  or  to  calculate  the  specific  coefficients. 

The  septa  and  tabulae  form  a  sub-quadrate  network  of  irregular  shape,  due  to  the 
slight  waving  of  the  intervallum  plates.  They  both  have  isolated,  simple,  round 
pores  arranged  in  quincunx.  The  outer  wall  also  has  simple  pores  in  quincunx  but 
with  the  majority  often  lengthened  horizontally.  The  inner  wall  is  smooth  on  the 
intervallum  side,  but  irregular  and  with  spines  around  the  pores  on  the  central  cavity 
face.  The  pores  are  arranged  in  horizontal  rows  and  alternate  from  one  row  to 
another. 

DIMENSIONS 

Lower  section  Upper  section 

(mm.)  (mm.) 

Cup: 

Height  25 

Diameter  n                                 16 

Intervallum  2-64                             2-64 

Interseptum  0-37-0 -50                        Q'37 

Intertabulum  o  •  30-0  -60 

Intervallum  coefficient  0-38                             0-2 

Parietal  coefficient  probably  3-1 

Outer  wall : 

3-4  rows  of  pores  per  intersept 

Diameter  of  pores  o  •  07-0  -15 

Partitions  o  •  1 1 

Thickness  0-05 
Inner  wall: 

2-3  rows  of  pores  per  intersept 

Diameter  of  pores  o  •  i  i-o  •  13 
Septa : 

20  pores  per  loculus 

Diameter  of  pores  0-03 

Partitions  o  •  1 1 

Thickness  0-05 
Tabulae : 

3  rows  of  20  pores  per  loculus 

Diameter  of  pores  0-03 

Partitions  0-06 

Thickness  o- 15  (with  the  spines) 

DISCUSSION.  The  peculiar  features  of  the  inner  wall  (smooth  in  the  intervallum. 
spined  in  the  central  cavity)  have  previously  been  described  for  E.  rugosus  (R.  &  W.  R. 
Bedford)  and  E.  equivallum  (Taylor) — two  species  that  differ  from  quadratus  in  other 
characters. 


LOWER  CAMBRIAN  ARCHAEOCYATHA  FROM  AUSTRALIA         329 

Okulitch  (1948  :  343)  compared  rhyacoensis  and  quadmtus,  as  both  have  subquad- 
rate  loculi.  However,  the  American  species  has  pore-tubes  in  the  outer  wall;  an 
inner  wall  which  is  typical  of  Coscinocyathus,  with  a  single  pore  series  to  each  inter- 
septum; and  an  inter  vallum  with  vesicular  tissue  as  well  as  tabulae. 

Erismacoscinus  textilis  (R.  &  W.  R.  Bedford) 
(PI.  9,  figs.  1-4) 

1934     Coscinocyathus  textilis  R.  &  W.  R.  Bedford  :  3,  pi.  2,  below  fig.  n. 
1964     Erismacoscinus  textilis  (R.  &  W.  R.  Bedford)  Debrenne  :  167. 

HOLOTYPE.     B.M.  (N.H.)  S  4155. 
PARATYPE.     B.M.  (N.H.)  S  4156. 

DESCRIPTION.  The  holotype  has  a  cylindro-conical  cap  that  tapers  gently  in  its 
lower  parts.  The  intervallum  consists  of  quadrate  to  rectangular  loculi,  limited  by 
radial  septa  and  flat,  irregularly-spaced  tabulae. 

The  outer  wall  has  a  single  vertical  series  of  pores  per  interseptum  (in  a  large  inter- 
septum there  are  two  rows,  but  these  soon  become  separated  by  a  new  septum).  The 
pores  of  the  outer  wall  are  much  larger  than  those  of  the  inner  wall,  they  are  also 
horizontally  lengthened  and  the  rows  alternate.  Pores  of  the  inner  wall  are  very 
small  and  arranged  in  quincunx.  Septa  and  tabulae  are  regularly  porous.  Skeletal 
tissue  is  well-developed. 

The  paratype  is  a  fragment  of  a  large,  bell-shaped  cup,  with  a  narrow  intervallum 
in  which  the  different  pore  characters  of  the  two  walls  can  easily  be  compared.  The 
inner  wall  has  a  regular  net  of  round  pores  in  quincunx,  whilst  the  outer  wall  has  1-2 
rows  of  pores.  Domes  of  vesicular  tissue  are  present  in  several  loculi. 

DIMENSIONS 

S  4155  (Upper)     S  4155  (Lower)       S  4156 

(mm.)  (mm.)  (mm.) 

Cup: 

Height :                                                                                   30  43 

Diameter  10  6  36  (Chord) 

Intervallum  1-32  1-32  2 

Interseptum  0-40  °'35  0-62 

Intertabulum  0-6-2-71  irregular  2-70-4-70 

Parietal  coefficient  4-2  3-3 

Outer  wall : 

No.  of  rows  of  pores  per  interseptum  1-2  1-2  1-2 

Diameter  0-34-0-15  0-34-0-15  0-15 

Skeletal  partitions  o-n  o-n  0-15 

Horizontal  partitions  0-07  ..  0-15 

Thickness  0-95  ..  0-15 

Inner  wall : 

No.  of  pore  rows  per  interseptum  3-4  .  .  4 

Diameter  0-07  ..  o-io 

Vertical  partitions  0-07  ..  o-io 

Horizontal  partitions  0-07  ..  o-io 

Thickness  o  •  n  .  0-07 


330  LOWER   CAMBRIAN    ARCH AEOCY ATH A   FROM   AUSTRALIA 

DIMENSIONS — continued 

S  4155  (Upper)     S  4155  (Lower)       S  4156 

(mm.)  (mm.)  (mm.) 

Septa : 

No.  of  rows  of  pores  8  . .  10 

Diameter  o-n  ..  o-io 

Vertical  partitions  0-15  ..  o-io 

Thickness  0-07  ..  o-io 

Tabulae : 

No.  of  pores  per  interseptum  4-5  4  5 

Diameter  0-07  0-07  0-05 

Vertical  partitions  0-07  0-07  0-07 

Horizontal  partitions  . .  . .  0-07 

Thickness  0-05  0-05  o  •  i  o 

Vesicular  tissue :  . .  . .  0-03 

DISCUSSION.  The  two  specimens,  a  cylindrical  one  and  a  bowl-shaped  one,  are 
placed  in  the  same  species.  More  material  is  needed  in  order  to  decide  whether  they 
belong  to  separate  species,  or  not.  Generally  the  outer  wall  of  Archaeocyatha  is 
thinner  than  the  inner  one.  However,  in  this  species  the  opposite  is  the  case.  It  is 
impossible  to  recognize  the  correct  position  of  a  fragment,  if  one  only  has  the  pore 
characters  of  a  wall  to  go  by.  Another  example  of  this  difficulty  is  the  case  of 
Coscinocyathus  unilinearis  R.  &  W.  R.  Bedford  (see  PI.  13,  fig.  4),  and  Coscinoptycta 
convoluta  (Taylor).  If  the  wall  with  a  single  pore,  is  the  inner  wall,  then  Coscino- 
ptycta is  a  synonym  of  Coscinocyathus  and  unilinearis  is  a  species  of  Coscinocyathus. 
On  the  other  hand,  if  it  is  the  outer  wall  that  has  one  pore,  then  Coscinoptycta  remains 
a  separate  genus  and  unilinearis  is  a  species  of  Erismacoscinus. 


Erismacoscinus  cellularis  (R.  &  W.  R.  Bedford) 

(PI.  8,  figs,  i  and  4) 
1934     Coscinocyathus  cellularis  R.  &  W.  R.  Bedford  :  3,  pi.  3,  fig.  16. 

HOLOTYPE.     S  4162  B.M.  (N.H.). 

DESCRIPTION.  The  holotype  is  a  longitudinally  broken  fragment  of  an  almost 
cylindrical  cup,  showing  the  porosity  of  the  different  skeletal  plates. 

The  outer  wall  is  a  thin,  regular  and  finely  porous  plate  with  horizontally  elongated 
pores.  The  inner  wall  is  thicker,  consisting  of  small,  regular  polygonal  tubes  that  face 
upwards;  its  central  cavity  face  has  several  spines,  or  other  outgrowths,  arising  from 
the  pore  walls. 

The  septa  have  small  evenly-spaced  pores.  The  tabulae  are  close  together  with 
their  convex  side  uppermost,  they  are  also  finely  perforated  but  the  pores  are  more 
numerous  than  those  of  the  septa. 


LOWER  CAMBRIAN  ARCH AEOC Y ATH A  FROM  AUSTRALIA         331 
DIMENSIONS 

(mm.) 

Cup: 

Height  55 

Diameter  20 

Intervallum  3 

Intertabulum  1-1-32 

Interseptum  o  •  30 

Intervallum  coefficient  o  •  75 

Parietal  coefficient  not  measurable 

Outer  wall : 

No.  of  pore  rows  per  interseptum  2 

Diameter  o  •  1 5-0  •  07 

Vertical  partitions  o  •  07 

Horizontal  partitions  o  •  1 1 

Thickness  o  •  07 

Inner  wall : 

No.  of  pore  rows  per  interseptum  3 
Diameter                                                         0-15;  o-u  ;  o-n  (nr. 

central  cavity) 

Vertical  partitions  o  •  1 1 

Thickness  0-26 

Septa : 

No.  of  pore  rows  per  interseptum  30  approx. 

Diameter  o  •  03 

Vertical  partitions  o  •  1 1 

Horizontal  partitions  o  •  08 

Thickness  o  •  07 

Tabulae : 

No.  of  pore  rows  per  interseptum  3-4 

Diameter  o  •  05 

Vertical  partitions  o  •  07 

Horizontal  partitions  o  •  07 

Thickness  o  •  07 

DISCUSSION.  R.  &  W.  R.  Bedford  (1934 : 3)  suggested  that  the  outer  wall  is 
covered  by  a  thin  membrane  that  has  very  minute  pores,  but  I  could  not  find  any 
evidence  of  this  structure.  Their  "  irregular  mosaic  "  of  the  inner  wall  is,  in  fact,  the 
result  of  the  modification  of  the  skeletal  tissue  ("  linteaux  ")  into  barbs  and  spines. 
The  regular  polygonal  tubes  can  be  seen  on  the  intervallum  face  of  the  inner  wall, 
as  in  E.  rugosus  and  E.  equivallum. 

Erismacoscinus  peter  si  (R.  &  W.  R.  Bedford) 

(PI.  10,  figs.  4  and  5) 

1934     Coscinocyathus  petersi  R.  &  W.  R.  Bedford  :  3,  pi.  3,  fig.  13. 
HOLOTYPE.     B.M.  (N.H.)  S  4158. 

DESCRIPTION.  The  holotype  is  a  unique,  small,  well-preserved  specimen.  The 
outer  wall  is  simply  porous  with  prominent,  sharply-ridged,  vertical  crests  between 
the  septa.  The  inner  wall  is  a  skeletal  plate  that  is  perforated  by  vertical  and 
horizontal  rows  of  regular  pores.  Consequently,  the  wall  tissue  between  neighbouring 


332 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A    FROM    AUSTRALIA 


horizontal  rows  of  pores,  forms  a  continuous  circular  bar.  A  small  tongue-like  plate 
arises  from  the  bar  under  each  pore,  but  it  was  not  possible  to  discover  whether  these 
tongues  are  separate  from  each  other,  or  connected  to  form  an  annular  ring.  The 
radial  septa  have  scattered  regular  pores,  while  the  tabulae  consist  of  a  thin  porous 
net. 


DIMENSIONS 


Cup: 

Height 

Diameter    ..... 

Intervallum         .... 

Interseptum  (outer  wall) 
(inner  wall) 

Intervallum  coefficient 

Parietal  coefficient 
Outer  wall : 

8  pores  (4  +  4)  per  interseptum 

Diameter  of  pores 

Thickness  of  linteaux  . 

Thickness  of  wall 
Inner  wall : 

No.  of  rows  of  pores  per  interseptum 

Diameter  of  pores 

Vertical  partitions 

Horizontal  partitions  . 

Thickness  of  wall 
Septa : 

No.  of  rows  of  pores  per  septa 

Diameter  of  pores 

Vertical  partitions 

Horizontal  partitions  . 
Tabulae : 

No.  of  rows  of  pores  per  interseptum 

Diameter  of  pores 

Partitions  . 


(mm.) 

9 

9 

i'5i 

0-83 

0-56 

0-25 


0-07 
O'Oy 
O'll 

2 

0-07 

O-II 

0-07 
0-26 

6 

0-03 

o-ii 

•  11-0-26 

12 

0-03 
O-II 


DISCUSSION.  The  horizontal  alignment  of  the  inner  wall  pores  is  an  important 
feature,  that  is  a  precursor  to  the  formation  of  annular  structures.  This  is  evident  in 
this  case,  for  the  horizontal  bars  are  already  slightly  modified  by  supplementary 
plates.  However,  as  the  stage  of  a  single  pore  per  interseptum  has  not  been  reached, 
this  species  should  be  retained  in  Erismacoscinus  and  not  placed  in  a  genus  of  the 
Salairocyathidae.  The  star-shaped  form  of  the  external  wall  is  similar  to  that  of  E. 
cancellatus  (Bornemann),  but  the  two  species  have  no  other  features  in  common. 

Erismacoscinus  retifer  (R.  &  W.  R.  Bedford) 
(PL  10,  figs.  2  and  3) 

1934     Coscinocyathus  retifer  R.  &  W.  R.  Bedford  :  3,  pi.  3,  fig.  14. 
1964     Erismacoscinus  retifer  (R.  &  W.  R.  Bedford)  Debrenne  :  167. 

HOLOTYPE.     B.M.  (N.H.)  S  4159. 


LOWER   CAMBRIAN    ARCH  AEOC  Y  ATM  A    FROM   AUSTRALIA  333 

DESCRIPTION.  The  species  is  based  on  a  single,  small,  incomplete  specimen,  but  it 
is  sufficiently  well-preserved  to  enable  one  to  see  the  details  of  each  skeletal  plate. 
The  thin  outer  wall  is  perforated  by  4-5  vertical  rows  of  pores  to  each  interseptum, 
these  pores  also  form  horizontal  rows.  The  septa  and  the  only  visible  tabula  are  of 
the  same  thickness  as  the  outer  wall,  and  have  their  round  pores  arranged  in 
quincunx;  the  skeletal  tissue  between  them  is  wider  than  the  pores  and  projects 
slightly. 

The  inner  wall  is  thicker  than  the  other  plates.  The  pores  are  elliptical,  with  the 
larger  diameter  horizontal  and  a  tongue  arising  under  each  pore.  One  vertical  row  of 
pores  is  only  just  separated  from  the  next  and,  at  times,  pores  from  neighbouring 
rows  may  join,  giving  a  single  pore  at  that  point  of  the  interseptum.  At  certain 
places,  the  pores  are  aligned  horizontally  and  this  involves  two  sporadic  modifications. 
Firstly,  the  coalescence  of  two  neighbouring  pores,  which  predicts  the  uniserially- 
pored  inner  wall.  Secondly,  the  development  of  small  tongue-like  plates  under  the 
pores,  forecasting  the  formation  of  annular  plates. 

DIMENSIONS 

(mm.) 

Cup: 

Height  (pars)       ......  9 

Diameter  ......  8 

Intervallum         .          .          .          .          .          .  1-96 

Intervallum  coefficient  .          .          .          .  0-5 

Parietal  coefficient        .          .          .          .          .  3-5 

Outer  wall: 

No.  of  pore  rows  per  interseptum .          .          .  4-5 

Diameter  of  pores         .          .          .          .          .  0-07 

Vertical  partitions        .          .          .          .          .  o-io 

Horizontal  partitions   .          .          .          .          .  o-io 

Thickness  .          .          .          .          .          .          .  0-13 

Inner  wall : 

Diameter  of  pores         .....          0-28-0-18 

Vertical  partitions        .          .  .          .  o-io 

Horizontal  partitions  .          .          .          .          .  0-15 

Length  of  plates .          .          .          .          .          .  0-20 

Septa : 

No.  of  pores  per  loculus         ....  9-10 

Diameter  of  pores         .          .          .          .          .  o  •  1 1 

Distance  between  horizontal  rows  .          .  o  •  1 8 

Distance  between  vertical  rows      .          .          .  0.22 

Tabulae : 

No.  of  pores  per  interseptum          .      .  .  6 

Diameter    .          .          .          .          .          .          .  0-04 

Partitions  .          .          .          .          .          .          .  0-05 

Thickness  .          .          .          .          .          .          .  0-05 


DISCUSSION.     This  species  continues  to  follow  the  evolution  already  outlined  under 
E.  peter  si  and  heralds  the  annular  forms  of  Salairocyathidae. 


334  LOWER   CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA 

Family  POLYCOSCINIDAE  Debrenne,  1964 

DIAGNOSIS.  Cup  with  porous  septa  and  tabulae.  Double,  porous  outer  wall. 
Simple  inner  wall. 

COMPOSITION  OF  THE  FAMILY.  Polycoscinus  R.  &  J.  Bedford  1937,  Tomocyathus 
Rozanov  1960 

Genus  TOMOCYATHUS  Rozanov,  1960 

1960  Tomocyathus  Rozanov  :  664,  figs,  i  d,  e. 

1964  Tomocyathus  Rozanov;  Debrenne  :  115 

1964  Tomocyathus  Rozanov;  Repina  et  al.  :  231. 

1965  Tomocyathus  Rozanov;  Hill  :  107 

TYPE  SPECIES.     Tomocyathus  operosus  Rozanov  1960,  by  original  designation. 

DIAGNOSIS.  Outer  wall  coarsely  porous,  covered  by  a  secondary  micro-porous 
sheath.  Straight  septa  and  convex  tabulae,  both  with  numerous  fine  pores.  Vesi- 
cular tissue.  Inner  wall  with  simple  pores,  and  stellate  in  transverse  section. 

DISCUSSION.  The  fluting  of  the  walls  is  often  considered  a  character  of  sub- 
generic  value  (Krasnopeeva  1955;  Debrenne  1964).  Recently,  Repina  et  al  (1964  : 
231)  described  Tomocyathus  without  any  mention  of  the  stellate  inner  wall.  Some 
species  that  they  included  in  this  genus,  compositus  (Zhuravleva)  and  shoriensis 
Rozanov,  have  little  to  no  vesicular  tissue,  rather  flat  tabulae  and  the  inner  wall  is  not 
fluted  but  possesses  spines  of  varying  size  and  shape. 

According  to  criteria  used  in  distinguishing  other  subgenera  e.g.  Ajacicyathus 
(Urcyathus)  Vologdin,  one  must  create  a  new  subgenus  for  the  smooth  forms  of 
Tomocyathus. 

COMPOSITION  OF  THE  GENUS.  Tomocyathus  (Tomocyathus)  operosus  Rozanov  1960, 
T.  (Tomocyathus}  michniaki  Rozanov  1966;  for  subgenus  Tomocyathus  (Erugato- 
cyathus} see  below. 

Subgenus  ERUGATOCYATHUS  nov. 

DERIVATION,     erugatus  —  unfolded. 

TYPE  SPECIES.     Coscinocyathus  papillatus  R.  &  W.  R.  Bedford,  designated  here. 

DIAGNOSIS.     Tomocyathus  with  non-folded  inner  wall. 

COMPOSITION  OF  THE  SUBGENUS.  Tomocyathus  (Erugatocyathus}  echinus  (Debrenne 
1964),  T.  (E.)  compositus  Zhuravleva  1955,  T.  (E.)  gini  Missarzhevsky  &  Rozanov 
1962,  T.  (E.)  kundatus  Rozanov  1966,  T.  (E.)  shoriensis  Rozanov  1964,  T.  (E.)  papilla- 
tus (R.  &  W.  R.  Bedford  1934). 

Tomocyathus  (Erugatocyathus)  papillatus  (R.  &  W.  R.  Bedford) 
(PL  n,  fig.  2,  Text-fig.  9) 

1934     Coscinocyathus  papillatus  R.  &  W.  R.  Bedford  :  3,  pi.  3,  fig.  12. 
1964     Erismacoscinus  papillatus  (R.  &  W.  R.  Bedford)  Debrenne  :  167 

HOLOTYPE.     B.M.  (N.H.)  S  4153. 
PARATYPE,    B.M.  (N.H.)  S  4154. 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A    FROM    AUSTRALIA 


335 


DESCRIPTION.  Fragments  of  cylindrico-conical  fossils.  Septa  with  remote  round 
pores.  Tabulae  reticular.  Loculi  rectangular  and  irregular  in  size.  The  circular 
pores  at  the  base  of  the  coarsely-porous  outer  wall  are  covered  by  a  microporous 
sheath,  in  which  each  group  of  micropores  consists  of  a  central  pore  surrounded  by 
five  others. 

A  skeletal  tongue  covers  each  simple  pore  of  the  inner  wall,  it  arises  under  the  pore, 
curves  upwards  and  rejoins  the  wall  above.  As  the  rows  of  pores  alternate,  each  pore 
is  surrounded  by  4  knobs,  corresponding  to  the  beginning  of  a  papilla.  This  feature  is 
well  shown  in  the  weathered  specimen  S  4154  (see  Text-fig.  9). 


FIG.  9.     Tomocyathus  (Erugatocyathus)  papillatus  (R.  &  W.  R.  Bedford) 

DIMENSIONS 


Cup: 
Height 
Diameter 

Intervallum  coefficient 
Parietal  coefficient 
Intervallum 
Interseptum 


S  4153  Upper  S  4153  Lower 

(mm.)  (mm.)  (mm.) 


13 
1-89 

3-8 
0-40 


o-34 


18 
10-5 


0-41 
2-5-4 


336 


LOWER   CAMBRIAN   ARCH AEOC Y ATH A    FROM    AUSTRALIA 


DIMENSIONS — continued 


Outer  wall: 

No.  of  rows  of  pores  per  interseptum 

Diameter 

Vertical  skeletal  partitions 

Horizontal  skeletal  partitions 

Thickness 
Inner  wall: 

No.  of  pore  rows  per  interseptum 

Diameter 

Vertical  skeletal  partitions 

Horizontal  skeletal  partitions 

Thickness 
Septa : 

No.  of  pore  rows  per  inter  vallum 

Diameter 

Vertical  skeletal  partitions 

Horizontal  skeletal  partitions 

Thickness 
Tabulae 

No.  of  pore  rows  per  interseptum 

Diameter 

Vertical  skeletal  partitions 

Thickness 


S  4153  Upper 
(mm.) 

84153  Lower 
(mm.) 

S4I54 

(mm.) 

2 

2 

0-15 
0-15-0-  18 

0-15-0-03 
0-15-0-03 

0-18 
0-15 

0-15-0-03 

0-15 

0-15 

O-II 

2 

2 

2 

O-22 

0-20 

0-20 

0-15 

0-15 

o-io  +  pap.  =  o-n  0-22 

10  8 

0-05  0-07 

o-io  o-ii 

0-07  0-07 

5-6  4-5 

0-05  0-05 

0-07  0-05 

0-05  0-05 


0-07 
o-ii 

O-II 

0-07 

4 

0-07 
0-07 
0-05 


DISCUSSION.     These  peculiar  curved  tongues  are  not  known  in  any  other  species 
of  the  Regularia. 

Family  SALAIROCYATHIDAE  Zhuravleva,  1955 

DIAGNOSIS.     Cups  with  intervallum  crossed  by  porous  septa  and  tabulae.     Simple 
outer  wall,  but  annular  inner  wall. 

COMPOSITION  OF  THE   FAMILY.    Salairocyathus   (Salairocyathus)   Vologdin   1940, 
Salairocyathus  (Polystillicidocyathus}  Debrenne  1959. 


Genus  SALAIROCYATHUS  Vologdin,  1940 
i94oa  Salairocyathus  Vologdin  :  89 

TYPE  SPECIES.     Salairocyathus  zenkovae  Vologdin  1940 :  89,  pi.  26,  fig.  6. 

DIAGNOSIS.  Simple  pores  in  outer  wall  and  also  in  septa  and  tabulae.  Annuli  on 
inner  wall,  v-shaped  in  section  and  open  towards  the  top. 

COMPOSITION  OF  THE  GENUS.  Salairocyathus  (Salairocyathus}  zenkovae  Vologdin 
1940,  5.  (S.)  pospelovi  Zhuravleva  1960,  ?  5.  (S.)  annulatus  (R.  &  W.  R.  Bedford 
1934).  Salairocyathus  (Polystillicidocyathus}  erbosimilis  Debrenne  1959. 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A    FROM   AUSTRALIA  337 

?  Salairocyathus  (Salairocyathus)  annulatus  (R.  &  W.  R.  Bedford) 

(PI.  ii,  fig.  4,  Text-fig.  10) 
1934     Coscinocyathus  annulatus  R.  &  W.  R.  Bedford  :  4,  pi.  3,  fig.  17. 

HOLOTYPE.     S  4163  in  British  Museum  (Natural  History). 

DESCRIPTION.     A  unique,  very  fragmentary  specimen,  that  has  part  of  the  inner 
wall,  as  seen  from  the  central  cavity,  well  preserved.     Porous  outer  wall,  but  not 


FIG.  10.     Salairocyathus  (Salairocyathus}  annulatus  (R.  &  W.  R.  Bedford) 


possible  to  distinguish  their  arrangement.  Septa  and  tabulae  frame  somewhat 
parallelepipedic  loculi.  The  rectangular  openings  of  the  inner  wall  are  bounded  by 
the  inner  edges  of  the  septa  and  a  vertical  series  of  horizontal  bars,  which  are 
annular  in  form.  The  ft  component  of  the  latter  is  thick  and  horizontal,  while  the  a 
component  is  much  thinner  and  curves  backwards  up  to  the  middle  of  the  following 
horizontal  row  of  openings,  which  are  thus  partly  closed,  (see  Text-fig.  10). 


338         LOWER  CAMBRIAN  ARCH AEOCY ATH A  FROM  AUSTRALIA 
DIMENSIONS 

(mm.) 

Cup: 

Height  27 

Diameter  1 1 

Intervallum  ? 

Interseptum  o  •  22-0  •  30 

Intertabulum  about  1-35 

Outer  wall: 

No.  of  rows  of  pores  per  interseptum  probably  4 

Thickness  0-13 

Inner  wall: 

No.  of  rows  of  pores  per  interseptum  i 

Diameter  of  pores  (lengthened  horizontally)  0-25-0 -15 

Skeletal  partitions  (constant  thickness)  o  •  06 

Septum : 

Diameter  of  pores  0-06 

Skeletal  partitions  0-13 

Thickness  0-13 

Tabulae : 

Diameter  of  pores  (radially  lengthened)  0-13-0-06 

Skeletal  partitions  o  •  06 

Thickness  0-15 

DISCUSSION.  This  annular  form  of  inner  wall  was  previously  unknown.  The 
specimen  is  described  as  Salairocyathus  with  reservation. 

Family  ALATAUGYATHIDAE  Zhuravleva,  1955 

DIAGNOSIS.  Cups  with  simply  porous  septa  and  tabulae.  Inner  wall  with  simple, 
or  S-shaped  pores.  Two  sub-families  are  distinguished  by  the  shape  of  the  pores  in 
the  outer  wall. 

Subfamily  TUMULOCOSCININAE  Zhuravleva,  1960 
DIAGNOSIS.     Outer  wall  with  simple  tumuli. 

COMPOSITION  OF  THE  SUBFAMILY.  Tumulocoscinus  Zhuravleva  1960,  Ethmoco- 
scinus  Simon  1939,  Asterotumulus  Rozanov  1964. 

Subfamily  ALATAUGYATHINAE  Zhuravleva,  1960 
DIAGNOSIS.     Outer  wall  with  knobbly  tumular  pores. 

COMPOSITION  OF  THE  SUBFAMILY.  Alataucyathus  (Alataucyathus)  Zhuravleva 
1955,  Alataucyathus  (Anaptyctocyathus)  subgen.  nov. 

Subfamily  TUMULICOSCININAE  Zhuravleva,  1960 

Genus  ETHMOCOSCINUS  Simon  1939 
1939     Ethmocoscinus  Simon  :  28 

TYPE  SPECIES.  Coscinocyathus  papillipora  R.  &  W.  R.  Bedford,  by  original  desig- 
nation (Simon  1939  :  28). 


LOWER  CAMBRIAN   ARCH AEOCYATH A   FROM   AUSTRALIA  339 

DIAGNOSIS.  Cylindro-conical  cups.  Outer  wall  with  simple  tumuli,  one  per 
interseptum.  Inner  wall  with  a  single  row  of  S-shaped  pore-tubes  per  interseptum. 
Septa  have  sparse  simple  pores.  Tabulae  with  polygonal,  somewhat  irregular 
pores. 

COMPOSITION  OF  THE  GENUS.     A  single  species  E.  papillipora  (R.  &  W.  R.  Bedford). 

Ethmocoscinus  papillipora  (R.  &  W.  R.  Bedford) 

(PL  3,  fig-  3) 

1934     Coscinocyathus  papillipora  R.  &  W.  R.  Bedford  :  18. 
1939     Ethmocoscinus  papillipora  (R.  &  W.  R.  Bedford)  Simon  :  28. 

HOLOTYPE.     B.M.  (N.H.)  S  4164. 

DESCRIPTION.  The  holotype,  the  only  known  specimen,  is  partly  weathered-out 
and  shows  the  structure  of  the  two  walls.  The  straight  septa  are  perforated  by  round 
regular  pores,  which  although  few,  are  arranged  in  quincunx.  A  single  tabula  is 
visible  (it  is  therefore  impossible  to  state  the  frequency)  and  consists  of  a  net  of  regular 
pores.  The  papillae  of  the  outer  wall  are  situated  between  the  septa  in  vertical  rows, 
which  are  just  separate  from  one  interseptum  to  the  next.  The  papillae  of  the  inner 
wall  occupy  a  much  larger  area  and  also  appear  to  be  in  alternate  vertical  rows. 

DIMENSIONS 

Cup:  (mm.) 

Height  (pars)  20 

Diameter  7  •  5 

Intervallum  i  •  2 
Interseptum                                                            from  0-41-0-45 

Number  of  septa  26 

Parietal  coefficient  3  •  4 

Intervallum  coefficient  0-2 

Outer  wall: 

No.  of  pore  rows  per  intersept  i 

No.  of  pore  rows  before  formation  of  septa  2 

Diameter  of  papilla  o  •  34-0  •  2  2 

Thickness  of  papilla  o  •  07 

Height  of  papilla  o  •  18 

Horizontal  partitions  0-15-0-18 

Vertical  partitions  o  •  1 1 

Inner  wall: 

No.  of  rows  of  pores  per  interseptum  I 

Diameter  of  pores  0-37 

Partitions  o  •  1 1 

Septa : 

No.  of  rows  of  pores  per  intervallum  4 

Diameter  of  pores  o  •  1 1 

Partitions  o  •  1 1 

Thickness  o  •  06 

Tabula : 

No.  of  rows  of  pores  per  interseptum  6 

Diameter  of  pores  o  •  07 

Partitions  o  •  03 

Thickness  o  •  03 


340  LOWER   CAMBRIAN    ARCH AEOCY ATH A   FROM   AUSTRALIA 

Subfamily  ALATAUCYATHINAE  Zhuravleva,  1960 

Genus  ALA  TA  UCYA  THUS  Zhuravleva,  1955 
1955     Alataucyathus  Zhuravleva  :  626,  figs.  la,  2d. 

TYPE  SPECIES.  By  original  designation  Alataucyathus  jaroschevitschi  Zhuravleva 
1955  :  626. 

DIAGNOSIS.  Intervallum  filled  with  simple  septa  and  tabulae.  Outer  wall  covered 
with  small  multi-perforate  knobs.  Inner  wall  simple  with  vertical  folds  between 
neighbouring  septa. 

DISCUSSION.  As  mentioned  in  the  discussion  on  Tomocyathus,  it  is  considered  that 
species  having  non-fluted  inner  walls,  should  be  placed  in  a  separate  sub-genus. 

Subgenus  ALATAUCYATHUS  (ANAPTYCTOCYATHUS)  nov. 

DERIVATION,     anaptyctos  =  unfolded. 

TYPE  SPECIES.     Coscinocyathus  cribripora  R.  &  W.  R.  Bedford  designated  here. 

COMPOSITION  OF  THE  GENUS.  A.  (A.)  cribripora  (R.  &  W.  R.  Bedford,  1934),  A .  (A .) 
minimiporus  (R.  &  J.  Bedford  1937) — included  with  some  doubt  as  the  structure  of 
the  walls  is  not  certain,  A.  (A.)  excellenthis  (Rozanov  1964),  A.  (A.)  verschkhovska- 
jae  (Zhuravleva  1961),  and  A .  (A  .}flabellus  nov. 

Alataucyathus  (Anaptyctocyathus]  cribripora  (R.  &  W.  R.  Bedford) 

(PL  n,  figs,  i  and  3) 
1934     Coscinocyathus  cribripora  R.  &  W.  R.  Bedford  :  3,  pi.  3,  fig.  15. 

HOLOTYPE.     B.M.  (N.H.)  S  4160. 

DESCRIPTION.  Cylindrical  cup.  Intervallum  with  straight  radial  septa,  in  which 
the  hexagonal  pores  occupy  a  much  larger  surface  in  proportion  to  the  skeletal  tissue, 
and  irregularly  distributed  flat  tabulae,  that  are  finely  perforated  by  very  narrow 
pores. 

The  inner  wall  is  simple,  with  two  rows  of  pores  per  interseptum,  which  are  only 
weakly  separated,  in  fact,  towards  the  upper  part  of  the  cup,  two  neighbouring  pores 
may  sometimes  coalesce.  The  pores  of  the  outer  wall  are  covered  by  knobs,  that  are 
perforated  by  a  central  pore  and  a  surrounding  circle  of  six  others. 

DIMENSIONS 

(mm.) 
Cup: 

Height  (pars)  36 

Upper  diameter  12 
Lower  diameter  6 

Intervallum  coefficient  0-28 

Interseptum  (Upper  diameter)  °'74 

(Lower  diameter)  o  •  56 

Intertabula  2-5 


LOWER   CAMBRIAN    ARCH AEOCY ATH A  FROM   AUSTRALIA             341 

DIMENSIONS — continued 

(mm.) 
Outer  wall: 

No.  of  rows  of  main  pores  per  interseptum  2 

(each  pore  covered  by  a  sheath  with  7  micropores) 

Main  pore  diameter  0-27 

skeletal  partitions  o  •  27-0  •  30 

Micropore  diameter  0-07 

skeletal  partitions  o  •  03 
Inner  wall : 

No.  of  pore  rows  per  interseptum  2 

(these  sometimes  coalesce) 

Diameter  of  pores  o  •  30 

Skeletal  partitions  o  •  07 
Septa: 

No.  of  hexagonal  pores  4-5 

Diameter  o  •  1 5-0  •  1 8 

Vertical  partitions  o  •  18 

Horizontal  partitions  0-18 
Tabulae : 

No.  of  pores  10-4 

Diameter  o  •  1 1  -o  •  06 

Partitions  o  •  07 

DISCUSSION.     A.  (A.)  cribripora  differs  from  the  two  Russian  species  by  having 
lower  tumuli,  coarse  micropores  and  a  thicker  inner  wall. 


Alataucyathus  (Anaptyctocyathus)  flabellus  nov. 
(PI.  10,  fig.  i) 

HOLOTYPE.     B.M.  (N.H.)  S  4161. 

DESCRIPTION.  Bowl-shaped  cup  with  narrow  intervallum.  Regularly  porous 
septa  in  alternate  vertical  rows.  The  tabulae  almost  join,  their  pores  are  much 
thinner  than  those  of  the  septa.  The  inner  wall  is  only  known  from  the  intervallum 
side  and  therefore,  it  is  not  possible  to  discover  whether  there  are  spines  on  the  central 
cavity  face.  Three  vertical  rows  of  pores  can  be  distinguished  per  interseptum.  The 
outer  wall  has  2-3  pores,  each  covered  by  a  small  microporous  tumulus  with  about  ten 
angular  pores  in  each. 

DIMENSIONS 

(mm.) 

Cup: 

Height  (pars)  20 

Span  of  visible  arc  78 

Intervallum  2 

Loculi  1/2/3 

Outer  wall: 

No.  of  rows  of  pores  per  interseptum  from  2-3 

Diameter  of  main  pores  0-40-0-27 


342  LOWER  CAMBRIAN   ARCH AEOCY ATH A   FROM  AUSTRALIA 

DIMENSIONS — continued 

(mm.) 

Vertical  skeletal  partition  o  •  16 

Horizontal  skeletal  partition  o  •  1 1 

Diameter  of  micropores  o  •  06 

Skeletal  partition  between  micropores  0-03 

Inner  wall: 

Diameter  of  pores  0-18 

Skeletal  partitions  o  •  22 

Thickness  0-20 

Septa : 

Diameter  of  pores  o  •  20 

Vertical  partitions  o  •  30 

Horizontal  partitions  0-20 

Thickness  o-n 

Tabulae : 

No.  of  rows  of  pores  per  interseptum  5 

Diameter  of  pores  0-06 

Skeletal  partitions  o  •  06 

Thickness  o  •  1 1 

DISCUSSION.  Differs  from  A.  (A.)  cribripora  by  its  bowl-shaped  cup,  the  specific 
coefficients  and  the  different  arrangement  of  the  outer  wall  pores. 

Class  IRREGULARIA  Vologdin,  1937 
Order  ARCHAEOCYATHA  Okulitch,  1935 
Family  DICTYOGYATHIDAE  Taylor,  1910 

DIAGNOSIS.  Cups  with  intervallum  containing  disorientated  short  rods  and  dis- 
sepiments. Two  simply  porous  walls. 

COMPOSITION  OF  THE  FAMILY.  Dictyocyathus  Bornemann  1891,  Spinosocyathus 
Zhuravleva  1960,  Pinacocyathus  R.  &  W.  R.  Bedford  1934,  Agastrocyathus  Debrenne 
1964,  Archaeopharetra  R.  &  W.  R.  Bedford  1936,  Chouberticyathus  Debrenne  1964. 

Genus  PINACOCYATHUS  R.  &  W.  R.  Bedford,  1934 

1934     Pinacocyathus  R.  &  W.  R.  Bedford  :  4,  fig.  21. 

1964  Pinacocyathus  R.  &  W.  R.  Bedford;  Debrenne  :  200 

1965  Pinacocyathus  R.  &  W.  R.  Bedford;  Hill  :  117,  fig.  22,  ya  and  b. 

TYPE  SPECIES.  By  monotypy  Pinacocyathus  spicularis  R.  &  W.  R.  Bedford 
1934  :  4. 

DIAGNOSIS.  Two-walled  cup  with  intervallar  rods.  The  outer  wall  is  formed  by  a 
scaffolding  of  vertical  pillars  connected  by  horizontal,  or  slightly  oblique  rods.  The 
inner  wall  is,  probably,  a  regular  net.  Oblique,  or  radial  horizontal  rods  in  the 
intervallum. 

COMPOSITION  OF  THE  GENUS.     A  single  species;  P.  spicularis. 


LOWER   CAMBRIAN    ARCH AEOCY ATH A   FROM   AUSTRALIA 


343 


Pinacocyathus  spicularis  R.  &  W.  R.  Bedford 

(PI.  12,  figs,  i,  2,  Text-fig,  n) 
1934     Pinacocyathus  spicularis  R.  &  W.  R.  Bedford  :  4,  fig.  21. 

HOLOTYPE.     By  monotypy  B.M.  (N.H.)  S  4169. 

DESCRIPTION.  Conical  tube.  The  outer  wall  consists  of  an  open  framework, 
which  is  mainly  composed  of  vertical  columns  arranged  around  the  cup  at  almost 
regular  intervals.  This  framework  is  completed  by  horizontal,  or  oblique  rods, 


FIG.  ii.     Pinacocyathus  spicularis  R.  &  W.  R.  Bedford 


which  connect  the  columns,  with  a  few  spines  occurring  at  the  junctions.  On  the 
lower  part,  the  net  is  more  compact  but  hidden  by  poor  preservation,  that  has 
swollen  the  structures.  Only  one  vertical  row  of  the  inner  wall  can  be  seen  and  this 
seems  to  be  composed  of  a  regular  mesh.  However,  it  was  not  possible  to  determine 
the  arrangement  of  the  intervallar  rods.  They  seem  to  arise  from  the  skeletal  junc- 
tions of  the  outer  wall  framework  and  continue  radially,  or  obliquely  towards  the 
inner  wall.  The  framework  is  probably  thicker  in  its  lower  parts. 

GEOL.  17,  7  27 


344         LOWER  CAMBRIAN  ARCH  AEOC  Y  ATM  A  FROM  AUSTRALIA 
DIMENSIONS 

(mm.) 

Cup: 

Height  (pars)       .          .          .  .  .  .  15 

Diameter    .          .          .          .  .  .  .  5  •  75 

Intervallum         .          .          .  .  .  .  1-42 

Central  cavity     .          .          .  .  .  .  0-6 

Outer  wall : 

Diameter  of  pores         .          .  .  .  .  o  •  55 

Distance  between  vertical  rods  .  .  .  o  •  67-1  •  o 

Distance  between  horizontal  rods  .  .  .  i  •  35-1  •  70 

Thickness  of  vertical  rods     .  .  .  .  °'33 

Thickness  of  horizontal  rods  .  .  .  0-33 

Thickness  of  oblique  rods      .  .  .  .  0-25 

Inner  wall : 

Diameter  of  pores         .          .  .  .  .  °'33 

Vertical  partitions        .          .  .  .  .  0-20 

Horizontal  partitions   •          •  •  •  •  0-20 

Thickness  .          .          .          .  .  .  .  0-20 

Intervallar  rods : 

Thickness  .          .          .          .  .  .  .  0-27 

DISCUSSION.  Only  one  specimen  of  this  curious  form  is  known.  It  is  included  in 
the  Dictyocyathidae  because  of  its  scaffolding  of  intervallar  rods,  but  no  other 
recognized  genus  has  such  a  loosely-spaced  framework. 

Family  FLINDERSICYATHIDAE  R.  &  J.  Bedford,  1939 

DIAGNOSIS.  Two-walled  cups,  simply  porous,  with  a  single  series  of  large  pores  in 
the  inner  wall.  A  scaffolding  of  rods  fills  the  intervallum,  consisting  of  taeniae,  that 
cross  the  intervallum,  joining  the  septa,  and  undulate  in  a  radial  plane,  together  with 
synapticulae,  which  connect  the  crests  of  neighbouring  taeniae.  Little  to  no  vesicular 
tissue  is  present.  Apex  of  Dictyocyathus-type. 

COMPOSITION  OF  THE  FAMILY.  Spirocyathella  Vologdin  1939,  Flindersicyathus 
(Flindersicyathus}  R.  &  J.  Bedford  1937,  Flindersicyathus  (Pycnoidocyathus}  (Taylor 
1910),  ?  Spirillicyathus  R.  &  J.  Bedford,  1937,  Copleicyathus  R.  &  J.  Bedford  1937. 

Genus  FLINDERSICYATHUS  R.  &  J.  Bedford,  1937 

1910  Pycnoidocyathus  Taylor  :  131. 

1937  Flindersicyathus  R.  &  J.  Bedford  :  28. 

1939  Flindersicyathus  R.  &  J.  Bedford;  Simon  :  30. 

1960  Archaeocyathus  Billings;  Zhuravleva  :  296  (pars). 

1965  Flindersicyathus  R.  &  J.  Bedford;  Hill  :  123. 

TYPE  SPECIES.  Flindersicyathus  decipiens  R.  &  J.  Bedford,  by  subsequent 
designation  R.  &  J.  Bedford  (i939(May)  :  78),  which  has  priority  over  Spirocyathus 
irregularis  Taylor  chosen  by  Simon  (1939  (Dec.)  :  30),  see  Hill  (1965  :  123).  Holo- 
type  material  No.  86670  in  Princeton  University,  U.S.A. 

DIAGNOSIS.  Hill  (1965  :  123)  writes:  "solitary  cups,  with  simply  porous  outer 
wall;  inner  wall  with  a  single  series  of  large  rounded  pores  per  intertaenial  loculus, 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A    FROM   AUSTRALIA 


345 


each  pore  bounded  by  the  taeniae  and  the  synapticulae  joining  them,  and  each  with  a 
louvre-like  plate  projecting  upwards  and  inwards  to  the  central  cavity  from  its  low 
synapticula".  [In  fact,  each  pore  is  a  short  tube  with  the  lower  part  projecting  into 
the  central  cavity,  rather  than  a  simple  pore  with  a  projecting  plate.]  Intervallum 
with  taeniae,  sparsely  porous  near  the  inner  wall,  coarsely  and  copiously  porous  else- 
where; the  taeniae  are  waved  in  the  radial  plane,  and  the  waves  have  angulated 
crests  and  troughs,  the  crests  and  the  trough-lines  curving  upwards  and  outwards 


FIG.  12.     Archaeocyathus  atlanticus  Billings X  4 


from  the  inner  wall ;  the  crests  of  neighbouring  taeniae  are  opposed  and  connected  by 
synapticulae.     Rare  dissepiments  may  occur.     No  tabulae  ". 

DISCUSSION.  R.  &  J.  Bedford  (1937  :  28)  and  Hill  (1965  :  123  and  128)  point  out 
that  Flindersicyathus  R.  &  J.  Bedford  and  Pycnoidocyathus  Taylor  have  a  very  similar 
structure.  The  latter  differs  only  in  having  much  stronger  transverse  annulations  of 
the  intervallum,  the  inner  wall  of  both  remaining  cylindrical.  The  two  genera  may 
be  synonymous,  but  I  have  not  seen  examples  of  the  type  species  of  Flindersicyathus, 
or  the  type  material  of  Pycnoidocyathus  and  therefore  cannot  settle  the  question.  My 
tentative  suggestion  is  that  Pycnoidocyathus  should  be  considered  a  subgenus  of 
Flindersicyathus,  due  to  its  different  outer  shape,  although  possibly  having  a  similar 
structure.  This  situation  is  comparable  to  the  distinction  between  Ajacicyathus  R.  & 


346  LOWER   CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA 

J.  Bedford  and  Orbicyathus  Vologdin,  which  are  considered  to  be  subgenera  distin- 
guished by  their  external  shape. 

Zhuravleva  (1960  :  296)  placed  Flindersicyathus  in  synonymy  with  Archaeocyathus 
Billings  1861.  Debrenne  (1964  :  117)  doubtfully  placed  Flindersicyathus  in  the  family 
Archaeocyathidae  (as  did  Hill  1965  :  123),  but  had  maintained  its  independence. 
However,  having  seen  the  holotype  of  Archaeocyathus  atlanticus  Billings  (1861,  fig.  10). 
I  can  state  that  the  taeniae  are  thickened  by  several  skeletal  layers,  have  few  and 
irregular  pores  (Text-fig.  12),  are  goffered  longitudinally  and  transversely,  and  are 
occasionally  connected  to  the  opposite  crests,  more  or  less  forming  elongated  pores. 
The  curved  hexagonal  tubular  mesh  of  Flindersicyathus  was  not  recognized.  Further 
characters  which  separate  the  two  genera  are  the  abundant  vesicular  tissue,  the  pore 
canals  of  the  inner  wall  and  the  irregular  net  of  the  outer  wall  of  Archaeocyathus. 

COMPOSITION  OF  THE  GENUS.  Flindersicyathus  (Flindersicyathus) :  F.  decipiens  R. 
&  J.  Bedford,  1937,  F.  circliporus  R.  &  J.  Bedford,  1937,  F.  contractus  Hill  1965,  F. 
graphicus  (R.  &  W.  R.  Bedford  1934),  F.  irregularis  (Taylor  1910),  F.  latiloculatus 
Hill  1965,  F.  major  (R.  &  W.  R.  Bedford  1934),  F.  multifidus  (R.  &  W.  R.  Bedford 
1936),  ?  F.  macdonnelli  R.  &  J.  Bedford  1937,  F.  rete  (R.  &  W.  R.  Bedford  1936), 
F.  simplex  (Taylor  1910),  F.  speciosus  (R.  &  W.  R.  Bedford  1934),  F.  tabulatus  R.  & 
J.  Bedford  1937. 

Flindersicyathus  (Pycnoidocyathus) :  F.  synapticulosus  (Taylor  1910),  F.  maximipora 
(R.  &  W.  R.  Bedford  1936),  F.  parvulus  (R.  &  W.  R.  Bedford  1936),  F.  ptychophragma 
(Taylor  1910),  F.  vicinisepta  (R.  &  W.  R.  Bedford  1936). 

\ 

Flindersicyathus  (Flindersicyathus)  graphicus  (R.  &  W.  R.  Bedford) 
(PI.  12,  figs.  3-5,  Text-fig.  13) 

1934     Protopharetra  graphica  R.  &  W.  R.  Bedford  :  4,  pi.  4,  fig.  22. 

1939     Dictyocyathus  graphicus  (R.  &  W.  R.  Bedford)  R.  &  J.  Bedford  :  73. 

1964     Metaldetes  graphica  (R.  &  W.  R.  Bedford)  Debrenne  :  220. 

HOLOTYPE.     B.M.  (N.H.).  S  4170. 
PARATYPES.     B.M.  (N.H.)  S  4171-4173. 
OTHER  MATERIAL.     B.M.  (N.H.)  S  4761,  S  7629. 

DESCRIPTION.  Slender  conical  cups,  undulating  exterior  form,  with  the  bulges  of 
the  inner  wall  following  those  of  the  outer  wall,  so  that  the  intervallum  remains  a 
constant  width.  Thin  outer  wall,  perforated  by  hexagonal  to  ellipsoidal  pores,  re- 
calling the  outer  wall  of  the  Regularia.  There  is  no  specific  inner  wall,  but,  instead, 
the  innermost  face  is  formed  by  the  inner  edges  of  the  taeniae  and  connecting  bars, 
which  enclose  apertures  that  are,  in  effect,  "  wall  pores  ". 

The  structure  of  the  intervallum  can,  perhaps  be  considered  as  an  assemblage  of 
septa  of  the  Volvacyathus-type,  with  regular,  somewhat  rounded,  hexagonal  pores  set 
in  a  radial  plane,  with  the  line  of  pores  curving  upwards  and  outwards  from  the  inner 
to  the  outer  wall.  These  pseudo-septa  (flat  taeniae)  are  joined  to  one  another  by 
horizontal  synapticulae  and  join  the  skeleton  at  each  pore  angle.  No  vesicular  tissue. 
Early  stages  of  the  species  unknown. 


LOWER   CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA 


347 


FIG.  13.     Flindersicyathus  (Flindersicyathus}  graphicus  (R.  &  W.  R.  Bedford) 


DIMENSIONS 


Cup: 

Height 

Diameter 

Intervallum 

Intervallum  coefficient 
Bars: 

Radial  distance 

Tangential  distance 

Vertical  distance 

Thickness 
Outer  wall : 

No.  of  pores  per 
intertaeniae 

Diameter  of  pores 

Partitions 

Thickness 
Inner  wall: 

No.  of  pores  per 
intertaeniae 

Diameter  of  pores 

Partitions 

Thickness 


S  4170 
(mm.) 

35 

12 

S  4171 
(mm.) 

45 

S  4172 
(mm.) 

15 
15 

sup. 
(mm.) 

7 

inf. 
(mm.) 

4 

S  7629 
(mm.) 

45 
20 

S  4761 
(mm.) 

30 
18 

1-9 

1-7 

2 

i  -5 

i  -13 

i  -5 

1  '5 

0-25 

0-17 

0-18 

o-3 

0-6 

o-  1 

O'  I 

0-26 

0-18 

o-34 

0-15 

0-15 

o-37 

0-22 

0-26 

0-26 

0-41 

O-22 

0-30 

0-30 

0-22 

o-37 

0-26 

o-37 

0-36 

O-22 

0-07 

0-07 

0-07 

0-07 

0-07 

0-07 

O-O7 

0-11-0-09 
0-03 

o-  ii 


I 
0-26 

0-13 

0-18 


3 

0-07 
0-03 
0-07 


i 

O-22 
0-07 
O-II 


o-37 

0-15 


0-05 


I 

0-18 

0-07 

O-II 


3 

o-ii 
0-05 
0-07 


0-18 
o-ii 


3 

o-ii 
0-07 
0-07 


0-18 
o-  ii 


0-07 


348  LOWER   CAMBRIAN    ARCH  AEOC  Y  ATH  A    FROM   AUSTRALIA 

DISCUSSION.  The  taeniae  (pseudosepta)  of  graphicus  are  not  waved  in  the  radial 
plane,  as  has  been  described  in  F.  decipiens  (R.  &  J.  Bedford)  probably  because  of  the 
narrowness  of  the  intervallum  and  the  density  of  the  taeniae  and  synapticulae.  The 
inner  wall  does  not  have  a  plate  on  the  lower  part  of  the  pore.  Thus  graphicus  is  much 
simpler  than  other  species  of  Flindersicyathus,  but  the  presence  of  well  defined  radial 
plates  (flat  taeniae)  indicates  a  more  complex  stage  than  that  of  having  sparse  rods  or 
plaquettes. 

The  species  was  first  described  as  Protopharetra,  subsequently  as  Dictyocyathus  and 
then,  in  1964,  removed  to  Metaldetes  because  of  the  pseudosepta.  However,  it  differs 
from  Metaldetes  in  having  a  very  regular  outer  wall,  numerous  synapticulae  and  the 
absence  of  connections  between  these.  The  scaffolding  of  the  intervallar  mesh  is  very 
like  that  of  Tabellaecyathus  Fonin  but  the  walls  are  simpler,  neither  tabello-reticulate 
nor  tabello-porous. 


Flindersicyathus  (Flindersicyathus}  irregularis  (Taylor) 
(PI.  14,  fig.  4) 

1910  Spirocyathus  irregularis  Taylor  :  148,  pi.  16,  figs.  93-94. 

1936  Spirocyathus  irregularis  Taylor;  R.  &  W.  R.  Bedford  :  14,  pi.  13,  fig.  64. 

1937  Flindersicyathus  irregularis  (Taylor)  R.  &  J.  Bedford  :  28. 
J937  Spirocyathus  atlanticus  Billings;  Ting  :  368-369,  pi.  13,  fig.  14. 

1961     Archaeocyathus  irregularis  (Taylor)  F.  &  M.  Debrenne  :  702,  pi.  20,  fig.  5. 

MATERIAL.     B.M.  (N.H.)  S  7625,  S  4763. 

DESCRIPTION.  Two-walled  cup  with  numerous  thin  radial  taeniae,  probably  quite 
porous  [a  deduction  made  from  Taylor's  comparison  with  F.  rete  in  his  original 
description  of  F.  irregularis  and  not  from  observation,  as  no  specimen  showing  a 
longitudinal  section  was  available],  wavy  and  connected  by  synapticulae  which 
form  a  network  of  more  or  less  regular  polygonal  apertures,  particularly  in  the  outer 
part.  Vesicular  tissue  sparse.  Skeletal  tissue  is  sometimes  thickened  by  secondary 
layers  and  occurs  mainly  near  the  inner  wall  (see  PI.  14,  fig.  4;  also  F.  &  M.  Debrenne, 
1961,  pi.  20,  fig.  5).  Thin  outer  wall,  with  numerous  small  pores;  inner  wall  with  one 
vertical  row  of  pores  per  intertaenial  space. 

DIMENSIONS 

Taylor  Ting  S  7625  S  4763 

(mm.)  (mm.)  (mm.)  (mm.) 
Cup: 

Height  (pars)                                         50  17  16  25 

Diameter                                                14  16-5  18  20 

Intervallum  coefficient                           0-5  0-8  0-8  0-9 

Intertaenial  space                                   0-5  0-4  ° '  45  ° '  5 
Intersynapticular  space 

Horizontal                                        . .  . .  0-70  0-6 

Vertical                                             . .  . .  irregular 

Taeniae  thickness                                   o  •  i  0-12  0-2  0-2 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A    FROM   AUSTRALIA  349 

DIMENSIONS — continued 

Taylor  Ting  S  7625  S  4763 

Outer  wall:  (mm-)  (mmO  (mm-)  (mm-) 

Diameter  of  pores  ..  0-2-0-4 

Partitions  ..  0-12 

Thickness  ..  0-12  0-13  0-15 

Inner  wall : 

Diameter  of  pores  . .  0-5  0-35  o  •  33 

Partitions  ..  0-3  0-27  0-3 

Thickness  . .  i  i  i 

DISCUSSION.  The  taeniae  are  thinner  and  more  numerous  than  in  other  species  of 
Flindersicyathus,  and  the  thickening  of  the  inner  wall  recalls  some  transverse  sections 
of  Copleicyathus  confertus  R.  &  J.  Bedford  (Hill  1965:  pi.  10,  figs.  4a-b),  but  at  the 
moment,  it  is  not  possible  to  take  the  comparison  any  further. 

Flindersicyathus  (Flindersicyathus)  major  (R.  &  W.  R.  Bedford) 

(PL  14,  fig.  2) 
1934     Spirocyathus  major  R.  &  W.  R.  Bedford  :  5,  pi.  4,  fig.  24. 

HOLOTYPE.     B.M.  (N.H.)  S  4174. 

DESCRIPTION.  Only  one  broken  specimen  known.  The  outer  wall  is  obscured  by 
silicification.  The  inner  wall  has  round,  shaft-like  pore  tubes,  that  are  limited  by 
neighbouring  taeniae  and  successive  vertical  synapticulae ;  the  lower  part  of  each  tube 
is  inclined  downwards  into  the  intervallum  and  projects  slightly  into  the  central 
cavity,  whereas  the  upper  part  is  flattened.  The  rows  of  pore-tubes  alternate.  The  pores 
are  more  crowded  at  the  base  of  the  cup,  because  the  taeniae  are  closer  together  there. 

The  intervallum  is  crossed  by  waved  taeniae,  with  the  crests  and  troughs  of  neigh- 
bouring taeniae  opposite  to  one  another.  Abundant  synapticulae  join  the  taeniae, 
particularly  in  the  central  and  outer  parts  of  the  intervallum.  The  ratio  diameter  of 
taenial  pores :  distance  apart  of  the  synapticulae  is  close  to  unity,  so  that  the  inter- 
vallum has  the  appearance  of  a  tubular  mesh  with  tubes  curving  upwards  and 
outwards. 

DIMENSIONS 

Cup:  (mm.) 

Height  (pars)     ......  40 

Diameter .......  about  20 

Intervallum       ......  10 

Interseptum       .          .          .          .          .          .  1-5 

Outer  wall : 

Thickness           .          .          .          .          .          .  0-20 

Inner  wall : 

Diameter  of  pores  :  horizontal     .          .          .  1-70 

vertical          .          .          .  i-I5 

Horizontal  partitions           .          .          .          .  i'i5 

Lateral  partitions       .          .          .          .          .  0-33 

Tubes : 

Irregular  diameter      .....  about  o  •  67 

Large  skeletal  plates 

Thickness           .          .          .          .          .          .  0-27 


350  LOWER   CAMBRIAN    ARCH  AEOC  Y  ATM  A    FROM   AUSTRALIA 

DISCUSSION.  The  septa  are  at  wider  intervals  than  in  other  species  of  Flinder- 
sicyathus. 

Flindersicyathus  (Flindersicyathus)  speciosus  (R.  &  W.  R.  Bedford) 

(PL  14,  figs,  i,  3) 
1934     Spirocyathus  speciosus  R.  &  W.  R.  Bedford  :  5,  pi.  4,  fig.  25. 

HOLOTYPE.     B.M.  (N.H.)  S  4175. 

DESCRIPTION.  Only  a  single  broken  specimen  known.  The  taeniae  are  thin,  radial, 
slightly  waved  and  connected  by  sparse  synapticulae,  which  are  chiefly  found  in  the 
outer  part  of  the  cup.  The  diameter  of  the  taenial  pores  is  less  than  the  width  of  the 
skeletal  tissue  between  them. 

Outer  wall  with  4  to  5  alternating  vertical  rows  of  round  pores  per  interseptum. 
The  inner  wall  has  a  single  tube  per  intertaenial  space,  the  lower  part  of  which  projects 
into  the  central  cavity,  giving  the  tube  a  crescentic  section.  The  rows  alternate. 

DIMENSIONS 

Cup:  (mm.) 

Height  (pars)     ......  30 

Upper  diameter           .....  about  13 -5 

Lower  diameter           .....  1 1 

Intervallum       .          .          .          .          .          .  4-5 

Intervallum  coefficient         ....  i 

Intertaenial  space ;  lower  end  of  cup     .          .  0-6 

upper  part  of  cup  .          .  o  •  75 

Outer  wall : 

Diameter  of  pores       .          .          .          .          .  0-15 

Vertical  skeletal  partitions .          .          .          .  0-07 

Horizontal  skeletal  partitions       .          .          .  0-18 

Inner  wall : 

Diameter  of  pores ;  horizontal      .          .          .  0-9 

vertical           .          .          .  0-4 

Vertical  skeletal  partitions .          .          .          .  0-13 

Crescentic  plates         .          .          .          .          .  0-33 

DISCUSSION.  The  taeniae  are  less  waved  and  not  so  distinct  as  those  in  Flinder- 
sicyathus major.  The  shaft-like  tubes  of  the  latter  are  not  present,  while  the  inner 
wall  pores  of  speciosus  are  crescentic  and  very  close  together.  Synapticulae  are  less 
numerous  than  in  other  Flindersicyathus  species. 

Subgenus  FLINDERSICYATHUS  (PYCNOIDOCYATHUS)  (Taylor),  1910 

1910    Pycnoidocyathus  Taylor  :  132. 

1939     Pycnoidocyathus  Taylor;  R.  &  J.  Bedford  :  78. 

1965     Pycnoidocyathus  Taylor;  Hill  :  128 

TYPE  SPECIES.  P.  synapticulosus  Taylor  (1910  :  132)  by  subsequent  designation 
R.  &  J.  Bedford,  (1939  :  78) ;  type  material  in  the  Univ.  of  Adelaide. 

DISCUSSION.  Although  Hill  (1965  :  128)  has  provisionally  treated  this  as  a  genus 
separate  from  Flindersicyathus,  I  regard  it  as  a  subgenus  (see  p.  345).  It  is  very  like 
Flindersicyathus,  but  with  much  stronger  expansions  and  contractions  of  the  inter- 


LOWER   CAMBRIAN    ARCH  AEOCY  ATM  A   FROM   AUSTRALIA  351 

vallum  which  do  not  affect  the  inner  wall.  The  problem  can  only  be  settled  by  exami- 
nation of  the  type  material,  in  order  to  discover  whether  dissepiments  occur  and  the 
exact  nature  of  the  inner  wall  pores. 

Flindersicyathus  (Pycnoidocyathus)  synapticulosus  (Taylor) 

(PI.  15,  fig-  i) 

1910    Pycnoidocyathus  synpaticulosus  Taylor  :  132,  pi.  12,  fig.  69. 

1936    Pycnoidocyathus  synapticulosus  Taylor;  R.  &  W.  R.  Bedford  :  15,  pi.  15,  fig.  69. 

HOLOTYPE.     Probably  at  Adelaide  University. 

MATERIAL.     B.M.  (N.H.)  S  208,  S  4825. 

DESCRIPTION.  Large  cup  with  annular  bulges.  On  the  specimens  examined  here, 
the  outer  part  of  each  bulge  is  made  of  successive  flanges.  Outer  wall  with  somewhat 
irregular  polygonal  fine  pores.  Inner  wall  with  a  vertical  series  of  short  tubes  per 
intertaenial  space,  the  tubes  lead  upwards  into  the  central  cavity.  Taeniae  are 
radial,  nearly  flat  and  close  together,  particularly  near  the  synapticulae,  so  that 
transverse  sections  appear  to  be  composed  of  irregular  polygonal  cells.  Abundant 
horizontal  and  vertical  synapticulae ;  generally  arranged  in  quincunx  from  one  inter- 
sept  to  the  next.  It  was  not  possible,  to  examine  the  size  and  arrangement  of  pores 
in  tangential  section,  but  Taylor's  description  (1910  :  132)  and  Bedford's  figure  (1936, 
fig.  69)  confirm  my  observations  from  transverse  sections  which  suggest  the  pores  are 
small,  isolated  and  without  any  important  modifications  in  the  lateral  bulges.  No 
vesicular  tissue. 

DIMENSIONS 

Cup:  (mm.) 

Diameter      .......  about  85 

Intervallum 

without  bulges  ......  8 

largest  part        .          .          .          .          .          .  19 

Interseptum           .          .          .          .          .          .  1-07 

Synapticulae 

horizontal           ......  1-2 

vertical     .......  0-6-2 

Outer  wall: 

No.  of  pores  per  interseptum.          ...  4 

Diameter  of  pores           .          .          .          .          .  0-13 

Partitions     .          .          .          .          .          .          •  0-06 

Inner  wall : 

No.  of  pores  per  interseptum  ....  i 

Diameter  of  pores           .          .          .          .          .  ° '  33 

Horizontal  partitions     .          .          .          .          .  0-67 

Thickness 0-67 

Taeniae : 

Thickness     .          .          .          .          .          .          .  o-i 

DISCUSSION.  This  is  the  largest  species  of  Flindersicyathus  (Pycnoidocyathus) 
known.  It  has  regular  annular  bulges,  the  skeletal  tissue  of  the  wavy  taeniae 
occupies  a  greater  area  than  the  pores,  and  the  synapticulae  are  more  numerous  than 
in  any  other  species  of  this  genus. 


352  LOWER   CAMBRIAN    ARCH  AEOC  Y  ATH  A    FROM    AUSTRALIA 

Flindersicyathus  (Pycnoidocyathus)  simplex  (Taylor) 
(PL  15,  fig.  2) 

1910     Pycnoidocyathus  simplex  Taylor  :  134,  pi.  2,  fig.  70. 

1936     Pycnoidocyathus  simplex  Taylor;  R.  &  W.  R.  Bedford  :  15,  pi.  15,  fig.  70. 

HOLOTYPE.     Probably  in  the  South  Australian  Museum. 
MATERIAL.     B.M.  (N.H.)  S  4824. 

DESCRIPTION.  Broken  piece  showing  well-preserved  inner  wall.  Taeniae  are 
radial,  not  waved  and  are  joined  with  the  synapticulae,  chiefly  towards  the  exterior. 
Outer  wall  is  only  see  on  part  of  a  transverse  section,  where  a  small  weathered-out 
surface  shows  that  the  pores  are  arranged  in  irregular  quincunx.  Inner  wall  has  a 
vertical  row  of  pores  per  intersept,  each  pore  having  a  raised  lower  edge  and  is 
vertically  elongated  but  flattens  towards  the  top.  The  rows  of  pores  alternate  in 
quincunx. 

DIMENSIONS 

(mm.) 

Cup: 

Height  (pars)     .....  40 

Diameter ......  about  22 

Intervallum       .          .          .          .          .  5-5 

Central  cavity      .....  13 

Intervallum  coefficient         .          .          .  0-4 

Intertaenial  space       ....  i 

Outer  wall : 

Diameter  of  pores       ....  0-40-0-33  (hor.) 

Skeletal  partitions      ....  0-20-0-20 

Thickness           .          .          .          .          .  0-13 

Inner  wall: 

Diameter  of  pores       ....  i -69-1 -42  (hor.) 

Vertical  partitions      .                               .  0-20 

Horizontal  partitions           .  o  •  47 

Taeniae : 

Diameter  of  pores       ...  o«6 

Vertical  partitions      .          .          .          .  0-13 

Horizontal  partitions           .          .          .  0-20 

Thickness           .          .          .          .          .  o  •  i 

DISCUSSION.  The  measurements  of  the  specimen  described  above  correspond  to 
those  given  by  Taylor  (1910).  R.  &  W.  R.  Bedford  (1936  :  15,  fig.  70)  pointed  out 
and  sketched  the  regular  annular  bulges  from  an  example  of  the  same  species.  F.  (P.) 
simplex  differs  from  typical  Pycnoidocyathus  by  the  scarcity  of  its  synapticulae  and 
the  straight  taeniae. 

Flindersicyathus  (Pycnoidocyathus)  maximipora  (R.  &  W.  R.  Bedford) 

(PI.  15,  fig.  3) 

1934     Pycnoidocyathus  maximipora  R.  &  W.  R.  Bedford  :  3,  pi.  2,  figs.  ga-c. 
1936     Pycnoidocyathus  maximipora  R.  &  W.  R.  Bedford  :  15,  pi.  15,  fig.  71. 

HOLOTYPE.     B.M.  (N.H.)  S  4150. 


LOWER   CAMBRIAN    ARCH  AEOC  Y  ATM  A   FROM   AUSTRALIA  353 

DESCRIPTION.  One  broken  specimen  with  a  single  annulate  bulge.  Intervallum 
crossed  by  un-waved  radial,  porous  taeniae,  with  the  skeletal  tissue  forming  a  greater 
area  than  the  pores,  and  sparse  synapticulae,  that  occur  mainly  in  the  lower  part  of 
the  fragment.  The  taenial  pores,  are  round  and  arranged  in  lines,  which  curve 
upwards  and  outwards,  near  the  inner  wall;  but  are  oval  and  bigger  near  the  outer 
wall  and  inside  the  annular  bulge.  The  thin,  irregular,  outer  wall  is  strengthened  by 
bars  springing  from  the  taeniae,  but  both  partial  weathering  and  preservation  give 
an  irregular  and  false  idea  of  its  original  structure. 

The  inner  wall  is  composed  of  very  short  piled-up  pipes,  leading  obliquely  upward, 
with  one  row  per  intertaenial  space,  which  alternates  with  the  next.  The  cross-sec- 
tion of  the  pipe  is  vertically  oval.  Each  pipe  is  withdrawn  from  the  one  immediately 
above  and  its  sides  are  stretched  slightly  to  join  with  its  neighbours.  Thus,  the 
inner  wall  does  not  present  a  flat  surface  to  the  central  cavity  but  has  the  appearance 
of  a  rasp.  The  various  sections  have  suggested  to  previous  authors  that  there  is  a 
louvre-like  plate  at  the  lower  part  of  each  pore,  but,  in  fact,  these  are  the  short  pipes. 

DIMENSIONS 

(mm.) 

Cup: 

Height  (pars)  35 

Diameter  22 

Intervallum  6 

Intertaenial  space  i  •  3 

Outer  wall: 

Diameter  of  the  pores  o  •  4 

Partitions  o  •  3 

Thickness  o  •  4 

Inner  wall: 

Diameter  of  the  pores  i  -49-2 '  71 

Lateral  partitions  0-13 

Horizontal  partitions  2  •  03 

Taeniae : 

Diameter  of  the  pores  near  the  inner  wall  o  •  6 

near  the  outer  wall  o  •  8-2  •  35 

Partitions  0-27-0-40 

DISCUSSION.  This  species  differs  from  others  by  the  greater  development  of 
tubes  in  the  inner  wall  and  its  relatively  narrower  taenial  pores. 

F Under sicyathus  (Pycnoidocyathus)  vicinisepta  (R.  &  W.  R.  Bedford) 

(PI.  15,  fig.  4) 

1936     Pycnoidocyathus  vicinisepta  R.  &  W.  R.  Bedford  :  15,  pi.  16,  fig.  72. 

HOLOTYPE.     Probably  in  the  South  Australian  Museum. 

MATERIAL.     B.M.  (N.H.)  S  4825. 

DESCRIPTION.  A  longitudinally  cut  fragment.  Several  annulate  horizontal  bulges 
are  visible,  they  are  rather  narrow  and  close  together.  Radial  un-waved  taeniae 


354  LOWER   CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA 

joined  by  numerous  synapticulae.  In  the  bulges,  the  taeniae  are  waved  with  opposite 
crests  connected  by  synapticulae,  so  that  the  intervallum  has  the  aspect  of  a  polygonal 
mesh.  A  very  small  part  of  the  outer  wall  is  preserved  in  which  the  pores  are  irregular 
and  polygonal.  The  inner  wall  has  one  vertical  row  of  pore-tubes  leading  up  into  the 
central  cavity,  in  each  intertaenial  space.  The  rows  alternate. 

DIMENSIONS 

(mm.) 

Cup: 

Height  (pars — in  two  pieces)  95 

Diameter  22 

Bulge  4 

Intervallum  without  bulge  4 

Central  cavity  14 

Intertaenial  space  o  •  33 

Intersynapticular  space  o  •  47-0  •  67 

Outer  wall : 

Diameter  of  pores  o  •  06 

Skeletal  tissue  o  •  05 

Thickness  not  known 

Inner  wall : 

No.  of  rows  of  pores  per  intertaenial  space  i 

Horizontal  partitions  0-27 

Vertical  partitions  o  •  40 

Diameter  of  pores  o  •  40 

Thickness  0-40 

Taeniae : 

Diameter  of  pores  0-27 

Horizontal  partitions  0-81 

Vertical  partitions  o  •  67 

Thickness  o-io 

DISCUSSION.  As  far  as  I  can  tell,  after  measuring  the  drawings  of  Bedford,  the 
coefficients  correspond  to  those  of  vicinisepta.  This  species  differs  from  other 
Pycnoidocyathus  in  having  many  more  taeniae. 

Family  METAGYATHIDAE  R.  &  W.  R.  Bedford,  1934 

DIAGNOSIS.  Cups  growing  from  an  apex  without  central  cavity,  occupied  by  rods, 
plates  and  dissepiments.  Adult  stages  with  more  or  less  definite  radial  septa,  the 
pores  of  which  are  arranged  in  rows  inclined  upwards  and  outwards  from  inner  to 
outer  wall.  Dissepiments  and  sometimes  synapticulae  present.  Simple  porous 
outer  and  inner  walls,  with  the  pores  sometimes  screened  by  a  microporous  sheath  or 
pellis. 

COMPOSITION  OF  THE  FAMILY.  Protopharetra  Bornemann  1887,  Volvacyathus 
Debrenne  1961,  Dendrocyathus  Okulitch  &  Roots  1947,  ?  Shidertycyathus  Krasnopeeva 
1959,  Metaldetes  Taylor  1910,  Cambrocyathus  Okulitch  1937,  Cambrocyathellus 
Zhuravleva  1960,  Okulitchicyathus  Zhuravleva  1960,  Pamnacyathus  R.  &  J.  Bedford 
1937,  ?  Ardrossacyathus  R.  &  J.  Bedford  1937,  Metafungia  R.  &  W.  R.  Bedford  1934. 


LOWER   CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA  355 

Genus  METALDETES  Taylor,  1910 

1910     Metaldetes  Taylor  :  151,  pi.  15,  figs.  86-88,  t.-figs.  n,  37  and  38. 
1934     Metacyathus  R.  &  W.  R.  Bedford  :  5. 
?i957     Bedfordcyathus  Vologdin  :  182  and  209. 

TYPE  SPECIES.  By  monotypy  Metaldetes  cylindricus  Taylor  1910,  the  holotype  of 
which  is  in  the  University  of  Adelaide. 

DIAGNOSIS.  Solitary,  or  sometimes  colonial  cups;  the  central  cavity  and  the  inner 
wall  slowly  develop  later.  In  the  lower  part  of  the  cup,  the  irregularly  arranged 
skeletal  structures  (bars,  rods,  plates  and  vesicular  tissue)  fill  the  whole  inner  space. 
Much  later,  the  central  cavity  is  defined  by  the  formation  of  a  regularly  porous  inner 
wall  and  the  structures  of  the  intervallum  are  arranged  in  radial  plates ;  the  tangential 
links  (synapticulae  and  dissepiments)  may  continue  into  the  adult  stage. 

Unfortunately,  the  type  specimen  of  the  genus  has  not  been  re-described  yet  and 
one  has  to  rely  on  the  original  account  and  illustrations.  According  to  Taylor 
(1910  :  151),  the  rugose  character  of  the  outer  wall  causes  a  transverse  section  to 
resemble  a  series  of  tridents.  R.  &  W.  R.  Bedford  (1934  :  5)  considered  the  outer  wall 
to  be  a  double  porous  sheath,  and  my  own  observations  agree  with  this  (Debrenne 
1964  :  219).  Hill  (1965  :  119)  mentions  that  the  longitudinal  ribs  between  the  rows 
of  pores  are  connected  together,  some  distance  behind  the  outer  edge,  by  transverse 
bars.  The  inner  wall,  which  is  absent  in  the  lower  part  of  the  cup,  was  described  by 
Taylor  (1910  :  152)  as  being  "  strongly  ridged  on  its  septal  side  ".  These  ridges 
correspond  to  the  beginnings  of  the  taeniae,  which  are  slightly  thickened  towards  the 
inner  wall.  The  number  of  pores  per  intersept  is  not  certain  (i,  2,  or  3),  nor  their 
shape  and  size. 

DISCUSSION.  A  complete  revision  of  the  type  material  is  desirable,  but  although 
Metaldetes  is  incompletely  known,  it  is  possible  to  regard  two  subsequent  genera, 
Metacyathus  Bedford  and  Bedfordcyathus  Vologdin,  both  found  at  the  same  locality 
and  level,  as  synonyms.  Okulitch  (1955  :  E  16),  Zhuravleva  (1960  :  283)  and  myself 
(Debrenne  1964  :  220)  have  previously  considered  Metacyathus  a  synonym  of  Metal- 
detes, but  Vologdin  (1957  :  43)  and  Hill  (1965  :  118)  thought  it  to  be  a  separate  genus. 
R.  &  W.  R.  Bedford  established  the  genus  to  include  those  species  in  which  the  taeniae 
occur  as  straight  radial  septa  in  the  inner  two-thirds  of  the  intervallum,  but  are 
separate  from  the  outer  wall,  as  in  the  type  M .  taylori  R.  &  W.  R.  Bedford 

Examination  of  the  holotype  of  M .  taylori  shows  that  the  structures  of  the  inter- 
vallum are  disturbed  by  the  occurrence  of  exothecal  lamellae,  but  that,  at  other 
levels,  the  plates  continue  from  one  wall  to  the  other.  The  porosity  of  the  outer  wall 
depends  on  the  presence  or  absence  of  the  exotheca;  the  inner  wall  has  one  vertical 
row  of  pores  per  intertaenial  space  in  the  lower  part  of  the  cup,  but  two  or  more  in 
the  upper  part,  with  probably  a  second  wall. 

Bedfordcyathus  was  established  by  Vologdin,  after  he  had  studied  Bedford's  figures, 
to  group  into  a  separate  genus  those  species  that  have  a  strong  development  of  vesi- 
cular tissue.  The  holotype  of  the  type  species  (M.  irregularis  Bedford),  now  in  the 
British  Museum  (Natural  History),  is  described  below.  It  shows  that  there  is  no 


356  LOWER  CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA 

appreciable  difference  from  Metacyathus,  in  the  structure  of  the  intervallum  (apart 
from  the  abundance  of  vesicular  tissue),  or  in  that  of  the  walls.  I  think  that  vesicular 
tissue  is  too  inconstant  and  variable  a  character  to  be  sufficient  grounds  for  establish- 
ing a  new  genus. 

The  genus  Metaldetes  is  therefore  denned  as  consisting  of  cups  which  grow  from  a 
base  that  does  not  have  a  distinct  central  cavity,  where  the  skeletal  elements  have 
not  attained  a  clear  radial  arrangement,  and  where  tangential  links  (synapticulae  and 
vesicular  tissue)  are  numerous.  Subsequently,  the  central  cavity  is  differentiated  by 
a  more  definite  development  of  the  inner  wall,  which  is,  at  first,  part  of  the  intervallum 
(with  one  pore  per  intertaenial  space),  but  then  becomes  a  more  distinct  structure 
(with  two  pores  per  intertaenial  space)  and  is,  perhaps,  protected  on  its  central  cavity 
side  by  a  microporous  sheath  or  vesicular  membrane.  At  this  same  level,  the  inter- 
vallar  elements  generally  form  strong,  compact,  radial  taeniae  with  high  oval  pores 
the  diameter  of  which  increases  along  upwardly  curving  lines  from  the  inner  to  the 
outer  wall;  the  largest  pores  being  close  to  the  outer  wall.  It  is  in  this  region  that 
positional  changes  and  external  influences  occur,  the  septa  become  less  regular  and 
sometimes  revert  to  curved  taeniea.  The  synapticular  links  may  persist  but  generally 
disappear  in  the  adult  stage,  whereas  vesicular  tissue  is  still  abundant.  The  outer 
wall  has  a  basic  layer  with  irregular  polygonal  pores,  overlapped  by  a  second  micro- 
porous  sheath,  which  disappears  when  the  cup  is  surrounded  by  exothecal  lamellae. 
Fossils  reach  considerable  size. 

RELATIONSHIPS  AND  DIFFERENCES 

Volvacyathus  Debrenne  1961  has  two  distinct  stages,  the  apex  being  quite  different 
from  the  adult.  The  walls  are  still  connected  with  the  intervallar  network,  the  septa 
are  not  clearly  denned  and  their  pores  are  wider  than  the  intervening  skeletal  tissue. 
This  genus  is  also  close  to  Protopharetra  Bornemann  1887. 

Cambrocyathus  Okulitch  1937  closely  resembles  Metaldetes  in  having  abundant 
vesicular  tissue,  but  the  adult  stage  is  reached  more  quickly.  The  central  cavity  is 
not  free  and  is  filled  by  thin  skeletal  rods  and  vesicular  tissue,  which  persists  as  a 
pellis  to  the  top  of  the  cup.  These  structures  are  also  found  as  an  exotheca.  The 
well-developed  taeniae  generally  have  a  laminated  structure  and  are  perforated  by 
pores,  that  are  almost  in  horizontal  lines.  Scarce  synapticulae  or  branching  septa, 
but  abundant  vesicular  tissue,  which  earlier  authors  have  interpreted  as  true  synap- 
ticulae. Reference  material  on  loan  from  the  Geological  Survey  of  Canada  and  a 
topotype  from  the  Yale  Peabody  Museum  showed  these  features  (Text-fig  14). 

Septa  of  Cambrocyathus-type  (see  Debrenne  1964  :  88)  are  the  final  stage  in  the 
process  of  radial  partition.  This  starts  with  Protopharetra,  leads  through  Volvacya- 
thus (see  Debrenne  1964,  fig.  51)  and  the  first  Metaldetes  (proteiis  and  dissutus),  with 
their  broken  septa,  to  the  complete  septa  of  Metaldetes  irregularis  and  finally,  to  the 
regular  septa  of  Cambrocyathus. 

COMPOSITION  OF  THE  GENUS.  M.  cylindricus  Taylor  1910,  M.  columbianus 
(Okulitch  1943),  M.  dissepimentalis  (Taylor  1910),  M.  dissutus  Debrenne  1964,  M. 
irregularis  (R.  &  W.  R.  Bedford  1934),  M.  ramulosus  R.  &  J.  Bedford  1937,  M.  solidus 


LOWER   CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA  357 


FIG.  14.     Cambrocyathus  profundus  (Billings)  X4 


358 


LOWER  CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA 


(Okulitch  1957),  M.  spiralis  R.  &  W.  R.  Bedford  1936,  M.  superbus  R.  &  W.  R. 
Bedford  1936,  M.  taylori  (R.  &  W.  R.  Bedford  1934),  M.  proteus  (Bornemann  1887). 


Metaldetes  dissepimentalis  (Taylor) 
(PI.  16,  figs.  1-3) 

1910     Archaeocyathus  dissepimentalis  Taylor  :  128,  pi.  10,  fig.  53. 
1934     Metaldetes  conicus  R.  &  W.  R.  Bedford  :  5,  figs  26,  28  and  31. 
1936     Metaldetes  conicus  R.  &  W.  R.  Bedford  :  18,  pi.  18,  fig.  77. 

HOLOTYPE.     Possibly  at  Adelaide  University. 
MATERIAL.     B.M.  (N.H.)  S  4176-4182. 

DESCRIPTION.  Large  conical  cups.  In  the  first  stages,  the  central  cavity  is  not 
formed  and  the  entire  cup  is  occupied  by  un-orientated  taeniae,  which  are  joined  by 
numerous  dissepiments  that  are  arranged  as  horizontal  vesicles.  Much  later,  the 
cavity  develops,  the  dissepiments  become  less  abundant  and  the  taeniae  form  regular 
radial  plates.  The  general  aspect  recalls  that  of  Cambrocyathus  profundus  Billings. 

The  outer  wall  is  twofold:  a  basal  wall  covered  externally  by  a  very  thin  micro- 
porous  sheath.  The  inner  wall  is  also  double;  the  main  wall  having  two  rows  of 
large  irregular  pores  per  intersept,  with  the  diameter  of  the  pores  being  much  greater 
than  the  width  of  the  skeletal  tissue  between  them;  with  a  microporous  sheath, 
similar  to  that  of  the  outer  wall,  occurring  on  the  central  cavity  side.  The  micro- 
porous  sheaths  are  not  well  known  for  they  are  often  eroded  and  no  good  tangential 
sections  have  been  seen.  Consequently,  the  arrangement,  size  and  shape  of  the 
micropores  is  not  certain,  they  appear  to  be  similar  to  those  of  the  secondary  wall  in 
Metafungia. 

DIMENSIONS 


Cup: 

Height  (pars) 

Diameter 

Intervallum 

Interseptum 

Intervallum  coefficient 
Outer  wall: 

No.  of  rows  of  pores  per 
interseptum 

Diameter 

Skeletal  partitions 

Micropores:  diameter 

skeletal  partitions 

Thickness 


S  4176 
(mm.) 

60 

24 
6 
0-4-0-7 


S  4178 

(mm.) 


13 
4 

o-5 
0-8 


S  4179  S  4180 
S  4180   up. 
low.   S  4181 

(mm.) 


(mm. 


13 


0-7 


15 

21 

6 
0-6 


S  4182 
(mm.) 

40 

45 
7'5 
i-35 
0-25 


dissepimentalis 
Taylor 
(mm.) 


22 
6 

7 
0-6 


I- 

-3 

1-3 

2 

2 

1-3 

2 

0 

•  17 

0'33 

O 

•2 

O 

•20 

O' 

•25-0 

•40 

O 

•30 

0 

•30 

O-2O 

O 

•15 

O 

•15 

O' 

•15-0 

•20 

O 

•20 

0 

°3 

O 

•06 

O 

•O6 

. 

. 

o 

•06 

O 

06 

.  . 

. 

. 

O-2O 


0-20 


LOWER  CAMBRIAN   ARCHAEOCYATH A   FROM   AUSTRALIA 

DIMENSIONS — continued 


359 


Inner  wall: 

No.  of  pore  rows  per 
interseptum 

Diameter 

Skeletal  partitions 

Micropores:  diameter 

skeletal  partitions 

Thickness 
Septa : 

Diameter 

Skeletal  partitions 

Thickness 


S  4176 

S  4178 

84179 
S  4180 
low. 

S  4180 
up. 
S  4181 

(mm.) 

(mm.) 

(mm.) 

(mm.) 

2 

2 

•• 

i-3 

0-67 
0-16 

0-47 
0-16 

•• 

o-io 

0-40 


0-13 


0-03 
0-03 


0-13 


S  4182 
(mm.) 
1-2 

o-33 
o-io 

0-03 
0-03 


0-40 
o • 40-0 • 60 

O-2O 


dissepimentalis 
Taylor 
(mm.) 


1-2 
0-30 

o-io 
0-03 
0-03 


DISCUSSION.  The  description  and  figures  given  by  Taylor  for  Archaeocyathm 
dissepimentalis  correspond  with  measurements  taken  from  the  photographs  of 
Metaldetes  conicus  Bedford.  The  two  species  are,  therefore,  considered  to  be  synony- 
mous and  dissepimentalis,  the  older,  is  the  name  conserved. 


Metaldetes  irregularis  (R.  &  W.  R.  Bedford) 
(PL  16,  fig.  4) 

1934  Metacyathus  irregularis  R.  &  W.  R.  Bedford  :  6,  pi.  5,  fig.  29. 

1957  Bedfordcyathus  irregularis  (Bedford)  Vologdin  :  43. 

1964  Metacyathus-Bedfordcyathus  irregularis  (R.  &  W.  R.  Bedford);  Debrenne  :  220  and  231. 

1965  Bedfordcyathus  irregularis  (R.  &  W.  R.  Bedford);  Hill  :  118,  fig.  22.9. 

It  has  not  been  possible  to  locate  the  original  description  by  Vologdin  of  his  genus 
Bedfordcyathus.  Both  Debrenne  (1964)  and  Vologdin  (1957,  1962)  give  the  year  1955 
for  this  genus,  but,  unfortunately,  there  is  no  mention  of  Bedfordcyathus  in  the  paper 
cited  under  that  year,  in  either  bibliography  (Dokl.  Akad.  Nauk  SSSR,  103,  i).  In 
subsequent  papers,  Vologdin  gives  the  year  as  1956,  but  the  only  paper  available  for 
that  year  merely  includes  the  genus  in  a  list  showing  the  classification  of  the  Archaeo- 
cyatha.  This  paper  appeared  in  a  slightly  more  elaborate  form  during  1957,  in  two 
journals:  Ada  paleont  sinica,  5:  173-222  and  Annls.  Cent.  Etud.  Docum  paleont., 
23  :  33-80  and  it  is  the  latter,  which  is  taken  as  the  basis  of  the  date  given  in  this 
paper  and  that  of  Hill  (1965).  However,  it  would  appear  that  the  paper  referred  to, 
is  included  in  the  yet  unpublished  Vol.  3  of  "  El  sistema  Cambrico  .  .  .  ",  Int.  Geol. 
Congr.  Mexico,  1955,  quoted  in  some  references  as  1961. 

HOLOTYPE.     B.M.  (N.H.)  S  4189,  by  monotypy. 

OTHER  MATERIAL.     Paratypes  B.M.  (N.H.)  S  4188,  S  4190. 

DESCRIPTION.  Conical  cup,  with  large  irregularly- waved  intervallum.  Double 
outer  wall;  first,  a  thick  basal  one,  pierced  by  funnel-shaped  pores  that  widen  towards 


GEOL.  17,  7 


28 


360  LOWER   CAMBRIAN   ARCH AEOCYATH A   FROM   AUSTRALIA 

the  exterior ;  this  is  covered  externally  by  a  very  thin  microporous  sheath.  The  inner 
wall  is  not  well  known,  but  from  a  small  poorly  preserved  surface,  appears  to  be 
covered  by  an  irregular,  microporous,  thin  wall,  which  screens  the  simple  pores  of  the 
intervallum. 

The  taeniae  in  the  upper  part,  are  straight,  radial  and  completely  cross  the  inter- 
vallum.  In  longitudinal  section,  the  pores  appear  circular  and  alternate  in  regular 
lines,  curving  shallowly  towards  the  exterior  from  the  inner  to  the  outer  wall.  The 
skeletal  tissue  and  the  pores,  occupy  equal  proportions  of  the  total  surface  area. 
Considerable  development  of  vesicular  tissue,  formed  of  imbricate,  horizontally 
elongated  vesicles,  that  cross  several  loculi,  but  do  not  penetrate  into  the  central 
cavity. 

DIMENSIONS 

(mm.) 

Cup: 

Height  (pars)  92 

Diameter  about  35 

Intervallum  10 

Intervallum  coefficient  0-6 

Interseptum  i'35 

Outer  wall : 

No.  of  rows  of  pores  per  interseptum  2 

Diameter  of  pores  0-27 

Partitions  0-20-0-40 

Micropores  o-io 

Inner  wall: 

Diameter  of  pores  o  •  35-0  •  67 

Partitions  0-13 
Micropores                                                        4  to  each  main  pore 

Diameter  of  micropores  o  •  05 

Thickness  o  •  06 

Taeniae : 

Diameter  of  pores  o  •  33-0  •  54 

Partitions  0-27-0-33 

DISCUSSION.  The  strong  development  of  vesicular  tissue  and  the  absence  of  exo- 
thecal  lamellae,  are  the  main  differences  between  this  species  and  others  in  the  genus. 

Metaldetes  taylori  (R.  &  W.  R.  Bedford) 
(PI.  13,  figs.  1-3) 

1934  Metacyathus  taylori  R.  &  W.  R.  Bedford  :  5,  pi.  5,  fig.  10. 

1936  Metacyathus  taylori  R.  &  W.  R.  Bedford;  R.  &  W.  R.  Bedford  :  18,  pi.  18,  fig.  80. 

1964  Metaldetes  taylori  (R.  &  W.  R.  Bedford)  Debrenne  :  220. 

1965  Metacyathus  taylori  R.  &  W.  R.  Bedford;  Hill  :  118,  pi.  9,  figs.  2a-d. 

HOLOTYPE.  By  monotypy,  B.M.  (N.H.)  S  4185-4187  (three  pieces  of  the  same 
specimen) . 

DESCRIPTION.  Although  the  three  pieces  are  said  to  come  from  the  same  cup,  it  is 
no  longer  possible  to  prove  this  with  any  certainty,  for  having  been  subjected  to 
various  palaeontological  techniques  (cutting,  etching  etc.),  they  do  not  fit  together. 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A   FROM   AUSTRALIA 


361 


Nevertheless,  there  is  little  chance  of  the  lower  part  belonging  to  another  specimen. 
This  is  pointed  out  because  the  lower  part  has  some  structures  obviously  different 
from  the  other  two  pieces  and  these  are  important  for  generic  determination. 

The  lower  part  is  conical ;  radial,  slightly  porous  taeniae  are  connected  by  synapti- 
culae,  which  develop  between  the  pores.  The  inner  wall  has  a  single  row  of  pores  per 
interseptum.  The  outer  wall  is  well  denned  and  separates  the  intervallum  from  the 
exothecal  lamellae.  Dissepiments  present. 

In  the  middle  and  upper  parts  of  the  cup,  the  inner  wall  has  two  rows  of  pores  per 
interseptum;  the  outer  wall  is  not  yet  separate  from  the  exothecal  structures,  which 
disturb  the  outer  part  of  the  intervallum  and  give  the  septa  the  form  of  curved  taeniae. 
No  synapticulae,  but  dissepiments  persist.  The  pseudo-septa  are  mainly  radial  at 
the  inner  and  middle  parts  of  the  intervallum,  but  are  interrupted  towards  the  outer 
wall  and  disappear  near  the  exothecal  lamellae. 

DIMENSIONS 

84185 
Apex 


Cup: 

Height  (pars) 
Diameter 

Intervallum 
Intervallum  coefficient 

Interseptum 
Outer  wall : 

Thickness 
Inner  wall : 

No.  of  pore  rows  per  interseptum 

Diameter 

Skeletal  partitions 

Thickness 
Taeniae 

Diameter  of  pores 

Thickness 


(mm.) 

35 
16 


0-72 


0-06 


S  4186 
(mm.) 


55 

35 
24 
ii 

7 

O' 

o 


upper 
lower 
upper 
lower 
upper 
lower 


S  4187 
(mm.) 

40 
35 


disappeared 

2 
o-33 

O-2O 

? 

o • 40-0 • 60 
0-40 

0-27 


disappeared 


2 
o-33 

O-2O 

? 

O-6o 
0-40 
O-25 


DISCUSSION.  Metaldetes  taylori  (R.  &  W.  R.  Bedford)  differs  from  M.  irregular-is 
(R.  &  W.  R.  Bedford)  by  its  narrow  central  cavity,  weak  development  of  vesicular 
tissue  and  the  abundance  of  exothecal  lamellae.  It  is  little  different  from  M. 
dissepimentalis  (Taylor),  which  has  straighter  septa,  a  well-defined,  double-porous 
outer  wall,  but  is  without  any  exothecal  lamellae. 


Genus  METAFUNGIA  R.  &  W.  R.  Bedford,  1934 

1934     Metafungia  R.  &  W.  R.  Bedford  :  5. 

1964  Metafungia  R.  &  W.  R.  Bedford;  Debrenne  :  219 

1965  Metafungia  R.  &  W.  R.  Bedford;  Hill  :  119. 

TYPE  SPECIES.     Metafungia  reticulata  R.  &  W.  R.  Bedford,  by  monotypy. 

GEOL.   17,  7  '* 


362  LOWER   CAMBRIAN   ARCH AEOCY ATH A    FROM    AUSTRALIA 

DIAGNOSIS.  Cup  with  central  cavity  that  becomes  free  of  skeletal  tissue  late  in  its 
development.  The  walls  are  double-porous.  The  intervallum  is  crossed  by  straight 
taeniae,  which  have  their  pores  arranged  in  lines  curving  upwards  and  outwards; 
numerous  synapticulae  join  the  taeniae.  The  vesicular  tissue  is  present  at  the  base 
and  in  contact  with  tersioid  outgrowths. 

COMPOSITION  OF  THE  GENUS.     Metafungia  reticulata  R.  &  W.  R.  Bedford. 


Metafungia  reticulata  R.  &  W.  R.  Bedford 
(PL  18,  figs.  1-3) 

1934     Metafungia  reticulata  R.  &  W.  R.  Bedford  :  5,  figs.  23  a-e. 

1965     Metafungia  reticulata  R.  &  W.  R.  Bedford;  Hill  :  119,  pi.  10,  fig.  2  and  text-fig.  22,  12  a-d. 

HOLOTYPE.     B.M.  (N.H.)  S  4184.  Other  Material  S  4183. 

DESCRIPTION.  Large  conical  cup,  the  apex  is  surrounded  by  exothecal  tersioid 
outgrowths  and  the  central  cavity  is  filled  with  endothecal  tissue.  The  intervallum  is 
crossed  by  straight,  radial  taeniae,  which  are  perforated  by  pores,  aligned  upwards 
and  outwards,  in  oblique  rows.  The  diameter  of  these  pores  increases  from  the  inner 
to  the  outer  wall,  giving  an  irregular  appearance.  The  numerous  synapticulae  con- 
nect at  the  skeletal  junctions  of  the  taeniae.  The  vesicular  tissue  is  only  developed  in 
the  exo-  and  endothecal  regions.  The  intervallar  mesh,  with  some  development  of 
skeletal  tissue  and  vertical  closing  of  the  apertures,  corresponds  to  an  outer  wall.  A 
regular  micro-porous  thin  wall  covers  this  externally.  The  inner  wall  may  also  be 
interpreted  as  a  double  wall.  The  inner  part  of  each  taenia  has  pores  no  larger  than 
0*7  mm;  these  taeniae  are  joined  by  small  plates,  resembling  widened  synapticulae,  to 
form  a  small  tube.  On  the  central  cavity  side,  a  thin  sheath  is  developed,  with  two 
pores  to  each  tube,  and  has  a  more  or  less  regular,  hexagonal  appearance. 

DIMENSIONS 

S  4183  Base  S  4184  Top 

(mm.)  (mm.)  (mm.) 
Cup: 

Height  ..  35 

Diameter  12  . .                        21 

Intervallum  2-5  . .                          5 

Intervallum  coefficient  0-3  . .                          0-4 

Interseptum  0-8  ..                           i-oi 

Synapticulae :  radial  distance  . .  . .                          o  •  61 

vertical  distance  . .  . .                          o  •  61 
Outer  wall: 

Diameter  of  main  pores  . .  0-70 

Partitions  ..  0-27 

Thickness  . .  0-2 

Micropores :  diameter  . .  0-07 

partitions  . .  0-06 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A   FROM    AUSTRALIA 


363 


DIMENSIONS — continued 


Inner  wall : 

Diameter  of  tubes 

Partitions 

Length  of  tubes 

Inner  pores :  diameter 
partitions 
Septa : 

Thickness 

Diameter  of  inner  pores 

Diameter  of  outer  pores 
Synapticulae : 

Thickness 


S  4183  Base 
(mm.) 


(mm.) 

o • 7-0 • 7 
0-27 
0-70 

O-2O 
0-13 

0-I3 
O-25 
0-40 

0-13 


S  4184  Top 

(mm.) 


DISCUSSION.  Metafungia  reticulata  has  an  intervallar  structure  similar  to  that  of 
Flindersicyathus  graphicus  (R.  &  W.  R.  Bedford).  However,  its  size  is  much  greater 
and  the  connecting  synapticulae  do  not  occur  at  every  skeletal  junction,  but  the 
essential  difference  is  in  the  structure  of  the  walls. 


Family  METAGOSCINIDAE  R.  &  W.  R.  Bedford 

DIAGNOSIS.     Two-walled  porous  cups.     Taeniae  and  sparse  convex  tabulae. 

COMPOSITION  OF  THE  FAMILY.  Metacoscinus  R.  &  W.  R.  Bedford  1934,  Pycnoido- 
coscinus  R.  &  W.  R.  Bedford  1936,  Paracoscinus  R.  &  W.  R.  Bedford  1936,  Claruscya- 
thus  Vologdin  1932,  Gabrielsocyathus  Debrenne  1964. 


Genus  METACOSCINUS  R.  &  W.  R.  Bedford 

1934     Metacoscinus  R.  &  W.  R.  Bedford  :  6,  pi.  5,  fig.  27. 

1965     Metacoscinus  R.  &  W.  R.  Bedford;  Hill  :  133,  pi.  12,  fig.  i,  text-fig.  5  a-d. 

TYPE  SPECIES.     Metacoscinus  reteseptatus  R.  &  W.  R.  Bedford,  by  monotypy. 

DESCRIPTION.  Conical  cups,  slightly  waved  externally.  Oblique  and  vertical 
rods  form  the  skeletal  tissue  of  taeniae  crossing  the  intervallum.  These  rods  have 
considerable  openings  between  them,  which  are  much  taller  than  wide  and  are 
roughly  polygonal  in  shape.  The  outer  pores  are  the  largest.  Less  frequent  horizon- 
tal structures  regarded  as  tabulae,  are  perforated  by  small  circular  pores,  that  are 
separated  by  "  linteaux  "  of  varying  size,  and  bear  little  irregular  tubercles  on  each 
side. 

Outer  wall  has  funnel-shaped  pores,  circular  on  the  intervallum  side,  larger  and 
more  irregular  on  their  open  outer  side.  Thin  inner  wall  with  two  rows  of  large 
polygonal  pores  per  intersept,  separated  by  a  thin  skeletal  mesh.  R.  &  W.  R. 
Bedford  figured  a  specimen  (1936  :  fig.  83),  which  had  several  tubercles  on  its  central 
cavity  side. 


364  LOWER   CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA 

COMPOSITION  OF  THE  GENUS.  Metacoscinus  reteseptatus  R.  &  W.  R.  Bedford  1934, 
?  Metacoscinus  insigne  R.  &  W.  R.  Bedford  1936. 

DISCUSSION.  Metacoscinus  insigne  is  retained  in  this  genus  with  some  misgiving, 
until  the  type  material  can  be  revised.  It  has  no  visible  tabulae  and  a  thick  outer 
wall  with  labyrinthoid  pores.  Nothing  is  known  about  the  structure  of  these  pores, 
whether  their  distorted  shape  is  due  to  the  fusion  of  several  tubercles,  as  seen  in  the 
tabulae  of  M .  reteseptatus,  or  results  from  lateral  connection  between  neighbouring 
pore-tubes. 

I  have  had  the  opportunity  to  examine  the  type  material  of  Metacoscinus  gabriel- 
sensis  (Okulitch  1955),  M.  deasensis  (Okulitch  1955)  and  M.  poolensis  (Kawase  & 
Okulitch  1957)  loaned  by  the  Geological  Survey  of  Canada  and  have  already  suggested 
(Debrenne  1964  :  248),  that  these  species  should  be  regarded  as  a  separate  genus, 
Gabrielsocyathus  Debrenne  1964,  with  G.  gabrielsensis  (Okulitch)  as  type  species.  This 
differs  from  Metacoscinus  in  having  simply-porous,  regularly-spaced  tabulae ;  branch- 
ing, or  waved  porous  taeniae ;  a  double,  porous,  outer  wall  with  exotheca ;  endothecal 
lamellae  partially  filling  the  central  cavity  and  extremely  abundant  vesicular  tissue. 

The  species  poolensis  (Kawase  &  Okulitch)  is  included  in  Gabrielsocyathus  with  some 
reservations.  The  septa  appear  to  undulate  and  bifurcate  and  have  fewer  pores  than 
the  other  two  species;  the  plates  thicken  at  the  junction  with  the  septa;  no  exotheca 
occurs  and  the  vesicular  tissue  is  less  abundant.  The  poor  preservation  of  the  speci- 
men prevents  confirmation  of  the  presence  of  fine,  irregular  pores  in  all  the  skeletal 
plates. 

Zhuravleva  (1960  :  311)  remarks  that  Paracoscinus  and  Metacoscinus  are  very 
close  to  each  other  and  goes  on  to  suggest  that  they  might  represent  one  genus,  but 
since  she  had  no  material,  it  was  difficult  to  make  a  definite  decision.  I  have  not  seen 
the  type  of  Paracoscinus,  but  from  the  figures  and  descriptions  of  R.  &  W.  R.  Bedford 
(1936  :  18,  pi.  20,  figs.  85-86),  I  consider  that  it  differs  from  Metacoscinus  in  its  wall 
structure  (microporous  and  not  a  large  open  mesh) ;  the  abundant  and  regular  plates, 
which  have  polygonal  pores  and  are  without  tubercles ;  the  thin  outer  wall  and  the 
inner  wall  with  a  single  row  of  pores  per  interseptum. 


Metacoscinus  reteseptatus  R.  &  W.  R.  Bedford 
(PL  17,  figs.  1-4) 

1910  Archaeocyathus  retesepta  Taylor  :  120,  pi.  7,  figs.  39-40. 

1934  Metacoscinus  reteseptatus  R.  &  W.  R.  Bedford  :  6,  pi.  5,  fig.  27. 

1936  Metacoscinus  reteseptatum  R.  &  W.  R.  Bedford;  R.  &  W.  R.  Bedford;  18,  pi.  19,  fig.  83. 

1965  Metacoscinus  reteseptatus  R.  &  W.  R.  Bedford;  Hill  :  132-3,  pi.  12,  fig.  i,  text-fig.  24,  5  a-d. 

MATERIAL.  Syntypes  B.M.  (N.H.)  S  4191-4195.  Other  material  S  4762,  S  4772, 
S  4743,  S  7633. 

DESCRIPTION.  Conical  cups  with  transverse  annular  bulges.  Outer  wall  has  two 
vertical  rows  of  funnel-shaped  pores  per  intersept,  the  larger  aperture  on  the  outside, 


LOWER   CAMBRIAN   ARCH AEOC Y ATH A   FROM   AUSTRALIA 


365 


with  an  irregular  rim.  Inner  wall,  also,  has  two  vertical  rows  of  pores,  but  these  are 
polygonal  and  separated  by  very  thin  skeletal  rods,  without  tubercles.  Regular, 
radial  pseudosepta,  perforated  by  numerous  hexagonal  pores  that  are  elongated 
upwards,  or  upwards  and  outwards  in  the  annular  bulges.  Little  vesicular  tissue 
except  in  the  lower  part,  where  the  pseudosepta  are  not  so  well  developed  and  skeletal 
elements  fill  both  the  central  cavity  and  the  intervallum.  Sparse  tabulae,  which  have 
tubercles  occurring  between  their  two  rows  of  quincunxial  pores  per  intersept.  The 
regular  pattern  of  the  pores  is  disturbed  by  the  coalescence  of  neighbouring  pores,  or 
by  anastomosed  tubercles,  giving  these  horizontal  plates  a  vermiculate  aspect,  when 
seen  from  above. 


DIMENSIONS 


Cup: 

Height  (pars) 

Diameter 

Intervallum 

Central  cavity 

Interseptum 

Intertabulum 
Outer  wall : 

No.  of  pore  rows 
per  interseptum 

Diameter 

Vertical  partitions 

Horizontal  partitions 

Thickness 
Inner  wall : 

No.  of  pore  rows 
interseptum 

Diameter 

Vertical  partitions 

Horizontal  partitions 

Thickness 
Septa : 

No.  of  pore  rows 

Diameter 

Vertical  partitions 
Horizontal  partitions 
Thickness 
Tabulae 

No.  of  pore  rows 
Diameter 
Vertical  partitions 
Horizontal  partitions 
Thickness 


S  4191 
(mm.) 


21 
12 

2 

8 


i-y-3'4 


2-3 
0-15 

0-06-0-10 
0-06-0- 10 

0-30 


0-07 
0-07 
O-2O 


0-15^-37 

0-07 
O-II 


2 
O-II 

o-  ii 

O-II 

0-15 


S  4192  & 

S  4193 
low 

(mm.) 

base 

7 

full 

O-22 


0-26 


S  4193 
upper 
(mm.) 


10 

2 

4 
0-26 


2 

0-07 

O-O7 

O-II 

6 

O  •  22-O • 40 
0-07 

0-07 


S  4195 
(mm.) 

25 
25-30 

6 
ii 

o-75 


2 

0-18 
0-15 
0-15 
0-18 


2-3 

0-18-0-37 
0-07 
0-07 

O-II 


0-56-1-51 
O-26-O-52 

0-18 
0-18 
o-ii 


84762 
(mm.) 

20 
15 

2-45 
unknown 

0-22 


0-15 

0-06-0-10 
o-io 

unknown 


1-2 

0-22-0-30 
0-07 

0-15 
unknown 

unknown 
0-26 

0-13 
0-07 


366 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A    FROM   AUSTRALIA 


DIMENSIONS — continued 


Cup: 

Height  (pars) 

Diameter 

Intervallum 

Central  cavity 

Interseptum 

Intertabulum 
Outer  wall: 

No.  of  pore  rows 
per  interseptum 

Diameter 

Vertical  partitions 

Horizontal  partitions 

Thickness 
Inner  wall: 

No.  of  pore  rows 
interseptum 

Diameter 

Vertical  partitions 

Horizontal  partitions 

Thickness 
Septa : 

No.  of  pore  rows 

Diameter 

Vertical  partitions 

Horizontal  partitions 

Thickness 
Tabulae : 

No.  of  pore  rows 

Diameter 

Vertical  partitions 

Horizontal  partitions 

Thickness 


84772 
(mm.) 

unknown 
16 
3'5 


S4743 
(mm.) 

12 
10 

2 
O-22 


2-3 

0-13 
0-07 

O-  II 


unseen 


$4743 
lower 
(mm.) 


84194 
lower 
(mm.) 


84194 
upper 
(mm.) 


7 

30 
9 

16 

i'35 
4  -3  clear 

i-45 
skel.  el. 

4-8 

o-33 

0-41 

o-37 

unseen  2  2 

0-26-0-15  0-26-0-26 
0-05        0-26—0- i 8 
o-n 
0-18 


2  unseen         unseen 

unseen  . .  0-18 

0-07 

0-07  . .  o-ii 


o-n 
o-io 

not  visible 
0-15 


87633 
(mm.) 


15 

4 


o-n 


2-3 

0-15 

O-II 
O-II 


7 

7 

5 

7 

o-45 

o-  13-0-22 

0-13 

0-49 

0-34-1-7 

0-26 

O-II 

0-07 

0-07 

0-07 

0-15 

O-II 

0-15 

0-15 

0-15 

O-II 

0-07 

O-II 

O-II 

O-II 

retesepta 
Taylor 
(mm.) 

90 
25 


2 

O-2O 

O-II 


2-3 

0-15 

o-io 

0-10 


0-15 
0-15 


DISCUSSION.  R.  &  W.  R.  Bedford  (1934  :  6)  compared  this  species  with  Archaeo- 
cyathus  retesepta  Taylor,  noting  the  likeness  between  the  taeniae,  but  no  horizontal 
structures  were  known  in  A.  retesepta.  Nevertheless,  from  Taylor's  figures  (PL  7, 
figs.  39  and  40),  one  can  perhaps  consider  that  the  structures  normal  to  those  in  the 
general  direction  are  the  beginnings  of  tabulae.  Amongst  all  the  specimens  in  the 
Bedford  collection  at  the  British  Museum  (Nat.  Hist.),  only  S  4195  and  S  4194  show 
these  rare  horizontal  structures.  The  dimensions  of  the  two  species  are  of  the  same 
order  and  they  are  probably  synonymous.  Before  joining  them  as  a  single  species  we 
need  more  material,  in  order  to  know  the  frequency  and  significance  of  these  horizon- 
tal structures  and  to  decide  whether  they  have  any  systematic  value,  or  are  true 
tabulae,  or  are  accidental  features. 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A    FROM    AUSTRALIA  367 

?  Family  METAGOSCINIDAE  R.  &  W.  R.  Bedford,  1936 
Genus  PYCNOIDOCOSCINUS  R.  &  W.  R.  Bedford,  1936 
1936     Pycnoidocoscinus  R.  &  W.  R.  Bedford  :  19,  pi.  20,  fig.  87. 

TYPE  SPECIES.  Pycnoidocoscinus  pycnoideum  R.  &  W.  R.  Bedford,  by  original 
designation. 

DIAGNOSIS.  Cups  with  annular  bulges  on  the  outer  wall,  with  the  inner  wall 
remaining  cylindrical.  Radial  septa  and  arched  tabulae.  The  outer  wall  is  a  porous 
mesh;  the  inner  wall  with  both  horizontal  and  vertical  rows  of  rectangular  pores. 

DISCUSSION.  As  the  early  stages  are  not  known,  it  is  difficult  to  classify  this  genus 
in  either  the  Irregularia — indicated  by  the  irregular  outer  wall  and  septa  with 
numerous  pores,  or  amongst  the  Regularia — as  suggested  from  the  type  of  inner  wall 
with  its  rectangular  apertures,  the  tabulae  with  the  axis  of  curvature  inside  the 
intervallum,  or  the  pore  pattern  of  Retecoscinus-type.  Only  having  examined  a 
single  broken  fragment,  it  is  not  possible  for  me  to  offer  any  solution  and  the  genus 
Pycnoidocoscinus  is  retained  in  the  Metacoscinidae,  but  with  some  reservations. 

COMPOSITION  OF  THE  GENUS.    Pycnoidocoscinus  pycnoideum  R.  &  W.  R.  Bedford. 

Pycnoidocoscinus  pycnoideum  R.  &  W.  R.  Bedford 
(PI.  18,  figs.  4-8) 

1936     Pycnoidocoscinus  pycnoideum  R.  &  W.  R.  Bedford  :  19,  pi.  20,  fig.  87. 

MATERIAL.  B.M.  (N.H.)  S  4832.  A  syntype  P  990  is  in  the  South  Australian 
Museum,  Adelaide. 

DESCRIPTION.  Only  one  weathered  and  poorly  preserved  specimen  has  been 
examined.  The  intervallum  is  crossed  by  radial  septa  and  arched  tabulae.  The 
septa  are  thickened  near  the  inner  wall  and  sometimes  near  the  outer  wall,  but  are 
very  thin  in  the  middle.  Septal  pores  are  not  visible.  Pores  on  a  small,  tangential 
surface  of  a  tabula  resemble  those  of  Retecoscinus  Zhuravleva  (1960),  with  two 
alternating  rows  of  oval  to  rectangular  pores. 

Outer  wall  eroded  away;  R.  &  W.  R.  Bedford  have  described  it  as  "  a  layer  of  fine 
pores  supported  by  an  irregular  mesh  ".  Inner  wall  has  two  rows  of  pores  per  inter- 
septum  and  two  rows  of  pores  per  intertabulum. 

The  vertical  partition  between  the  two  interseptal  pores,  is  the  beginning  of  a  new 
septum,  which  only  develops  in  the  inner  third  of  the  intervallum.  Sometimes 
opposite  this  crude  septa,  another  arises  from  the  outer  wall  to  meet  it. 

DIMENSIONS 

(mm.) 
Cup: 

Height  (pars)  30 

Diameter  22 

Interseptum  (at  the  inner  wall  side)  o  •  33 

Intertabulum  1-22 


368  LOWER   CAMBRIAN    ARCH AEOC Y ATH A   FROM   AUSTRALIA 

DIMENSIONS — continued 

(mm.) 

Outer  wall:  weathered  away 

Inner  wall : 

Diameter  of  pores  0-33 

Vertical  partitions  o  •  i 

Horizontal  partitions  o  •  06 

Septa 

Diameter  of  pores  o  •  06 

Vertical  partitions  o  •  2 

Thickness:  middle  0-06 

near  inner  wall  o  •  2 

Tabulae : 

No.  of  pores  per  loculus  2  x  22 

Diameter  of  pores  o  •  27-0  •  i 

Partitions  0-1-0-13 

Thickness  o  •  i 

DISCUSSION.  The  rows  of  rectangular  pores  of  the  inner  wall  and  the  presence  of 
"  rods  "  in  the  middle  of  the  interseptum  (interpreted  here  as  the  beginnings  of  new 
septa)  are  good  reasons  for  including  S  4832  in  P.  pycnoideum.  Unfortunately,  poor 
preservation  and  the  lack  of  an  apex  prevents  any  clarification  of  the  systematic 
position  of  Pycnoidocoscinus. 

III.    STRATIGRAPHICAL    CONCLUSIONS 

The  excellent  silicified  fossils  found  in  the  Australian  Cambrian,  as  well  as  being 
extremely  useful  for  anatomical  studies  of  the  Archaeocyatha,  are  also  of  considerable 
stratigraphical  interest. 

We  have  little  information  on  the  precise  horizon  in  the  Ajax  Limestone,  from  which 
the  Bedfords  collected  their  fossils.  According  to  Daily  (1956  :  129),  ".  .  .  Most,  if 
not  all,  of  the  species  described  from  the  Ajax  Mine  area  by  Taylor  and  the  Bedfords, 
are  believed  to  have  come  from  beds  containing  this  assemblage  [i.e.  faunal  assem- 
blage No.  i],  for  it  is  significant  that  no  fossils  characteristic  of  faunal  assemblage  No. 
2  have  been  reported  by  them.  Such  fossils  would  almost  certainly  have  been  noticed 
if  they  were  present  .  .  . ".  However,  Walter  (1967)  comments  (p.  145)  that  the 
rich  Ajax  fauna  described  by  the  Bedfords  &  Taylor,  cannot  be  placed  in  Daily's 
scheme  of  faunal  units.  This  is  because,  in  an  area  of  complex  structure,  it  was  not 
collected  in  sequence  and  the  additional  evidence  of  the  fauna  associated  with  the 
archaeocyathids  is  not  available.  As  a  result  it  is  of  very  limited  stratigraphical  use. 

The  out-dated  studies  of  Taylor  and  the  too  brief  publications  of  the  Bedfords,  have 
not  allowed  specialists  in  other  countries  to  make  precise  comparisons  and  possible 
correlations,  between  their  material  and  that  of  Australia.  An  attempt  was  made  on 
the  limited  material  available  in  the  Ting  collection  (F.  &  M.  Debrenne  1960).  It 
was  concluded,  from  the  majority  of  genera  appearing  at  Beltana,  that  the  fauna 
belonged  to  the  Aldanian  and  Lower  Lenian  stages.  Since  then,  I  have  become 
acquainted  with  Russian  work  that  amends  the  stratigraphical  correlation  between 
the  Southern  Siberia  geosyncline  and  the  Siberian  platform,  and  gives  detailed  lists  of 
revised  Archaeocyatha  faunas  from  level  to  level. 


LOWER   CAMBRIAN    ARCH AEOC Y ATH A    FROM   AUSTRALIA  369 

These  data,  together  with  the  present  study,  confirm  that  the  Ajax  Limestone  is 
situated  in  the  lower  half  of  the  Lower  Cambrian ;  to  be  precise,  the  fauna  is  similar  to 
that  of  the  Kameshki  horizon  and  also  has  a  certain  affinity  with  those  forms  found  in 
the  succeeding  Sanashtygkol  horizon.  This  opinion  is  based  on  the  presence,  in  the 
Ajax  fauna,  of  forms  having  advanced  evolutionary  stages  of  outer  and  inner  walls, 
or  intervallar  structures,  together  with  other  simple  forms  known  from  the  Bazaikha 
horizon,  such  as  Monocyathus,  Alphacyathus,  Ajacicyathus,  Robustocyathus  and 
Erismacoscinus . 

As  examples  of  the  advanced  forms,  one  can  show  the  presence  of  genera  with : 

1.  Outer  walls  with  simple  tumuli  (Tumuliolynthus ,  Ethmocoscinus] ,  or  multi- 
perforate  (A  lataucyathus] . 

According  to  Rozanov  (1963  :  8)  genera  with  tumuli  mainly  arise  in  the 
Kameshki  horizon. 

2.  Double  porous  outer  wall  (Tomocyathus,  Polycoscinus}. 

This  feature  appears  at  the  end  of  the  Bazaikha  horizon  and  genera  that 
possess  it  increase  in  numbers  through  the  Kameshki  and  Sanashtygkol  horizons. 

3.  Inner  wall  with  complex  annular  rings. 

This  is  seen  in  Cyathocricus  annulispinosus  (Vologdin)  found  in  the  Kameshki 
horizon  of  Eastern  Sajan,  and  the  Australian  species  annulatus  (Bedford) 
regarded  as  belonging  to  Salairocyathus,  which  occurs  in  the  Sanashtygkol 
horizon. 

4.  Inner  wall  with  branching  pore-tubes  (Zonacyathus). 

Three  Russian  species  probably  belong  to  this  genus ;  poletaevae,  vermiculatum 
and  flexum.  They  and  the  corresponding  colonial  genus  Sajanocyathus  are  of 
Sanashtygkol  age. 

5.  Radial  septa  with  few,  or  no  pores  (Cyathocricus,  Ethmocyathus,  Zonacyathus, 
Robustocyathus  subacutus  and  Archaeocyathellus  (Stapicyathus]  stapipora}. 

Zhuravleva  (1960  :  147)  and  Rozanov  &  Missarzhevsky  (1966  :  71)  consider 
that  the  decrease  or  loss  of  septal  porosity  is  an  evolutionary  character  of  the 
Regularia.  The  pores  of  the  plates  are  often  different  to  the  septal  pores  in 
Australian  Coscinocyathus,  whereas  in  specimens  found  at  lower  levels  in  the 
Cambrian  (e.g.  Soussian  horizon  in  Morrocco)  they  are  very  similar.  The  first 
appearance  of  any  difference  between  the  two  occurs  in  the  Sardinian  limestones, 
contemporaneous  with  the  Botoma  Stage  of  Southern  Siberia. 

LOWER  CAMBRIAN 

These  correlations  are  taken  from  Table  II  of  Walter  (1967),  who  used  information 
given  in  a  personal  communication  (Sept.  1966)  to  M.  F.  Glaessner  by  A.  Yu.  Rozanov. 

SIBERIAN  PLATFORM  GEOSYNCLINE  OF  SOUTHERN  SIBERIA 

Lower  subdivision 
f  Sunnagin  fKundatsk  horizon 

TOMMOT  STAGE  <   T,  I  ^         .,  ,       ,        . 

\Kenyada  ALDAN  STAGED  Bazaikha  horizon 
Atdaban  (^  Kameshki  horizon 

Tarynsk 

Sinsko-Kutorgina  BOTOMA  STAGE  Sanashtygkol  horizon 


370  LOWER   CAMBRIAN   ARCH  AEOC  Y  ATH  A    FROM    AUSTRALIA 

Upper  subdivision 

Ketema  LENA  STAGE      Solontsov  horizon 

Elanka  Obruchev  horizon 


Similar  studies  of  evolutionary  stages,  have  not  been  made  for  the  Irregularia,  but 
it  is  possible  to  make  the  following  points. 

1.  The  first  Irregularia  have  scattered  skeletal  elements,  rods,  or  plates,  as  in 
Dictyocyathus,  or  Protopharetra  and  are  of  Soussian  age.     They  then  follow  two 
particular  paths. 

a.  The  plates  of  Protopharetra  gave  rise  to  the  crude  pseudo-septa  of  Volva- 
cyathus-iype,  where  the  pores  occupy  a  greater  surface  area  than  the 
skeletal  tissue;  then  into  the  poorly  developed  pseudosepta  shown  in  the 
first  Metaldetes  of  Sardinia ;  followed  by  the  definite  radial  septa  of  the  true 
Metaldetes  of  Australia  and  ending  with  the  Cambrocyathus-type,  which 
occurs  in  the  upper  half  of  the  Lower  Cambrian. 

b.  The  rods  of  Dictyocyathus,  on  the  other  hand,  form  the  regular  scaffolding 
of  Flindersicyathus  and  Pycnoidocyathus  types,  which  are  comparable  to  the 
intervallar  structure  of  Tabellaecyathidae  Fonin,  characteristic  of  the 
Sanashtygkol  horizon. 

2.  Amongst  the  Ajax  fossils  one  finds  Syringocnema,  a  genus  characterized  by 
prismatic  loculi  in  the  intervallum.     In  the  USSR  the  species  referred  to  this 
genus  are  limited  to  the  Sanashtygkol  horizon. 

3.  The  walls  of  the  Irregularia  are  not  so  diverse  as  those  of  the  Regularia.     In  the 
older  forms  of  Dictyocyathus  and  Protopharetra  they  are  not  independent  of  the 
intervaller  structural  elements.     This  is  attained  in  the  genus  Flindersicyathus, 
with  its  simply  porous  outer  wall  and  inner  wall  having  pore-tubes,  the  lower 
parts  of  which  are  modified  and  stretched.     The  double  porous  inner  wall  is  seen 
in  the  genera  Metaldetes,  Metafungia  and  Metacoscinus.     Such  structures  indi- 
cate the  development  of  more  advanced  features,  e.g.  lamellae  and  tabulae,  that 
are  known  in  the  Sanashtygkol  horizon  (Fonin  1963). 

All  these  considerations  show  that  there  are  clear  affinities  between  the  Ajax  faunas 
and  those  of  the  Kameshki  and  Sanashtygkol  horizons.  The  absence  of  forms  with 
clathrate  outer  walls,  or  inner  walls  of  true  Ethmophyllum-type,  shows  that  the 
Sanashtygkol  fauna  was  not  well  established  in  Australia  at  the  time  of  the  Ajax 
limestone.  The  fauna  is,  therefore,  probably  of  Upper  Kameshki  and  Lower 
Sanashtygkol  age,  that  is  to  say  the  middle  part  of  the  lower  division  of  the  Lower 
Cambrian. 

Since  this  conclusion  was  first  written  Walter  (1967),  in  his  study  on  the  usefulness 
of  Archaeocyatha  for  zoning  and  correlation  in  the  Lower  Cambrian  of  the  Adelaide 
Geosyncline,  has  correlated  the  various  faunas  in  the  Hawker  Group  of  South 
Australia  with  those  of  Siberia.  Using  genera  as  the  basis  of  such  correlation,  he  has 
assigned  a  Sanashtygkol  age  to  the  Oraparinna  Shale;  the  latter  occurs  above  the 


LOWER   CAMBRIAN    ARCH  AEOC  Y  ATH  A   FROM    AUSTRALIA  371 


Hawker      Group 


Wilpena      Group 


S\\\\\' 


Wirrealpa      Lst. 
Billy  Creek   Form'n. 

Marina    Graywacke 
Oraparinna    Shale 
Bunkers     Sdst. 

Parara      Lst. 

Wilkawillina     Lst. 
Parachilna     Form'n. 

Pound     Qtzte. 


Ajax      Lst, 


FIG.  15.     Lower  Cambrian  Sediments,  Flinders  Ranges;  S.  Australia. 
After  Dalgarno  (1964)  &  Walter  (1967). 


Wilkawillina  Limestone  of  which,  in  the  Mt.  Scott  Range  area,  the  Ajax  Limestone  is 
said  to  be  the  equivalent  (Dalgarno  1964  :  136).  Commenting  on  the  Ajax  Limestone 
Walter  (p.  146)  suggests  that  the  presence  of  Syringocnema,  Flindersicyathus  and 
Pycnoidocyathus  indicate  correlation  with  the  upper  part  of  the  Hawker  Group  in  the 
Wilkawillina  Gorge  section  and  concludes  that,  in  the  Ajax  Mine  area,  the  Ajax  Lime- 
stone may  be  another  bioherm.  This  evidence  would  appear  to  support  my  con- 
clusions as  to  the  age  of  the  fauna  of  the  Ajax  Limestone. 

IV.    SUMMARY 

The  material  now  in  the  British  Museum  (Natural  History),  collected  and 
originally  described  by  R.  &  W.  R.  Bedford,  has  been  revised  according  to  current 
classification. 


372  LOWER   CAMBRIAN    ARCH AEOC Y ATH A   FROM   AUSTRALIA 

It  is  considered  necessary  to  create  a  separate  family  within  the  Ajacicyathacea  for 
the  genus  Ethmocyathus,  which  is  characterized  by  a  secondary  wall  of  annular  plates 
on  the  central  cavity  side  of  the  inner  wall.  A  new  genus,  Cyathocricus,  is  introduced 
for  Bronchocyathus  Bedford,  which  did  not  conform  to  the  rules  of  zoological  nomen- 
clature, and  includes  those  forms  belonging  to  the  Bronchocyathidae  that  have  a  more 
complex  inner  wall  than  Cyclocyathus.  Two  species  are  assigned  to  the  new  genus. 

A  new  subgenus  Loculicyathus  (Loculicyathellus]  is  distinguished  from  Loculicyathus 
(Loculicyathus}  by  the  longitudinal  corrugations  occurring  between  the  septa  on  the 
outer  wall.  Two  new  subgenera  are  introduced  for  the  smooth  inner  wall  forms  of 
Tomocyathus:  T.  (Erugatocyathus)  and  Alataucyathus  :  A.  (Anaptyctocyathus}.  A 
new  species  of  the  latter  is  erected,  A.  (A.}  flabellus,  which  is  distinguished  from  A. 
(A.}  cribripora  by  its  bowl-shaped  cup,  dimensions  and  arrangement  of  wall  pores. 

In  the  broader  classification,  the  family  Acanthinocyathidae  is  distinguished  from 
the  Dokidocyathidae,  and  the  family  Bronchocyathidae  is  re-introduced  to  take  the 
new  genus  Cyathocricus.  The  genus  Flindersicyathus  is  included  in  the  re-introduced 
family  Flindersicyathidae,  as  there  are  many  characters  of  this  genus  which  distin- 
guish it  from  Archaeocyathus.  Pycnoidocyathus  and  Flindersicyathus  are  treated  as 
subgenera  until  the  Ajax  Mine  material  in  both  Princeton  and  Adelaide  is  re-studied. 

Owing  to  both  the  poor  preservation  and  limited  number  of  some  specimens,  it  has 
been  impossible  to  see  sufficient  structural  details  for  exact  determination  and  the 
true  position  of  several  genera  remains  uncertain.  However,  the  good  condition  of 
other  material  has  enabled  a  thorough  study  of  the  internal  skeleton  to  be  made,  with 
the  result  that  new  types  of  structure  have  been  discovered,  e.g.  the  annular  form  of 
the  inner  wall  in  Salairocyathus  (S.)  annulatus. 

From  the  stratigraphical  evidence,  it  is  concluded  that  the  fauna  is  of  middle  Lower 
Cambrian  age,  i.e.  upper  Kameshki — lower  Sanashtygkol  horizons,  and  this  has  been 
borne  out  by  subsequent  work. 


V.    ACKNOWLEDGMENTS 

I  wish  to  thank  Dr.  E.  I.  White  and  Dr.  H.  W.  Ball  for  the  opportunity  of  studying 
the  Bedford  collection  in  the  Department  of  Palaeontology.  I  would  also  like  to 
thank  Dr.  Ball  and  Mr.  S.  Ware,  for  the  generous  help  and  information  given  to  me 
whilst  examining  this  collection.  I  am  also  extremely  grateful  to  Professor  M.  F. 
Glaessner  and  his  colleagues  of  the  University  of  Adelaide  and  at  the  Geological 
Survey  of  South  Australia,  for  allowing  me  to  reproduce  maps  and  stratigraphical 
information  in  text-figs.  13,  14  and  15.  In  addition,  thanks  are  also  due  to  my 
husband  M.  M.  Debrenne  and  Mr.  P.  Green  of  the  Photographic  Studio  at  the  B.M. 
(N.H.)  for  the  photographs.  Finally,  I  wish  to  thank  Dr.  Dorothy  Hill,  of  the  Uni- 
versity of  Queensland,  for  her  encouragement  and  advice,  and  for  permission  to 
reproduce  some  of  the  photographs  of  Bedford  material  included  in  her  monograph 
on  Antarctic  archaeocyathids. 


LOWER   CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA  373 

VI.     REFERENCES 

BEDFORD,  R.  &  J.  1936.  Further  notes  on  Archaeocyathi  (Cyathospongia)  and  other  organisms 
from  the  Lower  Cambrian  of  Beltana,  South  Australia.  Mem.  Kyancutta  Mus.,  Kyancutta. 
3  :  21-26,  6  pis. 

I937-  Further  notes  on  Archaeos  (Pleospongia)  from  the  Lower  Cambrian  of  South 

Australia.  Mem,  Kyancutta  Mus.,  Kyancutta.  4  :  27-38,  15  pis. 

—  1939.     Development  and  classification  of  Archaeos  (Pleospongia).     Mem.  Kyancutta  Mus., 
Kyancutta.     6  :  67-82,  n  pis. 

BEDFORD.  R.  &  W.  R.  1934.  New  species  of  Archaeocyathinae  and  other  organisms  from  the 
Lower  Cambrian  of  Beltana,  South  Australia,  Mem.  Kyancutta  Mus. ,  Kyancutta.  1  :  7,  6  pis. 

—  1936.     Further  notes  on  Archaeocyathi  (Cyathospongia)  and  other  organisms  from  the 
Lower    Cambrian    of    Beltana,    South    Australia.     Mem.    Kyancutta    Mus.,    Kyancutta. 
2  :  9-19,  14  pis. 

BILLINGS,  E.     1861.     New  species  of  Lower  Silurian  fossils:  On  some  new  or  little-known  species 

of  Lower  Silurian  fossils  from  the  Potsdam  Group  (Primordial  Zone).     In  Hitchcock,  E.  et 

al.  1861  "Report  on  the  geology  of  Vermont"  :  24,  Claremont,  N.H. 
BORNEMANN,  J.  G.     1887.     Die  Versteinerungen  des  Cambrischen  Schichtensystems  der  Insel 

Sardinien  nebst  vergleichenden  Untersuchungen  iiber  analogic  Vorkommnisse  aus  andern 

Landern.     Erste   Abtheilung.     iii.     Archaeocyathinae.     Nova  Acta  Acad.   Caesar.   Leop. 

Carol.,  Halle.     51,  i  :  28-78,  pis.  5-33. 
-  1891.     Zweite  Abtheilung.  Nachschrift.  iii.  Archaeocyathinae.     Nova  Acta  Acad.  Caesar. 

Leop.  Carol.,  Halle.     56,  3  :  495-500,  pis.  42-43. 
BROILI,  F.     1915.     Archaeocyathinae.     In  K.  A.  Zittel,  "  Grundzuge  der  Palaontologie"  1915, 

4th  edit.,  1  :  121,  Munich. 
CAMPANA  B.     1958.     The  Flinders  Ranges  in  "  The  geology  of  South  Australia  ".     See  under 

M.  F.  Glaessner,  &  L.  W.  Parkin,  (Edit.). 
DAILY,  B.     1956.     The  Cambrian  of  South  Australia.     In  "  El  sistema  Cambrico,  su  paleogeo- 

grafia  y  el  problema  de  su  base.     Part  2.     Australia,  America".     XXth  Int.  Geol.  Congr., 

Mexico  :  91-147. 
DALGARNO,  C.  R.     1964.     Lower  Cambrian  stratigraphy  of  the  Flinders  Ranges.     Trans.  Roy. 

Soc.     South     Australia.     Adelaide,     88  :  129—144. 
DEBRENNE,  F.     1958.      Sur  quelques  Archaeocyatha  du  Jebel  Taissa  (Anti- Atlas  occidental) 

Notes  Mem.  Serv.  Mines  Carte  geol.  Maroc.,  Rabat.     16,  No.  143  :  59-74,  3  pis.,  2  t.-figs. 

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—  1960.     Deux  nouveaux  genre  d'Archaeocyathides  du  Cambrien  marocain  (Geniculicyathus. 
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—  1960.     Archaeocyatha,  Porifera,  (Cambrian  System).     See  under  L.  L.  KHALFIN,  1960. 
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skfi  opornyi  razrez  nizhnego  Kembriya  v  Vostochnom  Sayane.     Mater.  Geol.  polez.,  Iskop., 

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1943-     North  American  Pleospongia.     Spec.  Pap.  geol.  Soc.  Am.,  Washington.     48  :  112, 

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1950.     Monocyathus    Bedford    versus    Archaeolynthus  Taylor.     J.  Paleont.,  Tulsa.     24, 

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1955-     Archaeocyatha  from  the  Mcdame  Area  of  northern  British  Colombia.     Proc.  Trans. 

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375 

ROZANOV,  A.  Yu.  1960.  [New  data  on  Archaeocyatha  of  the  Shorian  Highland.]  Dokl.  Akad. 
Nauk  SSSR,  131,  3  :  663-666.  [In  Russian,  Eng.  transl.  in  Dokl.  Acad.  Sci.  USSR.  Earth 
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-  1963.     [Some  problems  of  evolution  of  regular  Archaeocyathi.]     Paleont  Zh.,  Moscow.: 
1963,  1  :  3-12,  5  figs.     [In  Russian,  Eng.  transl.  in  Int.  Geol.  Rev.,  Washington.     6,    10 
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—  1964.     See  REPINA,  L.  N.  (Ed.)  1964. 

See  ZHURAVLEVA,  I.  T.,  KONYUSHOV,  K.  N.  &  ROZANOV,  A.  Yu. 


ROZANOV,  A.  Yu.  &  MISSARZHEVSKY,  V.  V.     1966.     [Biostratigraphy  and  fauna  of  Lower  Camb- 

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Kambrium  der  Sierra  Morena  (Spanien).     Abh.  senckenb.  naturforsch.  Ges.,  Frankfurt  a.M. 

448  :  87,  5  pis.,  5  figs. 
TAYLOR,  T.  G.     1908.     Preliminary  note  on  Archaeocyathinae  from  the  Cambrian  "  coral  reefs  " 

of  South  Australia.     Rep.  Australas.  Ass.  Advmt.  Sci.,  1907,  Adelaide.     11,  Sect.  C,  9  :  423 

437,  2  pis.,  8  figs. 
•  -  1910.     The  Archaeocyathinae  from  the  Cambrian  of  South  Australia  with  an  account  of  the 

morphology  and  affinities  of  the  whole  class.     Mem.  R.  Soc.  S.  Aust.,  Adelaide.     2,  2  :  55- 

188,  16  pis.,  51  figs. 
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—  —  1932.     The  Archaeocyathinae  of  Siberia.  2.     Fossils  of  the  Cambrian  Limestones  of  the 

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-  1937.     Arkheotsiaty  i  vodorosli  yuzhnogo   sklono   anaborskogo   massiva.     Trudy  arkt. 
nauchno-issled.  Inst.,  Leningrad.     91  :  9-46.     [French  translation  S.I.G.  1434.] 

-  1939.     Middle  Cambrian   Archaeocyatha  and  Algae  from  the   South  Urals.     Problem- 
Paleont.,  Moscow.     [In  Russian  with  English  translation.]     5  :  209-276,  12  pis.,  12  figs. 

—  1940.     Les  archeocyathes  et  les  algues  des  calcaires  cambriens  de  la  Mongolie  et  de  la  Tuva. 
Trudy  mongol'  Kom.,  34,  10  :  268.     [In  Russian  with  English  summary.] 

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1940  :  23-97,  31  pis.,  85  figs.     [In  Russian  with  French  transl.  SIG  1446.] 

-  1956.     Classification  du  type  Archaeocyatha.  Dokl.  Akad.  Nauk  SSSR.  Moscow.     Ill, 
4  :  877-880.     (In  Russian,  with  French  translation  SIG  No.  1510.) 

-  1957.     Les   Archaeocyatha   et   leur   signification   stratigraphique.     Annls.    Cent.    £tud. 
Docum.  paleont.,  Paris.     23,  2  :  34-73,  22  pis.     [French  translation  of  Acta  palaeontologica 
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VOLOGDIN,  A.  G.  1961.  Arkheotsiaty  i  ikh  stratigraficheskoe  znachenie.  In  "  El  sistema 
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VOLOGDIN,  A.  G.  1962.  [Archaeocyatha  and  algae  of  the  Cambrian  in  the  Baikal  Highlands.] 
Trudy  paleont.  Inst.,  Moscow  93  :  3-116,  21  pis.,  21  figs.  [In  Russian.] 

—  -  —  ig62a.     Tip  Archaeocyatha,  Arkheotsiaty.     In  ORLOV,  Y.  A.  (ed.  "  Osnovy  Paleontologii.  " 

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89-139.     [In  Russian.] 

---  ig62b.  [The  anatomy  of  the  Archaeocyathids.]  Paleont.  Zh.,  Moscow.  1962,  2  :  9-20. 
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376  LOWER  CAMBRIAN   ARCH AEOCY ATH A   FROM   AUSTRALIA 

VOLOGDIN,  A.  G.  1963.  [Late  Middle  Cambrian  Archaeocyathids  from  the  Amga  River  basin  (on  the 

Siberian  Platform)].     Dokl.  Akad.  Nauk  SSSR,  151,  4  :  946-949.     [In  Russian,  with  English 

translation  in  Dokl.  (Proc.)  Acad.  Sci.  URSS,  Earth  Sci.,  Washington.     151,  1963  :  199-202.] 
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Group,  South  Australia.     /.  geol.  Soc.  Aust.,  14,  i  :  139-152,  2  pis. 
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lynthus  Taylor  and  its  possible  genetic  connection  with  Echinodermata.]     Dokl.  Akad.  Nauk 

SSSR,  Moscow.     109,  4  :  855-857,  i  pi.     [In  Russian.] 
ZHURAVLEVA,  I.  T.     1949.     Certaines  donn6es  sur  la  structure  de  calice  chez  les  representants 

du  genre  Rhabdocyathus  Toll.  Dokl.  Akad.  Nauk  SSSR,  Moscow.     67,  3,:  547-550,  2  figs. 

[In  Russian.     French  translation  SIG  240.] 

-  1951.     Sur  1' individuality  des  calices  brises  d'Archeocyathes  Reguliers  et  de   « larves  » 
d'Archeocyathes.     Dokl.  Akad.  Nauk  SSSR,  Moscow.     80,  i  :  97-100,  3  figs.     [In  Russian, 
French  translation  SIG  No.  407.] 

—  1955.     Contribution  a  la  connaissance  des  Arch6ocyathes  de  Siberie.     Dokl.  Akad.  Nauk. 
SSSR,  Moscow.     104,  4  :  626-629,  i  pi-,  i  fig.     [In  Russian,  French  translation  SIG  1346.] 

1960.     Archaeocyathi  of  the  Siberian  Platform.     Izd.  Akad.  Nauk  SSSR,  Inst.  Geol.  Geofiz., 

Moscow.     1960  :  346,  33  pis.,  t.-figs.     [In  Russian.] 

—  i96oa     See  under  KHALFIN,  L.  L.  (Ed.).     1960. 

—  1961.     See  under  Musatov,  D.  I.  et  al.,  1961. 

-  1963.     Archaeocyatha  of  Siberia:  single-walled  archaeocyatha.     Orders  Monocyathida — 
Rhizacyathida.     Izd.  Akad.  Nauk  SSSR,  Inst.  Geol.  Geofiz.,  Moscow.     1963  :  139,  87  pis. 
[In  Russian.] 

-  1964.     See  under  REPINA,  L.  N.  (Ed.).  1964  :  166-251. 
1 964 a.     See  Below. 


ZHURAVLEVA,  I.  T.,  KONYUSHOV,  K.  N.  &  ROZANOV,  A.  Yu.  1964.  Archaeocyatha  of  Siberia: 
The  two-walled  Archaeocyatha.  Izd.  Akad.  Nauk  SSR,  Inst.  Geol.  Geofiz.,  Moscow. 
1964  :  1 66,  16  pis.,  75  figs.  [In  Russian.] 

APPENDIX 
GLOSSARY 

APEX  initial  part  of  cup. 

CENTRAL  CAVITY  space  inside  the  inner  wall. 

DISSEPIMENTS  non-skeletal  vesicular  connections  between  the  septa. 

ENDOTHECA  tissue  formed  on  the  central  cavity  side  of  the  inner  wall. 

EXOTHECA  diverse  external  outgrowths  from  the  intervallum  and  outer  wall. 

INTERSEPT/INTERSEPTUM       space  between  two  adjacent  septa. 

INTERTABULUM  space  between  2  successive  tabulae. 

INTERVALLUM  space  between  the  inner  and  outer  walls. 

LINTEAUX  skeletal  elements  around  the  pores. 

LOCULUS  space  limited  by  the  two  walls,  two  adjacent  septa  and  on  occasion 

by  two  successive  tabulae. 
PELLIS  thin  calcareous  sheath,  porous  or  not,  outside  the  walls  of  some 

genera. 

PORES  simple  perforations  in  the  skeletal  elements. 

SEPTA  porous,  radial  vertical  plates  connecting  the  inner  and  outer  walls. 

SYNAPTICULAE  horizontal  rods  between  2  neighbouring  septa. 

TABULAE  straight,   or  curved;   porous,   or  pectinate  plates  that  cross   the 

intervallum. 
TAENIAE  thick,   skeletal;   irregular,   or  waved;   sometimes  gondola-shaped, 

plates,  that  radially  subdivide  the  intervallum  and  are  only  found 

in  the  Irregularia. 
TUMULI  wall  protuberances. 


EXPLANATION    OF    PLATES 


All  the  figured  material  is  in  the  collections  of  the  British  Museum  (Natural 
History). 


PLATE    i 

FIG.  i.  Tumuliolynthus  irregularis  (R.  &  W.  R.  Bedford).  Holotype  84141.  Outer 
view  of  the  wall  and  tumuli.  X  15. 

FIG.  2.  Alphacyathus  cf.  annularis  R.  &  W.  R.  Bedford.  S  4822.  Etched  longitudinal 
section  showing  the  outer  wall,  the  inner  wall  and  the  horizontal  rods.  X  15. 

FIG.  3.  Monocyathus  porosus  R.  &  W.  R.  Bedford.  Lectotype  S  4140.  Cup  showing 
outer  view  of  the  wall.  X  6. 

FIG.  4.  Monocyathus  mellifer  R.  &  W.  R.  Bedford.  84821.  Outer  view  of  the  wall. 
X  6. 

FIG.  5.  Archaeocyathellus  (Stapicyathus)  stapipora  (Taylor).  84733.  Detail  of 
inner  wall.  X  25. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  i 


GEOL.  17,  7 


PLATE   2 
Acanthinocyathus  apertus  R.  &  W.  R.  Bedford 

FIG.  i.  Paratype  S  4167.  View  of  part  of  outer  wall  showing  pores  and  position  of  spines. 
X  10  approx. 

FIG.  2.  Paratype  S  4167.  View  of  part  of  outer  wall  showing  lateral  view  of  a  spine. 
X  6  approx. 

FIG.  3.  Holotype  S  4166.  Another  view  of  outer  wall  showing  the  occurrence  of  the  spines. 
X  10  approx. 

FIG.  4.     Paratype  S  4168.     View  of  inner  wall.      X  4. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  2 


GEOL.  17,  7 


PLATE   3 

Loculicyathus  (Loculicyathellus)  floreus  (R.  &  W.  R.  Bedford) 
Holotype  S  4144. 

FIG.  i.     Inner  wall  and  inner  part  of  septa,      x  10. 
FIG.  2.     Outer  wall  on  two  intersepts.      x  10. 
FIG.  4.     Upper  transverse  section,     x  10. 

Ethmocoscinus  papillipora  (R.  &  W.  R.  Bedford) 

Holotype  S  4164. 
FIG.  3.     Part  of  transverse  section,      x  15. 

Robustocyathus  tnagnipora  (R.  &  W.  R.  Bedford) 

Holotype  S  4146. 
FIG.  5.     Inner  wall,  septa  and  outer  wall  seen  from  the  central  cavity,      x  5. 

Robustocyathus  subacutus  (R.  &  W.  R.  Bedford) 
Holotype  S  4142. 

FIG.  6.     Transverse  section,      x  5. 

FIG.  7.     View  from  central  cavity  of  inner  wall,  septa,      x  5. 


Bull.  Br.  Mus.  nat.  Hist.  (GeoU  17,  7 


PLATE  3 


PLATE   4 
Zonacyathus  retezona  (Taylor) 

FIG.  I.     84731.     Detail  of  inner  wall,      x  12-5. 

FIG.  2.     S  4764.     Oblique  transverse  section,      x  6. 

FIG.  3.     S  4353.     Etched  longitudinal  section  showing  inner  wall,  septa  and  outer  wall,     x  6. 

Zonacyathus  retevallum  (R.  &  W.  R.  Bedford) 

FIG.  4.     Holotype  S  4147.     Detail  of  inner  wall.      X  15. 
FIG.  5.     84726.     Transverse  section,      x  6. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 
1 


PLATE  4 


PLATE   5 

Ethmocyathus  lineatus  R.  &  W.  R.  Bedford 
Holotype  S  4149. 

FIG.  i.     Detail  of  inner  wall  showing  lozenge  pores  and  tabellae.      x  10. 

FIG.  2.     Porosity  of  septa,      x  25. 

FIG.  3.     Inner  view  of  outer  wall.      X4O. 

Cyathocricus  tracheodentatus  R.  &  W.  R.  Bedford 
Holotype  S  4148. 

FIG.  4.     Lower  transverse  section,      x  6. 

FIG.  5.     Etched  longitudinal  section  showing  the  inner  wall,      x  6. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 
1 


PLATE  5 


4?'^: 

jf  ^ '  ^  *^*^  "•*» 

^T*~2*I-*- 
^V*"?!*&,. 

fsir 
^p. 


PLATE   6 
Cyathocricus  dentatus  (Taylor) 

FIG.  i.  S  4753.  Longitudinal  section  showing  the  rings  and  teeth  (note  that  the  septa  are 
only  porous  near  the  outer  wall)  and  outer  wall,  x  30. 

FIG.  2.     S  4753.     Etched  transverse  section,      x  10. 

FIG.  3.  S  4355.  Etched  longitudinal  section  showing  the  outer  wall,  the  septa  and  the  inner 
wall  (at  the  upper  part  of  the  figure),  x  8. 

Cyathocricus  tracheodentatus  (R.  &  W.  R.  Bedford) 
FIG.  4.     S  4754.     Detail  of  inner  wall,      x  25. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  6 


PLATE    7 

Erismacoscinus  equivallum  (Taylor) 
Holotype  S  198. 

FIG.  i.  Detail  of  one  tabula  [on  opposite  side  to  Fig.  2].      X  15. 

FIG.  2.  Transverse  section  showing  inner  wall  pores  and  tabula,      x  3. 

FIG.  3.  Detail  of  outer  wall,      x  8. 

FIG.  5.  Detail  of  inner  wall.      X  15. 

Erismacoscinus  quadratus  (R.  &  W.  R.  Bedford) 

Holotype  S  4157. 
FIG.  4.     Etched  oblique  longitudinal  section,      x  5. 


Bull.  Br.  Mus.  not.  Hist.  (Geol.)  17,  7 


PLATE  7 


PLATE   8 
Erismacoscinus  cellularis  (R.  &  W.  R.  Bedford) 

Holotype  S  4162. 

FIG.  i.     Detail  of  septa  and  inner  wall  (on  right).      X  15. 
FIG.  4.     Oblique  transverse  section,      x  6. 

Erismacoscinus  rugosus  (R.  &  W.  R.  Bedford) 
Holotype  S  4152. 

FIG.  2.     Outer  view  of  outer  wall,      x  10. 
FIG.  3.     Oblique  longitudinal  section,      x  10. 

Erismacoscinus  quadratus  (R.  &  W.  R.  Bedford) 
FIG.  5.     Holotype  S  4157.     Transverse  section  and  detail  of  the  inner  wall,      x  10. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 

1 


PLATE  8 


PLATE   9 
Erismacoscinus  textilis  (R.  &  W.  R.  Bedford) 

Paratype  S  4156. 

FIG.  I.     Detail  of  outer  wall  (lower  right),  septa,  tabulae,  vesicular  tissue  (centre)  and  inner 
wall  (above  left).      x6. 

FIG.  3.     Bowl  shape.      X  1-5. 

Holotype  S  4155. 

FIG.  2.     Cylindrical  shape,      x  4. 
FIG.  4.     Transverse  section,      x  5. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  9 


PLATE    10 
Alataucyathus  (Anaptyctocyathus)  flabellus  sp.  nov. 

Holotype  S  4161. 
FIG.  i.     Outer  view.     xa. 

Eristnacoscinus  retifer  (R.  &  W.  R.  Bedford) 
Holotype  S  4159. 

FIG.  2.     Etched  inner  wall  seen  from  central  cavity,      x  10. 
FIG.  3.     Transverse  section,      x  5. 

Eristnacoscinus  petersi  (R.  &  W.  R.  Bedford) 
Holotype  S  4158. 

FIG.  4.     Transverse  section,      x  10. 
FIG.  5.     Longitudinal  section,      x  10. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  10 


PLATE    ii 

Alataucyathus  (Anaptyctocyathus)  cribripora  (R.  &  W.  R.  Bedford) 

Holotype  S  4160. 

FIG.  i.     Detail  of  outer  wall,  external  view  of  pores  and  micropores.     X25. 
FIG.  3.     Longitudinal  section,      x  10. 

Tomocyathus  (Erugatocyathus)  papillatus  (R.  &  W.  R.  Bedford) 

Holotype  S  4153. 
FIG.  2.     Upper  transverse  section.     X5- 

Salairocyathus  (Salairocyathus)  annulatus  (R.  &  W.  R.  Bedford) 

Holotype  S  4163. 
FIG.  4.     Detailed  longitudinal  section  showing  the  inner  wall,     x  5. 


Bull.  Br.  Mus.  not.  Hist.  (Geol.)  17,  7 


PLATE  ii 


GEOL.  17,  7 


PLATE    12 

Pinacocyathus  spicularis  R.  &  W.  R.  Bedford 
Holotype  S  4169 

FIG.  i.     Transverse  section,      x  7. 

FIG.  2.     Outer  view  showing  the  scaffolding  of  rods,      x  7. 

Flindersicyathus  (Flindersicyathus)  graphicus  (R.  &  W.  R.  Bedford) 

FIG.  3.     Paratype  84171.  Longitudinal  section  through  an  unwaved  taenia  (with  central 
cavity  to  the  left),      x  6. 

FIG.  4.     Paratype  84171.  Longitudinal  section  showing  an  inner  view  of  the  inner  wall,     x  6. 

FIG.  5.     Holotype  S  4170.  Transverse  section,      x  6. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  12 


PLATE    13 

Metaldetes  taylori  (R.  &  W.  R.  Bedford) 
Holotype  S  4185-4187  (three  pieces  of  the  same  specimen) 

FIG.  i.     S  4185.  Lower  part;  external  view.      X2-2  approx. 

FIG.  2.     S  4186.  Upper  part;  transverse  section,      x  1-8. 

FIG.  3.     S  4186.  Upper  part;  oblique  and  longitudinal  sections  with  the  septa  running  N-S. 
X  2  approx. 

These  photographs  are  used  by  courtesy  of  the  British  Museum  (Nat.  Hist.)  and  Dr.  Dorothy 
Hill. 

tCoscinocyathus  unilinearis  (R.  &  W.  R.  Bedford) 
FIG.  4.     Holotype  S  4165.     View  of  both  inner  and  outer  walls  and  intervallum.      x  5. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  13 


f^  •   x»'.   j'if&itf^f^*)       ,     *  *  '''    '  *  i    *  ^^*^i*»     '* 

T*!'.  ',  *  •  v  ^x2v*4  '&*£*  ^>*^-t^**>  x  » 

'%.'.**»  '  *  <-./,<•  ^-^.^^^v*'*^^*.**.  «• 


PLATE    14 
Flindersicyathus  (F.)  speciosus  (R.  &  W.  R.  Bedford) 

Holotype  S  4175. 

FIG.  i.     Lower  transverse  section.      X5- 
FIG.  3.     Etched  longitudinal  section  showing  inner  wall  tubes,      x  5. 

Flindersicyathus  (F.)  major  (R.  &  W.  R.  Bedford) 

FIG.  2.     Holotype  S  4174.     Etched  longitudinal  section  showing  the  bending  of  flat  taeniae 
(on  right)  and  the  widened  tubes  of  the  inner  wall  (on  left),      x  6. 

Flindersicyathus  (F.)  irregularis  (Taylor) 
FIG.  4.     S  4763.     Part  of  transverse  section  showing  thickened  inner  side  of  taenia.     X  10. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  14 


PLATE    15 

FIG.  i.  Flindersicyathus  (Pycnoidocyathus)  synapticulosus  (Taylor)  8208.  Part  of 
transverse  section.  X  2-5. 

FIG.  2.  Flindersicyathus  (P.)  simplex  (Taylor).  Paratype  84824.  Part  of  longitudinal 
and  transverse  sections.  X  3- 

FIG.  3.  Flindersicyathus  (P.)  maximipora  (R.  &  W.  R.  Bedford).  Holotype  S  4150. 
Part  of  an  etched  longitudinal  section  showing  extended  pore-tubes  of  the  inner  wall  and  unwaved 
taeniae.  X  3- 

FIG  4.     Flindersicyathus  (P.)  vicinisepta  (R.  &  W.  R.  Bedford).  S  4825.     Part  of  longi- 
tudinal section  showing  the  inner  wall.      X  5. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  15 


PLATE    16 
Mctaldetes  dissepimentalis  (Taylor) 

FIG.  i.     84176.     Lateral  view  of  non- waved  taeniae.     x6. 
FIG.  2.     84182.     Part  of  transverse  section.      x6. 
FIG.  3.     8  4176.     Outer  view  of  outer  wall.      X  6. 

Metaldetes  irregularis  (R.  &  W.  R.  Bedford) 
FIG.  4.     Paratype  S  4190.     Outer  view  of  outer  wall,      x  6. 


Bull.  Br.  Mus.  not.  Hist.  (Geol.)  17,  7 


PLATE  16 


PLATE    17 
Metacoscinus  reteseptatus  R.  &  W.  R.  Bedford 

FIG.  i.  S  4194.  Outer  view  of  outer  wall,      x  6. 

FIG.  2.  S  4191.  Part  of  transverse  section  showing  a  tabula,      x  10. 

FIG.  3.  S  4195.  Inner  wall  seen  from  the  central  cavity.      x6. 

FIG.  4.  S  4194.  Section  along  a  taenia  (central  cavity  on  left).      x6. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  17 


PLATE    18 

Metafungia  reticulata  R.  &  W.  R.  Bedford 
Holotype  S  4184. 

FIG.  i.     Inner  wall  of  cup  (on  left).      X4- 

FIG.  2.     Outer  wall,      x  6. 

FIG.  3.     Transverse  section.      X3. 

Photographs  by  courtesy  of  the  British  Museum  (Nat.  Hist.)  and  Dr.  Dorothy  Hill. 

Pycnoidocoscinus  pycnoideum  R.  &  W.  R.  Bedford 
S  4832. 

FIG.  4.  Part  of  transverse  section  showing  thickened  taeniae.      X5. 

FIG.  5.  Longitudinal  section  showing  inner  wall,      x  5. 

FIG.  6.  Section  showing  area  of  fig.  7.      X5- 

FIG.  7.  Detail  of  tabulae  pores,      x  30. 

FIG.  8 .  Detail  of  taeniae  and  tabulae,      x  30. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  7 


PLATE  18 


PRINTED  IN  GREAT  BRITAIN 
BY  ADLARD  &  SON  LIMITED 
BARTHOLOMEW  PRESS,  DORKING 


SOME  BATHONIAN  OSTRACODA 
OF  ENGLAND  WITH  A  REVISION 
OF  THE  JONES  1884  AND  JONES 
&  SHERBORN  1888  COLLECTIONS 


R.  H.  BATE 


BULLETIN    OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
GEOLOGY  Vol.  17  No.  8 

LONDON:  1969 


SOME  BATHONIAN  OSTRACODA  OF  ENGLAND 

WITH  A  REVISION  OF  THE  JONES  1884  AND 

JONES  &  SHERBORN  1888  COLLECTIONS 


BY 

RAYMOND  HOLMES  BATE 


Pp.  377-437;  16  plates,  18  text-figures 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

GEOLOGY  Vol.  17  No.  8 

LONDON:  1969 


THE     BULLETIN    OF    THE    BRITISH    MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they  become 
ready.  Volumes  will  contain  about  three  or  four 
hundred  pages,  and  will  not  necessarily  be  completed 
within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  17,  No.  8  of  the  Geological 
(Palaeontological]  series.  The  abbreviated  titles  of 
periodicals  cited  follow  those  of  the  World  List  of 
Scientific  Periodicals. 


World  List  abbreviation 
Bull.  Br.  Mus.  nat.  Hist.  (Geol.). 


Trustees  of  the  British  Museum  (Natural  History)  1969 


TRUSTEES    OF 
THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 

Issued  16  May,  1969  Price  £3  55. 


SOME  BATHONIAN  OSTRACODA  OF  ENGLAND 

WITH  A  REVISION  OF  THE  JONES  1884  AND 

JONES  &  SHERBORN  1888  COLLECTIONS 

By  R.  H.  BATE 


CONTENTS 

Page 
I.     INTRODUCTION          .........         379 

II.     THE  T.  R.  JONES  COLLECTION  ......         380 

III.  THE  T.  R.  JONES  &  C.  D.  SHERBORN  AND  THE  H.  H.  WINWOOD 

COLLECTIONS         .........         394 

IV.  THE  J.  F.  BLAKE  COLLECTION  ......         427 

V.     REFERENCES   ..........         435 

SYNOPSIS 

Three  Bathonian  ostracod  Collections  from  the  Fuller's  Earth  Clay  of  Bath,  Somerset,  and 
one  of  similar  age  (the  T.  R.  Jones  Collection,  published  in  1884)  from  the  Richmond  borehole, 
Surrey,  are  described  and  illustrated,  that  of  J.  F.  Blake  for  the  first  time.  The  H.  H.  Win  wood 
Collection  is  identified  as  syntypic  material  of  the  T.  R.  Jones  &  C.  D.  Sherborn  Collection, 
published  in  1888.  Sixty- three  ostracod  species  are  described  of  which  six  are  new,  and  six 
are  left  under  open  nomenclature.  Nineteen  species  are  listed  as  lost.  Two  genera,  Hadro- 
cytheridea  and  Hekistocy there,  are  new. 

I.     INTRODUCTION 

IN  1884  T.  R.  Jones  described  the  foraminifera  and  Ostracoda  from  a  deep  boring  at 
Richmond,  Surrey.  The  ostracods  were  described  from  three  levels  and,  apart  from 
Pontocyprella  harrisiana  and  Schuleridea  jonesiana  which  are  Cretaceous  contami- 
nants of  the  collection,  are  Bathonian  in  age. 

Four  years  later,  Jones  &  Sherborn  (1888)  described  a  much  larger  ostracod  fauna 
from  the  Bathonian  of  Midford,  near  Bath.  The  samples  of  Blue  and  Yellow  Fuller's 
Earth  Clay  were  given  to  Jones  and  Sherborn  by  the  Rev.  H.  H.  Win  wood  (see  Jones 
&  Sherborn  1886). 

The  collections  of  Jones  1884,  and  of  Jones  &  Sherborn  1888,  are  of  considerable 
importance  for  present-day  work.  Because  of  this  it  is  proposed  here  to  re-describe 
and  re-illustrate  the  species  and  to  designate  types  where  necessary;  a  few  species 
originally  described  by  Jones  &  Sherborn  are  missing  from  the  collections  and  are 
presumed  lost. 

Two  additional  collections  from  the  Fuller's  Earth  of  Midford,  near  Bath,  are 
included  because  they  add  to  the  faunal  list  and  in  the  case  of  the  Blake  collection 
record  the  presence,  for  the  first  time  in  England,  of  a  number  of  species  previously 
only  described  from  N.W.  France  (Oertli  1959).  The  most  important  of  the  collec- 
tions, however,  is  that  presented  by  the  Rev.  H.  H.  Winwood.  A  label  stating  that 

GEOL.    17,  8  23 


380  REVISION    OF    SOME 

these  specimens  are  duplicates  of  the  originals  figured  in  1888  was  left  with  the  collec- 
tion by  C.  D.  Sherborn.  As  the  specimens  figured  by  Jones  &  Sherborn  were  ex- 
tracted from  material  given  them  by  Winwood,  there  would  appear  to  be  no  doubt 
that  this  collection  represents  syntypic  material. 

The  Fuller's  Earth  Clay  of  Midford,  near  Bath,  was  placed  by  Arkell  (1956  :  28) 
in  the  Oppelia  aspidoides  Zone.  Torrens  (1967  :  83),  however,  assigned  it  to  the  zone 
of  Prohecticoceras  retrocostatum,  basal  Upper  Bathonian  and  below  the  aspidoides 
Zone. 

The  terminology  and  classification  employed  throughout  this  paper  are  those  of 
Moore  (1961)  and  Bate  (1963).  All  the  material  described  is  in  the  Department  of 
Palaeontology,  British  Museum  (Natural  History). 


II.  THE  T.  R.  JONES  COLLECTION 

Order  PODOCOPIDA  Miiller  1894 

Suborder  PLATYCOPINA  Sars  1866 

Family  CYTHERELLIDAE  Sars  1866 

Genus  CYTHERELLA  Jones  1849 

Cytherella  symmetrica  Jones 

(PI.  I,  figS.  I,  2) 

1884     Cytherella  symmetrica  Jones  :  768,  pi.  34,  fig.  42. 
1884     Cytherella  subovata  Jones  :  773,  pi.  34,  fig.  43. 

DIAGNOSIS.  Cytherella  with  elongate-oval  carapace.  Left  valve  uniformly  over- 
reached by  larger  right  valve.  Greatest  height  of  carapace  in  posterior  third.  Shell 
surface  very  finely  punctate.  Dimorphic:  Adult  female  of  length  0-85  mm.,  male 
of  length  0-90  mm. 

LECTOTYPE.    IN.43503,  female  right  valve;  Richmond  boring,  depth  1,151  ft.  6  ins. 

PARALECTOTYPES.  IN.43496,  male  left  valve,  and  1.941,  female  carapace;  depth 
1,205  ft. 

DESCRIPTION.  Both  left  and  right  valves  are  elongate-oval  in  outline  with  well- 
rounded  anterior  and  posterior  margins.  Ventral  margin  straight  to  slightly  convex, 
dorsal  margin  strongly  convex  in  the  posterior  third  (region  of  greatest  height)  but 
straightens  out  anteriorly.  Greatest  length  through  mid-point.  Right  valve 
larger  than  the  left  which  it  overreaches  evenly  all-round.  Shell  surface  very  finely 
punctate,  though  the  state  of  preservation  does  not  always  show  this.  Selvage  in 
the  left  valve  strongly  developed  to  form  a  ridge  which  fits  into  a  corresponding 
groove  in  the  right. 

DIMENSIONS.  IN.43503.  Female  right  valve  and  lectotype  of  Cytherella  sym- 
metrica Jones,  length  0-85  mm. ;  height  0-52  mm.  11^.43496.  Male  left  valve  and 


ENGLISH    BATHONIAN    OSTRACODA  381 

lectotype  of  Cytherella  subovata  Jones,  length  0-90  mm. ;  height  0-50  mm.  1.941. 
Female  carapace,  length  0-77  mm. ;  height  0-49  mm. ;  width  0-36  mm. 

REMARKS.  The  figured  specimens  of  C.  symmetrical  and  of  C.  subovata  are  missing 
from  the  Jones  Collection  and  are  presumed  lost.  Unfigured  syntypes  have  therefore 
been  selected  as  lectotypes.  C.  subovata  has  been  recognized  as  the  male  dimorph 
and  C.  symmetrica  as  the  female  dimorph  of  the  species  Cytherella  symmetrica. 

Oval  species  of  Cytherella  look  very  much  alike,  and  C.  symmetrica  is  very  similar 
to  C.  suprajurassica  Oertli  (1957  :  649,  pi.  i,  figs.  i-io).  The  latter,  however,  is  a 
smaller  species  with  a  greater  mid-dorsal  projection  of  the  right  valve  over  the  left. 
In  C.  symmetrica  the  overreach  of  the  right  valve  is  much  more  uniformly  developed 
along  the  entire  dorsal  margin. 


Genus  CYTHERELLOIDEA  Alexander  1929 
Cytherelloidea  jugosa  (Jones) 

(PI.  i,  figs.  3,  4;  Text-fig,  i) 
1884     Cytherella  jugosa  Jones  :  773,  pi.  34,  fig.  44. 

DIAGNOSIS.  Carapace  sub-rectangular  in  outline  with  broadly  rounded  anterior 
and  posterior  margins.  Ventral  margin  strongly  incurved.  Ornamentation  consists 
of  a  broad  peripheral  ridge  and  a  thick,  central  sigmoid  ridge.  Shell  surface  finely 
punctate. 

LECTOTYPE.  ^.43497,  female  right  valve  (figured  Jones  1884) ;  Richmond  boring, 
1,205  ft. 

PARALECTOTYPE.     1.2311,  female  carapace;  Richmond  boring,  depth  not  recorded. 

DESCRIPTION.  Carapace  sub-rectangular  in  outline  with  well  rounded  anterior 
and  posterior  margins.  Dorsal  margin  broadly  convex  with  a  slight  concavity  just 
anterior  of  mid-point.  Ventral  margin  strongly  incurved.  Greatest  length  through 
mid-point ;  greatest  height  median,  despite  the  strong  concavity  of  the  ventral  margin. 
Greatest  width  at  the  posterior  margin.  The  two  specimens  available  are  female 
dimorphs  possessing  two  posterior  swellings,  of  which  the  lowermost  is  the  more 
prominent.  Shell  surface  finely  punctate,  further  ornamented  by  a  broad  peripheral 
ridge  extending  completely  around  each  valve  and  enclosing  a  sigmoid  inner  ridge 
which  commences  at  the  anterior  inner  edge  of  the  peripheral  ridge,  bends  upwards, 
then  curves  down  below  the  mid-dorsal  muscle  scar  depression,  finally  curving 
upwards  again  to  die  out  before  reaching  the  peripheral  ridge  in  the  region  ol  the 
postero-dorsal  angle.  Right  valve  larger  than  the  left,  which  it  overreaches  anteriorly 
and  overlaps  ventrally  and  dorsally,  especially  anterodorsally  where  there  is  a  slight 
concavity  in  the  margin.  Only  internal  details  of  the  right  valve  observed:  Muscle 
scars  situated  on  a  slightly  raised  dorso-median  boss,  are  typical  of  the  genus;  a 
groove  extends  around  the  posterior  and  along  the  dorsal  margin  for  the  reception 
of  the  left  valve.  Two  posterior  cavities  correspond  to  the  swellings  seen  on  the 
outside  of  the  valve. 


382  REVISION  OF  SOME 

DIMENSIONS. 

IN-43497.     Female  right  valve,  length  0-55  mm. ;  height  0-30  mm. 
1.2311.          Female  carapace,  length  0-56  mm. ;  height  0-34  mm. ;  width  0-23  mm. 
REMARKS.     C.  jugosa  is  similar  to  C.  paraweberi  Oertli  (1957  :  651,  pi.  i,  figs. 
12-15)  but  differs  in  being  rounded  rather  than  angular  postero-dorsally,  in  being 
more  noticeably  constricted  mid-dorsally,  and  in  being  narrower  anteriorly  where 
only  the  right  valve  tends  to  project.     Laterally,  the  ribbing  of  C.  jugosa  is  very 
much  broader.     A  weak  reticulation  may  be  observed  in  C.  paraweberi  on  the  lateral 
surface. 


FIG.  i.     Cytherelloidea  jugosa  (Jones).     Female  right  valve.     Lectotype  11^.43497.      X  135. 


FIG.  2.     Cytherelloidea  paraweberi  Oertli.     Right  side,  female  carapace,  10.3527.      x  130. 

Stratigraphically,  C.  jugosa  predates  C.  weberi  Steghaus  (1951  :  207,  pi.  14,  figs. 
3-6)  and  C.  paraweberi  Oertli,  both  of  which  appear  to  be  restricted  to  the 
Kimmeridgian.  The  close  similarity  in  ornamentation  between  these  species  suggests 
a  phylogenetic  relationship  (Text-figs,  i,  2).  This  is  probably  also  true  for  the 
Bajocian  species.,  Cytherelloidea  eastfieldensis  Bate  (19630  :  25,  pi.  i,  figs.  1-5,  Text- 
fig,  i)  which  is  easily  distinguishable  by  the  mid-dorsal  discontinuity  of  the  peripheral 
ridge. 


ENGLISH    BATHONIAN    OSTRACODA  383 

Suborder  PODOCOPINA  Sars  1866 

Superfamily  BAIRDIAGEA  Sars  1888 

Family  BAIRDIIDAE  Sars  1888 

Genus  BAIRDIA  McCoy  1844 

Bairdia  hilda  Jones 
(PL  i,  figs.  5,  6;  PL  4,  fig.  5) 

1884  Bairdia  hilda  Jones  :  771,  pi.  34,  fig.  20. 

1888  Bairdia  fullonica  Jones  &  Sherborn  :  253,  pi.  5,  fig.  ^a-c. 

1948  Bairdia  cf.  hilda  Jones;  Sylvester-Bradley  :  199,  Text-fig.  5. 

1963  Bairdia  hilda  Jones;  Bate  :  188,  pi.  2,  figs.  9-12,  pi.  3,  figs.  1-4. 

1964  Bairdia  hilda  Jones;  Bate  :  8. 

DIAGNOSIS.  Carapace  subdeltoid  laterally,  strongly  convex  dorsally.  Dorsal 
margin  high  with  steeply  inclined  anterior  and  posterior  slopes.  Posterior  end 
acuminate,  upturned.  Shell  surface  finely  punctate. 

LECTOTYPE.  IN.4I95I,  single  left  valve  figured  Jones  (1884)  Richmond  boring, 
depth  1,205  ft. 

PARALECTOTYPES.  10.3608-16  and  10.3620-6,  carapaces  and  single  valves  from 
a  depth  of  1,205  ft. 

REMARKS.  Full  descriptions  of  this  species  have  been  given  previously  by 
Sylvester-Bradley  (1948)  and  Bate  (1963).  However,  as  the  measurements  of  the 
types  were  not  included  these  are  given  here. 

DIMENSIONS.  IN.4I95I,  left  valve,  length  0-93  mm. ;  height  0-48  mm.  10.3608. 
carapace,  length  0-77  mm. ;  height  0-48  mm. ;  width  0-34  mm.  10.3610,  carapace, 
length  o-qo  mm. ;  height  0-47  mm. ;  width  0-34  mm. 

Bairdia  juddiana  Jones 
(PL  i,  fig.  7) 

1884     Bairdia  juddiana  Jones  :  767,  pi.  34,  fig.  18. 

DIAGNOSIS.  Carapace  globose,  convex  in  dorsal  view.  Shell  surface  finely 
punctate.  Antero-dorsal  and  postero-dorsal  slopes  convex.  Left  valve  distinctly 
overlaps  the  right  along  the  dorsal  and  ventral  margins,  particularly  mid-ventrally. 
Caudal  process  slightly  upturned.  Ventral  margin  angled  with  flattened  median 
portion  and  steeply  sloping  antero-ventral  and  postero- ventral  slopes. 

HOLOTYPE.  IN.435o6,  carapace,  figured  Jones  (1884),  Richmond  boring,  depth 
1,151  ft.-i,i5i  ft.  6  in. 

DESCRIPTION.  Carapace  rather  globose  in  lateral  view  with  convex  antero-dorsal 
and  postero-dorsal  slopes.  Anterior  end  rounded,  posterior  end  with  caudal  process 
slightly  upturned  although  damaged  in  the  specimen  available.  Dorsal  margin 


384  REVISION    OF    SOME 

with  convex  central  section  and  steeply  sloping  antero-dorsal  and  postero-dorsal 
slopes.  Greatest  length  of  carapace  through  the  mid-point;  greatest  height  and 
width  slightly  anterior  to  the  mid-point.  Left  valve  larger  than  the  right  which  it 
overlaps  along  the  entire  dorsal  and  ventral  margins  and  overreaches  slightly  at  the 
anterior  margin.  Shell  surface  finely  punctate.  Internal  features  not  known. 

DIMENSIONS.  IN.43506,  carapace,  length  0-85  mm. ;  height  0-59  mm. ;  width 
0-46  mm. 

REMARKS.  Bairdia  juddiana  is  sufficiently  distinct  for  the  single  carapace  to  be 
considered  a  separate  species;  it  differs  from  B.  hilda  in  being  shorter,  higher  and  more 
strongly  convex  in  dorsal  view.  The  postero-dorsal  slope  of  B.  hilda  is  more  strongly 
concave  than  that  of  B.  juddiana  which  lacks  the  slight  anterior  upturning  of  the 
antero-dorsal  slope  seen  in  the  former. 


Bairdia  jurassica  Jones 

(PI.  2,  figS.  I,  2) 

1884     Bairdia  jurassica  Jones  :  771,  pi.  34,  fig.  21. 

1884     Bairdia  jurassica  var  tenuis  Jones  :  771,  pi.  34,  fig.  22. 

DIAGNOSIS.  Carapace  elongate,  drawn  out,  with  acuminate  slightly  upturned 
caudal  process.  Dorsal  margin  broadly  arched  with  obliquely  sloping  antero-dorsal 
and  postero-dorsal  slopes.  Left  valve  larger  than  the  right  which  it  overlaps  antero- 
dorsally,  postero-dorsally  and  mid-ventrally.  Postero-ventrally,  the  left  valve 
turned  outwards  to  reveal  margin  of  right  valve.  Shell  surface  very  finely  punctate. 

LECTOTYPE.  IN.43494,  figured  Jones  1884,  pi.  34,  fig.  21,  Richmond  boring, 
depth  1,205  ft. 

PARALECTOTYPES.  ^.43495,  figured  Jones  1884,  pi.  34,  fig.  22,  and  10.3617, 
carapaces  from  depth  1,205  ft.,  Richmond  boring. 

DESCRIPTION.  Carapace  drawn  out,  elongate,  greatest  length  just  below  mid- 
point. Greatest  height  and  width  just  anterior  to  mid-point.  Dorsal  margin 
broadly  arched  with  the  antero-  and  postero-dorsal  slopes  obliquely  inclined  and 
only  slightly  convex.  Anteiior  end  rounded,  posterior  end  acuminate,  not  promi- 
nently upturned.  Ventral  margin  broadly  convex,  almost  straight.  Left  valve 
larger  than  the  right  which  it  overlaps  along  the  antero-dorsal  and  postero-dorsal 
slopes,  especially  towards  the  extreme  anterior,  and  posteriorly  along  the  caudal 
process.  Left  valve  overlap  prominent  mid-ventrally;  left  valve  very  noticeably 
turned  outwards  postero-ventrally  from  the  right  to  reveal  the  margin  of  the  right 
valve.  Shell  surface  finely  punctate.  Internal  features  not  seen. 

DIMENSIONS.  IN.43494,  carapace,  length  0-85  mm. ;  height  0-39  mm. ;  width 
0-31  mm.  IN.  43495,  carapace,  length  0-93  mm. ;  height  0-40  mm. ;  width  0-34  mm. 
10.3617,  carapace,  length  0-88  mm. ;  height  0-43  mm. ;  width  0-30  mm. 


ENGLISH    BATHONIAN    OSTRACODA  385 

REMARKS.  From  the  dimensions  given  above  it  can  be  seen  that  the  variety 
tenuis  is  more  elongate  than  the  other  two  specimens.  Some  variants  of  Bairdia 
hilda  Jones  are  similar  to  the  present  species  which  may  be  distinguished  on  the 
shape  of  its  carapace  and  the  type  of  overlap  by  the  left  valve. 

Superfamily  CYPRIDACEA  Baird  1845 
Family  PARACYPRIDIDAE  Sars  1923 
Genus  PONTOCYPRELLA  Ljubimova  1955 
Pontocyprella  harrisiana  (Jones  1849) 
(PL  2,  fig.  3) 

1849     Bairdia  harrisiana  Jones  :  25,  pi.  6,  figs.  ija-f. 
1884     Macrocypris  bradiana  Jones  :  766,  pi.  34,  fig.  23. 

For  a  full  synonymy  of  Cretaceous  forms  see  Kaye  1965  :  73. 

MATERIAL.  IN.435oo,  left  valve  figured  Jones  (1884),  depth  1,145  ft.  9  in.- 
1,146  ft.  6  in. 

REMARKS.  Jones  (1849  :  25)  first  recorded  this  species  as  Bairdia  harrisiana 
from  a  number  of  horizons,  but  typically  from  the  Cretaceous  (Gault,  Leacon  Hill; 
Chalk,  Gravesend  and  Charlton;  Detritus,  Charing  and  from  the  Speeton  Clay, 
Yorkshire) . 

The  presence  of  this  species  at  a  depth  of  1,145  ft.  9  in.-i,i46  ft.  6  in.  in  the 
Richmond  bore  indicates  that  some  contamination  of  the  cored  material  has  occurred. 
As  Jones  himself  originally  described  P.  harrisiana,  it  is  somewhat  surprising  that 
he  failed  to  recognize  it  here. 

DIMENSIONS.     IN. 43500,  left  valve,  length  0-88  mm.;  height  0-42  mm. 

Superfamily  CYTHERACEA  Baird  1850 
Family  SCHULERIDEIDAE  Mandelstam  1959 
Subfamily  SCHULERIDEINAE  Mandelstam  1959 

Genus  SCHULERIDEA  Swartz  &  Swain  1946 

Schuleridea  (Schuleridea)  jonesiana  (Bosquet) 

(P.I  2,  fig.  5) 

1884     Cytheridea  subperforata  Jones  :  772,  pi.  34,  fig.  26,  [not  p.  768,  pi.  34,  fig.  25]. 

For  the  synonymy  of  the  Cretaceous  forms  see  Kaye  1964  :  45. 

MATERIAL.     1X43490,  carapace,  figured  Jones  (1884),  Richmond  boring,  1,205  ft- 

REMARKS.  The  specimen  described  by  Jones  (1884  :  768)  fiom  1,151  ft.- 
1,151  ft.  6  in.  in  the  Richmond  boring  is  not  conspecific  with  that  described  from 
1,205  ft-  The  latter  is  a  Cretaceous  contaminant. 

DIMENSIONS.  1X43490,  carapace,  length  0-95  mm.;  height  0-60  mm.;  width 
0-43  mm. 


386  REVISION    OF    SOME 

Schuleridea  (Eoschulerided)  subperforata  (Jones) 
(PI.  2,  fig.  4) 

1884     Cytheridea  subperforata  Jones  :  768,  pi.  34,  fig.  25,  [not  p.  772,  pi.  34,  fig.  26], 

LECTOTYPE.  IN.43504,  right  valve,  figured  Jones  1884,  pi.  34,  fig.  25.  Richmond 
boring,  1,151  ft.- 1,151  ft.  6  in. 

REMARKS.  Male  right  valve  elongate-oval  in  outline  with  a  characteristic  promi- 
nence in  the  region  of  the  anterior  cardinal  angle.  Anteio-dorsal  slope  short, 
steeply  inclined,  and  concave.  Anterior  and  posterior  ends  rounded;  ventral  margin 
convex  with  a  prominent  antero- ventral  incurvature.  At  least  twenty-one  anterior 
radial  pore  canals  in  typical  fan-like  arrangement.  Hinge  broken,  terminal  teeth 
incomplete.  Duplicature  well  developed,  particularly  postero-ventrally.  Poor 
preservation  prevents  further  description. 

In  outline  this  species  is  similar  to  the  male  dimorph  of  Schuleridea  (Eoschuleridea) 
bathonica  Bate  (1967  :  41)  but  may  be  distinguished  by  the  antero-dorsal  prominence 
in  the  region  of  the  cardinal  angle  and  being  slightly  more  narrowly  rounded 
posteriorly. 

DIMENSIONS.     IN.435O4,  right  valve,  length  073  mm. ;  height  0-39  mm. 


Schuleridea  (Eoschuleridea)  trigonalis  (Jones) 
(PI.  2,  fig.  7) 

1884     Bairdia  trigonalis  Jones  :  767,  pi.  34,  fig.  19. 

DIAGNOSIS.  Carapace  trigonal  in  outline  with  high  dorsal  margin:  Dorsum  short, 
slightly  convex,  posteriorly  sloping  in  right  valve ;  umbonate  left  valve  which  promi- 
nently overreaches  the  right.  Greatest  length  of  carapace  below  mid-point. 
Greatest  height  anterior  of  mid-point  in  right  valve,  through  mid-point  in  left. 
Anterior  end  rounded,  posterior  strongly  acuminate.  Antero-  and  postero-dorsal 
slopes  steeply  inclined.  Anterior  radial  pore  canals  few  in  number. 

HOLOTYPE.  IN. 42373,  carapace  figured  Jones  1884,  pi.  34,  fig.  19;  Richmond 
boring,  depth  1,151  ft.-i,i5i  ft.  6  in. 

DESCRIPTION.  Carapace  strongly  trigonal  in  lateral  view  with  the  umbonate  left 
valve  projecting  noticeably  above  the  right.  Posterior  end  strongly  acuminate, 
anterior  end  rounded.  Greatest  length  of  carapace  below  mid-point ;  greatest  height 
of  the  left  valve  through  mid-point  and  of  the  right  valve  slightly  forward  of  this. 
Greatest  width  median.  Ventral  margin  broadly  convex;  dorsal  margin  in  the  right 
valve  short  and  slightly  convex,  sloping  to  the  posterior.  Dorsal  margin  in  the  left 
valve  strongly  umbonate.  Antero-dorsal  and  postero-dorsal  slopes  steeply  inclined, 
slightly  convex.  Shell  surface  very  finely  punctate.  Left  valve  larger  than  the  right 
which  it  overlaps  along  the  ventral  margin,  around  the  posterior  and  along  the 


ENGLISH    BATHONIAN    OSTRACODA  387 

antero-dorsal  and  postero-dorsal  slopes.  Mid-dorsally  the  left  valve  projects  strongly 
above  the  right.  Internal  details  not  seen.  Anterior  radial  pore  canals  not  clearly 
seen  but  appear  to  be  relatively  few. 

DIMENSIONS.  ^.42373,  carapace,  length  0-70  mm. ;  height  0-48  mm. ;  width 
0-36  mm. 

REMARKS.  If  the  species  is  dimorphic,  the  holotype  must  be  that  of  a  female 
carapace.  A  species  close  to  this  is  Schuleridea  (Eoschuleridea]  bathonica  Bate  (1967  : 
41)  from  which  it  may  be  distinguished  by  the  strongly  umbonate  left  valve  and  the 
more  strongly  acuminate  posterior. 


Genus  PRAESCHULERIDEA  Bate  1963 

Praeschuleridea  schwageriana  (Jones) 

(PI.  2,  fig.  6) 

1884     Cythere  schwageriana  Jones  :  766,  pi.  34,  fig.  27. 

DIAGNOSIS.  Carapace  oval  in  outline  with  the  left  valve  projecting  above  the 
right.  Postero-dorsal  slope  of  left  valve  rounded,  posterior  rounded.  Shell  surface 
finely  punctate. 

HOLOTYPE.  ^.43499,  female  carapace,  figured  Jones  1884,  pi.  34,  fig.  27.  From 
depth  1,145  ft-  9  in.-i,i46  ft.  6  in. 

DESCRIPTION.  Carapace  oval  in  outline  with  rounded  anterior  and  posterior 
margins.  Ventral  margin  broadly,  but  not  strongly  convex  in  left  valve,  somewhat 
flattened  in  the  right  valve.  Dorsal  margin  strongly  convex  in  the  left  valve, 
projecting  above  the  right.  Anterior  cardinal  angle  rounded,  posterior  angle  more 
sharply  acute.  Postero-dorsal  slope  steeply  inclined,  more  strongly  convex  in  the 
left  valve.  Antero-dorsal  slope  convex  in  both  valves.  Greatest  length  through 
mid-point.  Greatest  height  median;  greatest  width  slightly  behind  mid-point. 
Shell  surface  finely  punctate.  Left  valve  larger  than  the  right.  No  internal 
features  observed. 

DIMENSIONS.  1^43499,  female  carapace,  length  0-60  mm. ;  height  0-39  mm. ; 
width  0-31  mm. 

REMARKS.  Jones  (1884)  referred  to  but  a  single  specimen  for  this  species  and 
illustrated  a  left  valve.  However,  in  the  plate  description  he  referred  to  fig.  27  as  a 
right  valve.  The  actual  specimen  is  in  fact  a  complete  carapace,  and  in  the  absence 
of  any  evidence  to  the  contrary  is  considered  here  to  be  the  holotype. 

Praeschuleridea  schwageriana  is  similar  to  P.  subtrigona  (Jones  &  Sherborn  1888) 
but  is  larger  and  less  angular  posteriorly :  it  is  close  to  P.  subtrigona  intermedia  Bate 
(1965)  but  may  be  distinguished  by  the  more  uniformly  rounded  posterior  end. 
Given  more  material  showing  only  slight  variation  in  outline,  it  might  not  be  possible 
to  distinguish  P.  schwageriana  from  one  of  the  subspecies  of  P.  subtrigona. 


388  REVISION    OF    SOME 

Praeschuleridea  sp. 

(PI.  2,  fig.  8) 

DESCRIPTION.  Carapace  trigonal  in  outline  with  the  left  valve  strongly  umbonate 
and  projecting  above  the  right  valve.  Strongly  convex  when  viewed  dorsally. 
Greatest  length,  height  (left  valve)  and  width  pass  through  mid-point.  Right  valve 
somewhat  umbonate  in  the  region  of  the  anterior  cardinal  angle,  below  which  there 
is  a  deep,  oblique  groove.  Left  valve  larger  than  the  right.  Anterior  and  posterior 
margins  rounded;  ventral  margin  broadly  convex.  Internal  details  not  seen. 
Anterior  radial  pore  canals  in  part  seen  from  the  exterior  and  appear  typical  for  the 
genus. 

DIMENSIONS.  10.3619,  female  carapace,  length  0-59  mm. ;  height  0-43  mm. ; 
width  0-35  mm. 

REMARKS.  This  somewhat  globose,  triangular  ostracod  is  similar  in  general 
outline  to  Schuleridea  trigonalis  (Jones  1884) ,  but  differs  most  markedly  in  that  the  line 
of  greatest  length  passes  through  mid-point  and  not  below  it  as  in  the  latter  species. 

This  species  of  Praeschuleridea  is  almost  certainly  new  and  was  found  in  a  slide  of 
duplicate  material  obtained  from  the  Richmond  boring;  depth  1,151  ft.  6  in.  It  is 
considered  inadvisable  to  erect  a  new  species  on  only  a  single  specimen. 

Family  PROGONOGYTHERIDAE  Sylvester-Bradley  1948 

Subfamily  PROGONOCYTHERINAE  Sylvester-Bradley  1948 

Genus  GLYPTOCYTHERE  Brand  &  Malz  1962 

Glyptocythere  guembeliana  (Jones) 

(PI.  3,  figs,  i,  2 ;  PI.  4,  fig.  i) 

1884  Cythere  guembeliana  Jones  :  772,  pi.  34,  figs.  32,  33,  [not  fig.  31]. 

1888  Cytheridea  pulvinar  Jones  &  Sherborn  :  266,  pi.  3,  figs.  za-c. 

1888  Cytheridea  trapezoidalis  Terquem,  Jones  &  Sherborn  :  270,  pi.  4,  figs.  la,  b. 

1967  Glyptocythere  guembeliana  (Jones)  Bate  :  49,  pi.  13,  figs.  10-16,  pi.  14,  figs.  1-8. 

DIAGNOSIS.  Carapace  subquadrate,  elongate  in  male  dimorph.  Lateral  surface 
with  transverse  ridges  extending  down  from  dorsal  margin,  though  generally  poorly 
developed.  Marginal  borders  compressed.  Ventro-lateral  margin  evenly  convex 
in  female,  sharply  directed  upwards  posteriorly  in  male  right  valve.  Hinge  weakly 
entomodont. 

LECTOTYPE.  IN.43493,  male  right  valve  from  Richmond  boring,  depth  1,205  ft. 
Figured  Jones  (1884,  pi.  34,  fig.  33). 

PARALECTOTYPE.     10.3338,  male  carapace  from  same  depth. 

REMARKS.  This  species  was  revised  by  Bate  (1967  :  49).  The  specimen  illus- 
trated is  the  lectotype  (IN. 43493)  figured  by  Jones  1884,  pi.  34,  fig.  33,  and  recorded 
from  a  depth  of  1,205  ft.  The  specimen  for  fig.  32  is  missing  and  that  of  fig.  31  is 
not  considered  conspecific. 


ENGLISH    BATHONIAN    OSTRACODA  389 

DIMENSIONS.  Lectotype:  IN.43493,  male  right  valve,  length  0-96  mm. ;  height 
0-49  mm. 

Genus  FASTIGATOCYTHERE  Wienholz  1967 
Fastigatocy  there  juglandica  (Jones) 
(PI.  3,  figs.  4,  7,  8;  PI.  12,  fig.  3) 

1884  Cythere  juglandica  Jones  :  766,  768,  pi.  34,  figs.  36,  37. 

1888  Cythere  juglandica  var.  major  Jones  &  Sherborn  :  225,  pi.  4,  figs.  2a-b. 

1948  Progonocythere  juglandica  (Jones)  Sylvester-Bradley  :  193,  pi.  12,  figs.  5,  6,  pi.  13,  fig.  8. 

1963  Progonocythere  juglandica  juglandica  (Jones);  Grekoff  :  1731,  pi.  3,  fig.  55. 

1963  Progonocythere?  juglandica  (Jones) ;  Oertli  :  pis.  28,  29,  30. 

1967  Glyptocythere  juglandica  (Jones)  Bate  :  51. 

DIAGNOSIS.  Carapace  sub-rectangular  tapering  strongly  to  posterior  end.  Left 
valve  larger  than  right,  projecting  above  right  valve  dorsally,  except  mid-dorsally 
where  the  umbonate  right  valve  strongly  projects.  Ornamentation  of  transverse 
ridges  and  reticulae  radiate  down  from  dorsal  margin.  Ventro-lateral  margins 
convex;  carapace  swollen  medially.  Flattened  marginal  borders.  Hinge  entomo- 
dont. 

LECTOTYPE.  1X41947,  right  valve.  Richmond  boring,  1,146  ft.  Figured  Jones 
(1884). 

PARALECTOTYPES.     1X41948-9.     Two  carapaces  from  1,151  ft.  6  in. 

REMARKS.  This  species  was  described  by  Sylvester-Bradley  (1948)  who  placed  it 
in  his  new  genus  Progonocythere.  Brand  &  Malz  (1962)  removed  from  that  genus 
those  species  which  have  a  more  quadrate  outline,  which  possess  a  distinct  ornamenta- 
tion not  usually  present  in  species  of  Progonocythere  s.s.,  and  which  also  have  a  dorsal 
projection  of  the  dorso-median  part  of  the  right  valve.  The  last  feature  was  noted 
by  Sylvester-Bradley  (1948  :  194)  and  used  by  Bate  (1967  :  51)  to  justify  the  assign- 
ment of  this  species  to  Glyptocythere.  Subsequently  Wienholz  (1967  :  25)  separated 
from  Glyptocythere  those  species  which  have  a  more  elongate  carapace  outline,  a  more 
positive  antero-dorsal  furrow  and  strongly  diverging  ribs  radiating  from  the  dorsal 
margin.  For  these  species  Wienholz  erected  the  genus  Fastigatocy there,  to  which  she 
assigned  juglandica.  This  assignment  is  accepted  here. 

DIMENSIONS.  Lectotype :  1X41947,  right  valve,  length  0-93  mm. ;  height  047  mm. 
1X41948,  carapace,  length  0-87  mm. ;  height  0-51  mm. ;  width  049  mm.  1X41949, 
carapace,  length  0-77  mm. ;  height  049  mm. ;  width  049  mm. 

Genus  LOPHOCYTHERE  Sylvester-Bradley  1948 

REMARKS.  There  is  an  understandable  reluctance  on  the  part  of  some  ostracod 
workers  to  identify  a  genus  or  subgenus  simply  on  an  ornamental  variation.  However, 
some  ostracod  lineages  fall  naturally  into  morphological  groups  in  which  a  particular 
ornamental  trend  is  evident.  I  feel  that  within  the  Ostracoda  there  is  ornamentation 


390  REVISION    OF    SOME 

of  a  primary  kind  which  reflects  a  generic  or  possibly  subgeneric  status,  and  a  secon- 
dary ornamentation  of  a  more  specific  character.  The  latter  might  be  superimposed 
upon  the  former  or  simply  modify  it. 

Lophocythere  clearly  contains  two  distinct  morphological  groups :  the  first  centred 
around  Lophocythere  ostreata  (Jones  &  Sherborn  1888)  with  an  L-shaped  ridge  ex- 
tending around  the  anterior  margin  and  bending  back  along  the  ventro-lateral  margin 
as  the  diagnostic  character,  and  the  second  around  L.  bradiana  (Jones  1884),  in 
which  several  ridges  extend  across  the  lateral  surface  of  the  carapace. 

Whatley  (personal  communication)  informs  me  that  he  has,  in  manuscript,  sub- 
divided the  genus  Lophocythere  accordingly.  As  this  information  will  shortly  be 
published  by  him  it  is  not  my  intention  to  deal  further  with  this  generic  revision. 

Lophocythere  bradiana  (Jones) 
(PI.  3,  figs.  3,  5,  6;  Text-figs.  3,  4) 

1884     Cythere  bradiana  Jones  :  772,  pi.  34,  figs.  $8a-b. 

1888     Cytheridea  craticula  Jones  &  Sherborn  :  272,  pi.  4,  figs,  ga-c,  loa-c. 

1948     Lophocythere  bradiana  (Jones)  Sylvester-Bradley  :  196,  pi.  14,  figs.  7-10,  pi.  15,  figs.  8-n. 

DIAGNOSIS.  Carapace  subrectangular,  dimorphic.  Left  valve  larger  than,  and 
projects  dorsally  above,  the  right.  Anterior  end  broadly  rounded,  posterior  end 
triangular  with  greatest  length  of  carapace  through  mid-point.  Cardinal  angles 
prominent,  rounded.  Eye  swelling  situated  at  anterior  cardinal  angle.  Lateral 
surface  ornamented  by  four  major  carinae:  dorsal  carina  originates  at,  or  a  short 
distance  from,  the  anterior  margin,  curves  under  the  eye  swelling,  to  which  a  short 
offshoot  runs,  then  curves  back  over  dorso-median  part  of  valve  to  die  out  below 
posterior  cardinal  angle.  The  second  carina  originates  at  the  antero-ventral  margin, 
runs  parallel  to  the  dorsal  carina  for  a  short  distance  then  bifurcates,  the  two 
branches  running  parallel  almost  to  the  posterior  end  where  they  converge  and  almost 
meet.  The  fourth  carina  originates  at  the  antero-ventral  margin  then  follows  a 
course  roughly  parallel  to  lowermost  of  the  two  median  carinae.  A  short  carina 
occurs  between  the  ventral  carina  and  the  lowest  median  carina  in  the  anterior 
half  of  the  valve.  A  second  short  carina,  convex  dorsally,  occurs  between  the  dorsal 
carina  and  the  uppermost  of  the  two  median  carinae  in  the  posterior  half  of  the  valve, 
in  which  region  a  short,  straight  carina,  is  sometimes  developed  between  the  two 
median  carinae.  Shell  surface  reticulate  between  the  carinae. 

LECTOTYPE.  ^.42372,  male  carapace,  figured  Jones  (1884),  Richmond  boring, 
depth  1,205  ft. 

PARALECTOTYPES.  10.3627-9,  one  male  and  two  female  carapaces  from  depth 
1,205  ft. 

DESCRIPTION.  Carapace  subrectangular,  more  elongate  in  the  male  dimorphs. 
Dorsal  and  ventral  margins  virtually  parallel,  with  rounded  anterior,  and  triangular 
posterior  margins.  Cardinal  angles  prominent,  rounded,  especially  prominent  in  the 
larger  left  valve,  the  dorsal  margin  of  which  projects  above  the  right.  Ventral 


ENGLISH    BATHONIAN    OSTRACODA  391 

surface  ornamented  with  three  longitudinal  ridges  per  valve.  Left  valve  uniformly 
overlaps  the  right  along  the  ventral  margin,  more  strongly  so  antero-ventrally. 
Greatest  length  of  carapace  through  mid-point,  greatest  height  in  the  anterior  third, 
greatest  width  in  the  posterior  third.  Ornamentation  of  lateral  surface  as  in  the 
diagnosis.  Eye  swelling  prominently  situated  below  the  anterior  cardinal  angle. 
Details  of  hinge  not  known  from  the  type  material  although  it  is  entomodont  in 
comparative  material. 

DIMENSIONS.  IN.42372,  male  carapace,  length  o-6imm.;  height  0-32  mm.; 
width  0-29  mm.  10.3627,  male  carapace,  length  0-65  mm. ;  height  0-34  mm. ;  width 
0-30  mm.  10.3628,  female  carapace,  length  0-57  mm.  height  0-34  mm. ;  width 
0-29  mm.  10.3629,  female  carapace,  length  0-66  mm. ;  height  0-39  mm. ;  width 
0-34  mm. 

REMARKS.  Sylvester-Bradley  (1948)  placed  Cytheridea  bradiana  Jones  &  Sherborn 
in  synonymy  with  Cythere  bradiana  Jones.  However,  the  specimen  described  by 
Jones  &  Sherborn  is  not  conspecific  but  is  a  paralectotype  of  Lophocythere  septicostata 
Bate  (1967  :  52,  pi.  15,  figs.  7-13,  pi.  16,  figs.  1-4). 


FIG.  3.     Lophocythere  bradiana  (Jones).     Right  valve,  complete  carapace  of  lectotype  of 
Cytheridea  bradiana  Jones.     11^.42372.      x  140. 


FIG.  4.     Lophocythere  bradiana  (Jones).     Right  valve,  complete  carapace  of  lectotype  of 
Cytheridea  craticula  Jones  &  Sherborn.     1.1854.      x  T4°- 


392  REVISION    OF    SOME 

Sylvester-Bradley  (1948  :  197)  considered  Cytheridea  craticula  Jones  &  Sherborn 
1888  to  be  a  subspecies  of  bradiana  because  "  there  is  a  short  anterior  stem  from 
which  spring  the  dorsal,  the  two  median,  and  the  ventral  carinae  ".  Examination 
of  the  types  of  bradiana  and  of  craticula  show  this  is  not,  in  fact  so.  The  dorsal 
carina  does  not  in  all  specimens  reach  the  anterior  margin.  Where  the  reticulate 
ornamentation  is  particularly  well  developed  a  cross  reticulation  gives  the  impression 
that  the  dorsal  carina  is  joined  to  the  median  carina.  The  ventral  carina  originates 
at  the  antero-ventral  margin,  and  it  is  the  short  carina  situated  between  this  and  the 
lowermost  median  carina  which  sometimes  joins  the  stem  of  the  two  median  carinae. 

Although  the  lectotype  of  L.  bradiana  shows  the  development  of  a  low  ridge  or 
carina  between  the  two  median  carinae  in  the  posterior  part  of  the  valve,  many  speci- 
mens do  not  show  this  feature.  It  is,  however,  well  developed  in  the  male  paralecto- 
type,  10.3627,  and  should  not  be  regarded  as  being  a  feature  of  craticula  even  though 
the  lectotype  of  the  latter  shows  it  remarkably  well. 

To  summarize,  the  variations  found  within  a  population  of  Lophocythere  bradiana, 
such  as  the  occasional  imperfect  development  of  the  dorsal  carina  close  to  the  anterior 
margin  and  the  presence  or  absence  of  a  short  carina  posteriorly  between  the  two 
median  carinae,  do  not  indicate  either  a  separate  species  or  subspecies,  but  are  part 
of  the  normal  variation  present  in  the  species. 

L.  bradiana  is  followed  in  the  Oxfordian  of  France  by  L.  multicostata  Oertli  (1957), 
a  species  which  has  a  similar  ornamentation.  However,  L.  multicostata  does  not 
possess  the  anterior  carina  situated  between  the  ventral  and  lower  median  carina  of 
bradiana,  but  has  an  additional  anterior  ridge  in  front  of  the  eye  swelling. 

The  variations  within  Lophocythere  bradiana  are  illustrated  in  Text-figs.  3,  4. 


Genus  MICROPNEUMATOCYTHERE  Bate  1963 
Micropneumatocythere  subconcentrica  (Jones) 
(PL  4,  ng.  7) 

1884     Cy 'there  subconcentrica  Jones  :  768,  pi.  34,  figs.  28,  29. 

1967     Micropneumatocythere  subconcentrica  (Jones)  Bate  :  60,  pi.  21,  figs.  1-13. 

DIAGNOSIS.  Micropneumatocythere  with  oval  carapace  tapering  posteriorly. 
Ventro-lateral  margin  swollen.  Shell  surface  punctate  laterally.  Ventral  surface 
with  longitudinal  ridges  extending  on  to  ventro-lateral  margin  and  turning  upwards 
anteriorly  and  posteriorly.  Internal  details  as  for  genus. 

LECTOTYPE.  IN.43505,  left  valve  from  Richmond  boring,  depth  1,151  ft.  6  in. 
Figured  Jones  (1884,  pi.  34,  fig.  28). 

REMARKS.  This  species  has  been  revised  by  Bate  (1967  :  60)  and  need  not  be 
dealt  with  fully  here.  The  lectotype,  a  left  valve,  is  considered  to  be  the  specimen 
figured  by  Jones  although  the  caption  to  the  figure  states  that  it  is  a  right  valve. 

DIMENSIONS.     IN.43505,  left  valve,  length  0-56  mm.;  height  0-38  mm. 


ENGLISH    BATHONIAN    OSTRACODA  393 

Genus  TERQUEMULA  Blaszyk  &  Malz  1965 

Terquemula  blakeana  (Jones) 

(PI.  4,  fig.  8 ;  PI.  5,  fig.  i) 

1884  Cytheve  blakeana  Jones  :  772,  pi.  34,  figs.  34,  35. 

1888  Cytheridea  transversiplicata  Jones  &  Sherborn  :  267,  pi.  3,  fig.  4. 

?i888  Cytheridea  blakeana  (Jones),  Jones  &  Sherborn  :  265,  pi.  2,  fig.  n. 

1888  Cytheridea  egregia  Jones  &  Sherborn  :  267,  pi.  3,  fig.  5. 

1948  Progonocythere  blakeana  (Jones),  Sylvester-Bradley  :  191,  pi.  12,  figs.  3,  4,  pi.  13,  figs.  4,  5. 

1959  Progonocythere  ?  blakeana  (Jones),  Triebel  &  Klingler  :  339,  341. 

1963  Progonocythere  ?  blakeana  (Jones),  Oertli  :  36,  pi.  25,  fig.  c,  pi.  26,  fig.  c,  pi.  27,  fig.  c, 
pi.  28,  fig.  c,  pi.  30,  fig.  c. 

1965  Terquemula  blakeana  (Jones),  Blaszyk  &  Malz  :  445. 

DIAGNOSIS.  Carapace  oval-elongate,  noticeably  constricted  mid-dorsally  and 
mid-ventrally.  Anterior  end  rounded,  posterior  end  narrowly  rounded.  Postero- 
lateral  part  of  carapace  swollen,  overreaching  ventral  margin.  Greatest  length 
through  or  slightly  above  mid-point.  Shell  surface  with  large,  sunken  normal  pore 
canal  openings  and  a  broad  reticulation  of  wrinkles.  Muscle  scars  of  Type  A  (Bate 
1963  :  180).  Species  dimorphic.  Hinge  entomodont. 

LECTOTYPE.  11^.41950,  female  left  valve,  Richmond  boring,  depth  1,205  ft. 
Figured  Jones  (1884). 

REMARKS.  This  species  was  described  fully  by  Sylvester-Bradley  (1948).  The 
type  specimen  of  Cytheridea  egregia  Jones  &  Sherborn,  placed  by  Sylvester-Bradley 
in  synonymy  with  blakeana,  has  been  lost,  but  the  original  illustration  (pi.  3,  fig.  5) 
suggests  that  the  synonymy  is  justified. 

The  specimen  described  as  C.  blakeana  by  Jones  &  Sherborn  could  be  a  juvenile 
form  of  another  specimen  as  suggested  by  Sylvester-Bradley  (1948  :  193). 

DIMENSIONS.     Lectotype:  female  left  valve,  length  0-69  mm. ;  height  0-39  mm. 

Family  CYTHERURIDAE  Miiller  1894 
Genus  METACYTHEROPTERON  Oertli  1957 
Metacytheropteron  drupaceum  (Jones) 
(PL  5,  figs.  4-6) 

1884  Cy there  drupacea  Jones  :  772,  pi.  34,  fig.  30. 

I957«  Metacytheropteron  sp.  50,  Oertli  :  table  i. 

1963  Cytheropteron  jonesanum  Cory  ell  :  854. 

1967  Metacytheropteron  drupacea  (Jones) ;  Bate  :  44,  pi.  10,  figs.  1-9. 

DIAGNOSIS.  Metacytheropteron  with  elongate/sub-ovate  carapace.  Greatest 
height  at  anterior  cardinal  angle,  posterior  end  acuminate.  Dorsal  margin  broadly 
convex,  especially  in  .left  valve.  Shell  surface  strongly  ornamented  with  triangular 
arrangement  of  longitudinal  and  obliquely  transverse  ridges. 

GEOL.    17,  8  24 


394  REVISION    OF    SOME 

HOLOTYPE.  IN. 43498,  female  carapace  from  Richmond  boring,  depth  1,205  ft. 
Figured  Jones  (1884). 

REMARKS.  Coryell  (1963)  renamed  this  species  Cytheropteron  jonesanum  on  the 
assumption  that  C.  drupacea  mentioned  by  Richter  (1867  :  228)  and  Cy there  drupacea 
of  Jones  were  homonyms.  Reference  to  Richter  (1855  :  529),  however,  indicates 
that  the  species  referred  to  in  1867  had  been  placed  in  Cythereis  and  not  Cy  there  as 
supposed  by  Coryell.  There  is,  therefore,  no  nomenclatural  problem. 

DIMENSIONS.  Holotype.  1X43498,  carapace,  length  0-50  mm. ;  height  0-30  mm. ; 
width  0-29  mm. 

Family  TRACHYLEBERIDIDAE  Sylvester-Bradley  1948 
Genus  CYTHEREIS  Jones  1849 
Cythereis  cf.  blandu  Kaye  1963 
(PI.  5,  figs.  3,  7) 

1884     Cythere  (Cythereis)  quadrilatera  (Roemer),  Jones  :  766,  772,  pi.  34,  figs.  39,  40,  41. 

REMARKS.  Jones  recorded  Cythereis  quadrilatera  Roemer  from  two  horizons  in 
the  Richmond  boring  (1,145  ft.  9  in.-i,i46  ft.  6  in.,  and  1,205  ft.).  These  specimens 
are  probably  conspecific  with  Cythereis  blanda  Kaye  (1963),  although  the  median 
and  ventro-lateral  ridges  in  Jones's  material  tend  to  be  more  straight  and  less 
strongly  developed  than  in  C.  blanda.  But  this  may  be  within  the  species'  variation. 

Jones  referred  to  three  specimens,  here  regarded  as  Cretaceous  contaminants,  of 
Cythere  (Cythereis)  quadrilatera  but  only  two  now  remain ;  these  are  a  complete  cara- 
pace, IN.4349I,  figured  Jones  1884,  pi.  34,  fig.  40,  and  a  single  left  valve  IN.43502, 
figured  pi.  34,  fig.  39.  The  specimen  illustrated  by  fig.  41  is  missing  and  the  illustra- 
tion suggests  that  this  latter  specimen  was  not  conspecific  with  the  other  two. 

DIMENSIONS.  IN.4349I,  carapace,  length  0-77  mm. ;  height  0-46  mm. ;  width 
0-32  mm.  IN.435O2,  left  valve,  length  0-66  mm. ;  height  0-43  mm. 

Family  and  Genus  uncertain 

"  Cythere  "  tenella  Jones 

(PI.  5,  fig-  8) 

1884     Cythere  tenella  Jones  :  772,  pi.  34,  fig.  24. 

REMARKS.     This  small,  poorly-preserved  ostracod  is  probably  a  juvenile  instar. 
HOLOTYPE.     1X43492,  complete  carapace,  Richmond  boring,  depth  1,205  ft- 
DIMENSIONS.     Length  0-38  mm. ;  height  0-21  mm. ;  width  0-15  mm. 

III.  THE  T.  R.  JONES  &  C.  D.  SHERBORN  COLLECTION 
AND  THE  H.  H.  WINWOOD  COLLECTION 

The  following  figured  specimens  of  species  described  by  Jones  &  Sherborn  (1888) 
are  no  longer  represented  in  their  collection  and  may  be  presumed  lost.  Only  the 
first  five  are  known  to  have  been  presented  to  the  Museum. 


ENGLISH    BATHONIAN    OSTRACODA  395 

Bairdia  juddiana  Jones,  p.  253. 

Cytheridea  bicarinata  Jones  &  Sherborn,  p.  270,  pi.  4,  figs.  5#-c. 

Cytheridea  renoides  Jones  &  Sherborn,  p.  266,  pi.  3,  figs.  la-c. 

Cytheridea  winwoodiana  Jones  &  Sherborn,  p.  259,  pi.  I,  figs.  2a-d. 

Cytheridea  eximia  Jones  &  Sherborn,  p.  273,  pi.  5,  figs.  5«-c. 

Bairdia  trigonalis  Jones,  p.  253. 

Bythocypris  winwoodiana  Jones  &  Sherborn,  p.  252,  pi.  5,  figs.  xa-c. 

Cythere  juglandica  var.  minor  Jones  &  Sherborn,  p.  255,  pi.  4,  figs.  3#-c. 

Cythere  ?  speciosa  Jones  &  Sherborn,  p.  254,  pi.  2,  figs.  loa-c.     (This  species 

was  recorded  as  lost  by  Jones  &  Sherborn,  p.  254.) 
Cythere  trapezioides  Jones  &  Sherborn,  p.  256,  pi.  5,  figs.  ioa-c. 
Cythere  walfordiana  Jones  &  Sherborn,  p.  255,  pi.  5,  figs.  ga-c. 
Cytheridea  egregia  Jones  &  Sherborn,  p.  267,  pi.  3,  figs.  5«-c. 
Cytheridea  politula  Jones  &  Sherborn,  p.  265,  pi.  5,  figs.  ja-c. 
Cytheridea  pura  Jones  &  Sherborn,  p.  269,  pi.  3,  figs.  na-d. 
Cytheridea  retorrida  Jones  &  Sherborn,  p.  260,  pi.  i,  figs.  8a~c. 
Cytheridea  rugifera  Jones  &  Sherborn,  p.  271,  pi.  5,  figs.  na-c. 
Cytheridea  sedata  Jones  &  Sherborn,  p.  261,  p.  i,  figs.  loa-c. 
Cytheridea  subeminula  Jones  &  Sherborn,  p.  261,  pi.  5,  figs.  8a-c. 
Cytheridea  ventrosa  Jones  &  Sherborn,  p.  269,  pi.  3,  figs.  loa-c. 
Cythereis  walfordiana  Jones  &  Sherborn,  p.  257,  pi.  5,  figs.  I2a-c. 

Order  PODOCOPIDA  Miiller  1894 
Suborder  PLATYCOPINA  Sars  1866 
Family  GYTHERELLIDAE  Sars  1866 
Genus  CYTHERELLA  Jones  1849 
Cytherella  fullonica  Jones  &  Sherborn 
(PI.  5,  ng.  9;  PI.  6,  fig.  i) 

1888     Cytherella  fullonica  Jones  &  Sherborn  :  274,  pi.  i,  figs.  I2a—c. 
1963     Cytherella  fullonica  Jones  &  Sherborn;  Bate  :  184,  pi.  i,  figs,  i,  2. 

DIAGNOSIS.  Carapace  subrectangular  with  characteristically  steep  postero-dorsal 
slope.  Shell  surface  smooth,  with  dorso-median  muscle-scar  depression. 

LECTOTYPE.  1.1857,  left  valve,  figured  Jones  &  Sherborn  1888,  from  the  Blue 
Fuller's  Earth  Clay. 

PARALECTOTYPES.  1.4004  (Jones  &  Sherborn  Collection);  10.3636-41  (Winwood 
Collection),  left  and  right  valves.  Three  specimens  from  the  Blue,  and  four  from 
the  Yellow  Fuller's  Earth  Clay. 

REMARKS.     This  species  has  been  previously  described  in  Bate  (1963  :  184). 

DIMENSIONS.  Lectotype:  1.1857,  kft  valve,  length  0-64  mm.;  height  0-34  mm. 
Paralectotype  lo.  4004,  left  valve,  length  0-58  mm.;  height  0-32  mm. 


396  REVISION    OF    SOME 

Genus  CYTHERELLOIDEA  Alexander  1929 

Cytherelloidea  catenulata  (Jones  &  Sherborn) 

(PI.  6,  figs.  2,  3) 

1888  Cytherella  catenulata  Jones  &  Sherborn  :  274,  pi.  5,  figs.  6a-c. 

1948  ?  Cytherelloidea  catenulata  (Jones  &  Sherborn)  Sylvester-Bradley  :  200,  pi.  14,  fig.  n. 

1963  Cythetelloidea  catenulata  (Jones  &  Sherborn);  Bate  :  184,  pi.  i,  figs.  3-6. 

1963  Cytherelloidea  catenulata  (Jones  &  Sherborn) ;  Oertli  :  37,  pis.  27,  29. 

1964  Cytherelloidea  catenulata  (Jones  &  Sherborn) ;  Bate  :  8. 

DIAGNOSIS.  Carapace  with  crescent-shaped  swelling  situated  close  to  posterior 
margin  of  valve.  Dorsal  limb  of  crescent  bends  over  and  curves  forwards  close  to 
ventral  margin  to  produce  a  swelling  shaped  like  a  question  mark.  Shell  surface 
ornamented  with  longitudinal  and  terminal  (parallel  to  margins)  striae  producing  a 
reticulation. 

LECTOTYPE.  1.1876,  left  valve,  figured  Jones  &  Sherborn,  from  the  Blue  Fuller's 
Earth  Clay. 

PARALECTOTYPES.  1.1846  (left  valve)  and  10.3525  (right  valve),  Jones  &  Sherborn 
Collection;  10.3649-53,  left  and  right  valves  from  the  Winwood  Collection.  All 
specimens  from  the  Blue  Fuller's  Earth  Clay. 

REMARKS.     This  species  was  described  by  Bate  (1963  :  184-5). 

DIMENSIONS.  1.1876,  left  valve,  length  0-68  mm.;  height  0-37  mm.  1.1846, 
left  valve;  length  0-68  mm. ;  height  0-36  mm.  10.3525,  right  valve,  length  0-68  mm. ; 
height  0-38  mm. 

Cytherelloidea  refecta  (Jones  &  Sherborn) 
(PI.  6,  figs.  4-7) 

1888     Cytheridea  refecta  Jones  &  Sherborn  :  262,  pi.  2,  figs.  3«,  b. 

DIAGNOSIS.  Carapace  oval,  more  elongate  in  male  dimorph.  Anterior  and 
posterior  margins  rounded.  Posterior  cardinal  angle  prominent,  postero-dorsal  slope 
slopes  steeply  to  posterior.  Central  part  of  valve  with  broad  depression  bounded 
below  by  broad  crescentic  swelling.  Crescentic  furrow  delimits  lower  surface  of 
swelling.  A  small  rounded  swelling  is  situated  to  the  inside  of  the  anterior  part  of 
crescentic  swelling. 

LECTOTYPE.  1.1850,  female  right  valve,  figured  Jones  &  Sherborn  (1888),  from 
the  Blue  Fuller's  Earth  Clay. 

PARALECTOTYPES.  10.3931-2,  female  left  valve  and  male  left  valve  from  the 
Winwood  Collection.  Both  specimens  from  the  Blue  Fuller's  Earth  Clay. 

DESCRIPTION.  Carapace  oval  in  outline  with  rounded  anterior  and  posterior 
margins  and  prominent  posterior  cardinal  angle.  The  postero-dorsal  slope  is  steeply 
inclined,  but  slightly  convex.  Dorsal  margin  of  both  valves  slightly  concave  antero- 


ENGLISH    BATHONIAN    OSTRACODA  397 

medially.  Ventral  margin  convex  in  the  right  valve,  concave  in  the  left.  Shell 
surface  with  a  broad,  deep  furrow,  crescentic  in  shape  which  commences  postero- 
dorsally  and  terminates  antero-dorsally.  Above  this  furrow  a  broad  swelling  is 
developed,  also  crescentic  but  rather  irregular  in  outline.  Anteriorly  the  swelling 
appears  to  turn  back  sharply  upon  itself  for  a  short  distance  before  dying  out.  To 
the  inside  of  this  swelling  and  situated  in  the  angle  of  the  anterior  turn-back,  a  small 
circular  swelling  is  developed.  The  main  crescentic  swelling  is  delimited  on  its 
dorsal  side  by  a  broad  central  depression.  In  the  region  of  the  anterior  margin  the 
surface  of  the  shell  may  be  seen  to  be  weakly  reticulate.  Internally  the  dorsal  edge 
of  the  right  valve  possesses  a  groove  for  the  reception  of  the  valve  margin  of  the 
smaller  left  valve. 

DIMENSIONS.  1.1850,  female  right  valve,  length  0-60  mm.;  height  0-26  mm. 
10.3931,  female  left  valve,  length  0-52  mm. ;  height  0-28  mm.  10.3932,  male  left 
valve,  length  0-71  mm. ;  height  0-38  mm. 

REMARKS.  Cytherelloidea  refecta  has  a  distinctive  ornamentation  which  separates 
it  from  other  Jurassic  species  of  the  genus.  Within  the  Jurassic  the  predominant 
ornamentation  of  the  cytherelloideas  consists  of  either  a  peripheral  ridge  or  swelling 
and/or  a  central  ridge  or  swelling.  The  development  of  strong  lateral  ridges  extending 
from  the  posterior  region,  though  present  in  some  Jurassic  species  (Field  1966),  is 
predominantly  a  feature  of  Cretaceous  and  Tertiary  species. 

Suborder  PODOCOPINA  Sars  1866 

Superfamily  BAIRDIACEA  Sars  1888 

Family  BAIRDIIDAE  Sars  1888 

Genus  BAIRDIA  McCoy  1844 

Bairdia  Hilda  Jones 

(PI.  4,  fig-  5) 

1884     Bairdia  hilda  Jones  :  771,  pi.  34,  fig.  20. 

1888     Bairdia  fullonica  Jones  &  Sherborn  :  253,  pi.  5,  figs.  ^a-c. 

For  complete  synonymy  see  p.  383. 

REMARKS.  Sylvester-Bradley  (1948  :  199)  first  pointed  out  that  the  variation 
found  within  specimens  of  Bairdia  hilda  and  Bairdia  fullonica  was  such  that  there 
was  as  yet  no  evidence  for  regarding  these  species  as  distinct.  Accordingly  they 
were  placed  in  synonymy.  The  lectotype  of  B.  fullonica  (1.1873,  a  right  valve  from 
the  Blue  Fuller's  Earth  Clay,  figured  Jones  &  Sherborn)  was  in  the  Jones  &  Sherborn 
Collection,  whilst  the  paralectotype,  also  a  right  valve  (10.3554)  and  from  the  Blue 
Fuller's  Earth  Clay,  was  in  the  Win  wood  Collection. 

DIMENSIONS.  1.1873,  right  valve,  length  079  mm.;  height  0-42  mm.  10.3554, 
right  valve,  length  0-76  mm. ;  height  0-39  mm. 


398  REVISION    OF    SOME 

In  addition  to  the  types  there  are  four  right  valves  and  one  left  valve  (1.1848  from 
the  Jones  &  Sherborn  Collection  and  10.3911-2  and  10.271  from  the  Winwood 
Collection)  from  the  Blue  Fuller's  Earth  Clay  and  referred  to  by  Jones  &  Sherborn 
on  p.  253. 


Bairdia  sherborni  sp.  nov. 
(PI.  6,  fig.  8;  PL  7,  fig.  i) 

DIAGNOSIS.  Carapace  sub-rectangular  in  outline.  Dorsal  margin  very  slightly 
convex,  sloping  slightly  to  posterior.  Ventral  margin  strongly  incurved  medially. 
Anterior  end  obliquely  rounded.  Posterior  end  narrowly  rounded,  somewhat 
tapering,  but  not  upturned.  Left  valve  larger  than  the  right.  Muscle  scars  as  for 
the  genus.  Shell  surface  strongly  and  coarsely  pitted. 

HOLOTYPE.  10.3913,  complete  carapace  from  the  Yellow  Fuller's  Earth  Clay, 
Winwood  Collection. 

PARATYPE.  10.3557,  complete  carapace  from  the  Fuller's  Earth  Clay,  Mockler 
Collection. 

DESCRIPTION.  Carapace  sub-rectangular,  elongate  bairdioid  in  outline,  the  dorsal 
margin  very  slightly  convex  but  almost  straight,  sloping  gently  to  the  posterior. 
The  ventral  margin  is  typical  of  the  genus,  being  strongly  incurved  medially. 
Anteriorly  the  margin  is  obliquely  rounded,  sharply  cut  just  above  a  line  running 
through  mid-point  (line  of  greatest  length)  by  the  inclined  antero-dorsal  slope. 
Postero-dorsal  slope  slightly  concave,  postero-ventral  slope  convex.  Posterior  end 
tends  to  be  tapered  without  the  strong  upturning  of  most  bairdias.  Greatest  height 
of  carapace  through  the  anterior  cardinal  angle ;  greatest  width  in  the  posterior  third. 
Shell  surface  distinctly  and  evenly  pitted.  Left  valve  larger  than  the  right  which  it 
overlaps  mid-ventrally  though  not  antero-ventrally  and  postero-ventrally.  Antero- 
dorsally  and  postero-dorsally  the  gape  is  replaced  by  a  prominent  overlap  of  the 
right  valve  by  the  left.  Internal  details  not  observed  apart  from  the  muscle  scars 
which  may  be  observed  as  impressions  on  the  internal  cast  of  the  holotype.  They 
comprise  seven  adductor  scars  arranged  in  a  circle  with  an  eighth  situated  at  the 
centre  and  a  ninth,  probably  also  an  adductor  scar,  situated  dorsally  to  this  group. 
An  additional  dorsal  scar  is  placed  above  that  last  mentioned  whilst  antero-ventrally 
there  are  two  oval  "  mandibular  "  scars  and  antero-dorsally  a  single  "  antennal " 
scar. 

DIMENSIONS.  10.3913,  carapace,  length  0-58  mm. ;  height  0-31  mm. ;  width 
0-21  mm.  10.3557,  carapace,  length  0-71  mm.;  height  0-39  mm.;  width  0-27  mm. 

REMARKS.  Only  two  specimens  have  been  found,  one  in  each  of  two  major 
collections.  The  outline  of  the  carapace;  obvious  antero- ventral  and  postero-ventral 
gape  and  distinct  surface  pitting  distinguish  this  species  easily  from  all  other  Jurassic 
bairdiids. 


ENGLISH    BATHONIAN    OSTRACODA  399 

Superfamily  GYPRIDACEA  Baird  1845 
Family  PARACYPRIDIDAE  Sars  1923 

Genus  PARACYPRIS  Sars  1866 

Paracypris  terraefullonicae  (Jones  &  Sherborn) 

(PI.  7,  figs.  2,  4) 

1888     Macrocypris  terrae-fullonicae  Jones  &  Sherborn  :  252,  pi.  5,  figs.  $a-c. 

1888     Macrocypris  horatiana  Jones  &  Sherborn  :  252,  pi.  5,  figs.  za-c. 

1967     Paracypris  terraefullonica  (Jones  &  Sherborn)  Bate  :  27,  pi.  i,  figs.  1-6. 

DIAGNOSIS.  Carapace  elongate,  posteriorly  acuminate.  Anterior  end  rounded. 
Ventral  margin  almost  straight  in  the  larger  left  valve,  more  strongly  concave  in  the 
right.  Dorsal  margin  arched  with  antero-dorsal  slope  tending  to  be  slightly  concave, 
more  noticeably  so  in  the  right  valve.  Shell  surface  smooth.  Anterior  and  posterior 
vestibules  well-developed.  Radial  pore  canals  branching. 

LECTOTYPE.  1.1875,  left  valve,  figured  Jones  &  Sherborn  1888,  pi.  5,  figs.  3«-c, 
from  the  Blue  Fuller's  Earth  Clay. 

OTHER  MATERIAL.  1.1874,  right  valve  (lectotype  of  Macrocypris  horatiana  Jones 
&  Sherborn),  figured  Jones  &  Sherborn  1888,  pi.  5,  figs.  2a-c,  and  10.3548-53  (para- 
lectotypes  of  M.  horatiana  and  M.  terraefullonicae],  from  the  Blue  Fuller's  Earth 
Clay. 

REMARKS.  Paracypris  terraefullonicae  has  been  fully  described  in  Bate  (1967  :  27). 
The  additional  specimens  listed  above  (10.3548-53)  were  found  in  the  Winwood 
Collection  and  are  paralectotypes  of  Macrocypris  horatiana  and  Macrocypris  terrae- 
fullonicae which  "  species  "  are  the  right  and  left  valves  respectively  of  Paracypris 
terraefullonicae. 

DIMENSIONS.  1.1875,  left  valve,  length  0-60  mm.;  height  0-28  mm.  1.1874, 
right  valve,  length  0-58  mm. ;  height  0-26  mm. 

Superfamily  CYTHERACEA  Baird  1850 
Family  BYTHOCYTHERIDAE  Sars  1926 

Genus  MONOCERATINA  Roth  1928 
Monoceratina  visceralis  (Jones  &  Sherborn) 

(PI-  7>  fig-  3) 

1888     Cytheridea  visceralis  Jones  &  Sherborn  :  268,  pi.  3,  figs.  6a-c. 

DIAGNOSIS.  Carapace  sub-rectangular  in  outline  with  rounded  anterior  end;  the 
antero-dorsal  margin  tends  to  curve  obliquely  backwards.  Posterior  end  triangular 
with  greatest  prolongation  situated  in  dorsal  half  of  valve.  Dorsal  and  ventral 
margins  parallel.  A  distinct  median  sulcus  present  in  dorsal  half  of  valve  only. 


400  REVISION    OF    SOME 

Valve  convex,  prominently  swollen  in  postero-ventral  region.  Shell  surface  strongly 
pitted.  Postero-ventral  margin  slightly  serrated.  Hinge  with  strong,  straight, 
median  bar. 

HOLOTYPE.  1.1830,  left  valve,  figured  Jones  &  Sherborn  1888,  from  the  Blue 
Fuller's  Earth  Clay. 

DESCRIPTION.  The  carapace  is  sub-rectangular  in  outline  with  a  triangular 
posterior  end,  the  ventral  margin  of  which  is  longer  than  the  straight  postero-dorsal 
margin,  and  is  evenly  serrated  along  its  length.  At  the  postero-ventral  angle  there 
is  a  distinct  convexity  of  the  margin.  Anterior  cardinal  angle  prominently  rounded. 
Dorsal  and  ventral  margins  parallel.  A  distinct  vertical  median  sulcus  is  present  in 
the  dorsal  half  of  the  valve  only.  Carapace  strongly  swollen  along  the  ventro-lateral 
border  and  postero-ventrally  where  the  swelling  is  strongly  undercut.  Shell  surface 
coarsely  pitted,  the  pits  extending  uniformly  over  the  valve.  Line  of  greatest  length 
lies  above  mid-point,  whilst  the  line  of  greatest  height  passes  approximately  through 
the  centre  of  the  valve.  Internally  the  hinge  has  been  damaged,  only  the  posterior 
part  of  the  strong  hinge  bar  being  present.  No  other  internal  details  observed. 

DIMENSIONS.     1.1830,  left  valve,  length  0-66  mm.;  height  0-30  mm. 

REMARKS.  Only  the  holotype  is  known,  but  it  is  distinct  from  the  other  species 
of  Monoceratina  described  from  the  Jurassic.  M.  visceralis  is  close  to  M.  vulsa 
(Jones  &  Sherborn),  although  the  latter  has  a  much  coarser  ornamentation,  almost 
reticulate,  whilst  the  median  sulcus  is  much  more  strongly  developed,  as  is  the 
under-cutting  of  the  ventro-lateral  and  postero-ventral  swelling.  M.  vulsa  is  also 
distinguished  by  possessing  a  distinct  furrow  paralleling  the  anterior  margin. 
M.  ungulina  Triebel  &  Bartenstein  (1938,  pi.  I,  figs.  3,  4)  may  be  distinguished  on 
outline,  the  posterior  extremity  being  situated  higher  up  on  the  valve  because  of  the 
shorter  postero-dorsal  slope.  The  anterior  margin  is  also  uniformly  rounded  with 
a  flattened  marginal  border. 


Monoceratina  vulsa  (Jones  &  Sherborn) 
(H.  7,  ng.  5) 

1888  Cytheridea  vulsa  Jones  &  Sherborn  :  263,  pi.  2,  figs.  $a,  b. 

1938  Monoceratina  vulsa  (Jones  &  Sherborn)  Triebel  &  Bartenstein  :  516,  pi.  3,  figs.  17,  18. 

1960  Monoceratina  cf.  vulsa  (Jones  &  Sherborn) ;  Lutze  :  433,  pi.  37,  figs.  $a,  b. 

?I963  Monoceratina  sp.  juv.  aff.  vulsa  (Jones  &  Sherborn);  Plumhoff  :  48,  pi.  n,  figs.  167,  168. 

1963  Monoceratina  vulsa  (Jones  &  Sherborn);  Bate  :  189,  pi.  3,  figs.  5-12. 
19630  Monoceratina  vulsa  (Jones  &  Sherborn);  Bate  :  26,  pi.  i,  fig.  6. 

1964  Monoceratina  vulsa  (Jones  &  Sherborn) ;  Bate  :  9. 

DIAGNOSIS.  Shell  sub-rectangular  in  lateral  view,  convex  in  dorsal  view.  Vertical 
median  sulcus  deeply  incised,  surrounded  below  and  to  the  sides  by  a  prominent 
swelling  which  is  strongly  undercut  ventro-laterally.  Lateral  swelling  separated 
from  anterior  margin  by  a  furrow  which  runs  parallel  to  that  margin.  Shell  surface 
strongly  pitted,  the  raised  borders  of  the  pits  giving  the  surface  a  wrinkled  appearance. 


ENGLISH    BATHONIAN    OSTRACODA  401 

LECTOTYPE.  1.1842,  a  badly  damaged  left  valve,  figured  Jones  &  Sherborn  1888, 
from  the  Blue  Fuller's  Earth  Clay. 

REMARKS.  Only  a  single  damaged  valve  remains  of  the  two  specimens  originally 
placed  in  this  species  by  Jones  &  Sherborn  (p.  263),  and  this  is  the  figured  specimen. 
A  complete  description  was  given  by  Bate  (1963)  who  incorrectly  recorded  the 
lectotype  as  coming  from  the  Richmond  boring. 

DIMENSIONS.  1.1842,  length  0-64  mm.;  height  0-32  mm.  Both  measurements 
would  have  been  slightly  larger  if  the  specimen  had  not  suffered  slight  damage 
posteriorly  and  along  the  dorsal  margin. 

Family  TRACHYLEBERIDIDAE  Sylvester-Bradley  1948 

Subfamily  TRACHYLEBERIDINAE  Sylvester-Bradley  1948 

Genus  OLIGOCYTHEREIS  Sylvester-Bradley  1948 

Oligocythereis  fullonica  (Jones  &  Sherborn) 

(PI.  7,  fig.  6) 


1888     Cythereis  fullonica  Jones  &  Sherborn  :  256,  pi.  4,  figs, 

1948     Cythereis  cf.  fullonica  Jones  &  Sherborn;  Sylvester-Bradley  :  186,  pi.  12,  figs.  7,  8  [not 

figs.  9,  10],  pi.  13,  fig.  3  [not  fig.  9]. 

I948«  Oligocythereis  fullonica  (Jones  &  Sherborn)  Sylvester-Bradley  :  796. 
1963     Oligocythereis  fullonica  (Jones  &  Sherborn)  ;  Oertli  :  39,  pi.  25,  fig.  a?,  pi.  29,  fig.  a,  [not 

pi.  24,  fig.  a,  pi.  26,  fig.  a]. 
1967     Oligocythereis  fullonica  (Jones  &  Sherborn);  Bate  :  61,  pi.  21,  fig.  14,  [not  fig.  15]. 

DIAGNOSIS.  Carapace  subquadrate  in  outline  with  a  broadly  rounded  anterior  and 
broadly  triangular  posterior  end.  Anterior  and  posterior  margins  with  small 
denticles.  Ventral  surface  flattened,  V-shaped  in  outline;  ventral  margin  gently 
convex  anteriorly  and  posteriorly.  Dorsal  margin  almost  straight,  approximately 
paralleling  ventral  margin.  Anterior  cardinal  angle  extremely  prominent  with  a 
large,  rounded  eye  tubercle  below.  A  short,  thick  ridge  extends  obliquely  antero- 
ventrally  from  eye  tubercle.  Dorso-median  ridge  extends  back  from  eye  tubercle, 
enlarging  posteriorly  to  give  carapace  triangular  outline  in  dorsal  view.  Termination 
of  dorso-median  ridge  produces  a  sharply  pointed  tubercle,  angular  in  outline,  the 
two  sides  of  which  are  at  right  angles.  Postero-ventrally  a  short,  raised  ridge  almost 
vertically  directed  bends  round  at  an  oblique  angle  to  extend  forwards  ventro-laterally 
bending  upwards  antero-ventrally  to  terminate  in  a  prominent  tubercle.  This 
ventro-lateral  ridge  describes  a  broadly  crescentic  outline  around  a  prominent, 
irregular  tubercle.  Prominent  tubercle  developed  slightly  anterior  of  mid-point. 
Shell  surface  very  finely  punctate.  Left  valve  larger  than  right. 

LECTOTYPE.  1.1871,  complete  carapace,  figured  Jones  &  Sherborn  1888,  from  the 
Blue  Fuller's  Earth  Clay. 

DESCRIPTION.  Carapace  subquadrate  in  outline,  ornamented  as  diagnosed.  The 
angular  outline  of  the  shell  is  produced  dorsally  by  the  prominent  antero-dorsal 


402  REVISION    OF    SOME 

eye  tubercle  and  the  strong  postero-dorsal  tubercle  and  ventrally  by  the  postero- 
ventral  and  antero- ventral  terminations  of  the  ridge  which  extends  along  the  ventro- 
lateral  margin.  The  large  tubercle  situated  slightly  anterior  of  mid-point  is  the 
muscle  scar  node  characteristic  of  the  family.  The  anterior  and  posterior  margins 
possess  small  denticles  whilst  the  marginal  borders  bear  a  few  additional  nodes. 
Shell  surface  finely  punctate.  In  dorsal  view  the  outline  of  the  carapace  in  the 
median  part  is  V-shaped  as  is  the  ventral  surface.  This  outline  is  produced  by  the 
dorso-median  and  ventro-lateral  ridges  diverging  slightly  towards  the  posterior  end. 
Left  valve  larger  than  the  right  which  it  overlaps  along  the  ventral  margin  and  in 
the  region  of  the  cardinal  angles  and  overreaches  along  the  postero-dorsal  slope. 
Internal  details  not  observed  in  the  type  material,  but  the  entomodont  type  hinge 
has  been  described  by  Sylvester-Bradley  (1948  :  187). 

DIMENSIONS.  Lectotype,  1.1871,  carapace,  length  0-54 mm.;  height  0-32  mm.; 
width  0-27  mm. 

REMARKS.  Sylvester-Bradley  (1948  :  187)  noted  variation  within  the  ornamenta- 
tion of  specimens  which  he  placed  in  0.  fullonica  (Jones  &  Sherborn),  and  it  has  been 
current  practise  to  retain  these  within  the  species.  The  forms  possessing  a  postero- 
dorsal  tubercle  from  which  three  arms  radiate  (see  Sylvester-Bradley  1948,  pi.  12, 
figs.  9,  10)  should  be  assigned  to  a  new  species.  0.  fullonica  should  be  retained  only 
for  those  specimens  in  which  the  postero-dorsal  tubercle  has  two  arms  at  right  angles, 
and  in  which  the  muscle  scar  node  to  the  centre  of  the  valve  is  completely  isolated 
and  not  joined  to  the  anterior  termination  of  the  ventro-lateral  ridge. 

Family  SCHULERIDEIDAE  Mandelstam  1959 

Subfamily  SCHULERIDEINAE  Mandelstam  1959 

Genus  SCHULERIDEA  Swartz  &  Swain  1946 

Subgenus  EOSCHULERIDEA  Bate  1967 
Schuleridea  (Eoschuleridea)  horatiana  (Jones  &  Sherborn) 

(PI.  7,  fig.  7;  Text-fig.  5) 
1888     Cytheridea  horatiana  Jones  &  Sherborn  :  263,  pi.  2,  figs.  $a,  b. 

DIAGNOSIS.  Carapace  oval/elongate,  anteriorly  rounded,  posteriorly  tapering, 
narrowly  rounded.  Greatest  length  of  carapace  below  mid-point,  greatest  height 
at  anterior  cardinal  angle.  Elongate  eye  swelling  with  groove  beneath  present  in 
right  valve  below  anterior  cardinal  angle.  Left  valve  projects  slightly  above  right 
just  behind  anterior  cardinal  angle.  Shell  surface  punctate  with  widely  scattered, 
large,  normal  pore  canals.  Twenty-five  to  thirty  radial  pore  canals  splayed  fan-like 
around  anterior  margin.  Muscle  scars  type  C.  Hinge  paleomerodont. 

LECTOTYPE.  1.1852,  male  right  valve,  figured  Jones  &  Sherborn  1888,  from  the 
Blue  Fuller's  Earth  Clay. 

PARALECTOTYPE.  10.3940,  male  carapace  from  Blue  Fuller's  Earth  Clay,  Winwood 
Collection. 


ENGLISH    BATHONIAN    OSTRACODA  403 

DESCRIPTION.  Carapace  oval/elongate  in  outline  with  uniformly  rounded  anterior 
margin  and  narrowly  rounded,  tapered  posterior.  The  line  of  greatest  length  passes 
well  below  the  mid-point.  Greatest  height  at  the  anterior  cardinal  angle,  greatest 
width  just  behind  the  mid-point.  Dorsal  margin  convex,  slightly  umbonate  in  the 
left  valve  just  behind  the  anterior  cardinal  angle.  Ventral  margin  convex  with  an 
antero-median  incurvature.  Shell  surface  punctate  with  widely  scattered,  large, 


FIG.  5.     Muscle  scars,  x  200.     Schuleridea  (Eoschuleridea)  horatiana  (Jones  &  Sherborn), 

lectotype,  1.1852. 

rounded,  normal  pore  canals.  An  elongate  eye  swelling  is  situated  below  the 
anterior  cardinal  angle  in  the  right  valve  and  has  an  oblique  furrow  situated  beneath. 
Muscle  scars  as  viewed  externally  of  type  G  (Bate  1963).  Anterior  radial  pore 
canals  splayed  fan-like  around  the  anterior  margin,  largely  concentrated  below 
mid-height,  approximately  twenty-five  to  thirty  in  number.  Hinge  paleomerodont 
as  seen  in  the  right  valve.  Terminal  teeth  larger  anteriorly,  where  there  are  five, 
than  posteriorly,  where  there  are  eight.  Median  bar  smooth.  Left  valve  larger 
than  right.  Inner  margin  and  line  of  concrescence  coincide,  anterior  duplicature 
broad. 

DIMENSIONS.  1.1852,  right  valve,  length  0-85  mm.;  height  0-46  mm.  10.3940, 
carapace,  length  0-79  mm. ;  height  0-46  mm. ;  width  0-36  mm. 

REMARKS.  Both  available  specimens  of  this  species  are  male  dimorphs  as  indi- 
cated by  their  shell  outline.  No  female  dimorph  has  been  recognized.  This  species 
is  very  close  to  the  male  dimorph  of  Schuleridea  (Eoschuleridea)  bathonica  Bate  (1967) 
although  the  latter  has  fewer  anterior  radial  pore  canals  and  does  not  have  the  line 
of  greatest  length  as  ventrally  positioned  as  in  horatiana.  Male  dimorphs  of 
Schuleridea  species  are  notoriously  alike  and  extremely  difficult  to  tell  apart.  Until 


404  REVISIONOFSOME 

a  female  dimorph  has  been  identified  for  horatiana  from  the  Fuller's  Earth  it  is 
preferred  to  regard  these  two  species  as  distinct. 


Genus  ASCIOCYTHERE  Swain  1952 
Asciocythere  obovata  (Jones  &  Sherborn) 
(PI.  7,  fig.  8;  PI.  8,  figs.  2,  3,  7;  Text-fig.  6) 
1888     Cytheridea  obovata  Jones  &  Sherborn  :  264,  pi.  2,  figs.  6a-c. 

DIAGNOSIS.  Carapace  oval  in  lateral  view,  strongly  convex  in  dorsal  view. 
Dorsal  margin  broadly  arched,  curving  down  into  well-rounded  anterior  and  more 
narrowly  rounded  posterior  margin  without  change  in  slope  at  cardinal  angles. 
Ventral  margin  broadly  convex.  Line  of  greatest  length  slightly  below  mid-point. 
Shell  surface  very  finely  punctate. 

LECTOTYPE.  1.1836,  right  valve,  figured  Jones  &  Sherborn  1888,  pi.  2,  figs.  6a-c, 
from  the  Yellow  Fuller's  Earth  Clay. 

PARALECTOTYPE.  10.3938,  carapace  from  the  Yellow  Fuller's  Earth  Clay, 
Winwood  Collection. 

OTHER  MATERIAL.  10.3939,  left  valve  from  the  Blue  Fuller's  Earth  Clay,  Winwood 
Collection. 

DESCRIPTION.  Carapace  oval  in  outline  with  strongly  arched  dorsal  margin  and 
indistinct  cardinal  angles,  the  dorsal  margin  curving  down  without  a  break  into  the 
rounded  anterior  margin  and  the  more  narrowly  rounded  posterior  margin.  Ventral 
margin  broadly  convex.  Shell  surface  appears  smooth  but  is  in  fact  very  finely 
punctate.  Greatest  length  of  carapace  just  below  mid-point.  Greatest  height 
median;  greatest  width  just  behind  mid-point.  Left  valve  larger  than  right,  over- 
lapping the  right  along  the  ventral,  posterior  and  postero-dorsal  slopes.  Antero- 
dorsally  and  anteriorly  the  valves  tend  to  gape  and  there  is  no  overlap.  Hinge 
antimerodont :  right  valve  with  ten  dorsally  bifid  anterior  teeth  and  nine  dorsally 
bifid  posterior  teeth.  Median  groove  very  short  and  loculate.  Left  valve  hinge  not 
seen.  Inner  margin  and  line  of  concrescence  coincide,  duplicature  strongly 
developed.  Anterior  radial  pore  canals  widely  spaced  and  only  very  slightly 
curved,  almost  straight,  sixteen  in  number.  Muscle  scars  not  observed. 

DIMENSIONS.  1.1836,  right  valve,  length  0-54  mm.;  height  0-34  mm.;  10.3938, 
carapace,  length  0-49  mm. ;  height  0-35  mm. ;  width  0-30  mm.  10.3939,  left  valve, 
length  0-62  mm. ;  height  0-42  mm. 

REMARKS.  Only  two  species  of  Asciocythere  have  been  described  from  the  British 
Middle  Jurassic  so  far;  A.  lacunosa  Bate  1963^  and  A.  acuminata  Bate  1964,  both 
from  the  Bajocian.  The  present  species,  the  first  to  be  described  from  the  Bathonian, 
is  not  so  posteriorly  acuminate  as  A.  acuminata  and  not  so  strongly  pitted  as 
A.  lacunosa. 


ENGLISH    BATHONIAN    OSTRACODA  405 


FIG.  6.     Asciocythere  obovata  (Jones  &  Sherborn).     Internal  view,  right  valve,  lectotype, 

1.1836.      x  115. 

Genus  PRAESCHULERIDEA  Bate  1963 

Praeschuleridea  subtrigona  subtrigona  (Jones  &  Sherborn) 

(PL  8,  figs.  4-6) 

1888     Cytheridea  subtrigona  Jones  &  Sherborn  :  265,  pi.  2,  figs.  ga-c. 

1963  Praeschuleridea  subtrigona  (Jones  &  Sherborn)   Bate  :  207,  pi.   12,  figs.    12-16,  pi.   13, 
figs.  1-9. 

1963(3  Praeschuleridea  subtrigona  (Jones  &  Sherborn);  Bate  :  41. 

1964  Praeschuleridea  subtrigona  subtrigona  (Jones  &  Sherborn);  Bate  :  22. 

1965  Praeschuleridea  subtrigona  subtrigona  (Jones  &  Sherborn);  Bate  :  124. 

DIAGNOSIS.  Carapace  oval-subtrigonal  in  outline,  punctate.  Length  of  adult 
of  the  order  of  (female)  0-56  mm. ;  (male)  0-58  mm. 

LECTOTYPE.  1.1838,  male  right  valve,  not  the  figured  specimen  of  Jones  & 
Sherborn,  which  was  of  a  left  valve.  Blue  Fuller's  Earth  Clay. 

PARALECTOTYPES.  10.3935-7,  female  and  two  male  carapaces,  Blue  Fuller's 
Earth  Clay,  Win  wood  Collection. 

REMARKS.  Praeschuleridea  subtrigona  is  the  type  species  of  the  genus  and  as  such 
has  been  described  fully  in  earlier  publications.  Three  subspecies  are  recognized 
so  far,  subtrigona  subtrigona  being  the  smallest  and  having  the  longest  stratigraphical 
range:  Middle  Bajocian  (Sonninia  sowerbyi  Zone,  Hyperlioceras  discites  Subzone)  to 
Upper  Bathonian  (Prohecticoceras  retrocostatum  Zone) .  The  other  two  subspecies,  P. 
subtrigona  magna  Bate  (1964)  and  P.  subtrigona  intermedia  Bate  (1965),  have  much 
shorter  ranges  and  are  restricted  to  the  Bajocian.  The  measurements  of  the  type 
specimens  have  not  previously  been  given  and  are  included  here. 

DIMENSIONS.  1.1838,  male  right  valve,  length  0-53  mm.;  height  0-32  mm. 
10.3935,  male  carapace,  length  0-55  mm. ;  height  0-37  mm. ;  width  0-29  mm.  10.3936, 
male  carapace,  length  0-54  mm.;  height  0-36  mm.;  width  0-29  mm.  10.3937, 
female  carapace,  length  0-50  mm. ;  height  0-35  mm. ;  width  0-30  mm.  Comparison 
of  the  above  measurements  with  those  given  in  Bate  (1963  :  209)  shows  that  the  size 
of  the  P.  subtrigona  subtrigona  has  remained  stable  since  the  Bajocian. 


406  REVISION    OF    SOME 

Genus  EOCYTHERIDEA  Bate  1963 
Eocytheridea  sp. 

(PI.  8,  fig.  i) 

REMARKS.  Known  from  a  single  right  valve,  possibly  of  a  male  dimorph 
(Winwood  Collection).  The  surface  of  the  valve  is  strongly  pitted  by  rather  broad 
pits  which,  towards  the  centre  of  the  valve,  produce  a  reticulation  but  in  the  antero- 
dorsal  and  postero-dorsal  areas  are  elongated  obliquely  towards  the  centre  of  the 
dorsal  margin.  This  elongation  produces  a  series  of  ridges  which  radiate  down  from 
the  centre  of  the  dorsal  margin.  The  ornamentation  readily  distinguishes  this  species 
from  Eocytheridea  faveolata  Bate  (1964)  which  has  a  more  uniform  reticulate  orna- 
mentation without  the  development  of  obliquely  radiating  ridges  in  the  dorsal  part 
of  the  carapace.  The  hinge  is  hemimerodont.  Twelve  long,  slightly  curved  anterior 
radial  pore  canals  pass  through  the  broad  duplicature. 

This  is  the  first  record  of  the  genus  outside  the  Bajocian 

DIMENSIONS.     10.3906,  right  valve,  length  0-77  mm. ;  height  0*36  mm. 


Family  CYTHERIDEIDAE  Sars  1925 

Subfamily  CYTHERIDEINAE  Sars  1925 

Genus  HADROCYTHERIDEA  nov. 

DERIVATION  OF  NAME,     hadros,  Gr.,  well  developed  +  cytheridea. 

DIAGNOSIS.  Carapace  robust,  expanded  posteriorly  in  both  width  and  height. 
Hinge  straight,  antero-dorsal  and  postero-dorsal  slopes  steeply  angled.  Anterior 
end  rounded,  posterior  end  triangular  in  elongate  male  dimorph.  Female  with 
triangular  posterior  end,  slightly  upturned.  Line  of  greatest  length  lower  in  female 
dimorph.  Postero-ventral  part  of  carapace  characteristically  swollen,  particularly 
in  the  female.  Hine  antimerodont.  Inner  margin  and  line  of  concrescence  coincide; 
duplicature  broad.  Anterior  radial  pore  canals  long,  straight,  widely  and  uniformly 
spaced,  nine  in  the  type  species.  In  front  of  a  vertical  row  of  four  small,  oval 
adductor  scars,  there  is  a  bill-hook  or  broadly  heart-shaped  antero-dorsal  antennal 
scar  and  a  small  antero-ventral  mandibular  scar.  Left  valve  larger  than  the  right. 
Shell  surface  coarsely  pitted. 

TYPE  SPECIES.     Cytheridea  dolabra  Jones  &  Sherborn  1888. 

REMARKS.  This  new  genus  has  been  placed  in  the  Cytherideidae  on  the  basis  of 
carapace  morphology  and  muscle  scar  pattern.  It  is  easily  recognizable  by  its 
rectangular/quadrate  outline,  postero-ventral  convexity  and  position  of  greatest 
height  and  width  towards  the  posterior.  The  outline  of  the  male  dimorph  is  to  some 
extent  reminiscent  of  Fabanella  Martin  1961,  but  that  of  the  female  is  completely 
different. 


ENGLISH    BATHONIAN    OSTRACODA 

Hadrocytheridea  dolabra  (Jones  &  Sherborn) 
(PI.  9,  figs.  1-8;  PI.  10,  fig.  i;  Text-fig.  7) 


407 


1888  Cytheridea  dolabra  Jones  &  Sherborn  :  267,  pi.  3,  figs. 

1888  Cytheridea  puteolata  Jones  &  Sherborn.  :  259,  pi.  i,  figs.  ja-c. 

1888  Cytheridea  parallela  Jones  &  Sherborn  :  260,  pi.  i,  figs.  ga-c. 

1888  Cytheridea  pentagonalis  Jones  &  Sherborn  :  261,  pi.  2,  figs.  la-c. 

1888  PCytheridea  ignobilis  Jones  &  Sherborn  :  268,  pi.  3,  figs.  ga-c. 

DIAGNOSIS.  Hadrocytheridea  with  robust  dimorphic  carapace.  Shell  surface 
finely  punctate  between  distinct  circular  pits,  at  the  centre  of  which  is  a  large, 
circular  normal  pore  canal  opening.  Other  details  as  for  genus. 

LECTOTYPE.  1.1851,  female  right  valve,  figured  Jones  &  Sherborn  1888,  pi.  3, 
figs.  3«-c.  Blue  Fuller's  Earth  Clay,  Midford. 

PARALECTOTYPES.  1.1844,  male  left  valve,  figured  Jones  &  Sherborn  1888,  pi.  i, 
figs.  ja-c.  This  specimen  was  described  by  Jones  &  Sherborn  as  Cytheridea  puteolata 
and  is  selected  here  as  lectotype  of  that  species.  Additional  paralectotypes  of 
C.  puteolata  are  10.3921-2  from  the  Winwood  Collection. 

1.1859,  ma-le  carapace,  figured  Jones  &  Sherborn  1888,  pi.  i,  figs.  ga-c.  This  was 
described  as  Cytheridea  parallela  and  is  here  selected  as  lectotype  of  that  species. 
Additional  paralectotypes  of  C.  parallela  are  10.3924-8  from  the  Winwood  Collection. 

1.  1866,  female  left  valve,  figured  Jones  &  Sherborn  1888,  pi.  2,  figs.  la-c.  This  is 
the  holotype  of  Cytheridea  pentagonalis. 

All  the  above  paralectotypes  of  Hadrocytheridea  dolabra  apart  from  1.  1866,  which 
comes  from  the  Yellow  Fuller's  Earth  Clay,  come  from  the  Blue  Fuller's  Earth  Clay 
of  Midford,  near  Bath. 

OTHER  MATERIAL.  Jones  &  Sherborn  (1862  :  262)  stated  that  Cytheridea  penta- 
gonalis was  represented  by  a  single  specimen  only.  Two  single  right  valves  of  this 
species,  10.273  and  10.3929,  also  occur  in  the  Winwood  Collection  from  the  Blue 
Fuller's  Earth  Clay  but  cannot  be  regarded  as  paralectotypes. 

DESCRIPTION.  Carapace  quadrate  with  sharply  angled,  prominent  cardinal 
angles  and  steeply  sloping  antero-  and  postero-dorsal  slopes  in  the  female  dimorph, 
elongate,  more  rectangular  in  outline  in  the  male.  Dorsal  margin  straight,  ventral 
margin  medially  incurved.  Anterior  end  broadly  rounded  in  the  left  valve,  truncated 
dorsally  in  the  right  by  the  steeply  sloping  antero-dorsal  slope.  In  the  queried 
juvenile  right  valve  this  truncation  of  the  anterior  margin  is  not  present.  Posterior 
end  triangular,  the  postero-dorsal  slope  strongly  concave  in  the  female,  resulting  in 
a  distinct  upturning  of  the  posterior  end  in  both  valves  but  more  strongly  so  in  the 
right.  In  the  male  this  upturning  is  restricted  to  the  right  valve.  Greatest  length 
of  carapace  below  mid-point  in  both  dimorphs  but  more  ventrally  situated  in  the 
female.  Greatest  height  and  width  developed  in  the  posterior  third  of  the  carapace. 
In  the  male  the  increase  in  width  posteriorly  is  uniformly  developed  in  that  area.  In 
the  female  the  increase  in  width  tends  to  be  restricted  to  the  postero-ventral  region. 
Left  valve  larger  than  the  right,  with  the  left  overlapping  the  right  along  the  ventral 


408  REVISION    OF    SOME 


^^     tvv.'Vv'i-'.v. 

.-.••.•:'::'X  ttS.'::  :  •-•.••.•.•.•...'•;'< 

m  •  MW 


FIG.  7.     Hadrocytheridea  dolabra  (Jones  &  Sherborn).     Muscle  scars  of  lectotype  of 
Cytheridea  puteolata  Jones  &  Sherborn,  from  the  exterior.     1.1844.      X  500. 

margin.  Elsewhere  there  does  not  appear  to  be  any  overlap.  Shell  surface  very 
finely  punctate  in  between  the  large  circular  pits,  each  of  which  has  a  large  normal 
pore  canal  opening  at  its  centre.  Hinge  antimerodont,  strongly  developed  with  five 
anterior  and  five  posterior  teeth.  Median  groove  loculate.  Inner  margin  and  line 
of  concrescence  coincide  to  produce  a  broad  duplicature  through  which  pass  (anteri- 
orly) nine,  long,  straight  and  widely-spaced  radial  pore  canals.  Muscle  scars 
consist  of  a  vertical  row  of  four  small,  oval  adductor  scars,  a  small,  round  antero- 
ventral  mandibular  scar  and  a  large  antero-dorsal  antennal  scar  which  is  equal  in 
size  to  two  adductor  scars  and  is  either  heart-shaped  or  bill-hooked  in  outline. 

DIMENSIONS.  1.1851,  female  right  valve,  length  0-64  mm. ;  height  0-38  mm. 
1.1844,  male  left  valve,  length  0-71  mm.;  height  0-34 mm.  1.1859,  male  carapace, 
length  0-73  mm. ;  height  0-36  mm. ;  width  0-37  mm.  1. 1866,  female  left  valve, 
length  0-57  mm. ;  height  0-32  mm.  10.3294,  male  right  valve,  length  0-65  mm. ; 
height  0-31  mm.  1. 1868,  Pjuvenile  right  valve,  length  0-46  mm. ;  height  0-26  mm. 

REMARKS.  Jones  &  Sherborn  (1888)  described  four  different  species  which  were 
either  male  or  female  dimorphs  of  a  single  species.  Although  Cytheridea  puteolata 
had  page  preference  it  was  decided  to  select  a  female  dimorph  as  type  because  it  is 
the  female  outline  which  is  continuous  with  juvenile  instars.  Of  the  two  female 
specimens  available,  that  identified  as  Cytheridea  dolabra  was  the  better  preserved 
and  was  therefore  chosen  to  identify  the  species.  The  breakdown  of  the  synonymous 
species  is  as  follows: 

Female  right  valve  .  .  Cytheridea  dolabra 

Female  left  valve    .  .  .  Cytheridea  pentagonalis 

Male  carapace         .  .  .  Cytheridea  parattela 

Male  left  valve        .  .  .  Cytheridea  puteolata 

Pjuvenile  instar      .  .  .  Cytheridea  ignobilis 


ENGLISH    BATHONIAN    OSTRACODA  409 

Hadrocytheridea  dolabra  is  close  to  Cytheridea  punctiputeolata  Jones  &  Sherborn, 
as  far  as  the  male  right  valve  is  concerned,  but  C.  punctiputeolata  has  a  lophodont 
hinge,  more  rounded  posterior  end  and  a  more  strongly  developed  ornamentation  of 
punctae  and  pits.  Cytheridea  ignobilis  has  been  included  in  this  species  as  it  probably 
represents  a  juvenile  instar.  However,  because  of  the  slight  difference  between  the 
anterior  margin  of  C.  ignobilis  and  that  of  H .  dolabra  and  the  absence  of  other  juveniles 
to  establish  an  ontogentetic  sequence,  C.  ignobilis  is  doubtfully  referred  to  this  species 
and  is  not  placed  in  the  list  of  paralectotypes. 

Family  PROGONOCYTHERIDAE  Sylvester-Bradley  1948 
Subfamily  PROGONOCYTHERINAE  Sylvester-Bradley  1948 

Genus  RECTOCYTHERE  Malz  1958 

Rectocythere  sugillata  (Jones  &  Sherborn) 

(PL  10,  figs.  3-6) 

1888     Cytheridea  sugillata  Jones  &  Sherborn  :  262,  pi.  2,  figs.  -za-c. 

DIAGNOSIS.  Rectocythere  with  surface  ornamentation  of  irregular  ridges  and  raised 
areas  giving  parched  or  shrivelled-up  appearance.  Coarsely  pitted  in  well-preserved 
specimens. 

LECTOTYPE.  1.1855,  lert  valve  figured  Jones  &  Sherborn  1888.  Blue  Fuller's 
Earth  Clay,  Midford. 

PARALECTOTYPE.  10.3930,  right  valve  referred  to  by  Jones  &  Sherborn  1888  :  262, 
Blue  Fuller's  Earth  Clay,  Win  wood  Collection. 

DESCRIPTION.  Shell  robust,  with  high  anterior  and  posteriorly  sloping  dorsal 
margin.  Anterior  end  broadly  rounded;  posterior  end  narrowly  rounded  in  the  left 
valve,  triangular  in  the  right,  with  a  concave  postero-dorsal  slope  producing  an 
upturned  posterior  end.  Ventral  margin  slopes  upwards  towards  the  posterior  end, 
but  is  over-hung  postero-ventrally  by  the  prominent  convexity  of  the  ventro-lateral 
margin  in  that  region.  Anterior  and  posterior  marginal  borders  flattened,  the  convex 
central  part  of  the  carapace  is  strongly  ornamented  by  irregular  ridges  and  raised 
areas  which  give  a  dehydrated  appearance  to  the  ostracod.  Hinge  lophodont. 
Inner  margin  and  line  of  concrescence  coincide  producing  a  rather  broad  duplicature. 
To  the  outside  of  this  there  is  developed  a  narrow  flange  extending  around  the 
anterior  margin  and  along  the  ventral  margin  to  the  apex  of  the  posterior  end. 
Anterior  radial  pore  canals  long  and  straight,  widely  spaced  and  about  seven  to 
eight  in  number.  Muscle  scars  not  observed. 

DIMENSIONS.  1.1855,  left  valve,  length  0-50  mm.;  height  0-31  mm.  10.3930, 
right  valve,  length  0-51  mm. ;  height  0-32  mm. 

REMARKS.  Rectocythere  sugillata  possesses  the  same  basic  ornamentation  as 
R.  rugosa  Malz  (19660  :  405,  figs.  6-9)  but  may  be  distinguished  by  having  a  much 

GEOL.    17,  8  25 


4io  REVISION    OF    SOME 

finer  degree  of  surface  pitting  and  additional  irregular  swellings.     Dr.  Malz  kindly 
sent  material  of  R.  rugosa  for  comparison. 

The  ostracod  Camptocythere  lincolnensis  Bate  (1963  :  201,  pi.  10,  figs.  2-13)  also 
belongs  to  Rectocythere  on  its  similarity  of  carapace  outline,  ornamentation  and 
internal  details. 

Genus  CAYTONIDEA  Bate  1965 

DIAGNOSIS.  Progonocy thermae,  oval-rectangular  in  outline  with  well-rounded 
anterior  and  posterior  margins.  Low  eye  swelling  situated  at  anterior  cardinal 
angle.  Cardinal  angles  prominent,  broadly  rounded.  Shell  surface  with  reticulate 
ornamentation.  Hinge  antimerodont.  Muscle  scars  consist  of  sub  vertical  row  of 
four  adductor  scars,  rounded  antero-dorsal  antennal  scar  and  rounded  antero-ventral 
mandibular  scar.  Radial  pore  canals  long,  straight,  few  in  number.  Left  valve 
larger  than  right.  Dimorphic. 

REMARKS.  Since  the  publication  of  this  genus  a  second  species  has  been  identified 
which  enables  some  modification  to  the  diagnosis  to  be  made.  This  concerns  the 
identification  of  dimorphism  and  the  determination  of  a  reticulate  ornamentation  as 
a  constant  feature. 

A  re-examination  of  the  type  species,  Caytonidea  faveolata  Bate  (1965  :  100,  pi.  I, 
figs.  13-14,  pi.  2,  figs,  i-io),  has  shown  that  the  holotype  (10.1831)  is  a  female  dimorph 
and  the  paratype  (10.1834)  a  male. 


Caytonidea  terraefullonicae  (Jones  &  Sherborn) 

(PI.  10,  figs.  2,  7,  8;P1.  u,  figs.  1-3) 
1888     Cytheridea  terrae-fullonicae  Jones  &  Sherborn  :  258,  pi.  i,  figs.  5«-c. 

DIAGNOSIS.  Caytonidea  with  coarse  reticulate  ornamentation  of  five-  to  six-sided 
pits  covering  entire  shell  surface.  Extreme  postero-ventral  margin  swollen,  slightly 
overhanging  ventral  surface.  Males  more  elongate  in  outline  than  females. 

LECTOTYPE.  1.1869,  left  valve,  figured  Jones  &  Sherborn  1888.  Blue  Fuller's 
Earth  Clay,  Midford. 

PARALECTOTYPES.  10.3917-20,  male  and  female  specimens  from  the  Winwood 
Collection,  Blue  and  Yellow  Fuller's  Earth  Clay,  Midford. 

OTHER  MATERIAL.  10.4002-3,  right  valve  and  juvenile  carapace  from  the  J.  F. 
Blake  Collection,  Fuller's  Earth  Clay,  Bath. 

DESCRIPTION.  Carapace  rectangular  in  outline,  especially  in  the  male  dimorph, 
with  sub-parallel  dorsal  and  ventral  margins.  Anterior  end  high  with  greatest 
height  through  the  anterior  cardinal  angle  which  is  situated  close  to  the  anterior 
margin.  Anterior  end  broadly  but  slightly  obliquely  rounded  and  extended  ventrally 
below  the  ventral  margin.  Posterior  end  broadly  rounded  in  the  left  valve,  with 


ENGLISH    BATHONIAN    OSTRACODA  411 

oblique  postero-dorsal  slope  in  the  right  valve.  Postero- ventral  margin  slightly 
swollen  and  overhanging  ventral  surface.  Greatest  length  of  carapace  passes 
slightly  below  the  mid-point;  greatest  width  in  the  posterior  third.  There  are  no 
marginal  borders.  Left  valve  slightly  larger  than  the  right  which  it  overlaps  along 
the  ventral  margin  but  very  little  elsewhere.  Shell  surface  coarsely  reticulate,  the 
reticulations  producing  five-  to  six-sided  pits  which  cover  the  entire  shell  surface. 
In  the  juvenile  carapace,  the  reticulations  tend  to  form  longitudinal  ridges  along  the 
ventral  surface.  Normal  pore  canals  large  and  widely  spaced  over  the  surface  of 
the  carapace.  Inner  margin  and  line  of  concrescence  coincide  to  produce  a  broad 
duplicature  through  which  pass  a  small  number  (about  seven)  of  long,  straight,  and 
widely  spaced  anterior  radial  pore  canals.  A  narrow  flange  extends  around  the 
anterior  and  ventral  margins  and  along  the  postero-ventral  part  of  the  posterior 
margin.  Hinge  antimerodont,  the  median  element  being  long  and  finely  denticulate/ 
locellate.  Terminal  elements  dentate/loculate.  Muscle  scars  as  seen  from  the 
exterior  consist  of  a  small,  rounded  antero-dorsal  antennal  scar  and  a  vertical  row  of 
four  rounded  adductor  scars  (type  A).  An  oval  smooth  area  in  the  region  of  the 
anterior  cardinal  angle  is  indicative  of  an  eye  swelling.  This  feature  is  only  clearly 
seen  in  the  male  dimorph. 

DIMENSIONS.  1.1869,  female  left  valve,  length  0-54  mm. ;  height  0-30  mm. 
10.3919,  female  right  valve,  length  0-49  mm. ;  height  0-26  mm.  10.3917,  male  right 
valve,  length  0-54  mm. ;  height  0-26  mm.  10.3918,  male  left  valve,  length  0-55  mm. ; 
height  0-26  mm.  10.3920,  female  carapace,  length  0-52  mm. ;  height  0-29  mm. ; 
width  0-25  mm. 

REMARKS.  Caytonidea  terraefullonicae  differs  from  C.  faveolata  Bate  in  the  posses- 
sion of  a  much  stronger  reticulate  ornamentation,  straighter  dorsal  margin,  and 
postero-ventral  swelling  slightly  overlapping  the  ventral  surface.  Unfortunately, 
during  the  examination  of  this  material  10.3918  was  lost  and  10.3917  damaged. 


Genus  AC  AN  T  HOC  Y  THERE  Sylvester-Bradley  1956 
Acanthocythere  sphaerulata  (Jones  &  Sherborn) 
(PI.  n,  figs.  4-6) 

1888     Cythere  sphaerulata  Jones  &  Sherborn  :  253,  pi.  i,  figs.  6a-c. 

1956     Acanthocythere  sphaerulata  (Jones  &  Sherborn)  Sylvester-Bradley  :  12,  pi.  i,  figs.  1-4. 

DIAGNOSIS.     As  for  Sylvester-Bradley  1956  :  12. 

HOLOTYPE.  1.1835,  female  carapace,  length  0-52  mm. ;  height  0-31  mm. ;  width 
0-31  mm.  Figured  Jones  &  Sherborn  1888.  Blue  Fuller's  Earth  Clay,  Midford. 

REMARKS.  Cythere  sphaerulata  was  made  the  type  species  of  Acanthocythere 
(Sylvester-Bradley  1956),  additional  material  indicating  that  the  species  was 
dimorphic.  A  misprint  on  p.  12  stated  that  the  holotype  was  a  male  dimorph ;  it  is, 
in  fact,  a  female  dimorph. 


4i2  REVISION    OF    SOME 

Genus  FASTIGATOCYTHERE  Wienholz  1967 
Fastigatocythere  juglandica  (Jones) 

(PI-  12,  fig.  3) 
1888     Cythere  juglandica  var.  major  Jones  &  Sherborn  :  255,  pi.  4,  figs.  2a,  b. 

REMARKS.  The  synonymy  and  diagnosis  of  this  species  are  given  on  p.  389.  The 
variety  described  by  Jones  &  Sherborn,  of  which  the  specimen  1.1872  is  holotype 
was  correctly  placed  in  synonymy  by  Sylvester-Bradley  (1948). 

DIMENSIONS.     1.1872,  female  left  valve,  length  0-74  mm. ;  height  0-49  mm. 

Genus  GLYPTOCYTHERE  Brand  &  Malz  1962 

Glyptocythere  guembeliana  (Jones) 

(PI.  3,  fig.  2;  PI.  4,  fig.  i) 

1884  Cythere  guembeliana  Jones  :  772,  pi.  34,  figs,  32,  33  [not  fig.  31]. 

1888  Cytheridea  pulvinar  Jones  &  Sherborn  :  266,  pi.  3,  figs.  za-c. 

1888  Cytheridea  trapezoidalis  Terquem,  Jones  &  Sherborn  :  270,  pi.  4,  figs.  la,  b. 

1967  Glyptocythere  guembeliana  (Jones)  Bate  :  49,  pi.  13,  figs.  10-16,  pi.  14,  figs.  1-8. 

REMARKS.  This  species  has  been  described  fully  by  me  (Bate  1967)  although 
I  omitted  to  include  the  ostracod  identified  by  Jones  &  Sherborn  as  Cytheridea 
trapezoidalis  Terquem.  This  specimen  is  a  juvenile  instar  of  G.  guembeliana. 
Although  it  has  a  well-developed  ventro-lateral  keel,  the  outline  of  the  valve  is 
typically  juvenile  in  being  strongly  acuminate  posteriorly.  The  hinge,  in  being  anti- 
merodont,  reflects  an  early  stage  in  the  development  of  the  adult  entomodont  hinge. 

DIMENSIONS.  1.1858,  female  right  valve  (Cytheridea  pulvinar},  length  0-82  mm. ; 
height  0-47  mm.  1.1840,  juvenile  right  valve  (Cytheridea  trapezoidalis},  length 
0-54  mm. ;  height  0-30  mm. 

Glyptocythere  oscillum  (Jones  &  Sherborn) 
(PI.  5,  ng.  2;  PI.  12,  fig.  2;  Text-figs.  8,  9) 

1888     Cythere  oscillum  Jones  &  Sherborn  :  254,  pi.  3,  figs.  8a-c. 
1888     Cytheridea  striblita  Jones  &  Sherborn  :  268,  pi.  3,  figs.  ja-c. 

DIAGNOSIS.  Carapace  strongly  ornamented  with  series  of  grooves  and  irregular 
swollen  areas:  two  lateral  grooves  developed,  the  first  ventro-laterally,  the  second 
just  above  valve  middle,  both  joined  by  narrow  vertical  groove  passing  between 
two  swollen  areas.  Broad  dorso-median  sulcus  extends  down  to  median  groove. 
Area  below  ventio-lateral  groove  alate  in  right  valve,  swollen  in  both  valves.  From 
posterior  region,  two  swollen  areas  extend  towards  valve  centre  whilst  two  similar 
areas  extend  back  from  anterior  region.  Lower  of  anterior  swollen  area  terminates 
in  prominent  swelling.  Anterior  and  posterior  swollen  areas  do  not  unite.  Normal 
pore  canals  prominently  displayed  over  carapace. 


ENGLISH    BATHONIAN    OSTRACODA  433 

HOLOTYPE.  1.1849,  described,  figured  Jones  &  Sherborn  (1888).  Female  right 
valve  from  the  Blue  Fuller's  Earth  Clay,  Midford. 

OTHER  MATERIAL.  10.3943,  male  left  valve,  referred  to  Jones  &  Sherborn  (1888). 
Winwood  Collection.  Blue  Fuller's  Earth  Clay,  Midford.  This  is  the  lectotype  of 
Cytheridea  striblita  Jones  &  Sherborn.  10.3914-6,  two  male  left  valves  and  a  male 
right  valve,  Winwood  Collection,  Blue  Fuller's  Earth  Clay,  Midford. 

DESCRIPTION.  Carapace  subquadrate  to  sub-rectangular  in  outline,  the  more 
elongate  specimens  being  the  males.  Greatest  length  through  the  mid-point  in  the 
male  dimorph,  below  in  the  female  right  valve.  Greatest  height  in  the  posterior 
third.  Dorsal  margin,  in  the  left  valve,  concave  in  the  region  of  the  dorso-median 
sulcus,  convex  posteriorly.  The  dorsal  margin  of  the  right  valve  is  strongly  convex, 
especially  in  the  female.  Cardinal  angles  particularly  well  developed  in  the  right 
valve,  postero-dorsal  slope  concave  in  both  valves,  more  especially  so  in  the  female 
right  valve.  Antero-dorsal  slope  broadly  convex  in  the  male,  concave  in  the  female 
right  valve.  Anterior  end  broadly  rounded,  posterior  end  triangular.  Ventral 
margin  antero-medially  concave,  posteriorly  convex.  Ventro-lateral  margin  of 
right  valve  becomes  alate,  extending  below  ventral  surface.  This  feature  does  not 
appear  to  be  so  well  developed  in  the  left  valve.  Anterior  and  posterior  marginal 
borders  flattened.  Shell  surface  ornamented  as  described  in  the  diagnosis.  Normal 
pore  canal  openings  prominent  and  widely  scattered  over  the  carapace.  Hinge 
with  five  to  six  terminal  teeth  in  the  right  valve  and  a  median  loculate  groove.  In 
the  left  valve  the  terminal  loculate  sockets  are  separated  by  a  median  bar  which  is 
rather  coarsely  dentate,  particularly  in  the  anterior  part.  Although  not  as  well 
developed  as  in  some  species  the  hinge  is  regarded  as  entomodont.  Inner  margin 
and  line  of  concrescence  coincide.  Anterior  radial  pore  canals  long  and  straight, 
widely  spaced,  about  ten  in  number.  Muscle  scars  not  seen. 

DIMENSIONS.  Lectotype,  1.1849,  female  right  valve,  length  0-53  mm. ;  height 
0-30  mm.  10.3943,  male  left  valve,  length  0-62  mm. ;  height  0-32  mm.  10.3916, 
male  right  valve,  length  0-59  mm. ;  height  0-30  mm. 

REMARKS.  Jones  &  Sherborn  identified  the  female  specimens  of  the  present  species 
as  Cy there  oscillum;  the  males  they  placed  in  Cytheridea  striblita.  The  figured  speci- 
men of  the  latter  species  has  been  lost  but  the  lectotype  of  C.  striblita  was  found  in 
the  Winwood  Collection. 

Glyptocythere  rudimenta  Brand  &  Malz  (1962  and  1966)  is  similar  to  G.  oscillum  in 
ornamentation  but  differs  in  that  the  anterior  and  posterior  swollen  areas  take  the 
form  of  rather  thin  irregular  ridges,  not  nearly  so  broad  as  in  oscillum.  A  broad 
sulcus  is  developed  in  oscillum  in  the  dorso-median  area,  but  in  rudimenta  there  is  a 
series  of  irregular  ridges  radiating  down  from  the  dorsal  margin.  The  shell  outline 
in  the  right  valve  is  noticeably  different,  the  cardinal  angles  and  the  dorso-median 
extension  of  the  dorsal  margin  being  very  much  more  prominently  developed  in 
G.  oscillum.  The  similarities  in  ornamentation  between  the  two  species  suggests 
a  phylogenetic  relationship  which  is  supported  by  their  stratigraphic  position, 
G.  rudimenta  being  found  in  the  Upper  Bajocian  and  G.  oscillum  in  the  Upper 


REVISION    OF    SOME 


FIG. 


Glyptocythere  oscillum  (Jones  &  Sherborn).     Male  left  valve,  paralectotype, 
10.3943-      x  153. 


FIG.  9.     Glyptocythere  oscillum  (Jones  &  Sherborn).     Male  right  valve,  10.3916.      x  165. 


FIG.   10.     Glyptocythere  rudimenta  Brand  &  Malz.     Left  valve,  female  carapace.     10.3921. 

X  113. 


ENGLISH    BATHONIAN    OSTRACODA  415 

Bathonian.     The  kindness  of  Dr.  D.  E.  Brand  in  sending  me  specimens  of  G.  rudi- 
menta  is  acknowledged  here. 

Glyptocythere  persica  (Jones  &  Sherborn) 

(PL  12,  fig.  i;  Text-fig,  u) 
1888     Cytheridea  persica  Jones  &  Sherborn  :  270,  pi.  4,  fig.  4. 

DIAGNOSIS.  Carapace  with  broad  dorso-median  depression  in  anterior  half;  short, 
blade-like  ventral  keel  on  ventral  surface  and  ventro-lateral  margin  develops  into 
keel-like  ridge.  Lateral  surface  ornamented  by  neat  reticulation.  Ventral  surface 
with  low,  longitudinal  ridges.  Anterior  and  posterior  marginal  borders  smooth, 
compressed. 

HOLOTYPE.  1.1834,  right  valve,  figured  Jones  &  Sherborn  (1888),  Blue  Fuller's 
Earth  Clay,  Midford. 

OTHER  MATERIAL.  10.4006,  juvenile  carapace,  Mockler  Collection,  Fuller's  Earth, 
Midford. 

DESCRIPTION.  Carapace  with  greatest  height  at  the  anterior  end,  through  the 
anterior  cardinal  angle,  tapering  to  the  posterior  end.  Dorsal  margin  broadly 
convex,  sloping  to  the  posterior,  ventral  margin  with  a  broad,  median  incurvature, 
below  which  in  the  right  valve  there  is  lip-like  extension  of  the  flange.  The  flange 
is  continuous  around  the  anterior  margin  and  along  the  ventral  margin  to  the  tip  of 
the  triangular  posterior  end.  Anterior  end  broadly  rounded,  posterior  end  with 


FIG.   u.     Glyptocythere  persica  (Jones  &  Sherborn).     Internal  view  of  right  valve. 

Holotype,  1.1834.      x  I2°- 

concave  postero-dorsal  slope  and  convex  postero- ventral  slope.  Anterior  and 
posterior  marginal  borders  compressed  and  smooth,  distinct  from  the  more  strongly 
convex  part  of  the  carapace  which  is  ornamented  by  a  neat  reticulation.  The 
ornamentation  is  extended  along  the  ventro-lateral  margin  to  form  a  keel-like  ridge. 
Ventral  surface  ornamented  by  four  longitudinal  ridges,  one  of  which  is  developed 


416  REVISION    OF    SOME 

into  a  short,  blade-like  keel.  A  smooth  area  at  the  anterior  cardinal  angle  is  sugges- 
tive of  an  ocular  lens.  Just  anterior  of  valve  centre  and  in  the  dorso-median  part 
of  the  carapace  a  broad  shallow  sulcus  is  developed,  especially  noticeable  in  dorsal 
view.  The  posterior  part  of  the  valve  behind  the  sulcus  is  noticeably  more  convex, 
whilst  at  the  base  of  the  sulcus,  a  raised  area  locates  the  position  of  the  muscle  scars. 
The  left  valve  is  larger  than  the  right,  which  it  overlaps  along  the  ventral  margin  and 
slightly  overreaches  along  the  dorsal  margin.  The  inner  margin  and  line  of  con- 
crescence coincide.  The  hinge  is  weakly  entomodont. 

DIMENSIONS.  1.1834,  right  valve,  length  0-70  mm. ;  height  0-38  mm.  10.4006, 
juvenile  carapace,  length  0-52  mm. ;  height  0-28  mm. ;  width  0-24  mm. 

REMARKS.  Glyptocythere  persica,  with  its  neat,  reticulate  ornamentation,  distinct 
ventro-lateral  ridge,  and  well-developed  ventral,  blade-like  keel,  is  quite  distinct 
from  those  species  which  possess  well-developed  ventro-lateral  ridges  or  keels,  e.g. 
Glyptocythere  costata  Bate  (1965)  and  G.  guembeliana  (Jones)  (Bate  1967),  or  from 
those  which  have  a  much  stronger  reticulate  ornamentation,  e.g.  Glyptocythere 
tuberodentina  Brand  &  Malz  (19620). 


Genus  LOPHOCYTHERE  Sylvester-Bradley  1948 

BRADIANA  GROUP 

Lophocythere  acutiplicata  (Jones  &  Sherborn) 
(PI.  i,  fig.  8;  PI.  12,  figs.  4,  6) 

1888     Cytheridea  acutiplicata  Jones  &  Sherborn  :  271,  pi.  4,  figs,  'ja-b,  Sa-c. 

DIAGNOSIS.  Carapace  dimorphic,  tapering  posteriorly.  Eye  swelling  below 
anterior  cardinal  angle.  Shell  surface  weakly  ornamented  laterally,  the  most 
prominent  ridge  being  situated  ventro-medially  on  the  valve  and  extending  from 
posterior  margin  to  anterior  margin.  A  second  lateral  ridge  just  above  also  extends 
down  to  the  anterior  margin,  but  cannot  be  traced  backwards  beyond  the  anterior 
third  where  it  dies  out.  Two  short  ridges  branch  off  the  ventro-median  ridge  a  short 
distance  back  from  the  anterior  margin.  The  uppermost  ridge  is  very  short  and 
extends  into  valve  centre  at  an  acute  angle  whilst  the  lower  extends  below  the  ventro- 
median  ridge,  parallel  to  it  for  almost  three-quarters  of  its  length.  Three  short 
lateral  ridges  are  situated  in  the  dorso-median  part  of  the  posterior  third.  A  short 
ridge  extends  down  from  the  eye  swelling  to  die  out  above  mid-length.  Ventro- 
median  part  of  carapace  convex,  extended  below  ventral  surface.  On  this  extension 
the  left  valve  has  a  single  ridge  extending  almost  from  the  anterior  margin  to  the 
posterior  margin,  with  a  short  lateral  ridge  below  it  in  the  anterior  part  of  the  valve. 
In  the  right  valve  there  are  two  lateral  ridges  in  this  region,  both  convex  ventrally. 
The  upper  extends  the  length  of  the  posterior  half  of  the  valve  whilst  the  ridge 
slightly  offset  below  it  extends  the  length  of  the  anterior  half  with  some  overlap  of 
the  two  at  valve  middle.  Normal  pore  canals  prominent.  Hinge  antimerodont. 
Left  valve  larger  than  right. 


ENGLISH    BATHONIAN    OSTRACODA  417 

LECTOTYPE.     1.1863,  female  carapace,  figured  Jones  &  Sherborn   (1888),   Blue 
Fuller's  Earth  Clay,  Midford. 


PARALECTOTYPES.  1.1847,  female  right  valve,  figured  Jones  &  Sherborn 
Blue  Fuller's  Earth  Clay,  Midford.  10.3642-45,  two  male  and  one  female  right  valve 
from  the  Blue  Fuller's  Earth  Clay  and  one  female  right  valve  from  the  Yellow  Fuller's 
Earth  Clay,  Midford,  Winwood  Collection. 

DESCRIPTION.  Carapace  with  a  strong  posterior  taper  in  both  dimorphs,  of  which 
the  male  is  the  more  elongate.  Anterior  end  high,  broadly  rounded,  the  greatest 
height  being  through  the  anterior  cardinal  angle,  just  below  which  is  a  low  eye 
swelling.  Posterior  end  triangular,  narrow,  with  concave  postero-dorsal  slope  and 
convex  postero- ventral  slope.  Posterior  cardinal  angle  prominent.  Greatest  length 
passes  through  the  mid-point,  greatest  width  in  the  posterior  third  although  the 
width  in  the  anterior  third  is  almost  the  same.  Carapace  constricted  slightly  about 
centre  when  viewed  dorsally.  Normal  pore  canals  large  and  prominently  displayed 
over  the  carapace.  Shell  surface  ornamented  as  in  the  diagnosis.  Left  valve 
larger  than  the  right,  which  it  overlaps  along  the  ventral  margin  and  overreaches 
along  the  dorsal  margin.  Hinge  antimerodont,  only  seen  in  the  right  valve 
where  there  are  four  to  five  terminal  teeth  and  a  rather  long,  delicate,  loculate  median 
groove.  The  hinge  is  particularly  delicate,  and  the  carapace  as  a  whole  appears  to 
be  thin-shelled.  Inner  margin  and  line  of  concrescence  coincide.  Radial  pore 
canals  and  muscle  scars  not  observed.  A  narrow  flange  extends  around  the 
anterior  and  ventral  margins  and  along  the  ventral  part  of  the  posterior. 

DIMENSIONS.  1.1863,  female  carapace,  length  0-57  mm.;  height  0.31  mm.;  width 
0-25  mm.  1.1847,  female  right  valve,  length  0-54 mm.  (broken);  height  0-32  mm. 
10.3643,  male  right  valve,  length  0-65  mm. ;  height  0-33  mm.  10.3644,  female  right 
valve,  length  0-58  mm. ;  height  0-33  mm.  10.3645,  female  right  valve,  length 
0-56  mm. ;  height  0-31  mm. 

REMARKS.  The  carapace  morphology  and  surface  ornamentation  agree  closely 
with  the  basic  characters  of  those  ostracods  which  may  be  placed  in  the  bradiana 
grouping  of  Lophocy 'there.  Apart  from  the  details  of  the  radial  pore  canals  and 
muscle  scars,  which  are  not  so  far  known  for  L.  acutiplicata,  the  only  character  in 
disagreement  with  this  classification  is  the  hinge,  which,  being  antimerodont,  is  at 
once  at  variance  with  all  the  other  species  of  Lophocythere.  An  antimerodont  hinge 
is  normally  accepted  as  present  in  juvenile  instars  of  adults  having  an  entomodont 
hinge;  here  the  delicate  shell,  strong  posterior  taper  and  comparatively  small  size 
would  indicate  this  if  it  were  not  for  the  presence  of  strong  dimorphism  suggesting 
adult  status.  Sexual  dimorphism  is  known,  however,  in  pre-adult  instars.  As  only 
right  valves  are  available  for  the  examination  of  internal  features,  it  is  not  intended 
to  suggest  a  subgeneric  rank  for  this  species,  until  additional  material  becomes 
available.  In  ornamentation  L.  acutiplicata  may  be  distinguished  easily  by  the 
number  of  lateral  ridges,  their  tendency  to  fade  out  in  the  posterior  part  of  the 
carapace,  and  by  the  development  of  two  ventro-lateral  keels  in  the  right  valve. 


418  REVISION    OF    SOME 

Lophocythere  bradiana  (Jones) 
(PL  4,  figs.  2,  3;  Text-fig.  4) 

REMARKS.     The  species  has  already  been  described  (p.  390). 

DIMENSIONS.  1.1854  (lectotype  of  C.  craticula  figured  Jones  &  Sherborn,  pi.  4, 
figs.  loa-c),  length  0-62  mm.;  height  0-32  mm.;  width  0-32  mm.  1.1867  (paralecto- 
type  of  C.  craticula  figured  Jones  &  Sherborn,  pi.  4,  figs,  ga-c)  left  valve,  length 
0-60  mm.;  height  0-32  mm. 

Lophocythere  septicostata  Bate 

(PI.  12,  fig.  8) 

1888     Cytheridea  bradiana  Jones  &  Sherborn  :  272,  pi.  4,  figs.  na-c. 

1967     Lophocythere  septicostata  Bate  :  52,  pi.  15,  figs.  7-13,  pi.  16,  figs.  1-4. 

DIAGNOSIS.  Lophocythere  having  seven  longitudinal  ridges  on  lateral  surface  and 
two  on  ventral  surface.  Vertical  ridge  extends  downwards  from  oval  eye  swelling 
Interspaces  between  ridges  punctate,  with  large,  circular,  normal  pore  canal  openings. 
Anterior  and  posterior  marginal  borders  compressed. 

REMARKS.     For  remarks  relating  to  this  species  see  Bate  (1967). 
DIMENSIONS.     1.1843,  female  right  valve,  length  0-64  mm.;  height  0-33  mm. 

Lophocythere  sp. 

(Text-fig.  12) 

REMARKS.  A  single  left  valve  in  the  Winwood  Collection  represents  a  new  species 
of  Lophocythere.  It  comes  from  the  Blue  Fuller's  Earth  Clay  of  Midford  and  is  of 
the  Bradiana  Group.  The  arrangement  of  the  lateral  ridges  is  quite  different  from 
that  in  any  described  species.  The  dorsal  ridge  commences  at  the  anterior  margin, 
bends  sharply  upwards  to  pass  through  the  eye  swelling  and  then  in  an  undulating 


FIG.   12.     Lophocythere  sp.     External  view,  right  valve.     10.3900.      x  170. 


ENGLISH    BATHONIAN    OSTRACODA  419 

course  parallels  the  dorsal  and  postero-dorsal  margins  to  die  out  at  the  extreme 
posterior.  The  median  ridge  commences  at  the  anterior  margin  just  below  the  dorsal 
ridge  and  soon  bifurcates,  the  ventral  branch  extending  back  into  the  posterior 
third.  A  second  ridge  branches  from  the  median  ridge  a  short  distance  above  the 
first  bifurcation  and  extends  into  the  posterior  third  where  it  unites  with  the  first 
bifurcation.  Both  then  extend  to  the  posterior  margin  as  a  single  narrow  ridge. 
The  median  ridge  continues  its  dorsal  course  and  bifurcates  yet  again,  the  dorsal 
branch  continuing  until  it  fuses  with  the  dorsal  ridge  just  behind  the  eye  swelling; 
the  other  branch  passes  laterally  back  into  the  posterior  half  of  the  valve,  splits  into 
a  reticulate  complex  of  smaller  ridges  but  re-forms  to  fuse  with  the  dorsal  ridge  in 
the  posterior  quarter.  Two  ventral  ridges  originate  at  the  anterior  margin,  the 
lower  extending  back  into  the  posterior  half  of  the  valve  whilst  the  upper  is  restricted 
to  the  anterior  half  only.  Interspaces  between  the  ridges  are  reticulate.  Hinge 
strongly  entomodont. 

DIMENSIONS.     10.3900,  left  valve,  length  0-50  mm. ;  height  0-27  mm. 


OSTREATA  GROUP 

Lophocy there  fulgurata  (Jones  &  Sherborn) 
(PL  12,  fig.  7) 

1888     Cytheridea  fulgurata  Jones  &  Sherborn  :  273,  pi.  4,  figs.  i2a-c. 

DIAGNOSIS.  Carapace  sub-rectangular,  tapering  to  the  posterior.  Oval,  clear 
eye  swelling  at  anterior  cardinal  angle.  Anterior  and  posterior  marginal  areas 
flattened,  smooth.  Central  part  of  carapace  strongly  convex,  ornamented  by  two 
prominent  ridges.  The  first,  L-shaped,  commences  at  the  eye  swelling,  extends 
vertically  down  to  the  an tero- ventral  part  of  the  valve,  where  it  performs  a  right- 
angle,  and  runs  laterally  backwards  to  the  edge  of  the  posterior  marginal  area. 
A  second  lateral  ridge  arises  just  below  the  right-angled  bend  of  the  first  ridge  and 
extends  backwards,  paralleling  the  ridge  above  it,  to  die  out  in  a  similar  position  at 
the  edge  of  the  posterior  marginal  border.  The  area  inside  the  "  L  "  of  the  first 
ridge  is  occupied  by  strong  ridges  radiating  down  from  the  dorsal  margin  and 
occasionally  branching. 

LECTOTYPE.  1.1832,  right  valve,  figured  Jones  &  Sherborn  (1888),  Blue  Fuller's 
Earth  Clay,  Midford. 

PARALECTOTYPES.  10.3646-8,  right  and  left  valves  from  the  Blue  Fuller's  Earth 
Clay,  Midford,  Winwood  Collection. 

DESCRIPTION.  Carapace  sub-rectangular  with  dorsal  margin  sloping  to  posterior. 
Cardinal  angles  prominent,  situated  close  to  the  anterior  and  posterior  margins 
respectively.  Anterior  cardinal  angle  smooth,  indicating  the  presence  of  an  eye 
swelling.  Line  of  greatest  height  passes  through  this  point.  Line  of  greatest 
length  passes  through  the  mid-point,  or  slightly  below  it.  Anterior  end  high, 


420  REVISION    OF    SOME 

broadly  rounded;  posterior  end  small  and  triangular  with  a  concave  postero-dorsal 
slope  and  a  convex  postero-ventral  slope.  Ventral  margin  with  a  shallow  antero- 
median  incurvature.  Shell  surface  ornamented  as  in  the  diagnosis;  area  between 
the  ridges  appears  to  be  punctate.  Hinge  not  well  preserved,  the  median  element  in 
both  valves  having  been  damaged.  Terminal  elements  consist  of  dentate  ridges  in 
the  right  valve  and  loculate  sockets  in  the  left.  Median  bar  of  the  left  valve,  long, 
not  clearly  seen  to  be  dentate,  bounded  above  by  a  shallow  accommodation  groove. 
Median  groove  of  the  right  valve,  long  and  narrow,  not  seen  to  be  loculate  because  of 
damage.  Inner  margin  and  line  of  concrescence  coincide.  Radial  pore  canals  and 
muscle  scars  not  seen.  A  narrow  flange  extends  around  the  anterior  margin  and 
along  the  ventral  margin  to  the  tip  of  the  posterior  margin  in  the  right  valve. 

DIMENSIONS.  1.1832,  right  valve,  length  0-77  mm. ;  height  0-42  mm.  10.3648, 
left  valve,  length  0-66  mm. ;  height  0-36  mm. 

REMARKS.  Although  the  median  element  of  the  hinge  has  been  damaged,  the 
appearance  suggests  that  it  was  antimerodont  instead  of  the  entomodont  hinge 
typical  of  the  genus.  The  strongly  tapering  outline  and  the  hinge  suggest  that  the 
material  is  of  a  pre-adult  instar,  but  it  is  doubtful  whether  the  adult  ornamentation 
would  be  different  from  that  described  above.  Certainly  there  is  no  difference 
between  the  instars  examined. 


Lophocythere  ostreata  (Jones  &  Sherborn) 
(PI.  12,  fig.  5;  Text-fig.  13) 

1888     Cytheridea  ostreata  Jones  &  Sherborn  :  271,  pi.  4,  figs.  6a-c. 

1888     Cytheridea  bicarinata  Jones  &  Sherborn  :  270,  pi.  4,  figs.  50— c. 

1948     Lophocythere  ostreata  (Jones  &  Sherborn)  Sylvester-Bradley  :  195,  pi.  14,  figs.  1—4,  pi.  15, 

figs,  i,  2. 
1963     Lophocythere  ostreata  (Jones  &  Sherborn);  Oertli  :  42,  pi.  28,  fig.  o,  pi.  29,  fig.  o. 

DIAGNOSIS.  Species  of  Lophocythere  with  characteristic  L-shaped  ridge  com- 
mencing at  a  large,  clear  eye  node,  extending  down  to  the  antero-ventral  region,  to 
bend  back  at  a  right-angle  and  extend  to  the  edge  of  the  posterior  marginal  border. 
Second,  keel-like  ridge  starts  below  the  previous  ridge  in  the  antero-ventral  part  of 
the  valve  and  extends  backwards,  parallel  to  the  first  ridge  to  die  out  against  the 
edge  of  the  posterior  marginal  border.  Shell  surface  coarsely  reticulate,  the  reticula- 
tions strongly  upstanding  to  produce  an  oblique  row  of  three  prominent  areas,  the 
first  of  which  is  antero-central  in  position,  just  inside  the  angle  of  the  L-shaped 
ridge.  The  second  is  dorso-median  in  position,  just  behind  mid-point,  whilst  the 
third  is  postero-dorsal  in  position.  Two,  sometimes  three,  minor  raised  reticulations 
may  be  situated  below  and  behind  the  two  most  dorsal  of  the  major  raised  areas. 
Ventral  surface  with  minor  longitudinal  ridges. 

HOLOTYPE.  1.1833,  right  valve  figured  Jones  &  Sherborn  (1888),  Yellow  Fuller's 
Earth  Clay,  Midford. 


ENGLISH    BATHONIAN    OSTRACODA  421 

OTHER  MATERIAL.  10.3945-7,  two  carapaces  and  two  broken  right  valve  frag- 
ments, Winwood  Collection,  Blue  Fuller's  Earth  Clay,  Midford.  These  were  referred 
to  by  Jones  &  Sherborn  (1888)  and  are  syntypes  of  Cytheridea  bicarinata,  the  figured 
specimen  having  been  lost.  10.3945  is  here  selected  lectotype  of  C.  bicarinata. 


FIG.   13.     Lophocythere  ostreata  (Jones  &  Sherborn).     Internal  view,  right  valve. 

Holotype,  1.1833.      X  IO5- 


DESCRIPTION.  Carapace  sub-rectangular  with  sub-parallel  dorsal  and  ventral 
margins  in  the  adult.  Juvenile  specimens  have  distinct  posterior  taper.  Anterior 
end  high,  broadly  rounded,  posterior  end  triangular,  with  concave  postero-dorsal 
slope  and  convex  postero-ventral  slope.  Cardinal  angles  prominent,  rounded. 
Anterior  cardinal  angle  with  well-developed,  oval,  clear  eye  node  situated  just  below. 
Line  of  greatest  length  passes  through  the  mid-point,  line  of  greatest  height  through 
anterior  cardinal  angle  and  of  greatest  width  through  the  posterior  third  of  the 
carapace.  Ornamentation  as  in  the  diagnosis.  The  reticulations  continue  into  the 
lateral  ridges  and  pass  on  to  the  anterior  and  posterior  marginal  borders.  The 
postero-dorsal  raised  area  is  not  developed  in  juvenile  instars.  Left  valve  larger  than 
the  right,  which  it  overlaps  along  the  ventral  margin  but  overreaches  around  the 
anterior  and  posterior  margins  and  in  the  region  of  the  cardinal  angles.  Hinge 
weakly  entomodont,  seen  only  in  the  right  valve:  anterior  dentate  ridge  with  six 
teeth,  posterior  element  not  seen,  median  groove  poorly  preserved  and  not  definitely 
seen  to  be  loculate,  expanded  slightly  in  the  anterior  half.  Inner  margin  and  line 
of  concrescence  coincide  to  produce  a  broad  duplicature.  Selvage  distinct,  flange 
developed  around  the  anterior  margin,  extending  from  the  anterior  cardinal  angle, 
along  the  ventral  margin  to  the  tip  of  the  posterior  margin.  Radial  pore  canals 
and  muscle  scars  not  observed. 

DIMENSIONS.  1.1833,  right  valve,  length  0-81  mm. ;  height  0-40  mm.  10.3945, 
carapace,  length  0-85  mm. ;  height  0-43  mm. ;  width  0-42  mm.  10.3947,  juvenile 
carapace,  length  0-60  mm. ;  height  0-31  mm. ;  width  0-28  mm. 

REMARKS.  The  Ostreata  Group  of  Lophocythere  is  characterized  by  the  presence 
of  two  lateral  keel-like  ridges,  the  upper  of  which  turns  upwards  to  produce  a  reverse 
L-shaped  ridge.  The  ornamentation  within  this  "  L  "  varies  according  to  the 
species.  In  L.  ostreata  the  reticulate  ornamentation,  extended  into  a  number  of 
upstanding  areas,  is  quite  unlike  any  described  species.  Exaggerations  in  the 
ornamentation  of  species  within  this  group  tend  to  be  in  the  same  position  as  the 
areas  emphasized  in  L.  ostreata.  In  this  connection  the  closest  species  is  L.  scabra 
Triebel  (1951),  although  it  does  not  possess  the  strong  reticulate  ornamentation  of 
ostreata. 


422  REVISION    OF    SOME 

Genus  MICROPNEUMATOCYTHERE  Bate  1963 
Micropneumatocythere  limaciformis  (Jones  &  Sherborn) 

(PI.  13,  fig.  i) 
1888     Cytheridea  limaciformis  Jones  &  Sherborn  :  269  pi.  3   figs.  i2a-c. 

DIAGNOSIS.  Carapace  oval/elongate  with  dorso-median  area  swollen,  convex, 
projecting  above  dorsal  margin  in  right  valve;  left  valve  with  convex  dorsal  margin. 
Ventro-lateral  and  median  part  of  ventral  surface  swollen,  strongly  convex,  some- 
times with  weak  longitudinal  striae  on  ventral  surface  of  convexity.  Shell  surface 
punctate.  Normal  pore  canal  openings  laige,  circular,  evenly  spaced  over  the 
carapace. 

HOLOTYPE.  1.1831,  male  right  valve,  figured  Jones  &  Sherborn  (1888),  Blue 
Fuller's  Earth  Clay,  Midford. 

OTHER  MATERIAL.  10.276  and  4007,  and  10.3944,  female  right  and  left  valves 
from  the  Bradford  Clay,  Bradford-on-Avon,  and  male  right  valve  from  the  Blue 
Fuller's  Earth  Clay,  Midford.  Win  wood  Collection. 

DESCRIPTION.  Carapace  oval/elongate  in  outline,  the  more  elongate  specimens 
being  the  males.  In  the  right  valve  of  both  dimorphs,  especially  the  male,  the 
dorso-median  part  of  the  valve  is  distinctly  swollen,  the  convex  surface  projecting 
above  the  dorsal  margin.  Similarly  the  ventro-lateral  part  of  the  valve  is  also 
swollen  but  involves  the  ventral  surface  where,  in  some  specimens,  weak  longitudinal 
striae  are  present.  Left  valve  with  uniformly  convex  dorsal  and  ventral  margins. 
Anterior  end  unifoimly  rounded,  posterior  end  triangular  with  concave  postero-dorsal 
slope  and  convex  postero-ventral  slope,  the  posterior  extremity  tending  to  be 
upturned.  Cardinal  angles  particularly  distinct  in  the  right  valve,  apparently 
separated  from  the  central  convexity  of  the  valve  by  a  shallow  groove.  Greatest 
length  of  carapace  passes  through  the  mid-point  with  the  greatest  height  and  width 
also  at  valve  centie.  Shell  surface  finely  punctate  with  prominent,  large,  circular 
normal  pore  canal  openings  evenly  spaced  over  the  carapace.  Hinge  almost  certainly 
antimerodont,  the  terminal  dentate/loculate  elements  being  clearly  seen,  whilst  the 
median  bar  and  groove  is  not  sufficiently  well  preserved  to  state  that  it  is  a  dentate/ 
loculate  structure.  Inner  margin  and  line  of  concrescence  coincide,  anterior  radial 
pore  canals  straight,  evenly  spaced  around  the  anterior  margin,  exact  number  not 
observed  but  no  more  than  eight.  A  narrow  flange  extends  around  the  anterior 
margin  of  the  right  valve  and  along  the  median  part  of  the  ventral  margin.  Muscle 
scars  not  seen. 

DIMENSIONS.  1.1831,  male  right  valve,  length  0-71  mm. ;  height  0-37  mm. 
10.3944,  male  right  valve,  length  0-62  mm. ;  height  0-32  mm.  10.276,  female  right 
valve,  length  0-56  mm. ;  height  0-36  mm.  10.4007,  female  left  valve,  length 
(damaged)  0-61  mm. ;  height  0-37  mm. 

REMARKS.  The  closest  species  is  M.  quadmta  Bate  (1967),  particularly  in  the  case 
of  the  male  dimorph.  However,  the  characteristic  development  of  a  deep  postero- 


ENGLISH    BATHONIAN    OSTRACODA  423 

ventral  slope  in  M.  quadrata  and  the  less  strongly  developed  dorso-median  convexity 
of  the  right  valve  distinguish  this  species  from  M.  limaciformis. 

Genus  PROGONOCYTHERE  Sylvester-Bradley  1948 

Progonocythere  stilla  Sylvester-Bradley 

(PL  13,  ng.  2) 

1948     Progonocythere  stilla  Sylvester-Bradley  :  190,  pi.  12,  figs,  i,  2,  pi.  13,  figs,  i,  2. 

1956     Progonocythere  stilla  Sylvester-Bradley;  Grekoff  :  pi.  13,  figs.  325-7. 

1963     Progonocythere  stilla  Sylvester- Bradley ;  Oertli  :  44,  pi.  28,  fig.  n,  pi.  29,  fig.  n. 

DIAGNOSIS.  Progonocythere  with  finely  punctate  shell  surface,  large,  circular 
normal  pore  canal  openings,  somewhat  irregularly  scattered,  tending  to  be  con- 
centrated towards  centre  of  valve.  Anterior  end  high,  broadly  rounded.  Anterior 
cardinal  angle  close  to  anterior  margin.  Cardinal  angles  well  rounded  in  left  valve 
more  distinctly  angled  in  right.  Dorsal  margin  convex  in  right  valve,  straight  in  left, 
in  both  valves  steeply  sloping  to  posterior.  Posterior  rounded  in  left  valve,  slightly 
angled  in  right.  Anterior  marginal  border  distinct  from  convexity  of  valve,  ventro- 
lateral  margin  strongly  convex,  overhanging  ventral  margin.  Left  valve  larger 
than  the  right.  Hinge  entomodont.  Other  internal  features  as  for  genus. 
Dimorphic. 

REMARKS.  Progonocythere  stilla,  the  type  species  ol  the  genus,  was  described  by 
Sylvester-Bradley  from  the  Bathonian  Boueti  Bed  of  Langton  Herring.  A  single 
left  valve  (10.3657)  found  in  the  Winwood  Collection,  from  the  Blue  Fuller's  Earth 
Clay  of  Midford,  was  not  described  by  Jones  &  Sherborn.  Three  specimens 
(10.3584-6)  in  the  Mockler  Collection  are  from  the  Fuller's  Earth  of  Midford. 

DIMENSIONS.     10.3657,  left  valve,  length  0-60  mm.;  height  0-43  mm. 

Genus  TERQUEMULA  Blaszyk  &  Malz  1965 

Terquemula  blakeana  (Jones) 

(PL  4,  fig.  8) 

REMARKS.  For  synonymy  and  diagnosis  see  p.  393.  The  two  specimens  in  the 
Jones  &  Sherborn  Collection  which  have  been  placed  in  the  synonymy  are :  Cytheridea 
transversiplicata,  figured  Jones  &  Sherborn  1888,  pi.  3,  figs,  ^a-c,  1.1828  (holotype), 
right  valve,  length  0-59  mm.;  height  0-31  mm.,  and  Cytheridea  blakeana  (Jones) 
Jones  &  Sherborn,  figured  pi.  2,  figs.  na-c.  1.1853,  left  valve,  length  0-60  mm. ; 
height  0-31  mm.  In  addition  a  single  right  valve,  10.3941,  has  been  found  in  the 
Winwood  Collection  from  the  Blue  Fuller's  Earth  Clay,  Midford. 

INCERTAE  SEDIS 

The  following  ostracod  species  described  by  Jones  &  Sherborn  are  represented  by 
single  specimens  only.  They  are  retained  here  under  their  original  names  until 
comparative  material  enables  a  more  precise  determination. 


424  REVISION    OF    SOME 

"  Cythere  "  corrosa  Jones  &  Sherborn 

(PL  13,  fig-  4) 
1888     Cythere  corrosa  Jones  &  Sherborn  :  254,  pi.  2,  figs.  120— c. 

HOLOTYPE.  1.1865,  figured  Jones  &  Sherborn  (1888),  Blue  Fuller's  Earth  Clay, 
Midford. 

REMARKS.  This  small  ostracod  resembles  Caytonidea  in  outline  and  reticulate 
ornamentation,  but  differs  in  possessing  a  well-defined  anterior  marginal  border. 
In  Caytonidea  the  convex  outline  of  the  carapace  continues  without  a  break  on  to  the 
anterior  margin.  Internal  features  not  seen. 

DIMENSIONS.  1.1865,  carapace,  length  0-49  mm.;  height  0-28  mm.;  width 
0-28  mm. 


"  Cytheridea  "  aequabilis  Jones  &  Sherborn 

(PI.  13,  fig.  3) 
1888     Cytheridea  aequabilis  Jones  &  Sherborn  :  257,  pi.  i,  figs.  la-c. 

HOLOTYPE.  1.1864,  figured  Jones  &  Sherborn  (1888),  Yellow  Fuller's  Earth  Clay, 
Midford. 

REMARKS.  C.  aequabilis  has  a  finely  punctate  carapace,  rounded  anterior  and 
posterior  margins,  and  parallel  dorsal  and  ventral  margins.  In  outline  it  is  close  to 
the  unidentified  ostracod  figured  by  Bate  (1964,  pi.  13,  figs.  5-8).  Internal  features 
not  seen. 

DIMENSIONS.  1.1864,  carapace,  length  0-71  mm.;  height  0-30  mm  ;  width 
0-32  mm. 


"  Cytheridea  "  coarctata  Jones  &  Sherborn 

(PL  13,  fig-  5) 
1888     Cytheridea  coarctata  Jones  &  Sherborn  :  259,  pi.  i,  figs.  3«-e. 

HOLOTYPE.  1.1841,  figured  Jones  &  Sherborn  (1888),  Blue  Fuller's  Earth  Clay, 
Midford. 

REMARKS.  The  unornamented  carapace,  well-rounded  anterior  and  posterior 
margins,  parallel  dorsal  and  ventral  margins,  together  with  a  muscle  scar  pattern  in 
which  the  antero-dorsal  antennal  scar  is  crescentic  in  shape  (type  B),  suggest 
Homocytheridea  Bate  (i963«).  The  left  valve,  the  only  known  specimen,  has  been 
broken  posteriorly  and  a  more  definite  identification  is  not  attempted. 

DIMENSIONS.     1.1841,  left  valve,  length  (broken)  0-56  mm.;  height  0-26  mm. 


ENGLISH    BATHONIAN    OSTRACODA 


425 


"  Cytheridea  "  eminula  Jones  &  Sherborn 
(PI.  13,  fig.  6) 

1888     Cytheridea  eminula  Jones  &  Sherborn  :  261,  pi.  i,  figs.  na-c. 

LECTOTYPE.     1.1839,  figured  Jones  &  Sherborn  (1888),  Blue  Fuller's  Earth  Clay, 
Midford. 

REMARKS.     This  species  is  represented  by  a  single  left  valve,  although  two  speci- 
mens were  included  in  the  original  description. 

DIMENSIONS.     1.1839,  kft  valve,  length  0-54  mm.  ;  height  0-31  mm. 

"  Cytheridea  "  punctiputeolata  Jones  and  Sherborn 
(PI.  13,  fig.  8) 

1888     Cytheridea  punctiputeolata  Jones  &  Sherborn  :  258,  pi.  i,  figs. 


HOLOTYPE.  1.1856,  figured  Jones  &  Sherborn  (1888),  Blue  Fuller's  Earth  Clay, 
Midford. 

REMARKS.  Although  at  first  appearing  to  resemble  the  male  dimorph  of  Hadro- 
cytheridea  dolabra  the  present  species  differs  in  the  much  more  rounded  posterior 
margin,  in  the  greater  exaggeration  of  the  ornamentation,  which  here  consists  of  large, 
circular  pits  with  the  interspaces  between  distinctly  punctate,  and  in  the  lophodont 
hinge. 

DIMENSIONS.     1.1856,  right  valve,  length  0-73  mm.;  height  0-32  mm. 

"  Cytheridea  "  retorrida  Jones  &  Sherborn 

(PI.  15,  fig.  3;  Text-figs.  14,  15) 

1888     Cytheridea  retorrida  Jones  &  Sherborn  :  260,  pi.  i,  figs.  8a-c. 
MATERIAL.     10.3923,  Blue  Fuller's  Earth  Clay,  Midford.     Winwood  Collection. 

REMARKS.  The  figured  specimen  was  never  presented  and  is  presumed  lost. 
A  single  carapace  of  this  species  has  been  found  in  the  Winwood  Collection,  but 


FIG.  14.     Cytheridea  retorrida  Jones  &  Sherborn.     Right  valve,  complete  carapace. 

10.3923.      x  135. 

26 


GEOL.    17, 


426  REVISION    OF    SOME 

cannot  be  assigned  to  a  known  genus.  Like  C.  corrosa  it  has  some  affinities  with 
Caytonidea  but  has  been  excluded  on  the  presence  of  distinct  marginal  borders.  No 
internal  features  are  known. 


FIG.   15.     Cytheridea  retorrida  Jones  &  Sherborn.     Dorsal  view  of  carapace  with 
ornamentation  omitted.     10.3923.      x  135. 

DIMENSIONS.     10.3923,    carapace,    length    0-54  mm. ;    height    0-28  mm.;    width 
0-28  mm. 

"  Cytheridea  "  spinifastigiata  Jones  &  Sherborn 
(PI.  13,  ng.  7) 

1888     Cytheridea  spinifastigiata  Jones  &  Sherborn  :  264,  pi.  2,  figs.  8a-c. 

HOLOTYPE.     I.i86i,  figured  Jones  &  Sherborn  (1888),  Blue  Fuller's  Earth  Clay, 
Midford. 

REMARKS.     It  is  not  possible  to  assign  the  single  left  valve  to  any  known  genus. 
DIMENSIONS.     1. 1861,  left  valve,  length  0-60  mm.;  height  0-33  mm. 

"  Cytheridea  "  spinigyrata  Jones  &  Sherborn 

(PL  13,  fig.  9) 
1888     Cytheridea  spinigyrata  Jones  &  Sherborn  :  264,  pi.  2,  figs.  ja-c. 

HOLOTYPE.     1. 1860,  figured  Jones  &  Sherborn  (1888),  Blue  Fuller's  Earth  Clay, 
Midford. 


ENGLISH    BATHONIAN    OSTRACODA  427 

REMARKS.  Jones  &  Sherborn  (p.  264)  compared  this  species  with  Cythere  drupacea 
Jones  but  at  the  same  time  pointed  out  differences.  The  superficial  similarity  of 
C.  spinigyrata  to  C.  drupacaea  suggests  that  it,  too,  might  belong  to  Metacytheropteron 
Oertli  (1957). 

DIMENSIONS.     1. 1860,  right  valve,  length  0-61  mm.;  height  0-34  mm. 


IV.  THE  J.  F.  BLAKE  COLLECTION 

Order  MYODOCOPIDA  Sars  1866 

Suborder  CLADOCOPINA  Sars  1860 

Family  POLYCOPIDAE  Sars  1866 

Genus  POLYCOPE  Sars  1866 

Polycope  fungosa  sp.  nov. 

(PI.  16,  figs.  4,  9) 

DIAGNOSIS.  Carapace  oval  in  adults,  almost  circular  in  juvenile  instars.  Shell 
surface  strongly  pitted  with  weak  striae  near  and  parallel  to  the  free  margin. 
Anterior  end  flattened,  with  marginal  denticles.  Hinge  short,  sunk  in  dorsal  groove. 
Anterior  cardinal  angle  prominent,  antero-dorsal  slope  long,  broadly  convex  with 
prominent  overlap  of  left  valve  by  the  right.  Posterior  end  broadly  rounded,  ventral 
margin  strongly  convex.  Carapace  distinctly  angled  at  junction  of  antero-dorsal 
slope  and  anterior  margin.  Right  valve  larger  than  left. 

HOLOTYPE.     10.3951,  carapace,  Fuller's  Earth,  Midford,  near  Bath. 

PARATYPES.  10.3952-5,  one  adult  and  eighteen  juvenile  carapaces.  Horizon 
and  locality  as  above. 

DESCRIPTION.  Carapace  in  adult  specimens  oval  in  outline,  laterally  compressed. 
Lines  of  greatest  length  and  height  pass  through  the  mid-point.  Greatest  width  in 
the  holotype  situated  at  the  extreme  posterior  end.  A  shallow  central  depression 
on  the  lateral  surface  marks  the  position  of  the  muscle  scars.  Hinge  margin  situated 
in  the  posterior  half  of  the  carapace  and  sunk  into  a  dorsal  groove,  at  the  anterior 
end  of  which  the  cardinal  angle  is  prominently  developed.  Antero-dorsal  slope  long 
and  convex,  terminating  in  a  sharply  angled  junction  with  the  obliquely  flattened 
anterior  end.  Posterior  end  broadly  rounded,  ventral  margin  broadly  convex. 
Shell  surface  uniformly  and  distinctly  pitted;  striae  are  developed  in  the  anterior 
and  ventro-lateral  regions.  In  dorsal  view  one  stria  on  each  valve  can  be  seen  to 
project  beyond  the  anterior  margin.  Striations  are  arranged  parallel  to  the  free 
margins.  Anterior  margin  may  develop  small  denticles,  though  these,  like  the 
surface  ornamentation,  are  generally  absent  from  the  juvenile  instars.  Right  valve 
larger  than  the  left,  which  it  overlaps  strongly  along  the  antero-dorsal  slope  and  less 
strongly  around  the  posterior  end.  Internal  features  not  observed. 

DIMENSIONS.  10.3951,  carapace,  length  0-54 mm. ;  height  0-44 mm.;  width 
0-23  mm.  10.3952,  carapace,  length  0-46  mm.;  height  0-37  mm.;  width  0-20  mm. 


4z8  REVISION    OF    SOME 

10.3953,  juvenile  carapace,  length  0-32  mm. ;   height  0-27  mm. ;   width  0-09  mm. 

10.3954,  juvenile  carapace,  length  0-26  mm. ;  height  0-22  mm. ;  width  o-oi  mm. 
REMARKS.     A  number  of  species  of  Poly  cope  have  been  described  from  the  Lower 

Jurassic  (Apostolescu  1959;  Fischer  1961  and  Blake  1876),  Middle  Jurassic  (Terquem 
1885)  and  Upper  Jurassic  (Donze  1962  and  Oertli  1959).  P.  fungosa  differs  from 
them  all  in  the  type  of  ornamentation  present.  A  number  of  specimens  in  the  Blake 
collection  labelled  Poly  cope  sp.  are  probably  juvenile  specimens  of  the  present  species 
but  lack  the  specific  ornamentation. 

Family  PROGONOCYTHERIDAE  Sylvester-Bradley  1948 
Genus  TRACHYCYTHERE  Triebel  &  Klingler  1959 

REMARKS.  Trachycythere  Triebel  &  Klingler  (1959  :  343),  first  described  from  the 
Lias  of  Germany  and  later  recorded  from  the  Upper  Bathonian  of  France  (Oertli 
1963),  was  placed  in  the  Family  Trachyleberididae  (Moore  1961  :  341).  Later  Bate 
(19670  :  551)  transferred  it  to  the  Progonocytheridae,  a  classification  retained  in 
this  paper. 

Trachycythere  sp. 

(PL  16,  fig.  3) 

REMARKS.  A  single  left  valve  represents  this  Upper  Bathonian  species,  which 
differs  in  being  sub-rectangular  with  almost  parallel  dorsal  and  ventral  margins, 
from  two  of  the  three  species  originally  described  by  Triebel  &  Klingler,  which  have 
a  more  pronounced  posterior  taper.  Trachycythere  verrucosa  Triebel  &  Klingler 
(1959)  resembles  this  species  in  outline,  but  has  stronger,  more  massive  tubercles. 

DIMENSIONS.     10.3966,  left  valve,  length  0-45  mm. ;  height  0-25  mm. 

Genus  HEKISTOCY THERE  nov. 
DERIVATION  OF  NAME.     Hekistos  (Gr.),  least,  +  cythere. 

DIAGNOSIS.  A  small  genus  with  oval,  posteriorly  tapering  carapace.  Greatest 
height  close  to  the  anterior  margin.  Left  valve  larger  than  right.  Hinge  lophodont. 
Muscle  scars  of  type  A  although  antennal  scar  tends  to  be  almost  centrally  situated 
in  front  of  adductors.  Radial  pore  canals  straight,  widely  spaced,  ten  anteriorly, 
four  posteriorly.  Duplicature  narrow. 

TYPE  SPECIES.     Hekistocy there  venosa  sp.  nov. 

Hekistocy there  venosa  gen.  et  sp.  nov. 
(PI.  14,  figs.  2,  4,  9;  PI.  15,  fig.  7;  Text-figs.  16-18) 

DIAGNOSIS.  Hekistocy  there  with  strongly  ornamented  carapace.  Broad  diagonal 
ridge  extends  from  postero-dorsal  to  antero- ventral  region  of  valve.  Short,  thick, 


ENGLISH    BATHONIAN    OSTRACODA 


429 


subsidiary  ridges  given  off  on  dorsal  and  ventral  sides  of  main  ridge.  Smaller 
intermediary  ridges  producing  a  reticulate  ornamentation.  Diagonal  ridge  produced 
at  termination  of,  and  at  right-angles  to,  main  ridge  in  postero-dorsal  region. 

HOLOTYPE.     10.3995,  carapace,  Fuller's  Earth,  Bath. 

PARATYPES.     10.3996-9  (seven  specimens)  and  10.4011-3,  Fuller's  Earth,  Bath. 

DESCRIPTION.  Carapace  small  with  high,  rounded  anterior  end  and  narrow, 
rounded  posterior  end.  Carapace  parallel-sided  in  dorsal  view.  Line  of  greatest 
length  passes  through  the  mid-point.  Greatest  width  in  either  the  anterior  or 


18 


FIGS.   16-18.     Hekistocy there  venosa  sp.  nov.     FIG.   16.     Anterior  radial  pore  canals.     Para- 
type,     10.4013.      x  300.     FIG.   17.     Hinge,     right     valve,     paratype,     10.4011.      x  300. 

FIG.   18.     Muscle  scars,  left  valve,  paratype,  10.3997.      X  300. 


posterior  third,  the  carapace  being  slightly  constricted  medially.  Dorsal  margin 
convex,  especially  in  the  right  valve  where  it  projects  slightly  above  that  of  the  left 
valve.  Surface  ornamentation  very  coarsely  reticulate,  produced  by  a  series  of 
irregular  ridges,  the  major  of  which  extends  down  from  the  postero-dorsal  part  of  the 
valve  and  extends  to  the  antero- ventral  region.  From  this  ridge  a  number  of 
irregular  and  shorter  ridges  branch  off  on  its  dorsal  and  ventral  sides.  One  of  the 
secondary  ridges  is  particularly  noticeable  in  dorsal  view,  and  this  leaves  the  primary 
ridge  in  the  posterior  part  of  the  valve  to  extend  dorsally  in  an  antero-dorsal  direction 
until  it  fuses  with  the  dorsal  margin.  Left  valve  larger  than  the  right,  which  it 
overlaps  along  the  ventral  margin  and  overreaches  in  the  region  of  the  antero-  and 


430  REVISION    OF    SOME 

postero-dorsal  slopes.  Ventral  surface  flattened,  triangular  in  outline,  widening 
towards  the  posterior  end.  Hinge  lophodont:  right  valve  with  smooth  terminal 
elements  which  are  the  dorsal  terminations  of  the  selvage;  median  groove  deeply 
recessed,  smooth.  Left  valve  with  smooth  terminal  sockets  open  ventrally  to  the 
interior  of  the  valve.  Median  bar  long,  smooth,  convex,  the  convexity  of  the  bar 
facing  upwards.  Inner  margin  and  line  of  concrescence  coincide,  the  duplicature 
being  rather  narrow.  Radial  pore  canals  short  and  straight  and  widely  spaced, 
ten  anteriorly  and  four  posteriorly.  Muscle  scars  of  type  A  (Bate  1963),  with  four 
adductor  scars  in  a  slightly  crescentic  row;  the  second  dorsal  adductor  is  laterally 
elongate  with  the  rounded  antennal  scar  situated  in  front. 

DIMENSIONS.  10.3995,  carapace,  length  0-32  mm. ;  height  0-20  mm. ;  width 
0-15  mm.  10.3996,  left  valve,  length  0-32  mm. ;  height  0-21  mm.  10.4011,  right 
valve,  length  0-32  mm. ;  height  0-20  mm.  10.4012,  left  valve,  length  0-33  mm. ; 
height  0-21  mm. 

REMARKS.  Hekistocy there  has  a  distinctive  combination  of  lophodont  hinge,  small, 
oval,  posteriorly  tapered  carapace,  type  A  muscle  scars,  and  simple  radial  pore 
canals. 


Family  PROTOCYTHERIDAE  Ljubimova  1955 

Subfamily  KIRTONELLINAE  Bate  1963 

Genus  EKTYPHOCY THERE  Bate  1963 

Ektyphocythere  parva  (Oertli) 

(PI.  14,  figs,  i,  3) 

J959     Procytheridea  minuta  Oertli  :  122,  pi.  3,  figs.  37-40. 
1960    Procytheridea  parva  Oertli  :  70. 

REMARKS.  Two  complete  carapaces  represent  this  species,  one  male  the  other 
female. 

When  first  erected,  Ektyphocythere  Bate  (1963)  included  those  species  of  "  Pro- 
cytheridea "  which  possessed,  amongst  other  characters,  a  triangular  arrangement  of 
ridges  as  ornamentation  and  an  antimerodont  hinge.  P.  parva  was  one  of  the  species 
included. 

Subsequently  Malz  (1966)  placed  Ektyphocythere  in  synonymy  with  Kinkelinella 
Martin  (1960)  the  type  species,  K.  tenuicostati  Martin  (1960),  having  a  reticulate 
ornamentation.  Although  considered  here  to  be  closely  related  the  clear  distinction 
between  the  forms  having  a  triangular  rather  than  a  reticulate  ornamentation  is 
such  that  the  retention  of  the  former  in  Ektyphocythere  is  maintained  here. 

DIMENSIONS.  10.3982,  female  carapace,  length  0-41  mm. ;  height  0-23  mm. ; 
width  0-21  mm.  10.3983,  male  carapace,  length  0-45  mm.;  height  0-23  mm.;  width 
0-24  mm. 


ENGLISH  BATHONIAN  OSTRACODA  431 

FAMILY  UNCERTAIN 
Genus  PLATYCYTHERE  Bate  1967 
Platycythere  sp. 

(PI.  14,  fig.  5) 

REMARKS.  Represented  by  a  single  specimen  only.  In  outline  and  possession 
of  dorsal  and  ventro-lateral  swellings  this  species  exhibits  the  generic  features 
described  (Bate  1967),  but  differs  from  Platycythere  verriculata,  the  type  species,  by 
having  a  much  finer  reticulate  ornamentation  and  a  median  swelling  on  the  lateral 
surface  of  both  valves.  This  swelling,  positioned  about  the  centre  of  the  valve,  has 
an  oblique  postero-dorsal  antero- ventral  inclination. 

Gesoriacula  Oertli  (1959)  has  a  similar  external  morphology  but  a  more  primitive 
hinge  structure. 

DIMENSIONS.  10.3992,  carapace,  length  0-35  mm. ;  height  0-20  mm. ;  width 
0-16  mm. 

Family  CYTHERURIDAE  Miiller  1894 
Genus  LOONEYELLA  Peck  1951 

REMARKS.  The  type  species,  Cythere  monticula  Jones  (1893),  was  described  from 
the  Cretaceous  Bear  River  Formation,  Cokeville,  south-west  Wyoming,  in  association 
with  fresh-  to  brackish-water  ostracods.  Peck  (1951),  in  resampling  this  Formation, 
found  a  new  ostracod  species  which  he  considered  to  be  congeneric  with  C.  monticula, 
and  for  these  two  species  erected  the  genus  Looneyella,  C.  monticula  being  the  type. 

The  preservation  of  Peck's  material  is  considerably  better  than  that  of  the  holotype, 
which  is  in  the  Museum  collections.  The  latter,  1.5872,  figured  Jones  (1893,  pi.  15, 
fig.  13),  is  a  complete  carapace,  length  0-65  mm.;  height  0-40  mm.;  width  0-39  mm. 
(inclusive  of  spines).  The  holotype  is  figured  here  (PL  14,  fig.  6)  for  comparison  with 
the  Bathonian  species  L.  subtilis. 

To  date  the  following  species  have  been  placed  in  Looneyella : 

L.  monticula  (Jones)  from  the  Upper  Cretaceous  Bear  River  Formation. 
L.  quadrispina  Peck  1951,  also  from  the  Bear  River  Formation. 
L.  subtilis  Oertli,  from  the  Bathonian  of  the  Boulonnais,  France. 

Looneyella  subtilis  Oertli 
(PL  14,  figs.  7,  8) 

1959     Looneyella  ?  subtilis  Oertli  :  119,  pi.  3,  figs.  31-5. 

DIAGNOSIS.  Small  species  of  Looneyella  with  prominent  postero-dorsal  tubercle, 
an  antero-median  tubercle  somewhat  smaller  in  size,  and  a  marginal  ridge  which 
extends  around  the  anterior  margin  and  along  the  ventro-lateral  alar  extension. 
Both  tubercles  and  marginal  ridge  are  hollow  when  viewed  internally.  Antero- 


432  REVISION    OF    SOME 

medially  there  is  a  break  in  the  continuity  of  the  marginal  ridge.  Small  swelling 
may  be  present  below  the  postero-dorsal  tubercle,  and  at  the  termination  of  the  alar 
ridge.  Shell  surface  smooth  except  for  ventral  surface  which  has  four  longitudinal 
striae  per  valve.  Left  valve  larger  than  right.  Hinge  antimerodont.  Species 
dimorphic.  Eight  straight  anterior  radial  pore  canals;  inner  margin  and  line  of 
concrescence  coincide.  Low  eye  swelling  occurs  at  the  anterior  cardinal  angle  and  is 
connected  to  the  anterior  marginal  ridge. 

REMARKS.  Oertli  placed  this  species  only  tentatively  in  Looneyella,  but  com- 
parison with  the  holotype  of  the  type  species  convinces  me  that  L.  subtilis  is  con- 
generic. Dr.  Oertli  kindly  sent  me  specimens  of  his  material  for  comparison  and 
enabled  me  to  examine  internal  characters  not  available  in  my  material. 

Ecologically  this  is  an  interesting  genus  with  the  Cretaceous  species  recorded  from 
sediments  considered  to  be  non-marine  whilst  L.  subtilis  occurs  in  truly  marine 
conditions.  Possibly  the  Cretaceous  species  lived  under  brackish-water  conditions. 

The  material  from  the  Upper  Bathonian  Bradford  Clay  of  Bradford  consists  of 
a  single  female  carapace  in  the  Winwood  Collection  (10.3659),  and  two  male  carapaces 
and  a  juvenile  carapace  (10.3956-8)  from  the  Fuller's  Earth  of  Bath  in  the  Blake 
Collection. 

DIMENSIONS.  10.3659,  female  carapace,  length  0-37  mm. ;  height  0-22  mm. ; 
width  0-21  mm.  10.3957,  male  carapace,  length  0-35  mm.;  height  o-i8mm.; 
width  0-17  mm. 

Genus  ORTHONOTACYTHERE  Alexander  1933 
Orthonotacythere  sp. 

(PL  15,  ng.  i) 

REMARKS.  The  single  specimen  is  a  small  carapace  having  a  high  caudal  process, 
a  long,  straight  dorsal  margin  and  a  ventral  surface  triangular  in  outline.  The 
lateral  surface  is  weakly  reticulate  but  prominently  ornamented  by  three  large 
tubercles.  The  first  occurs  at  the  anterior  cardinal  angle  and  is  an  eye  tubercle. 
The  second  occurs  at  the  postero-dorsal  part  of  the  carapace  and  is  somewhat  elongate 
in  outline,  whilst  the  third  is  situated  directly  below  the  second  at  the  postero- ventral 
corner  of  the  carapace.  This  last  tubercle,  positioned  at  the  end  of  a  somewhat 
thickened  ventro-lateral  margin,  extends  slightly  beyond  the  carapace  posteriorly. 
A  weak  lateral  ridge  is  present  at  about  valve  centre,  but  does  not  estend  as  far  as  the 
terminal  margins. 

Orthonotacythere  nodosa  Plumhoff  (1963),  which  appears  in  the  Lias  of  Germany, 
differs  from  the  present  species  in  possessing  a  much  larger  number  of  tubercles. 
?0.  pulchella  Apostolescu  (1959)  from  the  Lias  of  France  does  not  belong  to  this 
genus.  Orthonotacythere  sp.  also  differs  from  the  small  number  of  species  described 
from  the  Upper  Jurassic  of  the  U.S.S.R.  (Ljubimova  1955),  Germany  (Triebel  1941) 
and  France  (Donze  1960). 

DIMENSIONS.  10.3993,  carapace,  length  0-30  mm. ;  height  0-15  mm. ;  width 
0-14  mm. 


ENGLISH    BATHONIAN    OSTRACODA  433 

Genus  PARARISCUS  Oertli  1959 

Parariscus  bathonicus  Oertli 

(PI.  15,  figs.  2,  8) 

1959     Parariscus  bathonicus  Oertli  :  118,  pi.  2,  figs.  20-29. 

REMARKS.  Eight  complete  carapaces  of  this  interesting  species  are  present  in  the 
Blake  Collection  of  Fuller's  Earth  Ostracoda.  Dr.  H.  J.  Oertli  kindly  sent  me 
comparative  material. 

DIMENSIONS.  10.3959,  carapace,  length  0-43  mm. ;  height  0-21  mm. ;  width 
0-20  mm. 

Genus  PARACYTHERIDEA  Miiller  1894 

Paracytheridea?  blakei  sp.  nov. 

(PI.  15,  figs.  4,  5,  9,  10) 

DIAGNOSIS.  Carapace  small,  reticulate,  with  dorso-median  constriction. 
Postero- ventral  angle  swollen,  extended  backwards  beyond  margin.  Eye  node 
developed  at  anterior  cardinal  angle.  Dorsal  margin  long,  straight.  Anterior  end 
high,  rounded.  Posterior  end  narrow,  triangular,  situated  high  on  carapace. 

HOLOTYPE.     10.4001,  carapace,  Fuller's  Earth,  Bath. 

PARATYPES.     10.4014-6,  three  carapaces,  Fuller's  Earth,  Bath. 

DESCRIPTION.  Carapace  small  with  coarse  reticulate  ornamentation.  Dorso- 
median  part  of  each  valve  with  broad  depression.  Ventro-lateral  margin  swollen, 
particularly  towards  the  posterior  end  where,  at  the  postero-ventral  angle,  there  is 
a  backward  projection  of  the  carapace  beyond  the  steeply  angled  postero-ventral 
slope.  Dorsal  margin  long  and  straight  with  sharply  angled  cardinal  angles.  An 
eye  node  is  situated  behind  the  anterior  cardinal  angle.  Anterior  end  high,  rounded. 
Posterior  end  situated  high  on  the  carapace,  sharply  triangular  with  short,  concave 
postero-dorsal  slope  and  long,  convex  postero-ventral  slope.  Line  of  greatest  length 
slightly  above  the  mid-point ;  greatest  height  at  the  anterior  cardinal  angle.  Greatest 
width  in  the  posterior  third.  Left  valve  slightly  larger  than  the  right,  which  it 
overlaps  along  the  ventral  margin.  Internal  features  not  seen  apart  from  the  muscle 
scars  which  are  visible  from  the  exterior :  these  consist  of  a  slightly  crescentic  row  of 
four  adductor  scars,  an  antero- ventral,  rounded,  manibular  scar,  and  an  antero-dorsal 
antennal  scar  which  is  composed  of  two  scars,  the  first  crescentic  in  shape,  the  second 
rounded  and  fitting  inside  the  crescent  of  the  first.  These  muscle  scars  are  situated 
low  down  on  the  carapace,  below  the  dorso-median  depression. 

DIMENSIONS.  10.4001,  holotype,  carapace,  length  0-30  mm. ;  height  0-16  mm. ; 
width  0-14  mm. 

REMARKS.  Paracytheridea?  blakei  was  named  after  J.  F.  Blake,  in  whose  collection 
the  species  was  found.  A  new  genus  might  be  preferred,  but  for  the  present  it  is 
sufficient  to  place  the  species  in  Paracytheridea  on  the  basis  of  the  external  shape. 
There  is  no  strong  nodose  ornamentation  as  in  Paracytheridea?  caytonensis  Bate 
(1965). 


434  REVISION    OF    SOME 

Genus  CYTHERURA  Sars  1866 

Cytherura  bathonica  sp.  nov. 

(PL  16,  figs.  5-8) 

DIAGNOSIS.  Cytherura  with  small,  elongate  carapace,  ornamented  with  slender, 
almost  parallel,  striae.  Short,  vertical  connecting  bars  between  striae  present  a 
slightly  reticulate  appearance.  Posterior  extremity  situated  above  the  line  of  the 
longitudinal  axis. 

HOLOTYPE.     10.4017,  carapace,  Fuller's  Earth,  Bath. 

PARATYPES.     10.4018-9  and  10.4120,  eight  carapaces.     Fuller's  Earth,  Bath. 

DESCRIPTION.  Carapace  small,  elongate,  with  line  of  greatest  length  very  slightly 
above  the  mid-point.  Greatest  height  at  the  extreme  anterior  end,  greatest  width 
in  the  posterior  third.  Shell  surface  ornamented  by  slender,  almost  parallel  striae 
with  connecting  vertical  bars  between.  A  slightly  reticulate  appearance  is  thereby 
presented  although  this  is  subordinate  to  the  sub-parallel  striae.  Dorsal  and  ventral 
margins  long,  straight  and  almost  parallel.  Anterior  end  broadly  rounded,  posterior 
end  with  narrow  triangular  margin  terminally  truncated  and  set  high  on  the  carapace : 
the  concave  postero-ventral  slope  is  much  longer  than  the  concave  postero-dorsal 
slope.  Left  valve  only  slightly  larger  than  the  right,  overlapping  it  ventrally  just 
anterior  of  median  and  at  the  cardinal  angles.  The  right  valve  overlaps  the  left 
along  the  dorsal  margin.  Internal  features  not  seen. 

DIMENSIONS.  10.4017,  carapace,  length  0-30  mm. ;  height  0-13  mm. ;  width 
o-io  mm.  10.4018,  carapace,  length  0-31  mm. ;  height  0-14  mm. ;  width  o-n  mm. 

REMARKS.  Cytherura  bathonica  is  a  true  Cytherura  as  far  as  the  external  details 
are  concerned.  The  presence  of  this  species  and  C.  mediojurassica  sp.  nov.  in  the 
Bathonian  indicates  that  the  genus  was  established  by  or  during  the  Middle  Jurassic. 
C.  bathonica  differs  from  previously  described  species  in  ornamentation  although  it 
comes  close  to  the  Recent  Cytherura  striata  Sars  (1866)  from  which  it  may  be  distin- 
guished by  the  position  of  the  posterior  extremity,  which  is  situated  below  the 
longitudinal  axis  in  C.  striata  and  above  in  C.  bathonica. 

Cytherura  mediojurassica  sp.  nov. 
(PL  15,  fig.  6;  PL  16,  figs,  i,  2) 

DIAGNOSIS.  Cytherura  with  reticulate  ornamentation.  Carapace  elongate,  dorsal 
and  ventral  margins  long,  almost  parallel  but  converging  slightly  towards  posterior 
end.  Short  caudal  process  set  high  up  on  carapace. 

HOLOTYPE.     10.4121,  carapace,  Fuller's  Earth,  Bath. 

PARATYPES.     10.4122-38,  sixteen  carapaces,  Fuller's  Earth  Clay,  Bath. 

DESCRIPTION.  Carapace  elongate  with  long,  sub-parallel  dorsal  and  ventral 
margins  which  converge  slightly  towards  the  posterior  end.  Greatest  length  of 


ENGLISH    BATHONIAN    OSTRACODA  435 

carapace  through  the  mid-point  whilst  the  greatest  height  is  at  the  extreme  anterior 
end,  through  the  anterior  cardinal  angle.  Greatest  width  in  the  posterior  third. 
Carapace  slender  in  dorsal  view.  Anterior  end  rounded,  posterior  end  with  short 
caudal  process  set  high  on  the  carapace,  slightly  above  the  line  of  greatest  length. 
Postero-dorsal  slope  short,  concave ;  postero- ventral  slope  long  and  concave,  although 
in  some  specimens  it  is  slightly  convex.  Termination  of  caudal  process  truncated. 
Shell  surface  ornamented  by  a  rather  coarse  reticulation.  Occasionally  a  weak 
longitudinal  stria  is  present  in  the  posterior  half  of  the  carapace  and  extends  along 
the  longitudinal  axis  as  far  as  the  valve  centre.  This  striation  is  really  an  exag- 
gerated development  of  the  reticulation  and  in  the  holotype  is  present  only  on  the 
right  valve.  Left  valve  slightly  larger  than  the  right  in  that  it  overlaps  the  right 
along  the  ventral  margin,  especially  along  its  anterior  half.  Dorsally,  however,  the 
right  valve  overreaches  the  left.  Internal  features  not  known. 

DIMENSIONS.  10.4121,  carapace,  length  0-33  mm. ;  height  0-15  mm. ;  width 
0-13  mm.  10.4122,  carapace,  length  0-31  mm. ;  height  0-15  mm. ;  width  0-12  mm. 

REMARKS.  Cytherura  mediojurassica  sp.  nov.  is  easily  distinguished  from  C. 
bathonica  by  its  strong  reticulate  ornamentation,  and  from  Cytherura?  liesbergensis 
Oertli  (1959)  by  its  more  elongate  carapace  in  which  the  dorsal  and  ventral  margins 
are  sub-parallel.  In  addition  the  ornamentation  of  C.?  liesbergensis  is  weakly 
reticulate,  with  four  lateral  striae  extending  along  most  of  the  length  of  the  carapace. 
These  striae  are  not  represented  in  the  present  species.  Dr.  H.  J.  Oertli  kindly  sent 
me  comparative  material. 

V.  REFERENCES 

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R.  H.  Bate,  B.Sc.,  Ph.D., 

Department  of  Palaeontology 

BRITISH  MUSEUM  (NATURAL  HISTORY) 

LONDON.  S.W.7 


EXPLANATION    OF   PLATES 

All  the  specimens  illustrated  are  in  the  Department  of  Palaeontology,  British 
Museum  (Natural  History).  With  the  exception  of  PL  5,  figs.  5,  6,  and  PL  16,  fig  4, 
the  photographs  were  taken  by  the  author  with  the  Stereoscan  scanning  electron 
microscope. 


PLATE    I 

FIG.  i.  Cytherella  symtnetrica  Jones.  External  view,  male  left  valve  and  lectotype  of 

Cytherella  subovata  Jones.  IN. 43496.  x  75. 

FIG.  2.  Cytherella  symmetrica  Jones.  External  view  of  lectotype,  female  left  valve. 

IN.43503.  x  85. 

FIG.  3.  Cytherelloidea  jugosa  (Jones).  External  view  of  lectotype,  female  right  valve. 

IN-43497.  x  120. 

FIG.  4.  Cytherelloidea  jugosa  (Jones).  External  view  of  paralectotype,  left  side  of  female 

carapace.  1.2311.  x  115. 

FIG.  5.  Bairdia  Hilda  Jones.  Right  valve  of  complete  carapace,  paralectotype,  10.3608. 
x  78. 

FIG.  6.  Bairdia  Hilda  Jones.     Left  valve,  lectotype,  IN. 4 1951.      x  73. 

FIG.  7.  Bairdia  juddiana  Jones.  Right  valve  of  complete  carapace,  holotype,  IN. 43506. 
X  65. 

FIG.  8.  Lophocythere  acutiplicata  (Jones  &  Sherborn).  Male  right  valve,  paralectotype, 

10.3643.  x  95- 


Butt.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE    i 


GEOL.    17,  8 


27 


PLATE   2 
FIG.  i.     Bairdia  jurassica  Jones.     Left  valve  of  complete  carapace,  lectotype,  1^43494. 

X  73- 

FIG.  2.  Bairdia  jurassica  Jones.  Right  valve  of  complete  carapace,  paralectotype. 
Specimen  figured  and  described  by  Jones  as  B.  jurassica  var  tenuis.  1^43495.  x  73. 

FIG.  3.  Pontocyprella  harrisiana  (Jones).  Left  valve  of  specimen  figured  and  described 
by  Jones  as  Macrocypris  bradiana.  IN. 43500.  x  70. 

FIG.  4.  Schuleridea  (Eoschuleridea)  subperjorata  (Jones).  External  view,  right  valve, 
lectotype,  originally  figured  as  Cytheridea  subperforata  (in  part)  by  Jones.  IN. 43504.  x  97. 

FIG.  5.  Schuleridea  (Schuleridea)  jonesiana  (Bosquet).  External  view,  left  valve  of 
complete  carapace  originally  figured  as  Cytheridea  subperforata  (in  part)  by  Jones.  IN-4349O. 

X  73- 

FIG.  6.  Praeschuleridea  schwageriana  (Jones).  Right  valve  of  complete  carapace, 

holotype.  ^.43499.  Originally  figured  as  Cythere  schwageriana  Jones,  x  99. 

FIG.  7.  Schuleridea  (Eoschuleridea)  trigonalis  (Jones).  Right  valve,  complete  carapace, 

holotype.  IN. 42373.  Originally  described  as  Bairdia  trigonalis  Jones,  x  85. 

FIG.  8.  Praeschuleridea  sp.     Right  valve  of  complete  female  carapace,  10.3619.      x  98. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE    2 


PLATE   3 

FIG.  i.  Glyptocythere  guembeliana  (Jones).  External  view,  male  right  valve,  lectotype, 
IN. 43493.  Originally  described  as  Cythere  guembeliana  by  Jones,  x  77. 

FIG.  2.  Glyptocythere  guembeliana  (Jones).  External  view,  juvenile  right  valve, 
1.1840.  Originally  described  as  Cytheridea  trapezoidalis  Terquem  by  Jones  &  Sherborn.  x  113. 

FIG.  3.  Lophocy there  bradiana  (Jones).  Left  valve  of  complete  male  carapace,  lectotype, 
IN. 42372.  Originally  described  as  Cythere  bradiana  by  Jones,  x  101. 

FIG.  5.  Lophocy  there  bradiana  (Jones).  Right  valve  of  complete  male  carapace,  para- 
lectotype,  10.3627.  x  95. 

FIG.  6.  Lophocy  there  bradiana  (Jones).  Left  valve  of  complete  female  carapace,  para- 
lectotype,  10.3628.  x  97. 

FIGS.  4,  7,  8.  Fastigatocythere  juglandica  (Jones).  Fig.  4,  external  view  right  valve, 
lectotype,  IN. 41947,  x  75.  Figs.  7,  8.  Normal  pore  canals  of  lectotype,  fig.  7  x  2,000, 
fig.  8  x  5,000.  Originally  described  as  Cythere  juglandica  Jones. 


Butt.  Br.  Mus.  not.  Hist.  (Geol.)  17,  8. 


PLATE    3 


PLATE   4 

FIG.  i.  Glyptocythere  guembeliana  (Jones).  Female  right  valve  and  lectotype  of 
Cytheridea  pulvinar  Jones  &  Sherborn.  1.1858.  x  85. 

FIG.  2.  Lophocythere  bradiana  (Jones).  Right  valve  of  complete  male  carapace,  1.1854. 
Lectotype  of  Cytheridea  craticula  Jones  &  Sherborn.  x  100. 

FIG.  3.  Lophocythere  bradiana  (Jones).  Male  left  valve,  paralectotype  of  C.  craticula 
Jones  &  Sherborn.  1.1867.  x  100. 

FIG.  4.  Lophocythere  bradiana  (Jones).  Ventral  view,  female  carapace,  paralectotype, 
10.3629  x  100. 

FIG.  5.  Bairdia  hilda  Jones.  Internal  view,  right  valve  of  paralectotype  of  Bairdia 
fullonica  Jones  &  Sherborn.  10.3554.  x  ?8- 

FIG.  6.  Bairdia  hilda  Jones.  External  view,  right  valve.  Lectotype  of  B.  fullonica 
Jones  &  Sherborn.  1.1873.  x  78. 

FIG.  7.  Micropneumatocythere  subconcentrica  (Jones).  Left  valve  of  lectotype 
IN. 43505.  Originally  described  as  Cythere  subconcentrica  Jones,  x  100. 

FIG.  8.  Terquemula  blakeana  (Jones).  Right  valve  originally  described  as  Cytheridea 
ransversiplicata  Jones  &  Sherborn.  1.1828.  x  100. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE   4 


PLATE   5 

FIG.  i.  Terquemula  blakeana  (Jones).  Female  left  valve,  lectotype,  IN. 41950.  Origi- 
nally described  as  Cythere  blakeana  Jones,  x  100. 

FIG.  2.  Glyptocythere  oscillum  (Jones  &  Sherborn).  Male  left  valve,  lectotype  of 
Cytheridea  striblita  Jones  &  Sherborn.  10.3943.  x  95. 

FIGS.  3,  7.  Cythereis  cf.  blanda  Kaye.  External  view,  left  valve,  IN. 43502  and  right 
valve  of  complete  carapace,  IN. 43491.  Both  specimens  originally  described  as  Cythere 
(Cythereis)  quadrilatera  (Roemer).  x  100. 

FIGS.  4-6.  Metacytheropteron  drupaceum  (Jones).  Left  valve  from  the  Blake  Collec- 
tion, 10.3977.  x  100,  and  left  and  right  views  of  complete  carapace,  holotype,  IN. 43498. 
X  85.  Originally  described  as  Cythere  drupacea  Jones. 

FIG.  8.  Cythere  ?  tenella  Jones.  Right  valve  of  complete  carapace,  holotype,  IN. 43492. 
X  100. 

FIG.  9.  Cytherella  fullonica  Jones  &  Sherborn.  Left  valve  of  paralectotype,  10.4004. 
x  105. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE    5 


PLATE   6 

FIG.   i.     Cytherella  fullonica  Jones  &  Sherborn.     Left  valve,  lectotype,  1.1857,      x  IO1- 

FIGS.  2,  3.  Cytherelloidea  catenulata  (Jones  &  Sherborn).  Right  valve,  paralectotype, 
1.1846,  x  100,  and  left  valve,  lectotype,  1.1876,  x  100.  Species  originally  described  as 
Cytherella  catenulata  Jones  &  Sherborn. 

FIGS.  4-7.  Cytherelloidea  refecta  (Jones  &  Sherborn).  Fig.  4,  female  left  valve,  10.3931 
paralectotype,  x  100.  Figs.  5,  6,  male  left  valve,  paralectotype,  10.3932,  fig.  5,  x  98,  fig.  6, 
anterior  ornamentation  of  same  specimen  x  5,000.  Fig.  7,  female  right  valve,  lectotype, 
1.1850,  x  95. 

FIG.  8.  Bairdia  sherborni  sp.  nov.  Left  valve,  complete  carapace,  holotype  10.3913. 
X  107. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE   6 


PLATE   7 

FIG.  i.  Bairdia  sherborni  sp.  nov.  Right  valve,  complete  carapace,  paratype,  10.3557. 
X  96. 

FIG.  2.  Paracypris  terraefullonicae  (Jones  &  Sherborn).  Right  valve  of  lectotype  of 
Macrocypris  horatiana  Jones  &  Sherborn.  1.1874.  x  I0°- 

FIG.  3.  Monoceratina  visceralis  (Jones  &  Sherborn).  Left  valve,  holotype,  1.1830, 
X  100.  Originally  described  as  Cytheridea  visceralis  Jones  &  Sherborn. 

FIG.  4.  Paracypris  terraefullonicae  (Jones  &  Sherborn).  Left  valve,  lectotype,  1.1875. 
Originally  described  as  Macrocypris  terrae-fullonicae  Jones  &  Sherborn.  x  100. 

FIG.  5.  Monoceratina  vulsa  (Jones  &  Sherborn).  Damaged  left  valve,  lectotype,  1.1842, 
X  100.  Originally  described  as  Cytheridea  vulsa  Jones  &  Sherborn. 

FIG.  6.  Oligocythereis  fullonica  (Jones  &  Sherborn).  Right  valve  of  complete  carapace, 
lectotype,  1.1871.  Originally  described  as  Cythereis  fullonica  Jones  &  Sherborn.  x  in. 

FIG.  7.  Schuleridea  (Eoschuleridea)  horatiana  (Jones  &  Sherborn).  Male  right  valve, 
lectotype,  1.1852.  Originally  described  as  Cytheridea  horatiana  Jones  &  Sherborn.  x  56. 

FIG.  8.  Asciocy there  obovata  (Jones  &  Sherborn).  Right  valve  of  complete  carapace, 
paralectotype,  10.3938,  x  100.  Originally  described  as  Cytheridea  obovata  Jones  &  Sherborn. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE    7 


PLATE   8 

FIG.  i.     Eocytheridea  sp.     Right  valve,  10.3906,  x  85. 

FIGS.  2,  7.  Asciocythere  obovata  (Jones  &  Sherborn).  Left  valve  of  complete  carapace, 
10.3939,  x  100  and  part  of  shell  surface  showing  normal  pore  canals,  x  650. 

FIG.  3.  Asciocythere  obovata  (Jones  &  Sherborn).  Right  valve,  lectotype,  1.1836. 
X  100. 

FIGS.  4-6.  Praeschuleridea  subtrigona  (Jones  &  Sherborn).  Male  right  valve,  lectotype, 
1.1838,  x  98;  right  valve,  female  carapace,  paralectotype,  10.3937,  x  100  and  left  valve,  male 
carapace,  paralectotype,  10.3936.  x  100. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE   8 


PLATE   9 

FIGS.  1-8.  Hadrocytheridea  dolabra  (Jones  &  Sherborn).  Figs,  i,  4.  External  and 
internal  views,  female  right  valve,  lectotype,  1.1851,  x  100.  (Lectotype  of  Cytheridea  dolabra 
Jones  &  Sherborn.)  Fig.  2.  Female  left  valve  Cytheridea  pentagonalis  of  Jones  &  Sherborn), 
10.3929,  x  100.  Fig.  3.  Internal  view  male  right  valve  (Cytheridea  parallela  of  Jones  & 
Sherborn),  paralectotype,  10.3926,  x  100.  Fig.  5.  Male  left  valve,  paralectotype,  1.1844 
(lectotype  of  Cytheridea  puteolata  Jones  &  Sherborn),  x  100.  Fig.  6.  Female  left  valve, 
paralectotype,  1. 1866  (holotype  of  Cytheridea  pentagonalis  Jones  &  Sherborn),  x  100.  Fig.  7. 
Male  right  valve,  complete  carapace,  paralectotype,  1.1859  (lectotype  of  Cytheridea  parallela 
Jones  &  Sherborn),  x  100.  Fig.  8.  Right  valve,  possibly  juvenile  instar  of  H.  dolabra  (lectotype 
of  Cytheridea  ignobilis  Jones  &  Sherborn),  1. 1868,  x  100. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


GEOL.   17,  8. 


PLATE    10 

FIG.  i.  Hadrocytheridea  dolabra  (Jones  &  Sherborn),  Dorsal  view,  male  carapace, 
paralectotype,  1.1859  (lectotype  of  Cytheridea  parallela  Jones  &  Sherborn).  x  100. 

FIGS.  2,  7,  8.  Caytonidea  terraefullonicae  (Jones  &  Sherborn).  Dorsal  view,  female 
carapace,  paralectotype,  10.3920,  x  100;  female  left  valve,  lectotype,  1. 1869,  x  100  and  female 
right  valve,  paralectotype,  10.3919,  x  100.  Species  originally  described  as  Cytheridea  terrae- 
fullonicae Jones  &  Sherborn. 

FIGS.  3-6.  Rectocy there  sugillata  (Jones  &  Sherborn).  Figs.  3,  6,  external  and  internal 
view,  left  valve  lectotype,  1.1855,  X  100.  Figs.  4,  5,  external  and  internal  view,  right  valve 
paralectotype,  10.3930,  x  100.  Species  originally  described  as  Cytheridea  sugillata  Jones  & 
Sherborn. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE    10 


PLATE    ii 

FIGS.  1-3.  Caytonidea  terraefullonicae  (Jones  &  Sherborn).  Internal  view,  female  left 
valve,  lectotype,  1. 1869,  x  100;  internal  view  x  100  and  surface  ornamentation  x  750  of 
female  right  valve,  paralectotype,  10.3919. 

FIGS.  4-6.  Acanthocythere  sphaerulata  (Jones  &  Sherborn).  Right  valve,  complete 
female  carapace  x  100  and  enlargement  of  surface  ornamentation  x  2,800  and  x  500.  Holo- 
type,  1.1835.  Species  originally  described  as  Cythere  sphaerulata  Jones  &  Sherborn. 


Bull.  Br.  Mus.  not.  Hist.  (Geol.)  17,  8. 


PLATE    ii 


PLATE    12 

FIG.  i.  Glyptocythere  persica  (Jones  &  Sherborn) .  Right  valve,  holotype,  1.1834,  X  IO°- 
Originally  described  as  Cytheridea  persica  Jones  &  Sherborn. 

FIG.  2.  Glyptocythere  oscillum  (Jones  &  Sherborn).  Female  right  valve,  lectotype, 
1.1849  (holotype  of  Cy there  oscillum  Jones  &  Sherborn),  x  105. 

FIG.  3.  Fastigatocy there  juglandica  (Jones  &  Sherborn).  Female  left  valve,  1.1872, 
X  92.  Originally  described  as  Cythere  juglandica  var  major  by  Jones  &  Sherborn. 

FIGS.  4,  6.  Lophocy there  acutiplicata  (Jones  &  Sherborn).  Left  valve,  female  carapace, 
lectotype,  1.1863,  X  112  and  internal  view,  female  right  valve,  paralectotype,  1.1847,  x  103. 

FIG.  5.  Lophocythere  ostreata  (Jones  &  Sherborn).  Right  valve,  holotype,  1.1833,  x  85. 
Originally  described  as  Cytheridea  ostreata  Jones  &  Sherborn. 

FIG.  7.  Lophocythere  fulgurata  (Jones  &  Sherborn).  Right  valve,  lectotype,  1.1832, 
X  85.  Originally  described  as  Cytheridea  fulgurata  Jones  &  Sherborn. 

FIG.  8.  Lophocythere  septicostata  Bate.  Female  right  valve,  1.1843,  x  100.  Originally 
described  as  Cytheridea  bradiana  Jones  &  Sherborn. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE    12 


PLATE    13 

FIG.  i.  Micropneutnatocy there  limaciformis  (Jones  &  Sherborn).  Right  valve, 
holotype,  1.1831.  Originally  described  as  Cytheridea  limaciformis  Jones  &  Sherborn.  x  85. 

FIG.  2.  Progonocy there  stilla  Sylvester-Bradley.  External  view,  left  valve,  10.3657. 
X  114. 

FIG.  3.  Cytheridea  aequabilis  Jones  &  Sherborn.  Left  valve  of  complete  carapace, 
holotype,  1.1864.  x  80. 

FIG.  4.  Cythere  corrosa  Jones  &  Sherborn.  Right  valve  of  complete  carapace,  holotype, 
1.1865.  X  I0°- 

FIG.  5.     Cytheridea  coarctata  Jones  &  Sherborn.     Left  valve,  holotype,  1.1841.      x  100. 

FIG.  6.     Cytheridea  eminula  Jones  &  Sherborn.     Left  valve,  lectotype,  1.1839.      x  100. 

FIG.  7.  Cytheridea  spinifastigiata  Jones  &  Sherborn.  Left  valve,  holotype,  I.i86i. 
X  100. 

FIG.  8.  Cytheridea  punctiputeolata  Jones  &  Sherborn.  Right  valve,  holotype,  1.1856. 
x  85. 

FIG.  9.  Cytheridea  spinigyrata  Jones  &  Sherborn.  Right  valve,  holotype,  1. 1860. 
X  100. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE     13 


PLATE    14 

FIGS.  1,3.  Ektyphocyt here  parva  Oertli.  Left  valve,  male  carapace,  10.3983,  x  115  and 
right  valve,  female  carapace,  10.3982,  x  114. 

FIGS.  2,  4,  9.  Hekistocy there  venosa  sp.  nov.  Left  valve,  paratype,  10.3996,  x  195; 
right  valve  of  complete  carapace,  holotype,  10.3995,  x  190,  and  internal  view,  left  valve, 
paratype,  10.4012,  x  150. 

FIG.  5.     Platycythere  sp.     Right  valve  of  complete  carapace,  10.3992,  x  no. 

FIG.  6.  Looneyella  monticula  (Jones).  Left  valve  of  complete  carapace,  holotype, 
1.5872,  from  the  Cretaceous  Bear  River  Formation,  Wyoming,  x  97. 

FIGS.  7,  8.  Looneyella  subtilis  Oertli.  Left  valve  of  complete  male  carapace,  10.3957, 
x  144,  and  right  valve  of  complete  male  carapace,  10.3956,  x  144. 


Bull.  BY.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE    14 


PLATE    15 

FIG.   i.     Or thonotacy there  sp.  nov.     Right  valve  of  complete  carapace,  10.3993,    x  144. 

FIGS.  2,  8.  Parariscus  bathonicus  Oertli.  Left  valve  of  complete  carapace,  10.3959, 
X  144  and  dorsal  view  of  complete  carapace,  10.3860,  x  144. 

FIG.  3.  Cytheridea  retorrida  Jones  &  Sherborn.  Left  valve  of  complete  carapace, 
10.3923.  x  108. 

FIGS.  4,  5,  9,  10.  Paracytheridea  ?  blakei  sp.  nov.  Fig.  4,  muscle  scars  of  paratype, 
10.4015,  as  viewed  externally,  x  250;  fig.  5,  dorsal  view,  complete  carapace,  paratype,  10.4016, 
X  200;  fig.  9,  right  valve,  complete  carapace,  holotype,  10.4001,  x  200  and  fig.  10,  left  valve, 
complete  carapace,  paratype,  10.4014,  x  200. 

FIG.  6.     Cytherura  mediojurassica  sp.  nov.     Dorsal  view  of  complete  carapace,    x  200. 

FIG.  7.  Hekistocy there  venosa  sp.  nov.  Dorsal  view  of  complete  carapace,  holotype, 
10.3995-  X  190. 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE    15 


PLATE    16 

FIGS,  i,  2.  Cytherura  tnediojurassica  sp.  nov.  Right  valve  of  complete  carapace, 
holotype^  10.4121,  x  190  and  left  valve  of  complete  carapace,  paratype,  10.4122,  x  195. 

FIG.  3.     Trachycythere  sp.     Left  valve,  10.3966,  x  114. 

FIGS.  4,  9.  Polycope  fungosa  sp.  nov.  Left  valve,  juvenile  carapace,  paratype,  10.3953, 
X  112,  and  left  valve  of  complete  adult  carapace,  holotype,  10.3951,  x  112. 

FIGS.  5-8.  Cytherura  bathonica  sp.  nov.  Fig.  5,  left  valve  of  complete  carapace,  para- 
type, 10.4018,  x  195;  fig.  8,  ornamentation  of  same  specimen,  x  1,200;  fig.  6,  dorsal  view  of 
complete  carapace,  paratype,  10.4019,  x  194;  fig.  7,  right  valve  of  complete  carapace,  holotype, 
10.4017,  x  195- 


Bull.  Br.  Mus.  nat.  Hist.  (Geol.)  17,  8. 


PLATE  16 


INDEX   TO   VOLUME   XVII 

New  taxonomic  names  and  the  page  numbers  of  the  principal  references  are  printed  in  Bold  type.  An 

asterisk  (*)  indicates  a  figure. 


Acanthinocyathidae  306,  372 
Acanthinocyathus  306,  308 

apertus  306,  307*-308;  PL  2,  figs.  1-4 

transiens  306 
Acanthistius  87 
Acanthocyathus  306 

apertus  306,  307 
Acanthocythere  411 

sphaerulata  411;  PL  u,  figs.  4-6 
Acanthuroidei  87,  92 
Achomosphaera  174 

ramulifera  174-175;  PL  10,  fig.  7 

reticulata  175 

sagena  175 
Acritarcha  107 
Acrogaster  73,  75 

brevicostatus  73 

daviesi  73 

heckeli  73* 

parvus  73 

Acrognathus  82,  83,  98 
Acroporidae  13 
Acrostichopteris  ruffordi  238 
Adnatosphaeridiaceae  171 
Adnatosphaeridium  171,  172 

chonetum  171-172;  PL  10,  figs,  n,  12 
Agastrocyathus  342 
Aipichthyidae  62,  92,  97,  100 
Aipichthys  49,  62-63,  65,  66,  84,  85,  87,  88,  93,  94, 

95.  97.  98,  99 

minor  63,  64*,  65* 

nuchalis  63 

pretiosus  62 

velifer  63,  66,  99 
Ajacicyathacea  372 
Ajacicyathida  305 
Ajacicyathidae  308 
Ajacicyathus  310,  345,  369 

walliseri  312,  314 

(Ajacicyathellus)  310 
Alataucyathinae  338,  340 
Alataucyathus  340,  369,  372 
Alataucyathidae  338 

(Urcyathus)  334 

jaroschevitshi  340 


(Alataucyathus)  338,  340 
(Anaptyctocyathus)  299,  372 

cribripora  340-341,  342,  372;  PL  n,  figs,  i,  3 

excellenthis  340 

flabellus  299,  340,  341-342,  372;  PL  10,  fig.  i 

minimiporus  340 

verschkhovskajae  340 
Albula  54 
Alectis  98 
Alepocephalus  84 
Alethopteris  goepperti  228 
Algites  221 

catenelloides  223 

valdensis  221 
Allothrissops  84,  85,  91 
Alphacyathus  305,  310,  369 

annularis  305 

cf.  annularis  305-306;  PL  i,  fig.  2 
Alveolites  heteropora  12 
Amphiblestrella  20 

elegans  20-21,  41;  PL  4,  figs.  1-3 
Amphiblestrum  21 

elegans  20 
Anabantidae  95 
Anabantoidei  89,  93 
Anasca  13 
Anemia  232 

adiantifolia  232 

cuneata  232 

Anomalopidae  68,  95,  96 
Anomozamites  schaumburgense  242 
Anoplogaster  68 
Anoplogasteridae  68,  99 
Antigonia  94,  96 
Aptea  polymorpha  171 

cf.  polymorpha  170,  171 
Apteodinium  130 

granulatum  121,  130-131;  PL  3,  figs.  5,  6 
Archaeocyatha  342,  359 
Archaeocyathellus  308,  310 

(Stapicyathus)  308 

stapipora  308,  309,  369;  PL  i,  fig.  5 

yukonensis  308 
Archaeocyathidae  346 
Archaeocyathus  344,  346,  372 


440 


INDEX 


acutus  312 

atlanticus  345*,  346 

dissepimentalis  358,  359 

floreus  310 

ichnusae  318 

infundibulum  318 

irregularis  348 

magnipora  311 

retesepta  364,  366 

retevallum  314,  315 

retezona  316 

robustus  311 

stapipora  308,  309 

subacutus  312 

trachealis  318 

tracheodentatus  318,  319 

tubavallum  318 

(Protocyathus)  stapipora  309 
Archaeolynthus  302 

porosus  303 

Archaeopharetra  312,  342 
Ardrossacyathus  354 
Areoligeraceae  166 
Argentina  54,  86 
Arripidae  87 
Arripis  87 
Asciocythere  404 

acuminata  404 

lacunosa  404 

o&owato  404-405*;  PI.  7,  fig.  8;  PI  8,  figs.  2,  3,  7 
Ascodinium  hialinium  no 
Ascophora  13 
Aspidistes  235 

sewardi  235,  236-237*;  PI.  6,  figs.  3,  4 

thomasi  235,  236 
Aspidium  235 
Asterocyathus  325 
Asterotumulus  338 
Aulacephalodon  286 
Aulolepididae  51,  83 
Aulolepis  53-54,  55,  63,  83,  84,  86,  87,  93,  98,  99 

typus  52*,  53* 
Aulopodidae  81 
Aulopus  54,  68,  81,  83,  84,  86,  88,  93,  98 

Bactrellaria  24 

rugica  24-25,  41;  PI.  2,  figs.  9-14 
Bairdia  383,  397 

fullonica  383,  397;  PI.  4,  figs.  5,  6 

harrisiana  385 

AiWa  383,  384,  385,  397-398;  PI.  i,  figs.  5,  6; 
PI.  4,  figs.  5,  6 

juddiana  383-384,  395;  PI.  i,  fig.  7 

jurassica  384-385;  PI.  2,  figs,  i,  2 
tennis  384,  385;  PI.  2,  fig.  2 

sherborni  398;  PI.  6,  fig.  8;  PI.  7,  fig.  i 

trigonalis  386,  395;  PI.  2,  fig.  7 
Bairdiacea  383,  397 
Biardiidae  383,  397 


Baltisphaeridium  armatum  153 

bifidum  165 

/ercuf  159 

longifurcatum  158 

multifurcatum  152 

pseudohystrichodinium  163 
Barroisina  trifossata  37,  38 
Barysoma  290 
Bathy master  91 
Bathysoma  49,  78-81,  96,  97,  100 

lutkeni  78,  79*,  81,  97 
Beaufort  series  291 
Becklesia  244 

anomala  244-245*,  248 

sulcata  247*-248,  249;  PI.  2,  fig.  6 
Bedfordcyathus  299,  355,  359 

irregularis  359 
Beisselina  39,  40 

flabellata  40 

5/nato  39-40,  41;  PI.  8,  figs.  3,  4 
Beisselinopsis  40 

flabellata  40,  41;  PI.  8,  figs,  i,  2 

marginata  40 
Belontiidae  95 
Bennettitales  211 
Berenicea  15 

sp.  15,  41 

Berycidae  68,  88,  94,  95,  96,  99 
Beryciformes  49,  56,  81,  83,  86,  87,  88,  92,  95,  96, 

97.  98,  99 
Berycoidei  49,  63,  67-69,  82,  84,  86,  87,  92,  93,  94, 

95.  96,  97.  99,  ioo 
Berycopsis  57-58,  59,  92 

elegans  57*,  58 

geramanus  58 
Beryx  94 
Biflustra  20,  22 

meudonensis  26 

radula  21 

simplex  20 

tesselata  25,  26 

unipora  19 
Brama  87 
Bramidae  87 
Branchiostegidae  87 
Branchiostegus  87 

Bronchocyathidae  317-318,  319,  372 
Bronchocyathus  317,  318,  319,  372 

dentatus  321 

trachealis  318 

tracheodentatus  319 
Brycon  84 

Bryophyta  210,  211,  218 
?  Bryophyte  leaf  218,  221,  222* 
Bythocypris  winwoodiana  395 
Bythocytheridae  399 

Callaiosphaeridium  151 

asymmetricum  152;  PI.  6,  fig.  6 


INDEX 


441 


Cambrocyathellus  354 
Cambrocyathus  354,  356,  370 

profundus  357*,  358 
Camptocythere  lincolnensis  410 
Canningia  140 

rotundata  141 
Cannosphaeropsis  172 

choneta  171 
Caproberyx  76-77,  78,  94,  95 

pharsus  77 

polydesmus  77 

superbus  76* 
Caproidae  93,  96 
Capros  96 
Carangidae  92 
Carpodinium  129 

obliquicostatum  129;  PI.  i,  figs.  3,  4 
Cassandra  82 
Gassiculosphaeridia  141 

reticulata  141,  142,;  PI.  3,  fig.  7;  PI.  4,  fig.  3 
Castanopora  8,  12 

guascoi  8 

jurassica  7-8,  n,  14,  19,  31,  37,  41,  PI.  7,  figs. 

4-7 
Catenella  223 

opuntia  223 
Cave  Sandstone  293 
Caytonidea  410,  424,  426 

faveolata  410,  411 

terraefullonicae  410-411;  PI.   10,  figs.  2,  7,  8; 

PI.  ii,  figs.  1-3 
Cellaria  9 

smithi  9 
Cellepora  9 

complanata  29 

davaiacensis  9 

polythele  9 

ricata  30 

(Dermatopora)  faujasi  8 

(Discopora)  subgranulata  30 

koninckiana  6 
Centrarchidae  92 
Centroberyx  affinis  68 
Centropomidae  87,  92 
Centropomus  87 
Ceratocorys  136 

veligera  136 
Chaetodon  87 
Chaetodontidae  87,  92 
Chanares  Formation  281 
Chanaria  281,  284 
Channiformes  89,  93,  95,  96 
Chanos  54 
Char  a  knowltoni  214,  215 

medicaginula  217 
Characeae  217 
Charales  214,  217 
Charixa  vennensis  13 
Cheilanthites  gopperti  229 


Cheilostomata  3,  4,  9,  12,  13,  14 
Chelmo  87 

Chinle  Formation  290 
Chlamydophorella  156 

wyei  155 

Chlorophthalmus  81 
Chlorophyceae  214 
Chondrus  crispus  223 
Chouberticyathus  342 
Chrysomonadinae  ?  140 
Chytroeisphaeridia  140 

chytroeides  141 

euteiches  141;  PI.  3,  figs.  8,  9 

sp.  141 
Cichlidae  87 
Girconitella  214-215 

knowltoni  215-216*-217;  PI.  i,  figs.  1-5 
Circulodinium  deflandrei  166 
Cirrhitidae  87 
Cirrhitus  87 
Cisternifera  9 
Cladocopida  427 
Cladocopina  427 
Cladograpsus  capillaris  185 

linearis  185,  186 
Cladophlebis  albertsii  237 

browniana  238 

dunkeri  238 

longipennis  237,  238 

(Klukia)  dunkeri  238 
Claruscyathus  363 
Cleistosphaeridium  152,  157 

ancoriferum  155 

armatum  no,  153-154;  PI.  8,  figs,  i,  2,  12 

heteracanthum  152;  PI.  7,  fig.  8 

huguonioti  109,  155-156;  PI.  7,  fig.  10 

var.  pertusum  156-157;  PI.  7,  figs.  6,  7,  9 

multifurcatum  no,  152-153;  PI.  8,  figs.  7,  10 

polypes  154,  155;  PI.  6,  figs.  7,  8 

var.  clavulum  154-155;  PI.  6,  figs.  9,  10 

?  aciculare  158;  PI.  6,  figs,  u,  12 

?  flexuosum  157;  PI.  7,  fig.  4 

?  parvum  157-158;  PI.  7,  figs,  n,  12 
Climacograptus  192 

minimus  191-193;  PI.  4,  fig.  3 

minutus  191*,  192*,  193-194;  PI.  4,  figs,  i,  2 

scalaris  miserabilis  193 ;  PI.  4,  fig.  2 

tangshanensis  linearis  193 

spp.  1 88 

Clonograptus  204 
Clupavidae  86,  98 
Clupavus  84,  85 
Coilostega  13,  22 
Cometodinium  obscurum  137 
Coniferales  211 
Copleicyathus  344 

confertus  349 
Coronifera  161 

oceanica  161,  162;  PI.  8,  figs.  8,  n 


442 


INDEX 


Corynoides  calicularis  195 
Coscinocyathus  325,  329,  330,  369 

annulatus  337 

cellularis  330 

cribripora  340 

equivallum  326 

papillatus  334 

papillipora  338,  339 

peter  si  331 

quadratus  328 

ye/i/er  332 

rhyacoensis  329 

rugosus  325 

textilis  329 

unilinearis  330;  PI.  13,  fig.  4 
Coscinopleura  29 

lamourouxi  29 

sp.  29,  41 
Coscinoptycta  325,  330 

convoluta  330 
Coscionteichus  325 
Cribroperidinium  121,  124,  125,  128 

edwardsi  127,  128 

intricatum     109,     no,    125-126*-127*-128; 
PI.  2,  figs.  1-3 

muderongensis  127,  128 

orthoceras  121,  127,  128-129,  134* 

sepimentum  125 
Crisisina  17 

carinata  17,  41;  PI.  i,  figs.  4,  5 
Cristivomer  84 
Cryptograptus  194 

tricornis  191*,  194-195;  PI.  4,  figs.  4,  5,  6 

schaeferi  195 
Cryptostomata  14 
Ctenostomata  14 
Ctenothrissa  54-56,  63,  81,  83,  84,  86,  87,  93,  98, 

99 

microcephala  55* 

radians  55 

signifer  54*.  55 

vexillifer  54 
Ctenothrissidae  54,  83 

Ctenothrissiformes  49,  50-51,  83,  86,  98,  99 
Cyathocricus  299,  318-319,  369,  372 

annulispinosus  318,  320,  322,  369 

dentatus  318,  319,  321*-322;  PI.  6,  figs.  1-3 

tracheodentatus  318,  319-320*,  322;  PI.  5,  figs. 

4,  5 ;  PI.  6,  fig.  4 
Cycadales  210,  211,  242 
C>»fls  246 

revoluta  246,  248 
Cyclocyathellidae  317 
Cyclocyathus  318,  319,  372 
Cyclonephelium  166,  168,  171 

compactum  167 

distinctum  166-167,  169*,  170;  PI.  n,  figs.  6,  7, 
8,  10 

eisenacki  166,  169*,  170-171;  PI.  8,  figs.  3,  4; 


PI.  9,  fig-  4 

membraniphorum  166,  167-168;  PI.  u,  fig.  9 
paucispinum  166,  170;  PI.  9,  figs,  i,  2 
vannophorum  166, 168,  169*,  170;  PI.  9,  fig.  3 

PI.  n,  figs,  n,  12 
uttadalicum  168 
Cyclostomata  9,  15 
Cymatiosphaera  cingulata  173 
Cynidiognathus  longiceps  291 
Cynognathus  Zone  257,  285,  290,  291,  292,  293 
Cypridacea  384,  399 
Cyrtograpsus  murchisoni  202 
Cyrtograptus  centtrifugus  202,  203 
murchisoni  202-2O3*,  PI.  5,  figs  8,  9 
bohemicus  202 
crassiusculus  203 
rigidus  203 
Cytheracea  385,  399 
Cythere  394 
blakeana  393 
bradiana  390,  391 
corrosa  424,  426  ;  PI.  13,  fig.  4 
drupacea  393,  394,  427 
guembeliana  388,  412 
juglandice  389 
major  389,  412 
minor  395 
monticula  431 

oscillum  412,  413;  PI.  12,  fig.  2 
schwageriana  387;  PI.  2,  fig.  6. 
speciosa  395 
sphaerulata  411 
subconcentrica  392 
?  teneWo  394;  PI.  5,  fig.  8 
trapezioides  395 
walfordiana  395 
(Cythereis)  quadrilatera  394 
Cythereis  394 
blanda  394 

cf.  Wawrfa  394;  PI.  5,  figs.  3,  7 
fullonica  401 
walfordiana  395 
Cytherella  380,  381,  395 
catenulata  396 

fullonica  395;  PL  5,  fig.  9;  PI-  6,  fig.  i 
jugosa  381 
subovate  380,  381 
suprajurassica  381 
symmetrica  380-381;  PI.  i,  figs,  i,  2 
Cytherellidae  380,  395 
Cytherelloidea  381,  396 

catenulata  396;  PI.  6,  figs.  2,  3 
eastfieldensis  382 
;«£osa  381-382*;  PI.  i,  figs.  3,  4 
paraweberi  382* 
ye/ec/a  396-397;  PI.  6,  figs  4-7 
weberi  382 

Cytheridae  acutiplicata  416 
aequabilis  424;  PI.  13,  fig.  3 


INDEX 


443 


bicarinata  395,  420,  421 

blakeana  393,  423 

bradiana  391*,  418 

coarctata  424;  PL  13,  fig.  5 

craticula  390,  391*,  392,  418;  PL  4,  figs.  2,  3 

dolabra  406,  407,  408 

egregia  393.  395 

eminula  425;  PL  13,  fig.  6 

eximia  395 

fulgarata  419 

horatiana  402 

ignobilis  407,  408,  409;  PL  9,  fig.  8 

limaciformis  422 

obovata  404 

ostreata  420 

parallels,  407,  408;  PL  9,  fig.  3;  PL  10,  fig.  i 

pentagonalis  407,  408;  PL  9,  figs.  2,  6 

persica  415 

politula  395 

pulvinar  388,  412;  PL  4,  fig.  i 

punctiputeolata  409,  425;  PL  13,  fig.  8 

£«ra  395 

puteolata  407,  408;  PL  9,  fig.  5 

refecta  396 

renoides  395 

retorrida  394,  425*-426*;  PL  15,  fig.  3 

rugifera  395 

sedata  395 

spinifastigiata  426;  PL  13,  fig.  7 

spinigyrata  426-427;  PL  13,  fig.  9 

striblita  412,  413;  PL  5,  fig.  2 

subeminula  395 

subperforata  385;  PL  2,  fig.  4 

subtrigona  405 

sugillata  409 

terraefullonicae  410 

transversiplicata  393,  423 

trapezoidalis  388,  412 

ventrosa  395 

visceralis  399 

vufca  400 

winwoodiana  395 
Cytherideidae  406 
Cytherideinae  406 
Cytheropteron  jonesanum  393,  394 
Cytherura  434 

bathonica  434;  PL  16,  figs.  5-8 

?  liesbergensis  435 

mediojurassica    434-435;     PL     15,    fig.    6; 
PL  1 6,  figs,  i,  2 

striata  434 

Cytheruridae  393,  431 
Cyttus  96 

Dactylopteriformes  96 
Daptocephalus  286 
Decurtaria  37 

allecta  38 

cornuta  37,  38 


cf.  cornuta  37-38,  41;  PL  7,  figs,  i,  2 
Deflandrea  no 
Dendrocyathus  354 
Dendrograptus  lentus  204 
Dermogenys  go 
Description  of  sections 

Compton  Bay,  Isle  of  Wight  112,  113* 

Devon  115,  117* 

Escalles  Borehole,  Pas  de  Calais  115,  118* 

Fetcham  Mill,  Surrey  in*,  112 

Hunstanton,  Norfolk  115,  116* 

Saskatchewan  115,  119* 

Speeton,  Yorkshire  112,  114* 

Texas  115 
Diademodon  259,  292 

polyphagus  292 

rhodesiensid  292 
Diastopora  9 

lamellosa  5 

ranvilliana  9 
Dicellograpsus  elegans  187 

moffatensis  189 
Dicellograptus  188,  190 

anceps  189 

duvaricatus  189,  190 

elegans  187-188*,  190;  PL  3,  fig.  i 

moffatensis  188*,  189-190;  PL  3,  figs.  5,  6 
alabamensis  190 

morrisii  188,  190 

pumilus  1 88 

vagus  190 
Dicentrarchus  87 
Dichopteris  delicatula  236 

sp.  cf.  £>.  laevigata  240 
Dicranograptus  190,  191 

brevicaulis  190 

clingani  190-191*;  PL  3,  figs.  2,  3,  4 
Dictyocyathidae  342,  344 
Dictyocyathus  342,  344,  370 

annularis  305 

graphicus  346,  348 
Dictyophyllum  roemeri  228 
Dictyopyxidia  141 
Didymograpsus  elegans  187 
Didymograpsus  moffatensis  189 
Dimorphostylus  32,  33 

tetrastichus  32,  33 
Dinanomodon  286 
Diniferophycidae  120 
Dinodontosaurus  281,  284 

turpior  283,  284 
Dinophyceae  120 
Dinopterygidae  62,  66-67,  92,  98 
Dinopterygoidei  62,  86,  87,  88,  97,  99 
Dinopteryx  67,  68,  84,  87,  88,  93,  95,  99 

spinosus  66 
Diphyes  161 
Diplograpsus  foliaceus  194 

minimus  191,  192*,  193 


444  INDEX 

minutus  192,  193 

pristis  192,  193 

tricornis  194 
Diplograptidae  194 
Diplopholeos  27 
Dipteridaceae  228 
Diretmidae  68,  88,  99 
Diretmus  68,  88,  99 

Diversograptus  ?  capillaris  198,  199*.  200 
Doidyx  142 

Dokidocyathidae  305,  372 
Dokidocyathus  305 
Drepane  87 

Esienackia  eras  sit  abulata  136 
Ektyphocy there  430 

parva  430;  PL  14,  figs,  i,  3 
Ellipsoidictyum  141 

cinctum  141 
Ellipsodinium  129-130 

rugulosum  130,  139*;  PL  3,  fig.  I 
Eops  54,  63,  83,  84,  85,  86,  88,  98 
Emydops  285 
Endothiodon  Zone  258 
Entalophora  g,  16 

benedeniana  16,  41 

madreporacea  var.  benedeni  16 
Eocytheridea  406 

faveolata  406 

sp.  406;  PL  8,  fig.  i 
Epelidosphaeridia  142 

spinosa  143;  PL  3,  figs.  10-12 
Ephippidae  87,  98 
Ephippus  87 
Equisetales  211,  227 
Equisetites  227-228 

/ye//i  227 

Er-ma-ying  Formation  290 
Erismacoscinidae  325 
Erismacoscinus  325,  332,  369 

cancellatus  332 

cellularis  330-331;  PL  8,  figs,  i,  4 

equivallum  326-327,  328,  331;  PL  7,  figs.  1-3,  5 

marocanus  325 

papillatus  334 

petersi  331-332,  333;  PL  10,  figs.  4,  5 

quadratus  328-329;  PL  7,  fig.  4;  PL  8,  fig.  5 

retifer  332-333;  PL  10,  figs.  2,  3 

rugosus  325-326,  328,  331;  PL  8,  figs.  2,  3 

textilis  329-330;  PL  9,  figs.  1-4 

(Pluralicoscinus]  325 
Erythrosuchus  291 
Escarpment  Grit  258,  293 
Escharidae  9 
Eschar a  9,  24 

bellona  35 

caecilia  36 

cepha  34 

gaimardi  24 


34 
40 

ranvilliana  9 

striata  39 

Escharifora  flabellata  40 
Escharinella  simplex  30 
Ethmocoscinus  338-339,  369 

papillipora  339;  PI.  3,  fig.  3 
Ethmocyathidae  299,  322-323,  372 
Ethmocyathus  322,  323,  369,  372 

lineatus  323-324*;  PI.  5,  figs.  1-3 
Ethmophyllidae  317 
Ethmophyllum  313,  314,  370 

dentatum  318,  319,  321 

flexum  315,  316 

poletevae  315 

vermiculatum  315 
Euparkeria  291 
Exellia  97,  98,  100 
Exochosphaeridium  131,  162,  166 

"  hirsutum  "  group  164,  165 

phragmites  163;  PI.  7,  fig.  5 

pseudohystrichodinium  163-164;  PI.  n,  figs.  4,  5 

spinosum  166 

var.  deflandrei  165,  166 

striolatum  162,  163,  164,  165 

var.  truncatum  no,  164-166;  PI.  7,  figs.  1-3 

(Hystrichosphaeridium)  cf.  hirsutum  165 

Fastigatocythere  389,  412 

juglandica  389,  412;  PI.  3,  figs.  4,  7,  8;  PI.  12, 

fig.  3 

Filicales  228 

Flindersicyathidae  344,  372 
Flindersicyathus  344-346,  350,  370,  371,  372 

(Flindersicyathus)  344,  345,  348,  349,  370,  371, 

372 

circliporus  346 
contractus  346 
decipiens  344,  348 

graphicus  346-347*-348,  363;  PI.  12,  figs.  3,  4,  5 
irregularis  346,  348-349;  PI.  14,  fig.  4 
latiloculatus  346 
macdonnelli  346 

mo/or  346,  349-350;  PL  14,  fig.  2 
multifidus  346 
yete  346,  348 
simplex  346 

speciosus  346,  350;  PI.  14,  figs,  i,  3 
tabulatus  346 
(Pycnoidocyathus)  344,  345,  350-351,  352,  354, 

37°.  3?i.  372 

maximipora  346,  352-353;  PI.  15,  fig.  3 
parvulus  346 
ptychophragma  346 
simplex  352;  PL  15,  fig.  2 
synapticulosus  346,  350,  351;  PL  15,  fig.  i 
vicinisepta  346,  353-354;  PL  15,  fig.  4 
Floridina  31 


INDEX 


445 


Floridinella  31 

scutata  31;  PI.  4,  fig.  7 
"  Flustra  "  flabelliformis  4-5 
Flustrella  20 

baculina  20 

convexa  25,  26 

irregularis  21 

marginata  25 

simplex  19 

Flustrellaria  incrassata  23 
Fromeaceae  140 
Fromea  140 

amphora  140;  PI.  3,  figs.  2,  3 
Frurionella  38,  39 

europaea  39;  PI.  8,  fig.  8 

fertilis  3,  n,  38-39,  41;  PI.  8,  figs.  5-7 

Gabrielsocyathus  363 

deasensis  364 

gabrielsensis  364 

poolensis  364 
Gasterosteiformes  96 
Gaterin  87 
Geniculicyathus  325 
Gephyroberyx  68 
Gesoriacula  431 
Gibberichthyidae  68,  96 
Gibberichthys  68 
Ginkgoales  210,  211,  248 
Gleichenites  cycadina  238 
Glossograptus  194 
Glyptocythere  388,  389,  412 

costata  416 

guembeliana  388-389,  412,  416;  PI.  3,  figs,  i,  2; 

PI-  4,  fig-  i 
juglandica  389 
oscillum  412-415  [414*];  PI.  5,  fig.  2;  PI.  12, 

fig.  2 

persica  415*-416;  PI.  12,  fig.  i 
rudimenta  413,  414*,  415 
tuberodentina  416 
Gnathoberyx  74-75 

stigmosus  74 

Gonyaulacysta  120,  121,  125,  128,  132 
ambigua  124 
cf.  ambigua  120 
apionis  127,  128 
cassidata  120 

delicata  123*-124;  PI.  i,  figs.  7,  8 
diaphinis  127,  128 
edwardsi  125 
exilicristata  no,  121,  122*,  125;  PI.  i,  figs. 

i,  2 

fetchamensis  120-121 
muderongensis  125 
obscura  128 
orthoceras  125 
wetzeli  128 
120 


Sp.  A.  122*,  124-5;  PI.  i,  figs.  9,  10 
Gonyaulacystaceae  120 
Gonyaulax  125 

edwardsi  128 

helicoidea  subsp.  cassidata  120 

muderongensis  128 

orthoceras  109,  128 

sp.  no,  127 
Grammascosoecia  19 

dichotoma  19 
Grammicolepidae  96 
Graptolithus  becki  195 

clingani  195 

hisingeri  197 

intermedius  196 

millepeda  195 

nilssoni  197 

tenuis  197 
Gymnospermae  242 
Gyrogona  217 

medicaginula  217-218 

Hadrocytheridea  379,  406 

dolabra  407-409  [408*],  425;  PI.  9,  figs.  1-8; 

PI.  10,  fig.  i 
Hagenowinella  23 

cf.  incrassata  23-24,  41  ;  PI.  2,  figs.  1-3 
Hausmannia  228 

dichotoma  228;  PI.  3,  figs,  i,  2 

pelletieri  228 
Hekistocythere  379,  428,  430 

venosa  428-429*^430;   PI.   14,  figs.  2,  4,  9; 

PI-  15,  fig-  7 
Heliodinium  voigti  175 
Heniochus  87 
Hepaticae  218-219 
Hepaticites  219,  220,  221 

amauros  223 

arcuatus  220 

haiburnensis  220 

laevis  220 


224 

rosenkrantzi  223 

ruffordi  220,  223;  PI.  2,  figs.  4,  5 

zeilleri  219-220,  223;  PL  2,  figs.  1-3 
Heterocella  27 
Heterocrisina  16 

abbottii  16 

communis  16-17,  41 
Heteropora  reticulata  18 
Hexasphaera  asymmetrica  152 
Hiodon  84 
Hippothoa  9 

simplex  13 
Histiocysta  138 

palla  138-139*-140;  PI.  i,  figs.  5,  6 
Histiopterus  87 

Holocentridae  67,  68,  75-76,  77,  88,  94,  95,  99 
Holocentrus  68,  75,  76 


446 

ascensionis,  68,  76 
Homocytheridea  424 
Homonotichthys  60-61,  88,  94,  95,  99 

dorsalis  60,  6 1 

pulchellns  60*.  6 1 

rotundus  61 
Hoplopteryx  69-72,  73,  75,  94,  95 

antiquus  70 

gephyrognathus  70,  71 

lewesiensis  70*,  71*,  72 

lewisi  71 

macracanthus  70,  71 

siwws  70,  71 

spinulosus  71 

syriacus  71,  72 

Hoplostethus  68,  70,  72,  74,  75,  94 

mediterraneus  67*,  68,  69 
Howesia  291 

Hymenopteris  psilotoides  238 
Hynnis  98 
Hystrichodiniwn  175 

dasys  175-176;  PI.  10,  figs.  8,  9 

pulchrum  109 

uoi^rt  109,  175;  PI.  10,  figs.  6,  10 
Hystrichokolpoma  159 

/m?#  159-160;  PI.  9,  figs.  5,  6,  7 
Hystrichosphaeraceae  172 
Hystrichosphaera  no,  172 

cingulata  var.  cingulata  173 
var.  perforata  173 
var.  polygonalis  174 
var.  recticulata  173-174 

crassimurata  174 

crassipellis  174 

furcata  172,  173 

ramosa  var.  gracilis  172 

var.  multibrevis  173;  PI.  10,  figs.  3,  4 
var.  ramosa  172;  PI.  10,  figs,  i,  2,  5 
var.  reticulata  173 

cf.  rowosa  174 
Hystrichosphaeridiaceae  143 
Hystrichosphaeridium  143 

ancoriferum  155 

anthophorum  147 

armatum  153 

asterigerum  no 

asymwe^ncwm  152 

bowerbanki  145;  PI.  5,  fig.  9 

complex  146 

costatum  146 

deanei  144;  PI.  4,  fig.  I 

145-146;  PI.  4,  figs.  2,  6,  7 
159 

heteracanthum  152 

hirsutum  165 

huguonioti  155 

longifurcatum  no,  158 

mantelli  109,  145;  PI.  4,  fig.  9 

multifurcatum  152,  153 


INDEX 


parvispinum  161 
polytrichum  no 
pseudohystrichodinium  163 
radiculatum  144-145;  PI.  4,  fig.  8 
readei  144 
recurvatum  151,  154 
subsp.  polypes  154 
reniforme  148 
cf.  salpingophorum  109 
siphoniphorum  148 
spinosum  var.  deflandrei  166 
stellatum  144 
cf.  striolatum  no 

tubiferum  143-144;  PI.  5,  figs.  5,  8 
sp.  22.  1 10 
sp.  23.  in 

I chthyodectes  84 
Idmidronea  16 

macilenta  16,  41 
Idmonea  abbottii  16 

communis  16 

disticha  16 

lichenoides  18 

macilenta  16 

pseudodisticha  16 

ramosa  16 

(Tubigera)  antiqua  16 
Irregularia  342,  370 
I schigualastia  257,  285,  290 
Ischigualasto  Formation  290 

Kannemeyeria  285,  290,  291,  292 
Kannemeyeriidae  290 
Kansius  sternbergi  78 
Kawinga  Formation  293 
Kingori  Sandstone  293 
Kinkelinella  430 

tenuicostati  430 
Kirtonellinae  430 

Kistecephalus  Zone  258,  285,  286,  293 
Korojonia  dubiosa  109 
Korsogasteridae  68 
Kuhliidae  87 
Kuhlia  87,  91 
Kyphosus  87 
Kyphosidae  87,  92 

Labiostomella  14 

Lampridae  98 

Lampridiformes  49,  78,  81,  96,  97,  98,  100 

Lampridoidei  78,  97 

Lampris  78,  81,  97 

Lateolabrax  87 

Leckenbya  valdensis  238 

Lepidorbitoides  10 

Leptograptus  185,  187 

capillaris  185-186;  PI.  i,  figs,  i,  2 

flaccidus  186 


INDEX 


447 


arcuatus  186 
Leptolepis  84,  91,  98 

coryphaenoides  85 

dubia  85 

normandica  85 
Leptosomus  82 
Lethrinidae  87 
Lethrinus  87 
Lingulodinium  165 
Lisso&ery*  72,  73,  75,  94,  95 

dayi  72* 
Litosphaeridium  148 

siphoniphorum  148-149*-150;  PI.  6,  figs.  3,  4 
Lobotes  87 
Lobotidae  87 

Loculicyathus  (Loculicyathus)  310,  372 
Loculicyathus  (Loculicyathellus)  299,  310,  372 

/Zoyews  310-311;  PI.  3,  figs,  i,  2,  4 
Lonchopteris  mantelli  239 
Looneyella  431 

monticula  431;  PI.  14,  fig.  6 

quadrispina  431 

subtilis  431-432;  PI.  14,  figs,  7,  8 
Lophocythere  389-390,  416;  417,  418,  421 

acutiplicata  416-417;  PI.  I,  fig.  8;  PI.  12,  figs. 
4,6 

bradiana  390-391*-392,  416,  417,  418;  PI.  3, 
figs,  3,  5,  6;  PI.  4,  figs.  2,  3,  4 

fulgurata  419-420;  PI.  12,  fig.  7 

multicostata  392 

ostreata  390,  419,  420-421*;  PI.  12,  fig.  5 

scabra  421 

septicostata  391,  418;  PI.  12,  fig.  8 

sp.  418*-419 

Lower   Cambrian   archaeocyathid    Stratigraphy 
368-371,  372 

Australia  368,  369,  370,  371 

Morocco  369 

Sardinia  369,  370 

Siberian  platform  368,  369,  370 

Southern  Siberia  368,  369,  370 
Luangwa  drysdalli  259,  292 
Luciocephalus  89 
Lunulites  34 

sp.  ii,  34,  41 
Lutjanidae  87 
Lut janus  87 
Lycopodiales  224 
Lystrosaurus  285 
Lystrosaurus  Zone  285,  291,  293 

Macristiella  perlucens  51 
Macristiidae  51 
Macristium  50 

chavesi  50 
Macrocypris  bradiana  385;  PL  2,  fig.  3 

horatiana  399;  PI.  7,  fig.  2 

terraefullonica  399 
Madumabisa  Mudstone  293 


Malacostega  13,  21,  22 

Manda  Formation  290,  291,  292,  293 

Marchantiales  218,  220 

Marchantiolites  219 

Marchantites  218,  219,  220 

hallei  220 

zeilleri  219 
Matoniaceae  228 
Matonidium  228 

goepperti  228 
Medialuna  87 
Melamphaeidae  68 
Melamphaes  68 

Membranilarnax  cf.  pterospermoides  140 
Membranipora  14,  19 

bipunctata  8 

constricta  13;  PI.  7,  fig.  8 

elliptica  13 

fragilis  13 

genucia  19,  20 

jurassica  3,  4,  6,  7 

neocomiensis  13 

?  obliqua  13 

simplex  20 

unipora  11,  19-20,  41;  PI.  2,  figs.  7,  8 
Membraniporae  13,  20 
Mene  80,  81,  97,  98,  100 

maculata  80*,  81 

oblongus  80 

phosphaticus  97 

rhombeus  80 
Menidae  80,  100 
Mesosuchus  291 
Metacoscinidae  363,  367 
Metacoscinus  363-364,  370 

insigne  364 

reteseptatus  363,  364-366;  PI.  17,  figs.  1-4 
Metacyathidae  354 
Metacyathus  299,  355,  356 

irregularis  355,  359 

tey/on  355,  360 
Metacytheropteron  393,  427 

drupaceum  393-394;  PI.  5,  figs.  4-6 

sp-  393 
Metafungia  354,  358,  361-362,  370 

reticulata  361,  362-363;  PI.  18,  figs.  1-3 
Metaldetes  299,  348,  354,  355-356,  358,  370 

columbianus  356 

conicus  358,  359 

cylindricus  355,  356 

dissepimentalis  356,  358-359,  361 ;  PI.  16,  figs. 

i-3 

dissutus  356 
graphica  346,  348 

irregularis  356,  359-360,  361;  PI.  16,  fig.  4 
proteus  356,  358 
ramulosus  356 
solidus  356 
spiralis  358 


448  INDEX 

superbus  358 

taylori  358,  360-361;  PI.  13,  figs.  1-3 
Micrhystridium  ambiguum  109 

veligerum  136 

sp.  ?  138,  140 
Microdiniaceae  132 
Microdinium  132,  135,  136,  137 

?  crinitum  137;  PI.  2,  figs.  7,  8 

distinctum  133,  134*,  135;  PI.  2,  figs.  9-11 

echinatum  133 

irregular  e  136 

ornatum  132,  133 

cf.  ornatum  132-133,  134*;  PI.  4,  fig.  5 

setosum  133,  134*;  PI.  2,  fig.  4 

variospinum  134*,  135;  PI.  2,  figs.  5,  6 

veligerum  136-137;  PI.  3,  fig.  4;  PI.  4,  fig.  4 
Micropneumatocythere  392,  422 

limaciformis  422-423;  PI.  13,  fig.  i 

quadrata  422,  423 

subconcentrica  392;  PI.  4,  fig.  7 
Micropora  32,  33 

rw^'ca  32,  33 

subgranulata  30 

transversa  32-33,  41;  PI.  7,  figs.  9-12 
Microporidae  13,  33 
Mohria  232 
Molteno  Beds  291,  293 
Monocentridae  68,  69,  95,  96 
Monocentris  63,  68,  69,  87,  88,  89,  99 

japonicus  69* 
Monoceratina  399,  400 

ungulina  400 

visceralis  399-400;  PI.  7,  fig.  3 

im/sa  400-401 ;  PI.  7,  fig.  5 
Monocyathida  302 
Monocyathidae  302 
Monocyathus  302-303,  305,  369 

absolutus  302 

bilateralis  302 

contractus  302 

copulatus  302 

kuzneskii  302 

lebedevae  302 

irregularis  304 

macrospinosus  302 

mellifer  302-304;  PI.  i,  fig.  4 

nalivkini  302,  303 

operculatus  302 

partibus  303 

polar  is  303 

porosus  302,  303;  PI.  i,  fig.  3 

robustus  303 

sibiricus  303 

simplex  303 

sparsipora  303 

spinosus  303 

tenuimurus  303 

to#t  303 

unimurus  303 


Monodactylidae  87,  92 
Monodactylus  87 
Monograptus  198 

attenuatus  198,  200 

capillaris  198 

clingani  195-196;  PI.  5,  figs.  1-5 

communis  197 

gemmatus  198,  199*,  200 

intermedius  196*-198;  PI.  5,  fig.  6 

involutus  197 

spiralis  203 

sp.  (atazms  or  acinaces  type)  192 

(Rastrites)  gemmatus  198 
M or one  87 
Mugil  88,  92 
Multicavea  19 
Multicrescis  18,  19 

laxata  7,  n,  14,  18-19,  31,  41 ;  PI.  i,  figs.  6-8 
Musci  218 
Myctophidae  83,  99 

Myctophoidei  49,  50,  81-83,  86,  88,  98,  99 
Myctophum  81 
Myodocopida  427 
Myosaurus  285 
Myripristis  68,  75,  76,  77 

adustus  75* 

Naiadita  lanceolate  218 

Nandidae  89,  93 

Nathorstia  valdensis  238 

Nemagrapsus  capillaris  185 

Nematonotus  63,  68,  82,  83,  84,  85,  86,  88,  93,  98 

fcottae  82*,  83 

longispinus  82* 
Nemipteridae  87 
Nemipterus  87 
Neocyttus  96 
Neomiodon  221 
Neoscopelus  81 
Neothunnus  89 
Nilssonia  242,  244 

revoluta  244 

schaumburgensis  237*,  242-243*-244 
Nitella  215,  217 

flexilis  21 7 

hyalina  217 

polygyra  217 
Nitellaceae  217 
Nitellites  214 

sahnii  217 

sp. 

Ntawere  Formation  257,  258,  259,  262,  283,  284, 
286,  290,  292,  293 

Okulitchicyathus  354 
Oligocythereis  401 

fullonica  401-402;  PI.  7,  fig.  6 
Oligosphaeridium  146 

anthophorum  147-148;  PL  5,  figs.  1-3 


INDEX 


449 


complex  no,  146-147;  PI.  5,  figs.  6,  7 

perforatum  148 

prolixispinosum  147;  PI.  5,  fig.  4 

pulcherrimum  148 

reniforme  148;  PI.  6,  fig.  i 

reticulatum  147 
Omosoma  58-59,  92 

intermedium  58 

pulchellum  58 

sahelalmae  58 
Omphalocyclus  10 
Oncorhynchus  92 
Onoclea  sensibilis  235 
Onychiopsis  219,  238 

elongata  238 

mantelli  238,  239 

psilotoides  211,  238-239 
Onychocella  4,  5,  12,  34 

bathonica  3,  4,  5-6,  7,  15,  25,  41 

bellona  n,  35-36,  41;  PI.  6,  figs.  5-7 

cf.  £e£Aa  ii,  34-35,  41;  PI.  6,  figs,  i,  2 

flabelliformis  6 

hagenowi  6 

koninckiana  6 

lamarcki  35 

wys#  34,  41 ;  PI.  6,  figs.  3,  4 

piriformis  5,  6,  n,  41;  PI.  5,  figs,  n,  12 
Onychocellaria  36 

caecilia  n,  36-37,  41;  PL  5,  figs.  1-6 
Onychocellidae  13 
Orbicyathus  346 
Orbitoides  10 
Oreosomatidae  96 
Orthograptus  cf.  whitfieldi  189 
Orthonotacythere  432 

nodosa  432 

pulchella  432 

sp.  432;  PI.  15,  fig.  i 
Osculipora  17 

truncata  17,  41 

Pachycormidae  98 
Pachypeteris  240 

lancsolata  240-241  *-242 ;  PI.  i,  fig.  6 
Pagruo  87 

Palaeocentrotus  81,  97 
Palaehystrichophora  infusorioides  109 

cf.  paucisetosa  109 
Palaeoperidinium  castanea  131 

spinosum  142,  143 

ventriosum  130,  131 
Palaeostomocystis  echinulata  157 
Paleschara  13,  14 
Palescharidae  13 
Paracoscinus  363,  364 
Paracycas  cteis  246 
Paracyclotosaurus  292 
Paracyprididae  385,  399 
Paracypris  399 


terraefullonicae  399;  PI.  7,  figs.  2,  4 
Paracytheridea  433 

blakei  433;  PI.  15,  figs.  4,  5,  9,  10 

caytonensis  433 
Parakannemeyeria  285,  290 
Paranacyathus  312,  354 

magnipora  311 
Parariscus  433 

bathonicus  433;  PI.  15,  figs.  2,  8 
Parasudis  81 
Paratrachichthys  68 
Pareodiniaceae  130 
Pareodina  sp.  109 
Parupeneus  91 
Pateroperca  51-52,  53,  54,  55,  83,  84,  86,  87,  93,  98 

libanica  51*,  52,  53 
Pavobeisselina  40 
Pecopteris  reticulata  239 
Pegasiformes  96 
Pelletieria  232 

valdensis  232-233*-234*-235;  PI.  6,  figs,  i,  2 
Pellia  220 
Peltostega  291 
Pentaceropsis  91 

recurvirostris  91*,  92 
Pentacerotidae  87,  91 
Percichthyidae  87,  89,  92 
Percidae  87 

Perciformes  49,  87,  88,  92,  93,  95,  96,  99 
Percoidei  86,  89,  92,  96 

Permo-Trias  continental  sediments  classification- 
Africa  293 

Petalopora  sp.  18,  41;  PI.  i,  figs.  1-3 
Pharmacichthyidae  63-65,  92,  97,  100 
Pharmacichthys  49,  63,  65,  66,  84,  87,  88,  93,  94, 

95.  97.  98,  99 

venenifer  63 
Pinacocyathus  342 

spicularis  342,  343*-344;  PI.  12,  figs,  i,  2 
Placerias  290 
P/a/a#  87 

Platycopina  380,  395 
Platycythere  431 

verriculata  431 

sp.  431 ;  PI.  14,  fig.  5 
Plecoglossus  86 
Pleurograpsus  linearis  186 
Pleurograptus  linearis  186-187*;  PI.  2,  figs.  1-3 

linearis  simplex  186,  187 
Pleuronectes  89 
Pleuronectiformes  93,  95 
Pleuronectoidei  89 
Pliophloea  37 

gluma  37 

sp.  7,  14,  15,  37,  41;  PI.  7,  fig.  3 
Pithodella  24 
Podocopida  380,  395 
Podocopina  383,  397 
Poly  cope  427 


45° 

fungosa  427-428;  PI.  16,  figs.  4,  9 
Polycopodae  427 
Polycoscinidae  334 
Polycoscinus  334,  369 
Polydactylus  87 
Polymixia  56,  57,  58,  59,  60,  61,  88,  94,  95 

japonicus  57 

nobilis  56*,  57 

Polymixiidae  56-57,  61,  92,  93,  95,  98 
Polymixioidei  49,  56,  63,  66,  84,  86,  88,  99 
Polynemoidei  87 
"  Polyphyma  bulbosa  "12 
Polypodiaceae  235 
Poly pr ion  87 
Polysphaeridium  151 

laminaspinosum  151;  PI.  4,  figs.  10,  n 

pumilum  151 
Polystichum  235,  236 
Polystillicidocyathus  318,  327 
Pomacanthus  87 
Pomadasyidae  87 
Pomatomidae  87 
Pomatomus  87 
Pontocyprella  385 

harrisiana  379,  385;  PI.  2,  fig.  3 
Porina  13 

cenomana  13 

flabellata  40 
Porinidae  13 

Praechara  symmetrica  215 
Praeschuleridea  387,  405 

schwageriana  387;  PI.  2,  fig.  6 

subtrigona  intermedia  387 

magna  405 

subtrigona  387,  405;  PI.  8,  figs.  4-6 

sp.  388;  PI.  2,  fig.  8 
Pristolepis  89 
Procytheridea  430 

ininuta  430 

parva  430 
Progonocythere  389,  423 

blakeana  393 

juglandica  389 

stilla423;  PI.  13,  fig.  2 
Progonocytheridae  388,  409,  428 
Progonocytherinae  388,  409 
Prelates  88 
Prolixosphaeridium  160,  161 

conulum  160-161;  PI  8,  figs.  5,  6 

granulosum  161 

mixtispinosum  161 
Prosoporella  37 

cornuta  37 

Protocheilostomata  14 
Protocytheridae  430 
Protopharetra  348,  354,  356,  370 

graphica  346,  348 
Psettodes  89 
Pseudoceratium  turneri  171 


INDEX 


Pseudocycas  dunkeriana  212 
Pseudotorellia  248,  250 

angustifolia  249 

crassifolia  249 

ensiformis  249,  250 

ephela  249,  250,  251 

grojecensis  249,  251 

heterophylla  248-250*-251 ;  PI.  6,  figs.  6,  7 

longifolia  249 

minuta  249,  250,  251 

nordenskioldi  248,  249,  250 
Pteridophyta  210,  211,  224 
Pteridospermae  210,  211,  240 
Pterophyllum  schaumburgense  242 
Puesto  Viejo  Formation  290 
Puncturiella  33 

cf.  superba  33-34,  41;  PI.  4,  figs.  8-10 
Pustulopora  benedeniana  16 
Pycnoidocoscinus  363,  367,  368 

pycnoideum  367-368;  PI.  18,  figs.  4-8 
Pycnosterinx  59-60,  99 

discoides  59 

dubius  59,  60,  61,  88 

gracilis  59 

/afws  59,  60 

russeggerii  59*,  60 
Pycnosteroides  65,  66,  67,  68,  84,  87,  88,  93,  95,  99 

levispinosus  65,  66* 
Pycnosteroididae  65-66,  92,  98 
Pyripora  texana  13 

Quadricellaria  29 

elegans  29,  41;  PI.  6,  figs.  8-10 
excavata  29 
pulchella  29  , 

Radulopora  3,  21-22 

minor  3,  21,  22-23,  31,  41;  PI.  3,  figs.  6-10 

radula  21,  22*;  PI.  3,  figs,  n,  12 
Rastrites  198 

capillaris  198,  199* 

carnicus  201 

distans  200 

gemmatus  198 

linnaei  200,  201* 

maximus  184,  199,  200-201*-202;  PI.  5,  fig.  7 

cf.  peregrinus  200 

spina  199 
Rectocythere  409 

lincolnensis  410 

rugosa  409 

sugillata  409-410;  PI.  10,  figs.  3-6 
Rectonychocella  27 
Red  Marl  (Beds)  258,  259,  292,  293 
Regularia  302,  312,  346,  370 
Reptescharinella  29,  30 
Retecava  abbottii  16 

lichenoides  18 

ramosa  16 


INDEX 


Retecoscinus  325,  367 
Retepora  lichenoides  18 

truncata  17 
Reteporidae  18 

lichenoides  15,  18,  41 
Reticrisina  12 

"  Reticulipora  dianthus  "12 
Retropinna  86 
Rhabdocnema  302 
Rhabdocyathus  302,  304 

tubexternus  304 
Rhabdolynthus  302 
Rhagasostoma  bellona  35 

simplex  30 
Rhammatopora  gaultina  1 3 

(?)  johnstoniana  13 

pembrokiae  13 

tn'wei  13 
Rhiniopora  8 

jurassica  7 

scabra  7,  8 
Rhytidosteus  291 
Robustocyathidae  311 
Robustocyathus  311,  312,  369 

ar genius  311 

annulatus  311 

artecaveatus  311 

biohermicus  311 

dewsws  311 

hupei  311 

levigatus  311 

magnipora  311-312;  PI.  3,  fig.  5 

moori  311 

novus  311 

polyseptatus  311 

proskurjakowi  311 

pseudotichus  311,  314 

salebrosus  311 

spinosus  311 

stapipora  309 

subacutus  311,  312-313*-314,  369;  PI.  3,  figs. 
6,7 

sucharichensis  311 

tennis  311 
Rosseliana  31 

thomasi  3,  7,  n,  14,  19,  31,  41;  PI.  4,  figs.  4-6 
Ruffordia  229 

goepperti  229-230*-231-232;  PI.  4,  figs.   1-7; 
PL  5,  figs.  1-4 
var.  latifolia  229 

Sajanocyathus  369 
Salairocyathidae  332,  333,  336 
Salairocyathus  336 

(Polystillicidocyathus)  336 

erbosimilis  336 

(Salairocyathus)  336 

annulatus  336,  337*-338,  369,  372 ;  PI.  n,  fig.  4 

pospelovi  336 


zenkovae  336 
Salmo  84 
Salmoniformes  81 

Sangusaurus  285,  286,  289,  290,  292 
Sangusaurus  edentatus  257,  286-287*-288*- 

289*-290 

Santa  Maria  Formation  281,  283,  284,  290 
Sardinioides  82,  83,  98 

attenuatus  83,  98 
Saurida  81,  89 
Scalenodontoides  291 
Schedophilus  87 

Schizaeaceae  229,  231,  232,  235 
Schuleridea  385,  402,  403 

(Eoschuleridea)  402 

bathonica  386,  387,  403 

horatiana  402-404.  [403*] ;  PI.  7,  fig.  7 

subperforata  386;  PI.  2,  fig.  4 

trigonalis  386-387,  388;  PI.  2,  fig.  7 

(Schuleridea)  jonesiana  379,  385 ;  PI.  2,  fig.  5 
Schulerideidae  385,  402 
Schulerideinae  385,  402 
Sciaena  87 
Sciaenidae  87 
Scolopsis  87 

Scorpaemiformes  95,  96 
Scorpididae  92 
Selaginella  224,  227 

dawsoni  224-225-226  *-227 

dichotoma  227 

emmeliana  227 

Aa//ei  224,  227 

sanguinolenta  227 
Selaginellaceae  224 
Selaginellites  224 

dawsoni  224 

nosikovii  227 

polaris  227 
Selene  98 

Semieschara  complanata  31 
Semiescharinella  29,  30 

complanata  n,  29-30,  41 ;  PI.  4,  figs.  11-15 
Semiescharipora  cornuta  37 
Semiophorus  97 
Serranidae  87,  92 
Sertulariadae  192 
Siderolites  10 
Siniperca  63,  89 

chuatsi  go* 

Sinokannemeyeria  285,  290 
Siphonella  elegans  20 
Smittipora  27 
Soleoidei  89 
Sparidae  87 
Sparus  87 
Spermatophyta  240 
Sphaerodon  87 

Sphenocephalidae  61-62,  92,  98 
Sphenocephalus  92,  93,  94,  95,  99 


452 

fissicaudus  61,  62* 

kurriana  212 

sternbergiana  212 
Sphenopteris  fittoni  238 

fontainei  238 

mantelli  238 

ruff  or di  238 
Sphyraena  87,  88,  92 
Sphyraenoidei  87 
Spinosocyathus  342 
Spirillicyathus  344  ^ 

Spirocyathella  344 
Spirocyathus  atlanticus  348 

irregularis  344,  348 

major  349 

speciosus  350 
Spirograptus  197 

intermedius  197 

planus  197 
Stahleckeria  257,  282,  283,  284 

potens  282* 
Stahleckeriidae  281 
Stamenocella  25 

inferaviculifera  25 

marginata  6,  n,  25-26,  41;  PI.  3,  figs.  1-5 

mediaviculifera  25 
Stephanoberycoidei  68 
Stichocentrus  77-78,  94,  95 

liratus  77,  78* 
Stichopora  regularis  16 
Stillicidocyathidae  317,  319 
Stillicidocyathus  318 
Stromateoidei  87 
Stomatopora  smithi  9 
Surculosphaeridium  no,  158 

longifurcatum  158-159;  PI.  8,  fig.  9 
Synbranchiformes  96 
Synodontidae  82 
Sy nodus  81 

foetens  57 
Syringocnema  301,  370,  371 

Tabellaecyathidae  370 
Tabellaecyathus  348 
Taeniopteris  beyrichii  var.  superba  244 
Teilhardia  valdensis  238 
Tannuolacyathidae  325 
Tannuolacyathus  325 
Tanyosphaeridium  151,  160 

variecalamum  151;  PI.  6,  figs.  2,  5 
Tarpon  54,  84,  85 
Terquemula  393,  423 

blakeana  393,  423;  PI.  4,  fig.  8;  PI.  5,  fig.  i 
Tetraodontiformes  95 
Tetrasphaera  132 
Thalamocyathidae  317 
Thalamocyathus  317,  318,  319 

flexuosus  318 

trachealis  318,  319 


INDEX 


Thallites  219,  220-221 

catenelloides  222*,  223-224 
uralensis  223 
valdensis  221-222*-223 

willsi  224 

yaiei  220 

zeilleri  219 

Thallophyta  210,  211,  214 
Thamnograptus  185 
Thecidea  40 

papillata  n,  40,  41,  42;  PI.  i,  figs.  9,  10 
Thrissops  84,  85 
Thyracella  26 

cf.  meudonensis  26,  41;  PI.  2,  figs.  4-6 
Tomocyathus  334,  340,  369,  372 

(Erugatocyathus)  299,  334,  372 

compositus  334 

echinus  334 

£*'»*  334 

kundatus  334 

papillatus  334-335*-336;  PI.  n,  fig.  2 

shoriensis  334 

(Tomocyathus}  michniaki  334 

operosus  334 
Tortula  218 

Trachichthyidae  67,  68,  69,  77,  94,  95,  96 
Trachichthyoides  ornatus  78 
Trachichthys  68 
Trachinocephalus  81 
Trachycythere  428 

verrucosa  428 

sp.  428;  PI.  1 6,  fig.  3 
Trachyleberididae  394,  401,  428 
Trachyleberidinae  401 
Trichodinium  131,  132,  162 

castaneum  131-132;  PI.  u,  figs,  i,  2,  3 

intermedium  132 

paucispinum  163 
Trichodium  castanea  131 
Triletes  224 

Truncatula  tetrasticha  17 
Truncatula  truncata  17 
Tselfatiidae  98 
Tubantia  74 

cataphractus  74* 
Tumuliolynthus  302,  304,  369 

irregularis  304-305;  PI.  i,  fig.  i 

karakolensis  304 

musatovi  304 

tubexternus  304 

vologdini  304 
Tumulocoscininae  338 
Tumulocoscinus  338 
Tunkia  302 
Tuvacyathus  325 


f/mo  karooensis  259 


INDEX 


453 


Velifer  Si,  97 
Veliferidae  78,  98 
Velumella  27 
Vincularia  27,  28 

canalifera  27-28,  41;  PI.  5,  figs.  7-10;  PI. 
figs.  9-12 

concinna  28;  PI.  8,  fig.  13 

flexuosa  27,  28;  PI.  8,  fig.  14 

fragilis  27 

rugica  32 

transversa  32 

undata  32 

Volvacyathus  354,  356,  370 
Vomer  98 
Vomeridae  98 
Vomeropsis  98 

Weichselia  236,  239 

mantelli  236,  239 

reticulata  236,  239;  PI.  6,  fig.  5 
W  etlugosaurus  291 
Wilbertopora  mutabilis  13 
Worthenopora  13,  14 
Worthenoporidae  13 

Xanthidium  ramosum  172 

spinosum  166 

tubiferum  143 

complex  146 
Xenichthys  87 


Xenistitius  87 
Xenocys  87 
Xenolepidichthys  96 


Zambiasaurus  257,  259,  260-261,  281,  282,  283, 
284,  287,  290,  292 

submersus    257,    262-263*-264*-265-266*- 
267-268*-269*-270*-271*-272*-273*- 
274*-275*-276*-277*-278*-279*-280*- 
281*-282*-286 
Zeidae  96 

Zeiformes  49,  93,  96,  97,  100 
Zews  96 
Zonacyathus  314-315,  369 

flexum  369 

poletaevae  369 

retevallum  314,  315-316,  317,  PI.  4,  figs.  4,  5 

retezona  315,  316*-317;  PI.  4,  figs.  1-3 

vermiculatum  369 
Zone  of  Climacograptus  wilsoni  189,  190,  195 

Dicranograptus  clingani  185,  188,  192,  193,  195 

Didymograptus  extensus  195 

Monograptus  convolutus  196 

Monograptus  cyphus  193 

Monograptus  gregarius  196 

Monograptus  riccartonensis  203 

Monograptus  sedgwickii  196 

Orthograptus  vesiculosus  193 

Pleurograptus  linearis  185,  188,  192,  193,  195 


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