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LI  B  RAR.Y 

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GEOLOGICAL  SERIES 

OF 

FIELD  MUSEUM  OF  NATURAL  HISTORY 

Volume  VI  Chicago,  May  15,  1935  No.  11 

DESCRIPTION  OF  A  SILURIAN  PHYLLOPOD 
MANDIBLE  WITH  RELATED  NOTES 

By  Sharat  Kumar  Roy 
Assistant  Curator  of  Gboixwy 

Messrs.  J.  Mann  and  A.  Lees,  of  Oak  Lawn,  Illinois,  have  recently- 
donated  to  the  Museum  two  specimens  (part  and  counterpart)  of 
a  mandible  referred  here  to  a  new  species  of  Ceratiocaris.  The 
mandible  was  found  in  the  Lecthaylus  shale.  Upper  Lockport  group 
(Niagaran),  near  Blue  Island,  Illinois,  within  one  hundred  feet  of 
Ceratiocaris  markhami  Roy  (1935,  fig.  29a-e).  The  specimen  is 
completely  carbonized  but  much  of  the  carbonized  material  has 
been  broken  away. 

The  writer  wishes  to  acknowledge  his  indebtedness  to  both 
the  donors  for  giving  him  the  opportunity  of  describing  this  rarely 
found  fossil,  and  takes  pleasure  in  naming  it  after  one  of  the  donors, 
Mr.  Lees. 

The  drawings  are  by  the  Staff  Illustrator  of  Field  Museum,. 
Mr.  Carl  Gronemann. 

Ceratiocaris  leesi  Roy  sp.  nov.     Fig.  33a  and  b. 

Description. — Transversely  elongated,  widest  anteriorly;  posterior 
end  roundly  pointed.  General  outline  somewhat  resembles  that  of 
a  mitre  with  the  "tooth"  row  forming  the  base.  Upper  and  lower 
margins  rounded,  giving  a  rim-like  appearance;  both  broadening 
anteriorly;  the  upper  one  arching  and  ending  in  a  narrow  pointed 
projection  which  overhangs  the  "tooth"  row,  the  lower  one  curving 
slightly  inward  and  meeting  the  anterior  margin  to  form  a  triangle. 
Anterior  or  oral  margin  thickest,  supporting  seven  toothed  ridges, 
the  uppermost  of  which  is  the  longest,  then  decreasing  uniformly  in 
size  downwards.  The  elongated  base,  representative  of  the  pro- 
topodite,  is  not  well  preserved.  Its  central  portion  is  missing  and 
its  posterior  end,  the  position  for  the  external  articular  process,  may 
or  may  not  have  been  outlined  (see  fig.  33a  and  b)  correctly. 

Dimensions. — Length  and  width  at  the  center  22.5  mm.  and 
10  mm.  respectively. 

No.  343  155 


f«r«tia:a]  Hiiit«rT  Survey 


156  Field  Museum  of  Natural  History — Geology,  Vol.  VI 

Remarks. — No  record  of  gastric  teeth  or  mandibles  of  phyllopod 
crustaceans  in  geologic  literature  is  to  be  found  prior  to  1843.  In 
that  year  J.  E.  Portlock  first  figured  the  gastric  teeth,  dextral  and 
sinistral,  of  Dithyrocaris  from  Tyrone  or  Londonderry,  Ireland  (pi. 
12,  fig.  6,  of  Portlock's  report  and  fig.  33c^  of  this  paper).  On  page 
315  (Portlock,  1843)  he  states,  "Fig.  6  represents  bodies  which  are 
frequently  found  on  the  specimens  of  this  crustacean,  and  in  this 
instance  together,  as  represented  in  the  figured  specimen,  they  each 
exhibit  a  single  row  of  tubercles,  and  were  in  all  probability  con- 
nected with  the  masticatory  apparatus,  which  it  is  probable,  there- 
fore, was  highly  developed  in  this  large  species." 

Following  the  report  of  Portlock,  several  other  teeth  of  Dithyro- 
caris, presumably  all  gastric,  have  been  found,  in  comparative 
abundance,  at  Campsie  and  East  Kilbride,  Lanarkshire,  Scotland, 
and  at  Orchard  Quarry,  near  Thornliebank,  Renfrewshire,  Scotland. 
These  teeth  have  been  described  and  illustrated  by  Woodward 
(1865,  pp.  401-404,  pi.  11)  and  more  adequately  by  Jones  and 
Woodward  (1898,  pp.  194-198,  pi.  26,  figs.  1-44). 

The  "teeth"  of  Ceratiocaris  (from  Lesmahagow,  Lanarkshire) 
were  first  reported  by  J.  W.  Salter  in  1860.  He  observes  (1860, 
p.  155):  "A  further  exploration  in  the  same  rich  deposits  at  Les- 
mahagow, where  Ceratiocaris  occurs  . . .  has  disinterred  the  rostrum, 
hard  jaws,  and  also  the  antennae  or  some  of  the  thoracic  appendages 
of  the  animal.  The  last  are  obscure,  but  were  detected  by  Professor 
Huxley,  who  also  found  a  clearly  articulated  hinge  uniting  the  two 
flaps  of  the  carapace.  This  is  a  very  important  character,  and  will 
remove  Ceratiocaris  from  any  very  near  alliance  with  Nebalia,  from 
which  also  the  solid  jaws,  like  those  of  Apus . .  .  tend  further  to 
separate  it." 

Salter's  brief  report  was  supplemented  by  more  adequate  explana- 
tion and  illustrations  by  Woodward  (1865,  pp.  401-404).  Referring 
to  plate  XI  of  his  paper  Woodward  states  (p.  401) :  "Fig.  1  represents 
a  perfect  carapace  of  Ceratiocaris,  at  the  anterior  end  of  which 
(at  a)  the  two  opposing  mandibles  may  be  seen  in  their  proper 
place,^  compressed  on  the  surface  of  the  left  valve.    A  detached 

*  Copied  from  a  figure  which  was  drawn  more  accurately  from  the  original 
specimen  by  Jones  and  Woodward  (1898,  pi.  26,  fig.  44).  A  facsimile  of  Portlock's 
figure  is  also  given  by  Woodward  (1865,  pi.  11,  fig.  8). 

^  The  mandibles  were  not  in  their  proper  place  as  was  explained  later  by  Jones 
and  Woodward  (1888,  p.  147).  In  normal  position  they  would  be  opposed  to  each 
other  horizontally.  That  they  appear  as  being  opposed  one  to  the  other  vertically 
is  due  to  the  compression  of  the  carapace,  and  to  their  being  squeezed  against 
the  inside  of  the  valves. 


1  K  I'i^  ,   .- 


L.      r 


V  .^ 


\       Description  of  a  Phyllopod  Mandible 


157 


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Fig.  33.  a,  h,  Ceratioearis  kegi  Roy  sp.  nov.  Mandible  (part  and  counterpart).  P.M.  No.  P23676. 
Natural  size,  c-e,  Gastric  teeth  of  Dithyroearis  (Jones  and  Woodward,  1898,  pi.  26,  fijfs.  23,  43,  and  44  ) :  c, 
Sinistral  lying  on  dextral;  d,  Sinistral;  e,  Dextral.  f-l,  "Teeth"  of  Ceratioearis?  f.  Chevron-shaped 
regular  cusps;  g,  h.  Cusps  longer  at  one  end;  t,  Cusps  with  more  flexuous  connecting  ridge;  i.  Cusps 
regularly  alternate;  k,  Cusps  irregularly  alternate;  I,  Cusps  in  single  row. 


158  Field  Museum  of  Natural  History — Geology,  Vol.  VI 

mandible  (fig.  2)  from  the  same  locality  as  fig.  1  (Upper  Ludlow  Rock, 
Lesmahagow,  Lanarkshire)  is  represented  in  the  plate  of  the  natural 
size."  On  page  402,  he  further  states:  "That  these  detached  fossil 
remains  are  the  teeth'  of  crustaceans  there  cannot  now  be  the  least 
doubt;  for,  in  addition  to  the  evidence  afforded  by  the  remains 
of  the  allied  genus  Ceratiocaris  (pi.  XI,  fig.  1),  and  by  the  fragment 
of  Dithyrocaris  (pi.  XI,  fig.  6),  one  needs  only  to  compare  the  gastric 
teeth  of  the  common  lobster  (Homarus  vulgaris),  figured  in  our 
plate  (fig.  12),  with  the  most  perfect  teeth  of  Dithyrocaris  (figs.  3, 
36),  to  see  the  striking  similarity  between  them,  both  in  general 
form  and  in  minuter  details." 

The  most  important  paper  on  the  present  subject,  "The  Gastric 
Teeth  of  Ceratiocaris"  was  published  by  Jones  and  Woodward  in 
1888.  The  paper  is  an  excellent  survey  of  all  the  important  specimens 
known  from  Bohemia,  Scandinavia  and  Great  Britain  and  contains 
two  important  conclusions  reached  by  the  authors  as  the  result  of 
their  comparative  studies.    These  may  be  stated  as  follows: 

I.     The  "teeth"  are  divisible  into  two  distinct  groups: 

(a)  Gastric  teeth.    Thick  and  solid. 

(b)  Mandible.    Trenchant  character  of  cutting  edges;  broad,  com- 

pressed, transversely  elongated  bases  of  attachment. 

II.     The  "teeth"  have  at  least  six  different  forms.^ 

(a)     Cusps,  chevron  shaped,  regular  in  size  (fig.  33/).' 

(6)     Cusps  longer  at  one  end  of  the  tooth  than  at  the  other  (fig.  33sf 
and  h).^ 

(c)  Cusps  with  a  more  flexuous  connecting  ridge  (fig.  33i).' 

(d)  Cusps  in  two  parallel  rows,  but  somewhat  alternate  (fig.  BSj).^ 

(e)  Cusps  irregularly  alternate  in  two  rows  (fig.  33 A:).' 
(/)  Cusps  in  a  single  row  (fig.  330-' 

As  to  the  first  conclusion,  my  own  inference  has  been  the  same 
but  I  have  hesitated  to  express  it  because  of  lack  of  actual  specimens 
to  make  a  comparative  study.  With  regard  to  the  second,  the 
authors,  it  appears,  have  based  their  groupings  on  criteria  which 
have  frankly  many  possibilities  of  error.     It  is  difficult  to  believe 

1  It  is  quite  apparent  that  up  to  this  time  at  least,  no  special  effort  was  made 
to  differentiate  between  mandibles  and  gastric  teeth.  Frequently,  both  organs 
were  included  in  the  general  term  "teeth." 

*  The  authors  have  included  mandibles  in  differentiating  these  forms.  See 
II  (6). 

3  These  figures  refer  to  the  figures  reproduced  in  the  present  paper.  Fig.  33/ 
(Jones  and  Woodward,  1888,  pi.  VI,  fig.  8);  g  and  h  (Jones  and  Woodward,  1888, 
text  figs.  5,  6);  i  (Jones  and  Woodward,  1888,  pi.  VI,  fig.  10c)  ;j  (Barrande,  1872, 
pi.  31,  fig.  21);  k  (Jones  and  Woodward,  1888,  text  fig.  4a);  I  (Jones  and  Wood- 
ward, 1888,  pi.  VI,  96). 


Description  of  a  Phyllopod  Mandible  159 

that  a  single  genus  would  possess  six  widely  different  forms  of  teeth. 
No  such  parallel  exists  among  living  forms.  The  gastric  teeth  or 
the  mandible  of  the  lobster,  Homarus,  or  of  the  crayfish,  Astacus, 
show  but  little  variation,  irrespective  of  the  species  to  which  they 
belong.  The  same  holds  true  of  fossil  forms.  The  gastric  teeth  of 
Dithyrocaris  (Jones  and  Woodward,  1898,  pi.  26,  figs.  1-44,  and 
fig.  SSc-e  of  this  paper)  found  at  widely  separated  localities  do  not 
show  any  appreciable  variations.  The  evident  conclusion,  there- 
fore, is  that  some  of  the  teeth  that  have  been  identified  with  Ceratio- 
caris  because  they  were  found  in  association  with  other  remains  of 
Ceratiocaris  are  not  referable  to  this  genus.  These  teeth  must  have 
come  from  other  crustaceans  which  have  not  left  any  remains  other 
than  teeth.  This  and  other  factors  which  have  impeded  proper 
identification  may  be  summed  up  as  follows: 

(1).     Commingling  of  gastric  teeth  of  more  than  one  genus. 

(2).  Commingling  of  gastric  teeth  and  mandibles  whose  bases  have  not  been 
preserved  or  whose  cutting  edges  have  been  worn  or  blunted,  thus 
causing  them  to  simulate  gastric  teeth. 

(3).     Relative  age  of  individuals. 

(4).     Conditions  under  which  fossilization  has  taken  place. 

The  above  factors  at  once  create  a  situation  which  does  not  admit 
of  identifications  that  can  be  relied  on.  They,  further,  definitely 
imply  that  unless  and  until  other  specimens  of  the  various  tjrpes  of 
teeth  hitherto  identified  with  Ceratiocaris  are  found  in  such  intimate 
association  with  remains  of  individuals  as  to  leave  no  doubt  that 
they  are  referable  to  those  individuals,  there  can  be  no  escape  from 
the  confusion  now  existing. 

It  has  been  mentioned  in  the  beginning  that  the  mandible  from 
Blue  Island  was  found  a  short  distance  from  C.  markhami.  I  have, 
nevertheless,  hesitated  to  identify  it  with  that  species  for  the  reason 
that  the  mandible  is  relatively  too  large,  and  hence  can  hardly  be 
referred  to  C.  markhami  unless  it  is  assumed  that  the  latter  has  not 
reached  its  adult  stage.  There  is,  however,  no  valid  reason  for  such 
assumption.  Structurally,  C.  markhami  shows  all  the  features  hither- 
to regarded  as  characterizing  an  adult  specimen  (S.  K.  Roy,  1935, 
pp.  142-144). 

Fossil  mandibles  of  Paleozoic  phyllopod  crustaceans  are  not  of 
common  occurrence.  The  mandible  here  figured  and  described  is 
the  only  one  yet  recorded  from  the  Silurian  of  North  America. 
Whitfield  (1896,  pi.  13,  fig.  5,  and  pi.  14,  figs.  1-4)  has  figured 
five  supposed  mandibles  from  the  Lower  Helderberg  (Devonian) 


160  Field  Museum  of  Natural  History— Geology,  Vol.  VI 

of  Wisconsin,  three  of  which  are  referred  to  Ceratiocaris  monroei 
Whitfield,  and  two  to  Entomocaris  telleri  Whitfield.  Whitfield's 
specimens  have  massive  cusps  and  lack  entirely  the  trenchant 
character  of  the  sharp  cutting  edges  of  mandibles.  There  is  a  possi- 
bility that  they  are  gastric  teeth  which  have  retained  their  zygo- 
cardiac  ossicles.  The  fact  that  mandible-like  organs  have  been 
found  attached  to  the  ventral  surface  of  the  anterior  region  of  the 
carapace  does  not  necessarily  imply  that  they  are  mandibles.  Gastric 
teeth,  sinistral  and  dextral,  may  easily  become  detached  from  the 
pyloric  ossicle  and,  dropping  out  of  the  stomach  (during  its  decom- 
position) become  lodged  inside  the  valves  of  the  carapace,  occupying 
the  position  of  mandibles. 

The  mandible  described  in  this  paper,  with  the  exception  of  its 
elongated  base,  compares  very  favorably  with  the  two  from  Les- 
mahagow,  Lanarkshire  (Jones  and  Woodward,  1888,  text  figs, 
and  6).  The  toothed  ridges  of  all  three  specimens  are  strikingly 
similar,  and,  judging  from  the  similarity  of  the  bases  of  the  Scottish 
specimens,  it  seems  highly  probable  that  the  base  of  the  present  one 
might  have  been  somewhat  distorted. 

Horizon  and  locality. — Lecthaylus  shale.  Upper  Lockport  group 
(Niagaran),  Blue  Island,  Illinois. 

P23676  Field  Museum. 

Holotype. 

REFERENCES 
Barrande,  J. 

1872.    Systeme  silurien  du  centre  de  la  Bohdme.     1,  Suppl. 
Jones,  T.  R.  and  Woodward,  H. 

1888.    The  Gastric  Teeth  of  Ceratiocaris.     Geol.  Mag.,  New  Ser.,  Dec.  3,  5, 

No.  4. 

1898.    The  Gastric  Teeth  of  Dithyrocaris  (Mong.  Brit.  Paleozoic  Phyllopoda, 
pt.  III).    Paleont.  Soc,  London. 

PORTLOCK,  J.  E. 

1843.    Report  on  the  Geology  of  the  County  of  Londonderry. 
Roy,  S.  K. 

1935.    A  New  Silurian   Phyllopodous   Crustacean.     Field   Mus.   Nat.   Hist., 
Geol.  Ser.,  6,  No.  9. 

Salter,  J.  W. 

1860.    On  New  Fossil  Crustacea  from  the  Silurian  Rocks.     Ann.  Mag.  Nat. 
Hist.,  3rd  Ser.,  5,  No.  27. 

Whitfield,  R.  P. 

1896.    Notice  and  Description  of  New  Species  and  a  New  Genus  of  Phyllo- 
caridae.    Bull.  Amer.  Mus.  Nat.  Hist.,  8. 

Woodward,  H. 

1865.    On  Some  Crustacean  Teeth  from  the  Carboniferous  and  Upper  Ludlow 
Rocks  of  Scotland.    Geol.  Mag.,  2.