Skip to main content

Full text of "Miscellaneous publication - University of Kansas, Museum of Natural History"

See other formats


HARVARD  UNIVERSITY 


LIBRARY 


OF  THE 


Museum  of  Comparative  Zoology 


,,  ^^e  Library 

••useu™  of  Ccnaratlve  Zoology 
Harvard  University 


'-    -  i,iU;^.  CU.ViP.  ZCCL 

LIBRARY 

UNIVERSITY  OF  KANSAS  miscellaneous 

MUSEUM  OF  NATURAL  HISTORY  JiW-il^Ajgpjj 

No.   55 

HARVARD^ 
UNlVERSl-ni 


Middle  American  Lizards  of 
the  Genus  Ameiva  (Teiidae)  with 
Emphasis  on  Geographic  Variation 


By 

Arthur  C.  Echternacht 


UNIVERSITY  OF  KANSAS 

LAWRENCE  1971  December  14, 1971 


Jk.. 


UNIVERSITY  OF  KANSAS  PUBLICATIONS 
MUSEUM  OF  NATURAL  HISTORY 

The  University  of  Kansas  Publications,  Museum  of  Natural  History,  beginning 
with  volume  1  in  1946,  was  discontinued  with  volume  20  in  1971.  Shorter  research 
papers  formerly  published  in  the  above  series  are  now  published  as  Occasional 
Papers,  Museum  of  Natural  History,  The  Miscellaneous  Publications,  Museum  of 
Natural  History,  began  with  number  1  in  1946.  Longer  research  papers  are  pub- 
lished in  that  series.  Monographs  of  the  Museum  of  Natural  History  were  initiated 
in  1970. 

Institutional  libraries  interested  in  exchanging  publications  may  obtain  the  Occa- 
sional Papers  and  Miscellaneous  Publications  by  addressing  the  Exchange  Librarian, 
University  of  Kansas  Library,  Lawrence,  Kansas  66044.  Individuals  may  purchase 
separate  numbers  of  all  series.  Prices  may  be  obtained  upon  request  addressed  to 
Publications  Secretary,  Museum  of  Natural  History,  University  of  Kansas,  Law- 
rence, Kansas    66044. 


University  of  Kansas 
Museum  of  Natural  History 


Miscellaneous  Publication  No.  55 
December  14,  1971 


Middle  American  Lizards 

of  the  Genus  Ameiva  (Teiidae) 

with  Emphasis  on  Geographic  Variation 


By 

Arthur  C.  Echternacht 

Department  of  Biology 

Boston  University 

Boston,  Massachusetts   02215 


University  of  Kansas 

Lawrence 

1971 


University  of  Kansas  Publications,  Museum  of  Natural  History 


Editors:   Craig  C.  Black,  William  E.  Duellman, 
Philip  S.  Humphrey 


Miscellaneous  Publication  No.  55 

pp.  1-86;  28  figures 

Published  December  14,  1971 


Museum  of  Natural  History 

University  of  Kansas 

Lawrence,  Kansas  66044 

U.S.A. 


Printed  by 

University  of  Kansas  Printing  Service 

Lawrence,  Kansas 


CONTENTS 

INTRODUCTION    4 

ACKNOWLEDGMENTS    5 

MATERIALS  AND  METHODS  6 

Characters   6 

Statistical  Procedures    11 

Sexual  Dimorphism  1 2 

Sources  of  Specimens  Examined  12 

SYSTEMATIC  ACCOUNT  OF  THE  SPECIES  13 

KEY  TO  THE  MIDDLE  AMERICAN  SPECIES  OF  AMEIVA 13 

Ameiva  ameiva  ( Linnaeus )  14 

A meiva  leptoph  njs  Cope  20 

Ameiva  f estiva  ( Lichtenstein  and  Von  Martens)  26 

Ameiva  quadrilineata  (Hallowell)  34 

Ameiva  undulata  ( Wiegmann )  40 

Ameiva  chaitzami  Stuart 59 

DISCUSSION    ■ 63 

INTERSPECIFIC   RELATIONSHIPS   63 

INTERSPECIFIC  PATTERNS  IN  GEOGRAPHIC  VARIATION  64 

HISTORY  OF  AMEIVA  IN  MIDDLE  AMERICA  67 

SUMMARY  71 

RESUMEN   72 

APPENDIX  A:    SUMMARY  OF  STATISTICS 75 

APPENDIX  B:    SPECIMENS  EXAMINED  76 

LITERATURE  CITED   83 


INTRODUCTION 


In  their  revision  of  the  Mexican 
species  of  Ameiva,  Smith  and  Laufe 
( 1946 )  prefaced  their  comments  with 
the  remark  that  interest  in  these  species 
had  "been  rising  to  a  crescendo  whose 
peak  surely  could  soon  be  anticipated." 
Little  could  they  have  foreseen  the 
surge  of  interest,  both  taxonomic  and 
ecological,  that  the  entire  genus  has 
enjoyed  over  the  past  five  years.  This 
attention  has  been  precipitated  by  the 
realization  that  Arneiva,  by  virtue  of  its 
wide  distribution  in  the  Neotropical 
Realm  and  the  extremes  of  variation 
exhibited  by  its  species,  is  especially 
well  suited  for  ecological,  biogeograph- 
ical,  and  evolutionary  studies.  Coupled 
with  this  has  been  the  discovery  that 
some  species  of  Ameiva  act  as  vectors 
for  disease-causing  organisms  actually 
or  potentially  dangerous  to  humans  (for 
example,  see  Schneider,  1965). 

There  are  three  natural  geographic 
groupings  of  Ameiva:  those  found  on 
the  various  islands  of  the  Caribbean, 
those  on  the  continent  of  South  Amer- 
ica, and  those  from  Middle  America 
(Mexico  and  Central  America).  Taxo- 
nomic studies  of  the  Caribbean  Arneiva 
have  been  conducted  by  Baskin  and 
Williams  ( 1966 ) ,  Heatwole  and  Torres 
(1967),  Schwartz  (1965,  1967,  1968), 
Schwartz  and  Klinikowski  ( 1966 ) ,  and 
Schwartz  and  McCoy  (1970).  Very 
little  recent  work  has  been  done  on  the 
Ameiva  of  South  America;  the  only 
study  is  that  of  Peters  (1964)  on  the 
Ecuadorian  species.  No  recent  compre- 
hensive taxonomic  studies  have  been 
made  of  the  Middle  American  species, 
although  Stuart  (1942)  and  Smith  and 
Laufe  ( 1946 )  contributed  to  our  knowl- 
edge of  certain  species.  The  most  re- 
cent revision  of  the  entire  genus  is  that 
of  Barbour  and  Noble  (1915).  Their 
conclusions  concerning  the  Middle 
American  species  of  Ameiva  were  based 
on  examination  of  only  41  specimens,  a 
situation  which  largely  masked  the  re- 


markable degree  of  geographic  varia- 
tion exhibited  by  some  of  the  species. 
It  is  the  purpose  of  this  study  to  present 
an  analysis  of  the  geographic  variation 
in  each  of  the  Middle  American  species 
of  Ameiva  as  well  as  to  summarize 
known  information  on  the  biology  and 
ecology  of  these  lizards. 

Some  persons  will  look  upon  the 
present  work  as  justification  for  the  rec- 
ognition of  subspecies  of  Middle  Amer- 
ican Ameiva.  The  controversy  over  sub- 
species has  been  debated  extensively 
and  heatedly  elsewhere  (Wilson  and 
Brown,  1953;  Peters,  et  ah,  1954;  Smith 
and  White,  1956;  Savage  and  Heyer, 
1967;  Smith,  1967;  and  others),  and  I 
will  not  dwell  on  it  here  other  than  to 
state  my  conviction  that  a  thorough  de- 
scription of  geographic  variation  may 
make  unwarranted  and  often  undesir- 
able the  arbitraiy  nomenclatural  frag- 
mentation of  highly  variable  species. 

There  is  a  growing  body  of  evi- 
dence indicating  the  congeneric  status 
of  Ameiva  and  Cnemidophorus.  Van- 
zolini  and  Valencia  ( 1965 ) ,  Estes 
(1969),  and  Gorman  (1970)  demon- 
strated the  closeness  of  the  relationship 
and  discussed  the  difficulties  encoun- 
tered in  trying  to  diagnose  the  two 
genera  on  the  basis  of  known  charac- 
ters. Gorman  (1970:240),  however,  was 
hesitant  to  make  a  decisive  statement 
on  the  matter  when  he  commented: 
"The  similarity  of  karyotypes  between 
Ameiva  and  the  southernmost  Cnemido- 
phorus species  group  does  not  neces- 
sarily demonstrate  close  relationship, 
but  there  seem  to  be  no  characters  that 
clearly  separate  the  two  genera."  I  be- 
lieve that  the  argument  will  not  be  re- 
solved until  increased  knowledge  of 
South  American  Ameiva,  especially  the 
perplexing  A.  lacertoides,  is  available. 
The  condition  of  the  base  of  the  tongue 
(free  and  heart-shaped  in  Cnemido- 
phorus, sheathed  in  Ameiva)  has  been 
considered     to     be     diagnostic     (Burt, 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


1931),  but  A.  lacertoides  has  a  Cnemi- 
dophorus-\ike  tongue  (Vanzolini  and 
Valencia,  1965;  personal  observation). 
I  prefer  not  to  comment  on  the  issue, 
but  instead  await  the  completion  of 
relevant  investigations  by  Richard  Estes, 
George  C.  Gorman,  William  P.  Mac- 
Lean  III,  and  William  Presch. 

ACKNOWLEDGMENTS 

During  the  course  of  this  study,  I 
have  built  up  a  huge  debt  of  gratitude 
to  many  people  who  have  provided  as- 
sistance in  one  way  or  another.  I  have 
probably  taxed  the  patience  of  a  num- 
ber of  museum  curators,  curatorial  as- 
sistants, and  collectors  with  requests  for 
the  loan  of  large  numbers  of  specimens 
and  locality  or  ecological  data  on  spe- 
cific specimens.  I  especially  acknowl- 
edge the  following:  Kraig  K.  Adler,  the 
late  Doris  M.  Cochran,  James  R.  Dixon, 
William  E.  Duellman,  Henry  S.  Fitch, 
George  Foley,  Jean  Guibe,  Giinther 
Peters,  Robert  F.  Inger,  Daniel  A.  Jan- 
zen,  John  M.  Legler,  Charles  H.  Lowe, 
Edmond  V.  Malnate,  Hymen  Marx, 
John  R.  Meyer,  Charles  W.  Myers, 
Thomas  Olechowski,  James  A.  Peters, 
Douglas  C.  Robinson,  Richard  D.  Sage, 
Jay  M.  Savage,  Norman  J.  Scott,  Hobart 
M.  Smith,  Dorothy  M.  Smith,  A.  G. 
Stimson,  L.  C.  Stuart,  Stephen  G.  Til- 
ley,  Donald  W.  Tinkle,  Robert  G.  Tuck, 
Jaime  Villa,  Ernest  E.  Williams,  John 
W.  Wright  and  Richard  G.  Zweifel. 

I  am  indebted  to  Richard  F.  John- 
ston, Irving  Pfau,  James  D.  Rising,  F. 
James  Rohlf,  Gerald  R.  Smith,  and 
especially  Dennis  M.  Power  for  advice 
on  statistical  methods  or  computer  pro- 
cedures. The  computer  programs  uti- 
lized in  this  study  were  provided  by 
Dennis  M.  Power. 

I  have  had  the  benefit  of  a  great 
deal  of  constructive  criticism  and  advice 
from  my  colleagues  at  the  University  of 
Kansas  Museum  of  Natural  History. 
William  E.  Duellman  has  been  a  con- 
stant source  of  encouragement,  and  his 
patience  with  me  has,   at  times,   been 


remarkable.  He,  along  with  Henry  S. 
Fitch  and  Ronald  L.  McGregor,  have 
reviewed  this  manuscript.  Donald  R. 
Clark,  Ko  Ko  Gyi,  John  D.  Lynch,  Rus- 
sell J.  Hall,  Charles  W.  Myers,  E.  H. 
Taylor  and  Linda  Trueb  have  been  will- 
ing to  discuss  special  problems  as  they 
arose,  and  their  help  is  gratefully 
acknowledged. 

Richard  Estes,  George  C.  Gorman 
and  James  A.  Peters  have  kindly  al- 
lowed me  to  read  their  unpublished 
manuscripts,  and  Estes  has  also  been 
gracious  with  his  time  and  knowledge 
of  fossil  teiids. 

E.  Raymond  Hall,  Philip  S.  Hum- 
phrey, and  William  E.  Duellman  of  the 
University  of  Kansas  Museum  of  Nat- 
ural History  have  allowed  me  free  ac- 
cess to  the  collections  and  facilities'  of 
the  museum.  The  Panamanian  collec- 
tions made  by  Duellman  and  Charles 
W.  Myers  have  been  especially  useful. 
Field  work  was  financed  in  part  by  Na- 
tional Science  Foundation  Summer  Fel- 
lowships for  Graduate  Teaching  As- 
sistants ( 1965  and  1966 )  and  by  travel 
grants  and  traineeships  provided  by  the 
Committee  fof  Systematics  and  Evo- 
lutionary Riology  from  funds  granted 
by  the  National  Science  Foundation 
(Grant  No.  GR-4446X)  to  the  Univer- 
sity of  Kansas,  Dr.  William  A.  Clemens 
and  Dr.  George  W.  Byers,  principal  in- 
vestigators (1966-1968).  A  grant-in-aid 
from  the  Society  of  the  Sigma  Xi  pro- 
vided funds  for  illustrations,  which  were 
executed  by  Mrs.  Susann  Jacobshagen. 
Computer-time  was  made  available  by 
the  University  of  Kansas  and  Boston 
University  Computer  Centers. 

To  my  wife  Sandra,  I  owe  my  great- 
est thanks.  She  has  been  a  constant 
source  of  encouragement  and  support 
and  has  proven  to  be  an  able  assistant 
in  both  field  and  laboratory.  She  has 
cheerfully  endured  the  many  incon- 
veniences experienced  by  the  families  of 
graduate  students,  and,  with  the  realiza- 
tion that  the  debt  can  never  be  repaid, 
I  dedicate  this  work  to  her. 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


MATERIALS  AND  METHODS 

This  study  is  based  on  the  examina- 
tion of  over  3000  specimens  of  Ameiva, 
2848  of  which  were  examined  in  detail 
with  respect  to  morphology,  color  and 
pattern.  Samples  were  drawn  from 
throughout  the  ranges  of  Ameiva  fes- 
tiva,  A.  leptophrys,  A.  quadriUneata,  A. 
tindulata,  and  A.  chaitzami.  Onlv  Pan- 
amanian  specimens  of  Ameiva  ameiva 
were  examined,  because  to  include 
specimens  from  throughout  the  range 
of  the  species  would  have  necessitated 
examination  of  material  from  South 
America,  where  inadequate  geographic 
representation  of  samples  would  have 
made  any  conclusions  more  than  usually 
speculative.  Ameiva  ameiva  will  be  in- 
cluded in  a  projected  study  of  South 
American  members  of  the  genus  which 
will  be  completed  only  after  field  work 
necessary  for  the  accumulation  of  sam- 
ples representative  of  the  entire  South 
American  range  of  the  species  is  pos- 
sible. South  American  samples  of 
Ameiva  festiva  have  been  included  in 
the  present  study  because  the  range  of 
the  species  is  primarily  in  Central 
America. 

For  purposes  of  statistical  analysis,  I 
attempted  to  amass  samples  of  at  least 
30  specimens  (ideally  15  males  and  15 
females)  from  relatively  restricted 
localities.  This  attempt  was  only  par- 
tially successful  in  that,  with  the  excep- 
tions of  Guatemala  and  parts  of  Mexico, 
large  series  of  Ameiva  from  single  lo- 
calities are  not  available.  Of  necessity, 
samples  of  ten  or  more  have  been  used; 
for  characters  showing  sexual  dimor- 
phism (Table  1),  the  samples  were  still 
smaller.  Greater  success  was  achieved 
in  accumulation  of  samples  representa- 
tive of  the  ranges  of  the  species,  al- 
though lack  of  collections  from  critical 
areas  in  Panama  has  precluded  exact 
statements  of  variation  in  A.  ameiva 
and,  especially,  A.  leptophrys. 

In  each  of  the  species  accounts,  a 
short  synonymy,  statement  of  distribu- 
tion, diagnosis,  and  description  are  fol- 


lowed by  an  extensive  discussion  of 
infraspecific  geograpliic  variation,  eco- 
logical notes,  and  general  remarks.  A 
discussion  of  interspecific  relationships 
and  the  history  of  the  genus  Ameiva  in 
Middle  America  follows  the  species 
accounts. 

Characters 

In  the  interests  of  standardization  of 
methods,  a  definition  of  each  of  the 
characters  analyzed  in  this  study  fol- 
lows. Unless  stated  otherwise,  charac- 
ters are  defined  in  the  same  manner  as 
by  Smith  ( 1946 ) .  An  asterisk  (  * )  indi- 
cates a  character  analyzed  for  geo- 
graphic variation  by  Power's  (1970) 
modification  of  Gabriel's  ( 1964 )  Sums 
of  Squares  Simultaneous  Test  Proce- 
dure (STP).  Interpretation  of  the  re- 
sults of  STP  analysis  is  summarized  in 
the  section  on  statistical  procedures.  It 
will  be  noted  that  certain  characters 
were  not  taken  for  some  species,  where- 
as in  other  instances  a  character  was 
taken  but  not  analyzed  by  the  STP 
method. 

Snoiit-Vent  Length  (SVL). — The 
distance,  to  the  nearest  millimeter,  from 
the  tip  of  the  snout  to  the  anterior  mar- 
gin of  the  vent.  Maximum  observed 
values  for  males  and  females  of  each 
species  are  given.  Collections  seem  to 
be  biased  in  favor  of  larger  (male)  in- 
dividuals. Measurements  were  taken 
with  a  flexible  plastic  rule. 

Head  Length. — The  distance  to  the 
nearest  0.1  millimeter  from  the  tip  of 
the  snout  to  the  posterior  surface  of  the 
quadrate  bone.  The  latter  point  was 
found  by  piercing  the  tympanum  with 
one  tip  of  the  dial  calipers  and  hooking 
the  tip  behind  the  bone.  That  the  quad- 
rate was,  in  fact,  the  bone  encountered 
was  verified  by  x-ray  photography. 

^Supralahials. — The  total  number  of 
supralabials  on  both  sides  of  the  head. 
The  posterior  supralabial  is  herein  de- 
fined as  the  last  enUu'ged  scale  in  con- 
tact with  both  the  lip  and  a  subocular 
scale  (Fig.  la). 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


'^Infralabiah. — The  total  number  of 
infralabials  on  both  sides  of  the  head. 
The  posterior  infralabial  is  determined 
by  the  point  at  which  the  masseter 
muscle  and  associated  membranes  pass 
from  the  lower  to  the  upper  jaw. 

'* Terminal  Siiblabials. — This  charac- 
ter was  noted  for  Ameiva  festiva  only. 
Stuart  (1943:21)  utilized  the  number 
of  terminal  sublabials  to  distinguish  A. 
festiva  festiva  from  A.  /.  edwarchi.  I 
counted  the  sublabials  present  in  the 
position  of  the  two  terminal  sublabials 
of  a  "typical"  A.  /.  festiva,  totaling  the 
numbers  obtained  for  both  sides  of  the 
head.  By  this  method,  a  "typical"  A.  /. 
festiva  (sensu  Stuart)  has  four,  whereas 
A.  /.  echvardsi  has  six  or  more. 

'*  Supraoculars. — The  total  number 
of  supraoculars  on  both  sides  of  the 
head.  Supraoculars  are  numbered  from 
front  to  back  (3a-c,  Fig.  lb);  there  are 
three  or  four  on  each  side  of  most 
specimens  of  Ameiva  from  Middle 
America. 

'^Loreals. — The  total  number  of 
loreals  on  both  sides  of  the  head.  If 
more  than  one  loreal  is  present  on  one 
side  of  the  head,  the  second  is  smaller 
and  usually  located  at  the  angle  formed 
by  the  contact  between  supralabials 
and  suborbitals  anterior  to  the  eye  (3, 
Fig.  la).  Occasionally,  there  are  two 
loreals  about  equal  in  size,  one  dorsal  to 
the  other.  If  a  third  scale  is  present  in 
the  area  encompassed  by  the  loreal(s), 
it  is  counted  as  a  third  loreal  only  if  it 
is  one-half  the  size  of  the  smaller  of  the 
two  other  loreals  or  larger. 

^Degree  of  Circumorbital  Granule 
Contact  With  the  Frontal  (COF).— The 
degree  of  contact  by  the  circumorbital 
granular  scales  with  the  frontal  was 
coded  for  purposes  of  statistical  analysis 
as  follows:  1)  circumorbital  granules 
not  in  contact  with  frontal-frontopari- 
etal  suture  (Fig.  lb),  2)  circumorbital 
granules  in  contact  with  frontal-fronto- 
parietal  suture,  but  not  extending  past 
it  (Fig.  Ic),  3)  circumorbital  granules 
extending    forward    past    the    frontal- 


frontoparietal  suture  (Fig.  Id).  The 
coded  values  obtained  for  both  sides  of 
the  head  were  totaled.  Thus,  the  low- 
est possible  value  for  a  given  specimen 
is  two;  the  highest  is  six. 

"^Circumorbital  Pattern  (COP). — The 
position  of  the  anteriormost  circum- 
orbital granule  relative  to  the  supra- 
ocular scales  was  coded  as  follows:  1) 
granules  extending  to  the  level  of  the 
middle  of  the  first  supraocular  scale, 
2)  granules  extending  to  the  level  of 
the  suture  between  the  first  and  second 
supraocular  scales,  3)  granules  reach- 
ing the  level  of  the  middle  of  the  sec- 
ond supraocular  scale  (Fig.  Id),  4) 
granules  extending  to  the  level  of  the 
suture  between  the  second  and  third 
supraocular  scales  (Fig.  Ic),  5)  gran- 
ules extending  to  the  level  of  the  mid- 
dle of  the  third  supraocular  scale  (Fig. 
lb),  6)  granules  extending  to  the  level 
of  the  suture  between  the  third  and 
fourth  supraocular  scales  or,  if  there  is 
no  fourth  supraocular,  granules  not 
reaching  the  level  of  the  middle  of  the 
third  supraocular,  7)  granules  extend- 
ing to  the  middle  of  the  fourth  supra- 
ocular scale.  Coded  values  for  both 
sides  of  the  head  were  totaled  for 
analysis. 

'^Extent  of  the  Double  Row  of  Gran- 
ules Between  the  Supraocular  and  Su- 
perciliary Scales  (SO-SC). — This  char- 
acter was  coded  as  follows:  1)  single 
row  of  granules  between  the  super- 
ciliary and  supraocular  series  ( Fig.  Id ) , 
2)  double  row  of  granules  extending 
foi-ward  to  the  level  of  the  middle  of 
the  third  supraocular  scale,  3)  double 
row  of  granules  extending  forward  to 
the  suture  between  the  second  and  third 
supraocular  scales,  4)  double  row  of 
granules  extending  forward  to  the 
level  of  the  middle  of  the  second  supra- 
ocular (Fig.  Ic),  5)  double  row  of 
granules  completely  separating  the  su- 
perciliary scales  from  the  second  and 
third  supraocular  scales  (Fig.  lb). 
Coded  values  for  both  sides  of  the  head 
were  totaled  for  analysis. 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


B 


Fig.  I.  Details  of  head  and  preanal  scutellation  of  Ameiva.  (A)  Lateral  view  of 
head:  1. — Postnasal;  2. — Anterior  loreal;  3. — Posterior  loreal;  4. — Suborbital 
scale  in  contact  with  posterior  supralabial  (small,  rectangular  scale  below  4). 
(B,  C,  and  D)  Dorsal  view  of  head,  left  supraorbital  region:  1. — Superciliary 
series;  2. — Supraocular-superciliary  granules;  3A,  3B,  and  3C. — Anterior,  mid- 
dle and  posterior  supraocular  scales,  respectively;  4. — Prefrontal  scale;  5. — 
Frontal  scale;  6. — Frontoparietal  sca'e;  7. — Circumorbital  granules.     (E)   Pre- 


anal  region:     1.- 


-Anterior 
scale;  3.- 


(1st)    preanal 
—Terminal  row 


scale;    2. — Posterior    (last)    preanal 
of  ventral  scutes. 


Gular  Scales. — The  central  gular 
scales  are  small  and  irregular  in  ar- 
rangement, large  and  irregular,  or  large 
and  oriented  longitudinally  in  the  mid- 
line. Intermediate  conditions  exist,  but 
most  specimens  can  be  assigned  without 


difRculty.  If  enlarged  midgular  scales 
are  present,  the  smaller  scales  around 
them  may  become  gradually  smaller  to- 
ward the  periphery  of  the  gular  region 
or  may  become  gradually  smaller 
laterally    and    anteriorly    but    abruptly 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


9 


very  small  posteriorly.  If  the  latter  is 
true,  the  gular  region  is  distinctly  di- 
vided into  pregular  and  postgular  areas 
(see  Fig.  5c,  A.  leptophrys) . 

MesoptychiaJ  Scales. — The  meso- 
pt\'ehial  scales  are  small  and  irregular, 
continuous  with  the  posterior  gular 
scales,  or  they  may  form  a  band  of 
abruptly  enlarged  scales  across  the 
throat. 

"^Granules  Around  the  Body  (GAB). 
— The  number  of  granules  around  the 
body  was  shown  by  Zweif el  ( 1959 )  to 
be  of  diagnostic  importance  in  distin- 
guishing species  of  Cnemidophorus,  and 
this  character  has  been  used  extensively 
by  students  of  that  difficult  genus.  In 
most  cases,  the  character  has  been  given 
as  "scales  around  midbody"  (I  believe 
the  term  "granules"  to  be  more  de- 
scriptive), but  rarely  is  there  a  state- 
ment as  to  what  "midbody"  refers.  This 
is  important  inasmuch  as  the  character 
is  intended  to  be  a  measure  of  granule 
size,  and  the  granules  vary  in  size  over 
the  length  of  the  body.  In  this  study, 
GAB  counts  were  made  starting  at  a 
point  adjacent  to  the  15th  transverse 
row  of  enlarged  ventral  scutes,  the  first 
transverse  row  of  ventral  scutes  is  taken 
as  the  first  row  extending  entirely  across 
the  chest  behind  the  gular  fold. 

*" Paravertebral  Granules  (PV). — The 
number  of  granules  between,  and  in- 
cluding, the  paravertebral  stripes  of 
Ameiva  leptophrys,  quadriUneata,  un- 
dulata,  and  chaitzami. 

'^Vertebral  Stripe  (VS). — The  num- 
ber of  granules  included  within  the 
vertebral  light  stripe  of  Ameiva  festiva. 

PV  and  VS  counts  were  omitted  for 
A.  ameiva  because  the  stripes  are  lack- 
ing or  very  faint  on  most  adults  of  that 
species. 

""PV/GAB  Ratio.— The  ratio  of  para- 
vertebral granules,  as  defined  above,  to 
granules  around  the  body  was  calcu- 
lated. 

^Granules  Occiput  to  Rump  (GOR). 
— The  number  of  granular  scales  be- 
tween the  enlarged  occipital  scales  and 


the  large,  keeled  dorsal  caudal  scales 
was  counted  for  all  species.  Although 
there  is  usually  a  high  correlation  be- 
tween values  obtained  for  this  count 
and  those  for  granules  around  the  body 
(correlation  coefficient=0.75  and  0.54 
for  males  and  females  of  A.  leptophrys, 
respectively),  GOR  sometimes  shows 
sexual  dimorphism  and  occasionally  is 
an  easier  and  more  accurate  count  than 
GAB.  The  latter  is  especially  true  in 
poorly  preserved  specimens  that  have 
been  coiled  or  that  have  numerous  folds 
in  the  skin  of  the  flanks.  With  experi- 
ence, neither  GAB  nor  GOR  counts  are 
difficult  or  time-consuming  to  make, 
and  error  is  not  excessive  (2-3%  based 
on  recounts  of  the  same  specimens). 
The  most  suitable  technique  for  count- 
ing dorsal  granules  seems  to  be  to  use 
a  fine  insect  pin  as  a  pointer  and  marker 
while  counting  granules  beneath  a  wide- 
field,  binocular  microscope.  Ideally,  the 
specimens  should  be  slightly  dry,  since 
fluid  between  the  granules  results  in 
glare  and  makes  counting  difficult. 

''GAB /GOR  Ratio.— This  ratio  was 
calculated  for  all  species. 

Prebrachials. — The  number  of  rows 
of  prebrachials  was  noted. 

Posthrachials. — T  h  e  postbrachials 
are  enlarged  or  not. 

Preantebrachials. — The  number  of 
rows  of  preantebrachial  scales  was  re- 
corded. 

Prefe moral  Scales. — Prefemoral  scales 
are  defined  herein  as  the  large  scales 
on  the  anterior  and  ventral  surfaces  of 
the  upper  hind  leg  as  far  back  as  the 
femoral  pore  series.  Thus  defined,  pre- 
femorals  include  the  infrafemoral  and 
prefemoral  scales  of  Smith  (1946:28). 
The  number  of  rows  was  counted. 

Infratibial  Scales. — The  number  of 
rows  of  infratibial  scales  was  counted. 

The  number  of  rows  of  preante- 
brachials, prebrachials,  prefemorals  and 
infratibials  is  variable  throughout  the 
length  of  the  series  on  a  single  speci- 
men. In  many  specimens,  there  is  a 
greater  number  of  scales  medially  than 


10 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


proximally.    This  variation  is   discussed 
for  each  species. 

'*  Femoral  Pores. — The  number  of 
femoral  pores  was  recorded;  counts 
given  in  the  text  and  tables  are  for  the 
total  number  of  pores  on  both  legs. 

'^Subdigital  Lamellae. — The  number 
of  subdigital  lamellae  on  the  anterior 
margin  of  the  fourth  (longest)  toe  of 
the  left  hind  foot  was  recorded.  The 
starting  point  for  this  count  is  the 
tubercule  immediately  proximal  to  the 
enlarged  tubercule  at  the  base  of  the 
first  phalanx.  If  there  is  no  tubercule 
in  this  position,  the  enlarged  basal 
terminal  is  itself  used  as  the  starting 
point. 

'*  Longitudinal  Rows  of  Ventral 
Scutes. — The  number  of  rows  of  en- 
larged ventral  scutes  was  counted  be- 
tween the  gular  fold  and  the  first  pre- 
anal  scale,  as  defined  below.  The  first 
row  is  the  anteriormost  row  extending 
entirely  across  the  chest  posterior  to 
the  gular  fold. 

'^Transverse  Rows  of  Ventral  Scutes. 
— The  number  of  transverse  rows  of 
ventral  scutes  was  counted  at  the  level 
of  the  15th  longitudinal  row  of  ventrals. 
The  outer  row  on  each  side  is  reduced 
in  size  in  some  specimens. 

'^ Total  Preanal  Scales. — The  number 
of  preanal  scales  from  just  posterior  to 
the  tenninal  row  of  longitudinal  ven- 
tral scutes  to  the  enlarged  scale(s)  pre- 
ceding the  vent  were  counted.  The 
first  preanal  scale  is  usually  single,  lies 
between  the  proximal  femoral  pores  of 
each  leg,  and  is  distinguished  from  the 
last  row  of  ventrals  by  its  unpaired  con- 
dition (1,  Fig.  le).  Smith  and  Laufe 
(1946:20)  encountered  difficulty  in  de- 
termining which  preanal  scale  is  termi- 
nal because  the  preanal  scales  occa- 
sionally grade  in  size  into  the  granular 
scales  immediately  anterior  to  the  vent. 
When  in  doubt  as  to  which  preanal  or 
row  of  preanals  is  terminal,  I  have  sim- 
ply recorded  the  last  row  having  three 
or  fewer  enlarged  scales.  In  most  cases, 
this  is  obviously  the  terminal  row  any- 


way. In  those  lizards  with  a  single 
terminal  preanal,  the  scales  lateral  to  it 
are  usually  very  much  smaller  and 
create  no  confusion  as  to  whether  or 
not  they  are  part  of  the  terminal  pre- 
anal series. 

'^Termitml  Preanal  Scales. — The 
number  of  scales  in  the  temiinal  posi- 
tion in  the  preanal  series  was  deter- 
mined by  the  method  outlined  above 
(2,  Fig.  le). 

''Scales  Around  the  Tail  (SAT).— 
The  number  of  scales  around  the  tail 
was  counted  at  the  level  of  the  15th 
verticil  of  caudal  scales;  the  first  verticil 
was  taken  as  that  row  of  caudal  scales 
which  first  completely  encircles  the  tail 
posterior  to  the  vent. 

Various  aspects  of  color  and  color 
pattern  were  noted  on  a  sample-by-sam- 
ple basis.  Recurring  pattern  compo- 
nents noted  were  as  follows: 

Vertebral  Stripe. — A  narrow  stripe 
originating  on  the  snout  or  occiput  and 
terminating  on  the  tail  unless  com- 
pletely or  partially  lost  in  adults. 

Middorsal  Field  or  Area. — A  broad 
stripe,  containing  a  vertebral  stripe  or 
not,  originating  on  the  occiput  and 
terminating  on  the  tail,  bordered  later- 
ally by  the  dorsolateral  light  stripes,  if 
present,   or  the   dorsolateral   dark  field. 

Dorsolateral  Light  Stripe. — A  nar- 
row, light-colored  stripe  originating  on, 
or  posterior  to,  the  superciliary  rows 
and  terminating  on  the  tail  and  which 
may  be  entirely  or  partially  lost  in 
adults. 

Secondary  Stripe. — A  narrow,  dark- 
colored  stripe,  the  lateral  border  of 
which  is  the  dorsolateral  light  stripe,  if 
present,  or  the  dorsolateral  dark  field 
and  the  medial  border  is  the  lighter  part 
of  the  middorsal  field  in  which  it  lies. 

Dorsolateral  Dark  Field. — A  dark, 
often  black  or  dark  brown,  area 
bounded  above  by  the  dorsolateral  light 
stripe,  if  present,  or  the  middorsal  field 
and  below  by  the  ventrolateral  light 
stripe,  if  present,  or  the  ventrolateral 
field;    originating   on   the   loreal   scales, 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


11 


terminating  on  the  tail  and  interrupted 
by  the  eye;  may  be  indistinct  or  lost  in 
adults. 

Ventrolateral  Light  Stripe. — A  nar- 
row, light-colored  stripe  originating  at 
the  posterior  margin  of  the  eye  or  pos- 
terodorsal  margin  of  the  ear  and  ex- 
tending along  the  body  to  the  groin, 
usually  continuing  behind  the  leg  and 
slightly  onto  the  tail. 

Ventrolateral  Field  or  Area. — Ven- 
trolateral one-half  or  two-thirds  of  the 
flanks  bordered  above  by  the  ventro- 
lateral light  stripe,  if  present,  or  the 
dorsolateral  dark  field  and  below  by  the 
enlarged  ventral  scutes;  distinguished 
from  the  dorsolateral  dark  field  by 
its  generally  lighter  color;  may  contain 
irregularly  arranged  spots,  blotches 
and/ or  bars  or  regularly  spaced  vertical 
bars. 

Lateral  Bars  or  Blotches. — Regular 
blotches  in  the  dorsolateral  dark  field 
or  bars  on  the  flanks. 

Head  scutellation  is  extremely  vari- 
able in  Ameiva.  This  is  especially  evi- 
dent in  Ameiva  leptophrys  and,  to  a 
lesser  extent,  in  festiva  and  quadriline- 
ata.  The  supraoculars  of  specimens  of 
quadrilineata  from  the  Bocas  del  Toro 
region  of  Panama  are  so  fragmented  as 
to  preclude  counting  them.  There  may 
be  three  or  four  supraoculars  per  side 
in  any  of  the  Middle  American  species 
of  Ameiva,  and  specimens  of  all  species 
are  known  which  have  four  on  one  side, 
three  on  the  other.  The  number  of 
supraoculars  is  thought  to  be  sufficiently 
constant  in  Cnemidophorus  that  the 
number  is  used  to  distinguish  species 
groups  ( Burt,  1931 ) .  The  posterior  dor- 
sal head  scales  of  Ameiva  leptophrys 
(Fig.  7)  and  some  festiva  are  variously 
fragmented,  so  that  in  extreme  cases 
parietal  scales  are  not  recognizable,  and 
the  frontoparietals  and  frontal  are  only 
slightly  less  disrupted.  It  might  seem 
that  such  variability  in  the  posterior 
head  scales  of  Ameiva  renders  them 
useless  for  taxonomic  purposes,  but  it 
seems  that  the  amount  of  variation  is  a 


useful  taxonomic  character.  For  exam- 
ple, the  presence  of  small,  irregular 
scales  separating  the  inteiparietal  from 
the  parietals  and  frontoparietals  can  be 
used  to  distinguish  leptophrys  from 
other  Middle  American  Ameiva. 

Statistical  Procedures 

Statistical  summaries  of  all  species 
examined  comprise  Appendix  A. 

Power's  ( 1970 )  modification  of  Gab- 
riel's (1964;  see  Gabriel  and  Sokal, 
1969)  Sums  of  Squares  Simultaneous 
Test  Procedure  ( STP )  provides  a  test 
of  significance  or  non-significance  of 
differences  among  sample  means  and  is 
employed  only  after  an  overall  analysis 
of  variance  demonstrates  significant  dif- 
ferences among  the  means.  Samples  are 
then  ranked  in  decreasing  order  of  their 
means  and  sums  of  squares  calculated 
by  sequentially  adding  means  (be- 
ginning with  the  largest)  until  a  maxi- 
mal non-significant  subset  is  delimited. 
The  procedure  is  then  repeated,  delet- 
ing one  or  more  of  the  larger  means  in 
the  previously  defined  subset,  until 
another  maximal  non-significant  subset 
is  described.  This  procedure  is  repeated 
until  all  samples  have  been  included  in 
at  least  one  such  subset.  A  constant 
probability  level  of  0.05  was  used  in  the 
study,  as  recommended  by  Power 
( 1970 ) .  Calculations  were  performed 
by  the  University  of  Kansas  Computa- 
tion Center  on  a  GE-625  computer,  and 
at  Boston  University  on  an  IBM  360-40. 

In  addition  to  STP,  a  number  of 
statistics  were  calculated  for  each  spe- 
cies and  each  sample.  Certain  of  these, 
along  with  the  STP  results,  are  sum- 
marized in  the  tables  accompanying 
each  species  account.  Information  per- 
taining to  the  total  number  of  femoral 
pores  of  Panamanian  Ameiva  ameiva 
will  serve  as  an  example  for  the  inter- 
pretation of  STP  results  (see  Appendix 
A).  The  first  column  of  numbers  in- 
cludes the  locality  code,  keyed  to  the 
range  map  of  Panamanian  A.  ameiva 
(Fig.  2).    The  localities  are  ordered  in 


12 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


TABLE   1. — Occurrence  of  sexual  dimorphism   among  Middle  American  Atneiva  as  determined 
by  STP  analysis.    A,  ameiva;   L,   Icptophnjs;   Q,   quadrilineata;    F,   festiva;    U,    undulata;    C,   chait- 

:ami:  N,  not  tested. 


Species 

Character  ~A      L      Q      F      U      C~  Character 

Supralabials  —      +      —      +      +      —  PV/GAB 

Infralabials  ______  GOR 

Supraoculars         ______  GAB/GOR 

COF  values  +_____  SAT 

COP  \'alues  ______  Total  Preanals 

SO-SC  values        N      —      N      N      —      —  Terminal  Preanals 

Loreals  N      —      N      N      —      —  Subdigital  Lamellae 

Sublabials  N      N      N      +      N       N  Trans.  Rows  of  Ventrals 

GAB  _      _      _      _^      _      _  Long.  Rows  of  Ventrals 

PV  N      —      —      +      +      —  Femoral  Pores 


Species 


Q 


u 


N 


-      -      +      - 


+ 


+ 


+ 

— 

+ 

— 

+ 

— 

— 

— 

+ 

— 

+ 

— 

— 

+ 

+ 

+ 

+ 

— 

+ 

+ 

+ 

+ 

— 

— 

— 

— 

+ 

— 

— 

— 

+    +    +    +    + 


terms  of  tlie  decreasing  magnitude  of 
their  means,  given  in  column  two.  Fol- 
lowing the  mean  values  are  the  standard 
deviation  of  the  mean,  sample  size  (in 
parentheses),  and  observed  range.  To 
the  right  of  each  block  of  statistics  are 
the  STP  results,  appearing  as  vertical 
lines.  There  are  no  STP  results  given  if 
the  overall  analysis  of  variance  indi- 
cated no  significant  differences  among 
the  sample  means,  and  in  these  in- 
stances the  samples  are  ordered  by  their 
locality  numbers  irrespective  of  their 
mean  values.  In  the  example,  samples 
6  and  5,  with  means  of  35.0  and  34.1, 
respectively,  form  the  first  maximal  non- 
significant subset  and  are  so  indicated 
by  the  left-most  line.  Localities  5,  7,  3, 
2  and  1  constitute  the  second  subset 
(middle  line),  and  localities  7,  3,  2,  1 
and  4  are  the  third  (right  line).  The 
mean  for  locality  6  is  significantly  dif- 
ferent from  all  other  means,  except  that 
of  locality  5  as  indicated  by  the  obser- 
vation that  the  mean  for  locality  6  ap- 
pears in  no  subsets  containing  means 
other  than  that  of  locality  5.  Likewise, 
locality  5  is  significantly  different  from 
locality  4.  Localities  5  and  7  are  not 
statistically  different,  although  it  would 
appear  so  if  only  the  third  subset  were 
considered,  but  in  this  subset  locality  5 
was    omitted    from    the    comparison    in 


order  to  allow  other  means  into  the  sub- 
set at  the  lower  end  of  the  scale.  Locali- 
ties 5  and  7  are  from  the  same  statistical 
population,  because  they  are  both  in- 
cluded in  the  second  subset. 

Statistical  significance  must  be  in- 
terpreted with  caution.  One  cannot 
infer  gene  pool  similarities  from  knowl- 
edge of  a  single  character;  for  example, 
it  would  be  improper  to  consider  liz- 
ards from  central  Mexico  and  Costa 
Rica  as  belonging  to  the  same  biological 
population,  even  if  they  are  shown  to 
belong  to  the  same  statistical  popula- 
tion. It  is  well  known  that  phenotypic 
expression  of  the  genotype  can  be  modi- 
fied by  environmental  pressures  and 
similarities,  or  differences  in  a  particu- 
lar character  among  samples  from  di- 
vergent localities  simply  may  be  an 
expression  of  this  pressure  on  a  rela- 
tively plastic  genome. 

Sexual  Dimorphism 

Presence  or  absence  of  sexual  dimor- 
phism for  individual  characters  was  de- 
termined by  analysis  of  variance.  This 
was  done  for  each  locality  sample  of 
Ameiva  festiva  and  undulata,  for  all 
males  against  all  females  of  A.  ameiva, 
chaitzami,  and  leptophrys,  and  for  At- 
lantic slope  males  versus  Atlantic  slope 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


13 


females  and  Pacific  slope  males  versus 
Pacific  slope  females  of  quadrUineata. 
If  any  sample  as  so  defined  was  sevually 
dimorphic  for  a  character,  the  entire 
If  any  sample  as  so  defined  was  sexually 
dimorphic  for  that  character  for  pur- 
poses of  statistical  analysis.  Presence  or 
absence  of  sexual  dimorphism  for  most 
of  the  characters  utilized  in  this  study  is 
indicated  in  Table  1. 

Sources  of  Specimens  Examined 

A  complete  list  of  specimens  exam- 
ined, along  with  locality  data  for  each, 
is  included  in  Appendix  B.  The  follow- 
ing abbreviations  have  been  used  in 
referring  to  individual  specimens  and 
indicate  the  museum  or  private  collec- 
tion in  which  the  specimens  are  housed. 

ANSP  Academy   of   Natural   Sciences   of 

Philadelphia 

AMNH  American  Museum  of  Natural 

History 

ARP  A.  R.  Philips  Collection 

(University  of  Arizona) 


BMNH 

British  Museum   (Natural  History) 

FMNH 

Field  Museum  of  Natural  History 

JRM 

John  R.  Meyer  (Private 

Collection) 

LACM 

Los  Angeles  County  Museum 

MCZ 

Museum  of  Comparative  Zoology 

MNHN 

Museum   National  d'Histoire 

Naturelle,  Paris 

RDS 

Richard  D.  Sage  (Private 

Collection) 

TCWC 

Texas  Cooperative  Wildlife 

Collection 

USNM 

United  States  National  Museum 

UCR 

Universidad  de  Costa  Rica 

UCLA 

University  of  California  at  Los 

Angeles  (Specimens  now 

housed  in  the  Los  Angeles 

County  Museum) 

UIMNH 

University  of  Illinois  Museum  of 

Natural   History 

KU 

University   of  Kansas   Museum  of 

Natural   History 

UMMZ 

University  of  Michigan   Museum 

of  Zoology 

USC(CRE) 

University   of   Southern   California 

(Costa  Rican   Expedition) 

UU 

University  of  Utah 

ZMB 

Zoologisches   Museum,   Berlin 

SYSTEMATIC  ACCOUNT  OF  THE  SPECIES 

I    have    recognized    six    species    of  KEY  TO  THE  MIDDLE  AMERICAN 

Ameiva  in  Middle  America.   All  of  these  SPECIES  OF  AMEIVA 

are  easily  recognizable  in  the  field  by  ,    „                   i       i       .     t     i                r 

the  experienced  student,  but  are  easily  ^-  ^^"   ^?   *^^^^^^   longitudmal   rows   of 

confused  when  dealing  with  preserved  ^^^^''''^  /^'"^f^;    "^^^optychial    scales 

specimens.     As    is    often    the    case,    the  subequal;  adults  spotted    _. ..  amewa 

species  cannot  be  distinguished  readily  Eight    longitudinal    rows    of    ventral 

on  the  basis   of  single   characters,   and  scutes;    transverse    row    of    enlarged 

identification    of    preserved    specimens  mesoptychial        scales;        principally 

can  be  difficult  even  if  a  suite  of  char-  striped   2 

acters  is  used.    The  following  artificial  2.  Posterior    gular    scales    smaller    than 

key  should  be  used  in  conjunction  with  anterior  gular  scales                              3 
the  tables  in  the  analyses  of  geographic 

variation   and   with  the   descriptions   of  Anterior    and    posterior    gular    scales 

color  and  pattern  variation.   No  attempt  subequal  m  size  4 

has  been  made  to  place  the  key  into  a  3.  Postnasal  scales  not  in  contact  with 

phylogenetic    framework.      Fortunately,  prefrontals;  lateral  parietal  and  fron- 

the  situation  is  not  so  bad  as  that  which  toparietal  scales  separated  by  one  or 

caused  Stuart  (1955),  in  discussing  his  more  rows  of  small,  irregular  scales; 

key  to  Guatemalan  Anolis,  to  comment  midgular  scales  greatly   enlarged,   ir- 

that  ".  .  .  the  worker  who  knows  what  regular  in  arrangement        leptophrys 

species   he   has   before   him   should   ex-  Postnasal  scales  in  contact  with  pre- 

perience  few  difficulties  in  its  use."  frontals;    parietal    and    frontoparietal 


14 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


82° 


78° 


50      100 


KILOMETERS 


82° 


78° 


Fig.  2.    Map  showing  locality  records  for  Ameiva   ameiva  in  Panama.    To  avoid  crowding  of  sym- 
bols, some  localities  have  been  omitted.    Samples  used   in   statistical   analyses  were    ( 1 )    Vicinity  of 
Panama  City,  (2)   Vicinity  of  Colon,   (3)   Vicinity  of  Niievo  Gorgona,   (4)   Isla  San  Jose,   (5)   Prov. 
Herrera,    (6)    Prov.   Los  Santos,   and    (7)   Vicinity  of  Cerro  Campana. 


.scales  in  contact;  midgular  scales 
slightly  to  moderately  enlarged,  ir- 
regular or  in  a  single  longitudinal 
row   quadrilineata 

Midgular  scales  much  enlarged,  ir- 
regular in  arrangement;  narrow, 
light-colored  vertebral  stripe  present 
except  in  large  adults  festiva 

Midgular  scales  much  enlarged,  in 
longitudinal  arrangement  or  not, 
moderately  enlarged  and  irregular  in 
arrangement;  no  vertebral  stripe      .  5 

Small  (maximum  observed  SVL  85 
mm  for  males,  75  mm  for  females); 
paravertebral  stripe  narrow  (mean 
PV  =  37.1);  dorsolateral  blotches  in 
males  fused  to  dorsolateral  light 
stripe  chaitzami 

Moderately  large  (maximum  ob- 
served SVL  129  mm  for  males.  111 
for      females ) ;      paravertebral 


mm 


stripe  broad  (mean  PV  =  47.4  for 
males,  46.4  for  females);  dorsolateral 
blotches  in  males,  if  present,  do  not 

fuse  with  dorsolateral  light  stripe  

undulata 


Ameiva  ameiva  (Linnaeus) 

Lacerta  ameiva  Linnaeus,  Systema  naturae,  ed. 
12,  1:362,  1766.— America. 

Cnemidophorus  praesignis  Baird  and  Girard, 
Proc.  Acad.  Nat.  Sci.  Philadelphia,  6:129, 
1852  [Syntypes:  USNM  5519  and  UMMZ 
3823,  Chagres,  Panama;  C.  B.  Adams,  col- 
lector]. 

Ameiva  praesignis — Cope,  Proc.  Acad.  Nat. 
Sci.  Philadelphia,  14:67,   1862. 

Ameiva  praesigna — Bocourt,  Mission  Scienti- 
fique  au  Mexique  et  dans  I'Amerique  Cen- 
trale;  fitudes  sur  les  reptiles,  livr.  3,  p.  265, 
pi.  20b,  fig.  9-9d,  1874. 

Ameiva  surinamensis — Boulenger,  Catalogue  of 
the  Lizards  in  the  British  Museum  (Natu- 
ral History),  2nd  Ed.,  vol.  2,  p.  352,  1885 
(part).  Giinther,  Biologia  Central!- Ameri- 
cana; Reptilia  and  Batrachia,  p.  22,  pi.  19, 
1885   (part). 

Ameiva  ameiva  praesignis — Barbour  and 
Noble,  Bull.  Mus.  Comp.  Zool.,  59(6):468, 
1915.  Dunn,  Proc.  Acad.  Nat.  Sci.  Phila- 
delphia, 92:113,  1940. 

Distribution  in  Tanaind. — Grasslands 
and  savannas  of  the  Pacific  slopes  near 
Chepo,  Panama,  to  David,  Chiriqui,  and 
in  open  areas  across  the  Canal  Zone  to 
Colon  and  Portobelo,  Colon,  on  the 
Atlantic  side  (Fig.  2). 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


io 


Diagnosis. — Ameiva  ameiva  can  be 
distinguished  from  all  other  Middle 
American  Ameiva  by  the  following  com- 
bination of  characters:  Size  large,  max- 
imum observed  SVL  197  mm  for  males 
and  157  mm  for  females;  mesoptychial 
scales  small,  irregular;  scales  around  the 
tail  more  than  30;  ventrals  at  midbody 
usually  more  than  eight;  supraoculars 
usually  eight;  color  pattern  in  adult 
males  emphasizes  spots,  rather  than 
longitudinal  stripes  or  lateral  bars  and 
blotches. 

Description. — Maximum  observed 
SVL  197  mm  for  males,  157  mm  for 
females;  11-16  (12.9)  supralabials;  8-12 
(10.1)  infralabials;  6-10  (8.0)  supraocu- 
lars; COF  values  2-6  (2.1),  but  note 
geographic  variation  discussed  below; 
COP  values  8-16  (10.6);  nostril  in  pre- 
nasal-postnasal  suture;  prefrontals  in 
contact  with  postnasals;  loreals  2;  GAB 
128-204  (146.0),  204  an  exceptional 
value — the  second  highest  value  re- 
corded was  169;  GOR  229-324  (287.8) 
for  males,  225-394  (295.5)  for  females; 
GAB/GOR  0.45-0.68  (0.51)  for  males, 
0.41-0.67  (0.49)  for  females;  SAT  34-45 
(39.5);  total  preanal  scales  7-16  (10.8); 
terminal  preanal  scales  2-3  (2.4)  for 
males,  2-3  (2.2)  for  females;  one  row  of 
enlarged  prebrachials  with  irregularly 
arranged  accessory  rows  on  either  side; 
postbrachials  slightly  to  moderately  en- 
larged, irregularly  arranged;  preante- 
brachials  enlarged,  one  (occasionally 
two)  rows;  prefemoral  scales  enlarged, 
in  four  or  more  rows;  infratibial  scales 
in  three  rows;  femoral  pores  25-41 
(32.4);  subdigital  lamellae  26-36  (30.9); 
longitudinal  rows  of  ventral  scutes  28- 
33  (31.2);  transverse  rows  of  ventral 
scutes  6-12  (11.0);  dorsal  caudal  scales 
weakly  keeled.  For  details  of  scutella- 
tion  see  figure  3. 

Color  and  Pattern  in  Alcohol. — Juve- 
nile males:  Top  and  upper  half  of  side 
of  head  gray-green  to  brown  with  black 
markings  immediately  anterior  to  eye; 
lower  one-half  of  head  from  eye  to  post- 
nasal  white;    supralabials   black    above, 


Fig.  3.  Head  and  preanal  scutellation  of 
Ameiva  ameiva  from  Panama  (KU  108258): 
( A )  Lateral  view  of  head,  ( B )  Dorsal  view  of 
head,  (C)  Ventral  view  of  head.  X  L3. 
(D)   Preanal  region.    X   2. 


white  below  to  lip;  infralabials  black 
below,  white  above  to  lip;  chin  and 
gular  region  white  or  cream  with  small, 
scattered  flecks  and  often  a  transverse 
row  of  black  flecks  between  larger  me- 
dian gulars  and  smaller  posterior  gulars; 
middorsal  area  between  occiput  and  tail 
black  or  dark  brown  with  brown  or 
gray-green  middorsal  stripe,  anterior 
one-fifth  of  which  as  broad  as  paraver- 
tebral field,  gradually  narrowing  to  a 
width  of  about  10  granules  near  base  of 
tail  and  with  irregular  margins  through- 
out; cream  or  gray-white  paravertebral 
stripes  from  (and  including)  supercili- 
ary series  well  onto  tail;  dorsolateral 
dark  stripe  black  with  or  without  a 
brown  median  stripe  and  with  many 
small,  white  or  blue- white  spots;  lateral 
light  stripe  cream,  originating  at  dorso- 
posterior  margin  of  ear  and  extending 


16 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


to  point  of  insertion  of  hind  leg  and 
onto  anterior  surface  of  thigh,  beginning 
again  on  posterior  surface  of  thigh  and 
extending  well  onto  tail;  area  between 
lateral  light  stripe  and  ventral  scutes 
black  with  small,  white  spots  (some  in- 
distinct); venter  white  or  cream  medial- 
ly with  lateral  scutes  and  (occasionally) 
chest  marked  with  black  Hecks;  dorsal 
surfaces  of  fore  and  hind  limbs  black 
with  blue,  blue-white  or  pale  brown 
spots  and  blotches,  this  color  and  pat- 
tern grading  to  uniform  cream  or  white 
ventrally  with  or  without  black  Hecks; 
pre-  and  postfemoral  light  stripes  cream. 
Tail  brown  with  black  or  blue-gray 
flecking  above,  flecks  sometimes  ar- 
ranged to  give  the  appearance  of 
whorls;  white  dorsolateral  stripe;  sides 
black  with  white  flecking;  ventrolateral 
light  stripe  white,  blending  into  white 
or  cream  ventral  coloring;  subcaudal 
area  sometimes  with  black  blotches; 
short,  black  postanal  stripes  extending 
from  each  corner  of  the  vent. 

Through  ontogeny  there  is  a  gradual 
loss  of  stripes  and  an  increased  empha- 
sis on  spotting,  so  that  in  adult  males 
the  dorsal  pattern  is  one  of  transverse 
rows  of  white  or  blue-white  spots  on  a 
black  background  with  indistinct  gray- 
green  or  pale  brown  areas  between  the 
rows  of  spots  (Fig.  4).  Anteriorly,  the 
pattern  breaks  down  altogether,  leaving 
only  fine,  light  brown  reticulations  on  a 
black  background.  This  pattern  carries 
over  onto  the  dorsal  surfaces  of  the 
forelimbs  and  the  granules  of  the  side 
of  the  head.  The  hind  limbs  become 
black  with  blue  flecks,  again  sometimes 
arranged  to  give  the  appearance  of 
whorls.  Ventrally,  the  body,  tail  and 
hind  limbs  are  blue;  the  chin,  gular 
region  and  forelimbs  white  or  cream 
with  black  flecks.  In  adult  males  the 
chin,  gular  region,  and  the  forepart  of 
the  chest  may  be  pale  orange.  Males 
from  eastern  Panama  retain  a  bright 
orange  or  reddish-tan  vertebral  stripe  as 
adults. 

Females:    Juvenile  females  resemble 


juvenile  males,  but  there  is  strong  sex- 
ual dimorphism  in  adult  color  patterns 
( Fig.  4 ) .  Adult  females  retain  longi- 
tudinal stripes,  and  spotting  does  not 
become  as  pronounced  as  in  adult 
males.  Adult  females  have  brownish- 
white  to  white  dorsolateral  and  lateral 
stripes  bounded  by  black.  Middorsally, 
there  may  be  a  remnant  of  the  vertebral 
stripe  (this  is  always  true  of  specimens 
from  eastern  Panama)  with  the  remain- 
ing area  mottled  gray-green  or  brown 
on  black  with  scattered  white  spots.  A 
short  cream  stripe  originating  at  the 
posteroventral  corner  of  the  ear  and 
ending  at  the  shoulder  may  be  present. 
The  dorsolateral  dark  stripe  is  black 
with  white  spots  and  may  contain  a  dif- 
fuse, brown  secondary  stripe.  Between 
the  lateral  light  stripe  and  the  enlarged 
ventral  scutes  there  is  a  broad,  light 
brown  stripe.  Some  females  have  no 
spotting  at  all,  the  areas  between  the 
paravertebral  and  lateral  light  stripes 
appearing  uniformly  green  or  pale 
brown.  All  combinations  of  plain  and 
spotted  patterns  in  middorsal,  dorsolat- 
eral and  ventrolateral  fields  may  be 
found. 

Color  and  Pattern  in  Life. — The  fol- 
lowing description  of  color  in  life  of 
Atneiva  ameiva  is  quoted  (with  modi- 
fications to  eliminate  abbreviations) 
from  the  field  notes  of  C.  W.  Myers  and 


Fig.  4.  Ontogenetic  Change  and  Variation  in 
Pattern  of  Ameiva  ameiva  from  Panama. 
Males:  (A)  KU  107521,  SVL  68  mm,  from 
Guanico,  Los  Santos  Prov.;  (B)  KU  107542, 
107  mm,  from  Pese,  Herrera  Prov.;  (C)  KU 
107532,  160  mm,  from  Guanico,  Los  Santos 
Prov.  Females:  (D)  KU  76106,  45  mm,  from 
S  Slope  Cerro  Campana,  Panama  Prov.;  (E) 
KU  95546,  92  mm,  from  the  Rio  Salado, 
Chiriqui  Prov.;  (F)  KU  107524,  132  mm,  from 
Guanico,  Los  Santos  Prov.;  (G)  KU  108259, 
150  mm,  from  Juan  Mina,  Chagres  River, 
Canal  Zone.  Small  males  resemble  small  fe- 
males as  exemplified  by  KU  76106  (D  above). 
KU  108259  (G  above)  is  included  to  illustrate 
the  presence  of  a  well  defined  vertebral  stripe 
in  adult  lizards  from  eastern  Panama  ( see 
te.xt  for  discussion). 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


17 


18 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


pertains  to  specimens  collected  at 
Guanico  Arriba,  Los  Santos,  Panama: 
(KU  107521-3;  juveniles)  ".  .  .  Mid- 
dorsum  brown  with  darker  brown  marks 
and  bounded  by  pale  yellow  dorsolat- 
eral stripe.  Sides  rich  brown  or  blackish 
brown,  with  pale  tan  dots,  bordered 
below  by  a  creamy  stripe."  ( KU  107524- 
7;  adult  females)  ".  .  .  Dorsolateral  and 
lateral  stripes  remain  conspicuous,  and 
the  dorsum  takes  on  a  greenish  tinge  or 
not;  a  variable  amount  of  tan  to  yellow 
dorsal  spots.  Throat  light  orange,  and 
undersides  of  arms  orange  tinged  to 
light  dull  red;  rest  of  venter  white  in 
one  and  pale  blue  in  the  others;  rear  of 
the  thighs  white  or  pinkish  white.  Iris 
light  brown."  (KU  107528-31;  adult 
males)  ".  .  .  Body  stripes  absent  or  in- 
conspicuous and  body  with  many  con- 
spicuous light  spots  which  tend  to  be 
greenish  yellow  above  and  pale  yellow 
on  sides.  Venter  as  in  females  but  there 
seems  to  be  a  tendency  for  the  colors, 
especially  the  orange,  to  be  brighter 
(but  the  belly  of  one  is  bluish  white). 
Iris  light  brown." 

Geographic  Variation. — Sample  lo- 
calities are  indicated  in  figure  2.  Of  the 
characters  analyzed  by  STP,  the  follow- 
ing show  no  among-sample  statistical 
significance  (ranges  of  sample  means  in 
parentheses ) :  Total  infralabials  ( 9.8- 
10.2),  total  supraoculars  (8.0-8.1),  COF 
values  for  females  (2.0),  GOR  (males, 
282.5-295.8;  females,  288.5-318.2),  and 
GAB/GOR  (males,  0.50-0.51;  females, 
0.47-0.50).  As  previously  stated,  A. 
ameiva  from  eastern  Panama  differ  from 
individuals  from  western  Panama  in 
their  retention  of  a  distinct  vertebral 
stripe  as  adults.  Specimens  collected 
west  of  a  line  between  Penonome, 
Code,  and  the  mouth  of  the  Rio  Estan- 
cia,  Code,  lack  the  stripe  as  adults. 

On  the  basis  of  STP  results,  the 
sample  from  Isla  San  Jose  in  the  Bay  of 
Panama  (Sample  4)  can  be  distin- 
guished from  other  samples  in  three 
characters  (within  sample  ranges  fol- 
lowed  by   sample   means   in  parenthe- 


ses): Total  supralabials  14-15  (14.4), 
other  samples  11-16  (12.3-12.8);  COF 
values  for  males  2-6  (4.5),  other  sam- 
ples 2  (2.0);  COP  values  8-12  (9.2), 
other  samples  8-15  (10.0-12.0).  Addi- 
tionally, the  Isla  San  Jose  sample  has 
the  largest  sample  means  for  the  fol- 
lowing characters,  although  the  sample 
does  not  differ  statistically  from  main- 
land samples  for  these  characters:  total 
supraoculars,  GAB,  GOR  for  females, 
GAB/GOR  for  males,  terminal  preanals 
for  females,  and  transverse  rows  of  ven- 
trals.  The  sample  values  were  lowest 
among  the  localities  compared  for  GAB/ 
GOR  for  females,  femoral  pores,  ven- 
trals  at  midbody  and  COP  values.  With 
respect  to  the  total  number  of  supra- 
labials, the  significant  difference  be- 
tween sample  4  and  other  samples  is 
probably  a  function  of  the  few  large 
samples  available  for  analysis.  That  in- 
clusion of  other  samples  in  the  compari- 
son might  alter  the  picture  is  suggested 
by  values  obtained  for  this  character 
from  small,  mainland  samples  and  from 
small  samples  from  other  islands  in  the 
Bay  of  Panama.  Values  intermediate 
between  that  obtained  for  Isla  San  Jose 
and  mainland  samples  used  in  STP 
analysis  were  obtained  from  Isla  Taboga 
(mean  =  13.0,  n  =  5);  Rio  Mamoni, 
Panama  (13.6,  10);  vicinity  of  Nata, 
Code  (13.6,  5);  Rio  Chorcha  east  of 
Chiriqui,  Chiriqui  (13.8,  5);  and  Isla 
San  Miguel  (14.0,  6).  A  sample  of  five 
specimens  from  Tres  Puntas,  Herrera, 
has  a  mean  of  14.6,  higher  than  that  of 
the  Isla  San  Jose  sample  for  total  num- 
ber of  supralabials. 

Samples  5  and  6  from  the  Azuero 
Peninsula  can  be  distinguished  statisti- 
cally as  a  unit  from  other  samples  for 
COP  values,  12  (12.0)  and  10-13  (11.9), 
respectively;  other  samples  8-15  (9.2- 
10.5).  These  samples  possess  the  great- 
est (although  not  statistically  signifi- 
cant) mean  values  for  total  number  of 
femoral  pores  and  number  of  subdigital 
lamellae. 

On  the  mainland  of  Panama,  distinct 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


19 


geographic  trends  are  apparent  only  for 
the  number  of  femoral  pores,  number  of 
termhial  preanals  of  males  and  COP 
values.  All  of  these  characters  show  in- 
creasing values  from  east  to  west. 
Values  obtained  for  small  samples  not 
used  in  STP  analysis  indicate  the  same 
trends.  Other  characters  demonstrate 
no  geographic  trends  although  certain 
localities  differ  from  others  statistically. 
Ecological  Notes. — Ameiva  ameiva 
is  characteristic  of  open  areas  in  Pana- 
ma. It  is  common  in  the  savannas  of  the 
lower  Pacific  slopes  and  in  cleared  areas 
across  the  Canal  Zone.  Cochran  ( 1946 ) 
reported  it  in  the  same  habitat  on  Isla 
San  Jose.  It  has  adapted  well  to  the 
presence  of  man  and  is  common  in  resi- 
dential parts  of  Panama  City.  Heatwole 
(1966)  reported  A.  ameiva  to  be  absent 
in  the  Darien  Gap  and  noted  that  the 
Panamanian  population  of  the  species  is 
disjunct  from  South  American  popula- 
tions. He  postulated  that  the  species 
was  once  continuously  distributed  over 
the  isthmus  but  that  post-Columbian 
reforestation  isolated  the  Panamanian 
population.  Heatwole  further  demon- 
strated that  A.  ameiva  is  presently  ex- 
panding its  range  eastward  in  Panama 
by  moving  into  areas  recently  cleared 
by  man. 

Remarks. — It  is  difficult  to  explain 
the  presence  of  two  geographically  dis- 
tinct color  types  (presence  or  absence 
of  a  reddish  vertebral  stripe  in  adults) 
of  Ameiva  ameiva  in  Panama.  Present 
environmental  conditions  are  similar 
east  and  west  of  the  Penonome-Rio 
Estancia  line.  Bennett  ( 1968 )  provided 
a  clue  to  solution  of  the  problem  with 
his  suggestion  that  the  eruption  of  Vol- 
can  El  Valle,  which  may  have  occurred 
during  the  Pleistocene  (Terry,  1956), 
was  an  event  of  biogeographic  im- 
portance. The  ejecta  from  El  Valle  may 
have  covered  many  square  miles  and 
could  have  divided  the  Panamanian 
population  of  Ameiva  ameiva  for  suf- 
ficient time  for  divergence  of  color  pat- 
tern  to   occur.     Detailed   ecological   in- 


vestigations of  A.  ameiva  might  yield 
infomiation  on  this  point,  as  well  as  on 
the  origins  of  east-west  clines  for  certain 
other  characters,  but  no  such  studies 
have  been  undertaken.  Even  statistical 
studies  aimed  at  correlating  geographic 
variation  in  characters  with  environ- 
mental factors  cannot  be  carried  out 
until  samples  of  adequate  size  for  analy- 
sis can  be  accumulated  from  through- 
out the  range  of  the  species  in  Panama. 
Taxonomic  recognition  of  segments  of 
the  Panamanian  population  of  Ameiva 
ameiva  would  be  premature  until  analy- 
sis of  character  variation  throughout  the 
range  of  the  species  in  South  America 
can  be  completed. 

Two  specimens  of  Ameiva  suritia- 
mensis  ('=A.  ameiva)  were  listed  by 
Boulenger  (1885:353)  as  collected  in 
Costa  Rica  by  "Mr.  Geale."  There  are 
no  records  at  the  British  Museum  that 
these  lizards  (BMNH  80.6.21.1-2)  were 
collected  by  Ceale  himself,  although  the 
museum  bought  a  number  of  specimens 
from  him  between  1866  and  1880  (Mr. 
A.  F.  Stimson,  pers.  comm. ) .  The  speci- 
mens are  indeed  A.  ameiva  and  can  be 
referred  to  the  subspecies  praesignis 
which  ranges  through  northwestern 
Venezuela,  northeastern  Colombia  and 
Panama,  but  it  is  doubtful  that  they 
were  collected  in  Costa  Rica.  Both 
specimens  possess  relatively  well-de- 
fined middorsal  stripes,  suggesting  that 
they  may  have  come  from  eastern 
Panama,  if  they  were  collected  in  Cen- 
tral America  at  all.  There  is  presently 
no  suitable  habitat  for  A.  ameiva  in 
Costa  Rica  with  the  possible  exception 
of  the  dry  Guanacaste  region  where 
the  species  is  not  now  known. 

Barbour  and  Noble  ( 1915 )  examined 
specimens  of  Ameiva  ameiva  ostensibly 
from  Acapulco,  Mexico.  Re-examina- 
tion of  the  specimens  (MCZ  2728  and 
2730)  confirms  the  identification,  but 
the  locality  information  is  obviously 
erroneous. 

Yunker  and  Radovsky  ( 1966 )  and 
Brennan   and   Yunker    ( 1966 )    reported 


20 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


10' 


8' 


50      100 

I         -I 


KILOMETERS 


Fig.  5.    Map  showing  locality  records  of  Ameiva  Jeptophnjs.    To  avoid  crowding  of  symbols,  some 
localities  have  been  omitted.    Samples  used  in  statistical   analyses   were    ( 1 )    Osa   Peninsula,   Punta- 
renas  Prov.,  Costa  Rica,   (2)   Palmar-Golfito  Region,  Puntarenas  Prov.,  Costa  Rica,  (3)  Barro  Colo- 
rado Island,  Canal  Zone,  Panama,  and   (4)   Eastern  Darien  Prov.,  Panama. 


the  acarinids  Dracomjsus  belgicae 
Yunker  and  Radovsky,  Eutrombicula 
alfreddugesi  (Oudemans)  and  E.  goeldi 
(Oudemans)  parasitic  on  Ameiva  hi- 
frontata  from  Panama.  Examination  of 
the  series  of  Uzards  from  which  these 
parasites  were  taken  proves  them  to  be 
A.  ameiva. 

Ameiva  leptophrys  Cope 

Amiva  leptophrijs  Cope,  Proc.  Amer.  Phil.  Soc, 
31:341-342,  1893  [Holotype:  Apparently 
lost;  given  by  Cope  as  "No.  318."  Type 
Locality:  "Buenos  Ayres"  (=  Buenos  Aires, 
Puntarenas  Prov.,  Costa  Rica)]. 

Ameiva  leptophrys — Dunn,  Proc.  Acad.  Nat. 
Sci.  Philadelphia,  92:114,  1940.  Stuart, 
Proc.  Biol.  Soc.  Washington,  55:146,  1942. 
Taylor,  Univ.  Kansas  Sci.  Bull.,  38(1): 
257-260,  1956. 

Ameiva  nithveni  Barbour  and  Noble,  Bull. 
Mus.  Comp.  Zool.,  59(6) :471-473,  1915 
[Holotype:  MCZ  10927.  Type  Locality: 
near  Panama  City.  Collector:  W.  W. 
Brown,  Jr.].  Burt  and  Burt,  Trans.  Acad. 
Sci.  St.  Louis,  28(1  ):53,  1933. 

Ameiva  festiva — Wettstein,  Sitzungsli.  Akad. 
Wiss.  Wien,  math.-natiuw.  Kb,  Abt.  I,  143. 
Bd.,  1.U.2  Heft.  p.  30,  1934  (part). 

Distribution. — Forested       areas       of 
Pacific  slopes  from  Parrita,  Puntarenas 


Prov.,  and  San  Isidro  del  General,  San 
Jose  Prov.,  Costa  Rica  to  Jaque,  Darien 
Prov.,  and  the  Rio  Mono,  Darien  Prov., 
within  a  few  kilometers  of  the  Colom- 
bian border  in  Panama;  across  the 
Canal  Zone  and  east  along  the  Atlantic 
slopes  at  least  as  far  as  the  Rio  Sasardi, 
San  Bias  Prov.,  Panama  (Fig.  5).  The 
species  probably  occurs  in  extreme 
northwestern  Colombia. 

Diagnosis. — Ameiva  leptophrys  can 
be  distinguished  from  its  Middle  Amer- 
ican congeners  by  the  following  com- 
bination of  characters:  Separation  of 
parietal  and  frontoparietal  scales  from 
the  interparietal  by  one  or  more  rows  of 
small,  irregular  scales;  abrupt  reduction 
in  size  of  small  scales  immediately  pos- 
terior to  the  enlarged,  median  gular 
scales;  large  number  of  femoral  pores 
(mean  number  for  males,  49.1;  females, 
44.7);  a  transverse  row  of  abruptly  en- 
larged mesoptychial  scales;  eight  longi- 
tudinal rows  of  ventral  scutes;  color 
pattern  emphasizing  longitudinal  stripes 
rather  than  spotting,  but  lacking  a  light 
vertebral  stripe. 

Description. — Maximum        observed 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


21 


Fig.  6.  Head  and  preanal  scutellation  of 
Arneiva  Icptophrijs:  (A)  Lateral  view  of  head, 
KU  96827,  (BJ  Dorsal  view  of  head,  KU 
96827,  (C)  Ventral  view  of  head,  KU  96827. 
X   1.2.    (D)  Preanal  region,  KU  96815.    X   1. 

SVL  133.0  for  males,  129.0  for  females; 
supralabials  11-15  (12.5)  for  males,  12- 
15  (12.7)  for  females;  infralabials  10-12 
(10.1);  supraoculars  6-8  (6.2);  COF 
values  2-6  (3.5);  COP  values  5-10 
(8.1);  SO-SC  values  2-10  (5.9);  loreals 
2-6  (3.7);  nostril  in  prenasal-postnasal 
suture;  prefrontal  usually  not  in  contact 
with  postnasal  (96.3%,  N  =  136);  GAB 
127-182  (153.8);  PV  28-63  (41.0);  PV/ 
GAB  0.19-0.35  (0.26);  GOR  182-285 
(235.6);  GAB /GOR  0.54-0.74  (0.66); 
SAT  18-25  (21.0);  total  preanal  Scales 
4-10  (6.6)  for  males,  6-10  (8.1)  for  fe- 
males; terminal  preanal  Scales  1-3  (2.1) 
for  males,  1-3  (1.9)  for  females;  one 
row  of  enlarged  prebrachials;  postbra- 
chials  not  enlarged  to  moderately  en- 
larged, irregularly  arranged  or  in  one  or 
two  rows;  preantebrachials  enlarged,  in 


two  rows  proximally,  one  row  distally; 
3  (usually)  or  4  rows  of  enlarged  pre- 
femoral  scales;  two  rows  of  enlarged 
infratibials;  femoral  pores  42-61  (49.1) 
for  males,  34-53  (44.7)  for  females;  sub- 
digital  lamellae  24-35  (28.1);  longi- 
tudinal rows  of  ventral  scutes  26-31 
(28.8);  transverse  rows  of  ventral  scutes 
8-10  (8.1).  For  details  of  scutellation 
see  figure  6. 

Individual  variation  in  dorsal  head 
scutellation  is  extreme  (Fig.  7).  There 
is  no  predictable  pattern  to  scutellation 
posterior  to  the  frontal,  and  even  this 
scale  may  be  divided.  However,  usually 
two  frontoparietal  and  three  parietal 
scales  can  be  recognized.  The  degree 
of  variation  in  posterior  head  scales  is 
unequalled  by  other  species  of  Middle 
American  Ameiva,  although  it  is  ap- 
proached in  the  closely  related  A.  fes- 
tiva. 


Fig.  7.  Variation  of  dorsal  head  scutellation  of 
Ameiva  leptophrys.  (A)  KU  96820,  X  2, 
(B)  KU  96826,  X  2.2,  (C)  KU  96814,  X  2.3, 
(D)  KU  95577,  x  3.8,  (E)  KU  25641,  x  4.1. 


22 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


Color  and  Pattern  in  Alcohol. — Juve- 
nile males:  Top  of  head  light  brown; 
rostral  and  anterior  part  of  frontonasal 
pearl-white  with  a  slight  greenish  tinge; 
side  of  head  dark  brown  above  grading 
to  gray-brown  below;  area  between 
orbit  and  ear  brown  with  darker  brown 
postorbital  bar  dorsally;  white  crescent 
beneath  and  slightly  behind  eye;  chin, 
infralabials,  gular  region  and  throat  im- 
maculate. Middorsal  stripe  broad, 
slightly  irridescent,  gray-brown,  bor- 
dered laterally  by  white  paravertebral 
stripes  originating  on  superciliary  scales 
and  extending  well  onto  tail;  paraverte- 
bral stripes  (and  edges  of  middorsal 
stripe)  undulating  on  posterior  three- 
quarters  of  body;  velvet-black  dorso- 
lateral stripe  from  above  shoulder  to 
insertion  of  hind  legs,  bordered  below 
by  broken  white  line  originating  on 
shoulder  and  extending  to  insertion  of 
hind  limbs  and  onto  anterior  surface  of 
femur;  area  below  this  lateral  light 
stripe  brown,  grading  to  gray-brown 
above  ventral  scutes.  Venter  immacu- 
late. Dorsal  surfaces  of  fore  and  hind 
limbs  dark  brown  with  lighter  brown 
blotches,  spots  or  mottling;  no  post- 
femoral  light  stripe;  ventral  surfaces  of 
limbs  immaculate  to  gray-white.  Tail 
brown  with  black  Hecks  above,  brown 
laterally  (below  dorsolateral  light 
stripe)  blending  to  white  or  gray- white 
ventrally;  short  postanal  white  stripe 
bounded  by  brown  below  for  about  six 
scales,  then  joining  white  ventral  pat- 
tern. 

Through  ontogeny  ( Fig.  8 )  the  mar- 
gins of  the  middorsal  stripe  become 
more  undulating  and  the  paravertebral 
stripes  are  lost  except  where  the  mid- 
dorsal stripe  remains  broadest.  Gray- 
brown  or  dark  brown  bars  or  large  spots 
appear  laterally  overlaying  the  dorso- 
lateral and  ventrolateral  dark  fields.  In 
some  large  adult  males  the  lateral  light 
stripes  are  lost  entirely,  and  the  venter 
becomes  blue-white  or  coppery  from 
chin  to  vent  and  onto  the  ventral  sur- 
face of  the  tail. 


Females:  Juvenile  females  are  simi- 
lar to  juvenile  males.  Through  ontogeny 
( Fig.  8 )  the  middorsal  stripe  often  be- 
comes interrupted  by  narrow,  dark 
brown  cross-bars.  The  adult  pattern  of 
females  is  similar  to  that  of  adult  males, 
faint  to  relatively  distinct  bars  appear- 
ing in  the  dorsolateral  and  ventrolateral 
dark  fields.  However,  some  females  lack 
lateral  bars,  and  the  lateral  light  stripes 
are  retained  throughout  life. 

Color  and  Pattern  in  Life. — The  dor- 
sal coloration  in  life  of  Ameiva  lepto- 
phry.s  does  not  differ  markedly  from 
that  seen  in  preserved  specimens.  The 
middorsal  stripe  is  metallic  brown  and 
may  have  a  greenish  tint.  Paravertebral 
and  lateral  light  stripes  are  yellowish  to 
yellowish-gray,  and  lateral  bars  or 
blotches,  if  present,  are  yellowish-gray 
to  rust  colored.  Ventral  coloration  is 
variable.  The  entire  ventral  surface  may 
be  white,  cream  or  copper  colored  in 
juvenile  males  and  females.  Adults  of 
both  sexes  may  retain  these  colors,  but 
some  adult  males  become  pale  blue 
over  the  entire  ventral  surface,  or  they 
may  have  greenish-yellow,  orange  or 
brick  red  on  chin,  gular  region  and 
throat  with  the  ventral  surfaces  of  body, 
limbs  and  tail  blue.  Some  adult  males 
exhibit  a  variation  of  the  latter  color 
pattern  in  which  the  hind  limbs  pos- 
terior to  the  femoral  pores  and  the 
ventral  surface  of  the  tail  is  copper- 
colored.     Variability   in   ventral    colora- 


FiG.  8.  Ontogenetic  Change  and  Variation  in 
Pattern  of  Anieiva  Jeptophnjs.  Males:  (A) 
KU  107555,  SVL  85  mm  from  N  slope  Cerro 
Cambutal,  Los  Santos  Prov.,  Panama;  (B)  KU 
76142,  115  mm  from  Barro  Colorado  Island, 
Canal  Zone,  Panama;  (C)  KU  95582,  118  mm 
from  6  km  W  Palmar  Norte,  Puntarenas  Prov., 
Costa  Rica  (included  for  comparison  with  KU 
76142).  Females:  (D)  KU  95578,  44  mm 
from  6  km  W  Palmar  Norte,  Puntarenas  Prov., 
Costa  Rica;  (E)  KU  76144,  89  mm  from  Barro 
Colorado  Island,  Canal  Zone,  Panama;  (F) 
KU  76163,  112  mm  from  El  Valle,  Code  Prov., 
Panama.  Small  males  are  similar  to  small 
females  as  exemplified  by  KU  95578  ( D 
above ) . 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


23 


24 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


tion  might  be  correlated  with  reproduc- 
tive condition  in  these  Hzards,  but  this 
aspect  of  their  biology  has  not  been 
investigated.  It  \\'ould  be  instructive  to 
retain  specimens  in  captivity  in  order  to 
ascertain  not  only  when  color  changes 
occur,  but  whether  or  not  greenish-yel- 
low and  orange  pigmentation  represent 
stages   leading  to  brick  red   coloration. 

Geographic  Variation. — Any  analysis 
of  geographic  variation  of  characters  of 
Ameiva  leptophnjs  must  be  considered 
preliminary,  pending  the  acquisition  of 
additional  large  samples  of  specimens 
from  throughout  the  range  of  the 
species.  Only  four  such  samples  have 
been  available  for  this  study  (Fig.  5). 
There  is  a  noticeable  lack  of  specimens 
from  western  Panama,  and  the  follow- 
ing results  should  be  viewed  with  both 
the  paucity  of  large  series  of  specimens 
and  the  lack  of  good  geographic  repre- 
sentation in  mind. 

The  following  characters  exhibit  no 
significant  geographic  differences  among 
sample  means  (ranges  of  sample  means 
in  parentheses):  Supralabials  (males, 
12.1-12.9;  females,  12.4-13.2),  infra- 
labials  (10.0-10.3),  SO-SC  values  (5.2- 
6.3)  and  terminal  preanal  scales  of 
males  (1.9-2.3). 

Samples  1  and  2  (Costa  Rica)  are 
significantly  different  from  samples  3 
and  4  (Panama)  with  respect  to  the 
following  characters  (locality  number 
followed  by  sample  mean  in  parenthe- 
ses): Number  of  subdigital  lamellae 
(Sample  1,  29.2;  2,  29.4;  3,  26.5;  4,  27.2), 
paravertebral  granules  (1,  45.9;  2,  48.9; 

3,  38.1;  4,  37.4)  and  PV/GAB  (1,  0.28; 
2,  0.30;  3,  0.25;  4,  0.25).  Samples  1,  2 
and  3  differ  significantly  from  sample  4, 
but  not  from  each  other,  in  mean  COF 
values  (1,  7.4;  2,  7.3;  3,  8.0;  4,  8.7).  One 
character,  total  number  of  femoral 
pores,  shows  a  significant  difference  be- 
tween samples  1  and  2  ( 1,  43.6;  2,  48.4 ) . 
In  addition  to  COF  values,  mentioned 
above,  sample  3  differs  significantly 
from  sample  4  in  GAB/GOR   (3,  0.68; 

4,  0.65),  but  sample  4  is  not  significantly 


different  from  samples  1  and  2  (both 
0.66).  There  are  a  number  of  charac- 
ters for  which  mean  values  of  either 
sample  1  or  2  differ  significantly  from 
that  of  sample  3,  but  not  from  sample 
4,  and  a  few  (COP,  GAB,  Longitudinal 
Rows  of  Ventrals)  for  which  mean 
values  of  sample  1  and/ or  sample  2  are 
not  significantly  different  from  sample 
3,  but  do  differ  from  the  mean  value  for 
sample  4. 

East-west  clines  are  apparent  for  a 
number  of  characters.  In  terms  of  mean 
values,  there  is  a  decrease  from  east  to 
west  ( infralabials,  longitudinal  rows  of 
ventrals)  or  an  increase  from  east  to 
west  (COF  values,  terminal  preanals  of 
females,  total  preanals  of  females).  If 
samples  1  and  2  are  combined  (it  is 
legitimate  to  do  so  according  to  STP 
results),  additional  characters  show  the 
same  clinal  tendencies.  An  increase  in 
mean  value  from  east  to  west  is  noted 
for  number  of  supralabials  of  males, 
number  of  supraoculars,  GAB,  PV  and 
PV/GAB;  decreasing  mean  values  from 
east  to  west  are  noted  for  COP  values. 
A  small  sample  (n  =r  10)  of  leptophnjs 
from  near  the  Rio  Sasardi  in  San  Bias 
Prov.,  Panama,  was  collected  by  C.  W. 
Myers  and  arrived  too  late  to  be  in- 
cluded in  the  STP  analysis.  Mean 
values  calculated  for  each  character  for 
the  Rio  Sasardi  sample  are  in  accord 
with  the  east-west  trends  discussed 
above  for  (Rio  Sasardi  means  in  paren- 
these):  COF  values  (2.8),  total  pre- 
anal scales  of  females  (8.4),  number  of 
supralabials  of  males  (12.2)  and  COP 
values  (9.1).  The  Rio  Sasardi  sample 
runs  counter  to  east-west  trends  estab- 
lished on  the  basis  of  mean  values  of 
samples  1-4  for  number  of  terminal  pre- 
anals of  females  (2.0),  number  of  in- 
fralabials (10.0),  longitudinal  rows  of 
ventrals  (28.7),  number  of  supraoculars 
(6.3),  GAB  (160.1)  and  PV/GAB 
(0.24).  These  results  once  again  call 
attention  to  the  necessity  of  obtaining 
additional  samples  from  intermediate 
localities    before    attempting    a    really 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


25 


definitive  analysis  of  geographic  varia- 
tion of  leptophrys. 

With  the  possible  exception  of  color 
pattern,  none  of  the  characters  not  ana- 
lyzed by  STP  appears  to  vary  geo- 
graphically. Male  leptophrys  from  cen- 
tral and  eastern  Panama  tend  to  have 
more  emphasis  on  spotting  (rather  than 
bars)  in  the  lateral  dark  fields  then  do 
specimens  from  western  Panama  and 
Costa  Rica.  With  the  acquisition  of 
more  specimens  from  the  hiatus  in  west 
and  central  Panama  this  difference  may 
also  prove  to  be  clinal.  In  addition, 
specimens  from  the  Osa  Peninsula  of 
Costa  Rica  ( sample  1 )  appear  darker  in 
overall  coloration  than  those  collected 
at  other  localities,  including  those  from 
the  adjacent  non-peninsular  mainland  of 
Costa  Rica.  Since  my  sample  from  the 
Osa  Peninsula  was  collected  over  a 
short  period  of  time  and  by  the  same 
collectors,  it  may  be  that  the  darkness 
of  the  specimens  can  be  attributed  to 
preservation  in  warm  formalin  or  to  an 
unusually  long  period  of  time  in  fomia- 
lin  before  transfer  to  ethyl  alcohol. 

Ecological  Notes. — Amewa  lepto- 
phrys is  a  forest  species;  at  various  lo- 
calities throughout  its  range  it  is  found 
sympatrically  with  all  other  species  of 
Middle  American  Ameiva,  except  chait- 
zami.  Hillman  (1969)  studied  niche 
specificity  of  leptophrys,  festiva  and 
quadrilineata  near  Rincon  on  the  Osa 
Peninsula  of  Costa  Rica.  He  found  that 
adult  leptophrys  forage  deeper  into  the 
forest  than  its  sympatric  congeners;  the 
latter  search  for  food  in  more  open 
areas.  Roth  leptophrys  and  festiva  seem 
to  utilize  the  same  areas  for  basking, 
but  there  is  little  overlap  with  the  more 
heliophilous  quadrilineata.  Hillman 
found  hatchling  leptophrys  occupying 
more  open  areas  than  the  adults  but 
virtually  identical  to  areas  utilized  by 
adult  festiva.  Presumably  the  size  dif- 
ference between  juvenile  leptophrys  and 
adult  festiva  acts  to  lessen  direct  com- 
petition. In  the  vicinity  of  Palmar 
Norte,    Puntarenas    Prov.,    Costa    Rica, 


festiva   is    not   abundant,    and    I    found 
leptophrys      and      quadrilineata      more 
closely      associated      ecologically     than 
either   is    to   festiva.     The   two    species 
were  seen  basking  within  a  few  meters 
of  one  another,  but  whereas  quadriline- 
ata utilized  clearings,  leptophrys  basked 
in    the    sun-flecked    areas    beneath    the 
dense,    low   vegetation.     Ameiva    lepto- 
phrys would  allow  approach  to  within 
a  meter  before  bolting  when  so  hidden. 
Tree-fall    clearings    in    a    swamp    forest 
near   Palmar   Norte   were   occupied   by 
juvenile    quadrilineata    and    leptophrys, 
but  I  never  found  the  young  of  both 
species     utilizing     the     same     clearing. 
Dunn    (1940a)    found    leptophrys    and 
festiva    sympatric    on    Rarro    Colorado 
Island    but    reported    that    festiva    oc- 
cupied the   more   shaded   parts   of  the 
habitat,     whereas     leptophrys     utilized 
more   open   areas,   the   opposite   of   the 
situation    in    Costa    Rica.     Charles    W. 
Myers  (personal  communication)  found 
leptophrys    and    ameiva    sympatric    at 
Guanico  Arriba,  Los  Santos  Prov.,  Pana- 
ma.    There    leptophrys    occupies    dry, 
partly    disturbed   forest   and   ameiva    is 
common  in  open  pastures.    Recently  col- 
lected material  from  near  Parrita,  Pun- 
tarenas Prov.,  Costa  Rica,  confirms  the 
sympatry    of    leptophrys    and    undulata 
hesitantly  reported  by  Dunn  (in  Stuart, 
1942). 

Remarks. — Rarbour  and  Loveridge 
(1929b:  213)  listed  two  paratypes  of 
Ameiva  ruthveni  (^=A.  leptophri/s): 
MCZ  10925-10926.  Rarbour  and  Noble 
(1915:471-472)  acknowledged  examina- 
tion of  only  two  specimens  in  their  de- 
scription of  A.  ruthveni:  MCZ  9931  (re- 
catalogued  MCZ  10927;  E.  E.  Williams, 
in.  litt.),  the  holotype,  and  MCZ  9932 
(recatalogued  MCZ  10926).  The  third 
hzard,  MCZ  10925  (originally  MCZ 
9928),  is  not  a  paratvpe.  One  paratype 
(MCZ  10926)  was  not  located. 

Brennan  and  Yunker  (1966:235-236, 
258)  reported  Ameiva  undulata  from 
Panama  as  host  for  the  acarinids  Eu- 
tromhicula     alfreddugesi     (Oudemans) 


26 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


and  E.  goeldi  (Oudemans).  Examina- 
tion of  the  specimens  of  Ameiva  prove 
them  to  be  leptophrys.  There  are  no 
vahd  records  of  A.  undiihta  from 
Panama. 

Ameiva  festiva  (Lichtenstein  and 
Von  Martens) 

Cnemidophorus  festivus  Lichtenstein  and  Von 
Martens,  Nomenclator  Reptilium  et  Am- 
philiiorum  Musei  Zoologici  Berolinensis,  p. 
13,  1856  [Lectotype,  herein  designated: 
ZMB  881a;  Paralectotype,  formerly  syn- 
type:  ZMB  881b.  Type  Loca]it>-:  "Vera- 
gua"   =   Veragiias,  Panama]. 

Ai7iewa  eutropia  Cope,  Proc.  Acad.  Nat.  Sci. 
Philadelphia,  14:62,  1862  [Syntypes: 
ANSP  9071,  USNM  4320(3).  Type  lo- 
cality: "Region  of  the  Triiando,  New  Gre- 
nada" (=Colombia).  Collector:  Arthur 
Schott]. 

Aiyjeiva  edwardsi  Bocourt,  Ann.  Sci.  Nat.  17 
(17):  1-2,  1873  [Holotype:  MNHN  5480. 
Type  Locality:  Izabel  and  Santa  Maria  de 
Panzos  near  Rio  Polochic,  Guatemala.  Col- 
lector :    unknown] . 

Ameiva  festivus — Bocourt,  Mission  Scientifique 
au  Mexique  et  dans  TAmerique  Centrale; 
fitudes  sur  les  reptiles,  livr.  3,  pp.  260-263, 
pi.  20A,  Fig.  10;  pi.  20D,  Fig.  3;  pi.  20, 
Fig.  2;  1874. 

Ameiva  festiva — Giinther,  Biologia  Centrali- 
Americana;  Reptilia  and  Batrachia,  p.  24, 
1885.  Boulenger,  Catalogue  of  the  Lizards 
in  the  British  Museum  (Natural  History), 
vol.  2,  p.  347,  1885.  Barbour  and  Noble, 
Bull.  Mus.  Comp.  Zool.,  59(6):473-4, 
1915.  Burt  and  Burt,  Bull.  American  Mus. 
Nat.  Hist.,  61(5)  :309,  1931.  Burt  and 
Burt,  Trans.  Acad.  Sci.  St.  Louis,  28(1): 
53,  1933.  Wettstein,  Sitzungsb.  Akad. 
Wiss.  Wien,  math.-naturw.  Kl.,  Abt.  I,  143. 
Bd.,  l.u.  2  Heft.,  p.  30,  1934. 

Ameiva  festiva  festiva — Dunn,  Proc.  Acad. 
Nat.  Sci.  Philadelphia,  92:114,  1940.  Tay- 
lor, Univ.  Kansas  Sci.  Bull.,  38(1):260- 
264,  1956. 
Ameiva  festiva  edwardsi — Stuart,  Occ.  Pap. 
Mus.  Zool.,  Univ.  Michigan,  No.  471,  p. 
21,  Figs.  6-7,  1943.  Smith  and  Laufe, 
Univ.  Kansas  Sci.  Bull.  31(2):64-66,  1945. 
Smith  and  Taylor,  U.  S.  Nafl.  Mus.  Bull, 
199:174,  1950.  Stuart,  Misc.  Publ.  Mus. 
Zool.,  Univ.  Michigan,  No.  122,  p.  77, 
1963. 
Ameiva  festiva  occidentalis  Taylor,  Univ.  Kan- 
sas Sci.  Bull.,  38(l):260-264,  1956  [Holo- 
type: FMNH  120236.  Type  Locality:  8.0 
km  E  San  Isidro  del  General,  San  Jose 
Prov.,  Costa  Rica.  Collector:  Richard  C. 
Taylor] . 


Distribution. — Ameiva  festiva  is 
found  in  the  forests  of  the  Caribbean 
lowlands  from  Tabasco,  Mexico  to 
Depto.  Boyaca,  Colombia  and  on  the 
Pacific  lowlands  from  the  vicinity  of 
San  Isidro  del  General,  San  Jose  Prov., 
Costa  Rica,  at  least  to  the  Condoto- 
Andagoya  area  in  the  Intendencia  de 
Choco,  Colombia.  The  species  also  is 
known  from  the  vicinity  of  Tilaran, 
Guanacaste  Prov.,  Costa  Rica,  just  west 
of  the  continental  divide  (Fig.  9). 

Diagnosis. — Ameiva  festiva  can  be 
distinguished  from  other  Middle  Amer- 
ican Ameiva  by  the  following  combina- 
tion of  characters:  Central  gular  scales 
much  enlarged,  irregular  in  arrange- 
ment; gradual  reduction  in  size  of  scales 
radiating  outward  from  enlarged,  cen- 
tral gular  scales;  many  femoral  pores 
(mean  for  males,  40.8;  females,  40.0); 
a  transverse  row  of  abruptly  enlarged 
mesoptychial  scales;  eight  longitudinal 
rows  of  ventral  scutes;  a  distinct,  light 
colored  vertebral  stripe  present  in  all 
but  large  adults.  No  other  species  of 
Ameiva  from  Middle  America  possesses 
such  a  well-defined  vertebral  stripe,  al- 
though several  South  American  Ameiva 
do  have  such  a  stripe  (edracantha,  or- 
cesi,  bridgesi,  and  septemlineata). 

Description. — Maximum  obsei"ved 
SVL  114.0  mm.  for  males,  129  mm.  for 
females;  supralabials  12-18  (14.2)  for 
males,  12-17  (13.9)  for  females;  infra- 
labials  8-15  (10.8);  sublabials  4-9  (4.7) 
for  males,  4-7  (4.5)  for  females;  supra- 
oculars 4-10  (6.5);  COF  values  2-6 
(2.6);  COP  values  2-14  (9.4);  loreals 
2;  nostril  in  prenasal-postnasal  suture; 
prefrontals  in  contact  with  postnasal 
(41.1%)  or  no  (54.5%),  4.5  percent  in 
contact  on  one  side  but  not  the  other 
(n  =  448);  GAB  145-221  (182.5)  for 
males,  143-213  (179.8)  for  females;  VS 
5-29  (12.8)  for  males,  4-27  (11.2)  for 
females;  VS/GAB  0.02-0.14  (0.06); 
COR  209-324  (261.1)  for  males,  215- 
312  (259.5)  for  females;  GAB/GOR 
0.57-0.85  (0.70);  SAT  16-22  (18.3); 
total  preanal  scales  4-10  (6.6)  for  males, 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


27 


Fig.  9.  Map  showing  locality  records  of  Aineiva  festiva.  To  avoid  crowding  of  symbols,  some 
localities  haxe  been  omitted.  Samples  used  in  statistical  analyses  were  ( 1 )  Piedras  Negras,  Depto. 
El  Peten,  Guatemala,  (2)  Vicinity  of  Chinaja,  Depto.  Alta  Verapaz  and  Depto.  El  Peten,  Guate- 
mala, (3)  Finca  Chama,  Depto.  Alta  Verapaz,  Guatemala,  (4)  North  Coast  Localities,  Honduras, 
(5)  Bonanza,  Depto.  Zelaya,  Nicaragua,  (6)  Cukra  and  Kanawa,  Depto.  Zelaya,  Nicaragua,  (7) 
Rio  Mico  and  vicinity,  Depto.  Zelaya,  Nicaragua,  (8)  Tuli  Creek,  Depto.  Rio  San  Juan,  Nicaragua, 
(9)  Puerto  Viejo,  Heredia  Prov.,  Costa  Rica,  (10)  Turrialba,  Cartago  Prov.,  Costa  Rica,  (11) 
Tortuguero,  Limon  Prov.,  Costa  Rica,  (12)  Los  Diamantes  and  vicinity,  Limon  Prov.,  Costa  Rica, 
(13)  Almirante,  Bocas  del  Toro  Prov.,  Panama,  (14)  Isla  Escudo  de  Veraguas,  Panama,  (15) 
Barro  Colorado  Island,  Canal  Zone,  Panama,  (16)  Rio  Mono,  Darien  Prov.,  Panama,  (17)  Golfito 
Region,  Puntarenas  Prov.,  Costa  Rica,  (18)  Vicinity  of  Rincon,  Osa  Peninsula,  Puntarenas  Prov., 
Costa  Rica,   (19)   Rio  Ure  and  vicinity,  Depto.  Cordoba,   Colombia,    (20)    Condoto-Andagoya-Tado 

Area,  Intendencia  de  Choco,  Colombia. 


4-9  (6.4)  for  females;  terminal  preanal 
scales  1-3  (2.0)  for  males,  1-3  (2.2)  for 
females;  one  row  of  enlarged  prebra- 
chials;  postbrachials  enlarged,  in  a 
single  short  row;  preantebrachials  en- 
larged, two  rows  proximally,  one  dis- 
tally;  three  rows  of  enlarged  prefemoral 
scales;  two  rows  of  enlarged  tibial 
scales;  femoral  pores  33-55  (40.8)  for 
males,   32-50    (40.0)    for   females;    sub- 


digital  lamellae  23-34  (28.4);  longitu- 
dinal rows  of  ventral  scutes  25-33 
(27.8);  transverse  rows  of  ventral  scutes 
6-8  (8.0). 

Individual  variation  in  posterior 
head  scutellation  is  great,  second  only 
to  that  of  Ameiva  leptophrijs,  but  two 
frontoparietal  and  three  parietal  scales 
are  usually  recognizable  (Fig.  10). 

Color    and    Pattern    in    Alcohol. — 


28 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


Fig.  10.  Head  and  preanal  scutellation  of 
Ameiva  festiva  (ZMB  881a;  Lectotype):  (A) 
Lateral  view  of  head,  ( B )  Dorsal  view  of 
head,  (C)  Ventral  view  of  head.  X  1.5.  (D) 
Preanal  region.    X  1-6. 

There  is  extensive  geographic  variation 
in  color  and  pattern  over  the  range  of 
Ameiva  festiva.  As  a  basis  for  discuss- 
ing this  variation,  the  following  com- 
plete description  of  color  and  pattern 
of  specimens  from  Puerto  Viejo,  Here- 
dia  Prov.,  Costa  Rica,  is  presented. 

Juvenile  males:  Blue-gray  vertebral 
stripe  5-13  granules  wide  at  midbody 
extending  from  rostral  scale  well  onto 
tail,  the  margins  slightly  scalloped  on 
the  posterior  three-fourths  of  body;  top 
of  head  lateral  to  vertebral  stripe  dark 
gray;  side  of  head  dark  gray  above  be- 
coming lighter  below,  anterior  to  eye; 
ear  encircled  by  interrupted  white  line; 
chin,  infralabials,  gular  region  and 
throat  light  gray  or  white;  dorsal  sur- 
face of  body  very  dark  brown  lateral  to 
vertebral  stripe;  dorsolateral  light  stripe 
white,  narrow  and  interrupted,  originat- 
ing on  canthus  and  extending  onto  tail; 
dorsolateral    dark    stripe    velvet    black, 


beginning  immediately  posterior  to  eye 
and  extending  onto  tail,  bordered  below 
by  white,  narrow,  interrupted  ventro- 
lateral light  stripe  originating  beneath 
eye,  passing  over  ear  to  the  groin, 
thence  onto  anterior  surface  of  femur  at 
juncture  of  dorsal  granular  scales  and 
enlarged  ventral  scales;  vertical  bars  ex- 
tending ventrally  from  ventrolateral 
light  stripe  immediately  anterior  and 
posterior  to  forelimbs;  ventrolateral 
field  dark  brown  with  irregular  blue- 
gray  spots  or  narrow,  regularly  spaced 
blue-gray  bars,  or  both.  Ventral  sur- 
faces of  body,  limbs  and  tail  gray-white. 
Dorsal  surface  of  forelimbs  and  femur 
dark  brown  to  black;  shank  dark  brown 
with  blue-gray  spots  above. 

Through  ontogeny  (Fig.  11)  the 
vertebral  stripe  becomes  sinuous  and 
less  well  defined.  It  may  be  lost  entirely 
on  the  head,  posterior  part  of  the  body 
and  tail.  A  light-colored  rostral  scale  is 
retained,  as  usually  is  the  vertebral 
stripe  on  the  anterior  third  of  the  body. 
The  head  becomes  uniformly  brown 
above  and  the  pale  markings  above  and 
below  the  eye,  on  the  side  of  the  neck 
and  around  the  ear  are  lost.  The  velvet- 
black  dorsolateral  dark  stripe  is  re- 
tained. In  larger  individuals,  the  mid- 
dorsal  area  immediately  lateral  to  the 
position  occupied  by  the  vertebral 
stripe  becomes  regularly  marked  with 
dark  and  light  brown  blotches,  the 
darker  occupying  positions  where  the 
sinuous  or  scalloped  margin  of  the  ver- 
tebral stripe  curved  inwards  toward  the 
midline;  this  gives  the  back  a  ladder- 
like pattern.  The  dorsal  surface  of  the 
tail  becomes  imiformly  brown  or  brown 
with  darker  brown  blotches  which  may 
be  paired  or  nearly  so.  The  dorsolateral 
and  ventrolateral  light  stripes  are  re- 
tained posteriorly  only.  Most  males  be- 
come blue-gray  overall  ventrally,  the 
lateral  ventral  scutes  and  enlarged  fem- 
oral scales  marked  with  black.  The  in- 
fralabials may  also  be  flecked  with 
black. 

Females:    The  color  and  pattern  of 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


29 


Fig.    11.     Ontogenetic   Pattern   Change   of   Ameiva   f estiva   from    the   Atlantic 

Slopes  of  Costa  Rica.    Males:     (A)  AMNH  89210,  SVL  43  mm;   (B)   AMNH 

89191,  80  mm;   (C)  AMNH  89203,   106  mm.    Females:     (D)   AMNH  89212, 

47  mm;  (E)  AMNH  89204,  69  mm;   (F)  AMNH  89193,  90  mm. 


30 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


juvenile  females  is  similar  to  that  of 
juvenile  males.  Females  differ  from 
males  in  that  the  juvenile  color  and  pat- 
tern is  retained  with  little  fading  in  all 
but  very  large  adults  and  adult  females 
do  not  develop  blue  colors  ventrally. 

Color  and  Pattern  in  Life. — The  fol- 
lowing notes  refer  to  specimens  col- 
lected by  me  on  1  August  1965  at  Puerto 
Viejo,  Heredia  Pro  v.,  Costa  Rica.  Adult 
male  (KU  95558),  SVL  99  mm:  Neck 
and  chest  reddish-orange;  chin  and 
gular  region  pale  blue;  belly  pale  blue 
medially  and  pale  blue  broken  by  black 
blotches  laterally;  middorsal  area  brown 
with  indications  of  a  faint  yellow  verte- 
bral stripe  extending  from  occiput  to 
base  of  tail;  dorsolateral  dark  field 
black,  bordered  above  and  below  by 
narrow,  interrupted  yellow  stripes;  ven- 
trolateral field  olive  with  narrow  yellow 
bars.  Adult  female  (KU  95559),  SVL 
79  mm:  Venter  pale  bronze;  dorsal  pat- 
tern similar  to  KU  95558  except  verte- 
bral stripe  more  pronounced  and  ex- 
tending from  rostral  well  onto  tail;  yel- 
low on  head  and  body  becoming  pale 
blue  on  tail. 

Juvenile  Ameiva  festiva  from  near 
Bluefields,  Depto.  Zelaya,  Nicaragua 
(KU  101214-101215)  possess  vertebral 
stripes  which  are  yellow  anteriorly  be- 
coming blue-green  on  the  tail.  The  dor- 
solateral and  ventrolateral  light  stripes 
are  yellow  to  yellowish-orange.  An 
adult  female  (KU  101213)  from  the 
same  locality  has  the  same  color  and 
pattern  as  KU  95559  from  Puerto  Viejo. 

As  in  other  species  of  Ameiva  I  have 
examined,  gular  coloration  varies  in 
adult  males.  Collections  are  not  such 
that  this  can  be  correlated  with  season. 
I  suspect  the  color  to  be  indicative  of 
reproductive  state,  at  least  in  males,  but 
lack  evidence  to  substantiate  this  hy- 
pothesis. Field  notes  taken  by  W.  E. 
Duellman,  C.  W.  Myers,  and  myself,  as 
well  as  color  photographs  provided  by 
D.  H.  Janzen  disclose  that  the  gular 
region  and  chest  of  adult  male  festiva 
may  be  uniformly  blue  (if  so,  the  entire 


venter  is  blue),  copper,  dull  brick-red, 
yellow-orange,  golden-yellow  or  yellow- 
green  (all  commonly  associated  with  a 
blue  venter,  although  individuals  hav- 
ing a  copper  gular  region  often  have  the 
entire  venter  that  color).  There  seems 
to  be  no  geographic  pattern  associated 
with  these  colors  or  none  that  I  can  de- 
tect from  the  field  notes  available  to 
me.  The  ventral  surfaces  of  females 
may  be  uniformly  copper  or  cream 
(often  with  a  pinkish  tint),  or  may  be 
one  of  these  colors  with  the  gular  region 
and  chest  gray. 

Geographic  Variation. — Only  the 
number  of  transverse  rows  of  ventral 
scutes  and  the  number  of  terminal  pre- 
anals  of  males  show  no  significant  dif- 
ferences among  sample  means.  A  num- 
ber of  characters  exhibit  variation 
among  samples,  but  show  no  marked 
geographic  trends.  These  characters 
are:  GOR,  GAB,  number  of  femoral 
pores,  number  of  longitudinal  rows  of 
ventrals,  total  number  of  preanal  scales 
and  SAT  (although  SAT  values  from 
samples  in  the  Atlantic  lowlands  may 
tend  to  be  higher  than  values  from 
Pacific  samples). 

A  striking  aspect  of  geographic  vari- 
ation among  samples  of  festiva  is  the 
apparent  "center"  of  variation  for  cer- 
tain characters  in  the  Caribbean  low- 
lands of  Costa  Rica  and  Nicaragua. 
Values  for  VS,  VS/GAB,  and  number  of 
sublabials  of  males  gradually  increase 
in  samples  to  the  north  and  east  of  these 
lowlands.  A  decrease  in  number  of 
supralabials  and  in  COF  values  is  ap- 
parent in  the  same  directions. 

A  clinal  increase  in  mean  values 
from  north  (Guatemala)  to  southeast 
( Panama  and  Colombia )  is  expressed 
for  GAB/GOR,  but  samples  17  (Golfito 
Region  of  Costa  Rica)  and  18  (Osa 
Peninsula  of  Costa  Rica)  have  high 
means,  nearer  to  values  noted  for  Sam- 
ples 16  (Rio  Mono,  Darien  Prov.,  Pana- 
ma) and  20  ( Condoto-Andagoya-Tado 
Area,  Intendencia  de  Choco,  Colombia) 
to  the  south  and  east.    The  mean  value 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


31 


for   sample    19    ( Rio    Ure   and   vicinity, 
Depto.    Cordoba,    Colombia)    is    lower 
than  would  be  predicted  by  the  trend. 
A  clinal  decrease  from  north  to  south- 
east  is   expressed   in   number   of   infra- 
labials   ( Samples   1  and  19  have  lower 
and    higher   mean    values,    respectively, 
than  anticipated  by  the  trend),  number 
of  sublabials  of  females  (although  high 
values  are  noted  for  samples  14  and  20 ) , 
COP  values  (locality  20  has  a  very  low 
mean  and  represents  a  separate  subset) 
and  the  number  of  subdigital  lamellae. 
As  is  evident,  these  clinal  tendencies  are 
never  perfectly  expressed:    one  or  more 
samples  disrupt  the  trend.    Samples   19 
and  20   (in  South  America)   counter  in 
some  respects  geographic  trends  estab- 
lished in  the  more  northerly  parts  of  the 
range   of  festiva.    These   two   localities 
are  separated   by  high   mountains   and 
differ  from  one  another  in  a  statistically 
significant  way  for  a  number  of  charac- 
ters.   Samples   15    (Barro   Colorado   Is- 
land) and  16  (Rio  Mono,  Darien  Prov., 
Panama)  both  show  greater  similarities 
to  Sample  20  than  to  Sample  19.    Sam- 
ples  17  and  18,  in  southeastern  Costa 
Rica,  show  similarities  to  Sample  19  for 
some  characters,  to  sample  20  for  others. 
It  would  be  instructive  to  have  addi- 
tional samples  from  the  Pacific  slopes 
of    western    Panama    in    order    to    see 
whether   there   is    further   evidence   for 
the   "Panamanian    crossover"   suggested 
by  Dunn  (1940b). 

The  more  northerly  samples  of  fes- 
tiva (Samples  1-4)  differ  from  others  in 
a  number  of  characters,  none  of  which 
set  these  samples  apart  statistically  from 
all  others.  The  samples  from  Guatemala 
(Samples  1-3)  and  Honduras  (Sample 
4),  taken  as  a  group,  differ  from  adja- 
cent samples  in  a  statistically  significant 
way  for  number  of  supralabials  of  males 
and  females,  VS  of  males  (data  from 
Sample  4  unavailable)  and  total  pre- 
anal  scales  of  females. 

As  alluded  to  earlier,  there  is  marked 
geographic  variation  in  the  color  and 
pattern  over  the  range  of  festiva.   There 


are  four  distinct  pattern  types  with 
variation  to  a  lesser  degree  within  two 
of  these.  The  four  patterns  are  dis- 
cussed separately  below  relative  to  the 
description  of  specimens  from  Puerto 
Viejo,  Costa  Rica,  given  above. 

Pattern  Type  A:  The  color  and  pat- 
tern described  for  festiva  from  Puerto 
Viejo  is  characteristic  in  major  aspects 
of  specimens  collected  in  the  Caribbean 
lowlands  of  Costa  Rica,  western  Pana- 
ma, and  northward  into  Mexico.  Liz- 
ards comprising  Samples  1-7  and  pos- 
sibly 8  (Nicaragua  northward)  tend  to 
be  browner  in  overall  coloration  than 
those  from  Costa  Rica  and  Panama 
(Samples  9-13),  although  this  is  a  sub- 
jective observation  and  not  based  upon 
quantitative  evidence.  The  first  impres- 
sion one  gets  upon  seeing  living  festiva 
from  Costa  Rica  and  Panama  is  one  of  a 
black  lizard  with  a  prominent  vertebral 
stripe.  The  lizards  from  Honduras  and 
Guatemala  have  a  broader  vertebral 
stripe  than  others  with  Pattern  Type  A; 
this  is  especially  striking  in  specimens 
from  Guatemala. 

Pattern  Type  B:  A  second  widely 
distributed  pattern  type,  largely  allo- 
patric  to  Pattern  Type  A,  is  peculiar  to 
lizards  from  the  Pacific  slopes  of  south- 
eastern Costa  Rica  and  western  Pana- 
ma, the  San  Bias  Coast  of  Panama  and 
Colombia  east  of  the  Cordillera  Central 
(Samples  15-19).  Specimens  from  this 
area  are  characterized  by  a  relatively 
broader  vertebral  stripe  than  that  of 
other  pattern  types  and  by  having  the 
ventrolateral  light  stripe  either  lacking, 
present  or  present  only  posteriorly.  The 
vertebral  stripe  broadens  posteriorly  and 
is  broadest  in  specimens  from  eastern 
Panama  and  Colombia.  Lizards  from 
the  San  Bias  Coast  of  Panama  and  from 
east  of  the  Cordillera  Central  in  Colom- 
bia possess  vertebral  stripes  that  may 
become  as  broad  as  the  middorsal  area 
at  the  base  of  the  tail  (Fig.  12B). 
Ameiva  festiva  from  the  Osa  Peninsula 
of  Costa  Rica  (Sample  18)  are  darker 
in  overall  coloration  than  those  from  the 


32 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


Fio.  12.  Patterns  of  Ameiva  f estiva  from  Colombia.  (A)  FMNH  165748,  male,  SVL  44  mm,  from 
the  Upper  Rio  Ure,  Depto.  Cordoba.  (B)  FMNH  165227,  male,  101  mm,  from  the  Upper  Rio 
Ure,  Depto.  Cordoba.    (C)  FMNH  43828,  female,  81  mm,  from  Pizzaro,  Intendencia  de  Choco  (?). 


adjacent  mainland.  Thi.s  is  not  the  re- 
sult of  preservation,  for  color  photo- 
graphs taken  by  Daniel  H.  Janzen  be- 
fore the  lizards  were  preserved  show 
them  to  be  darkly  pigmented.  Two 
specimens  of  f estiva  ( KU  80210  and  KU 
110745)  from  Almirante,  Bocas  del  Toro 
Prov.,  Panama,  exhibit  a  pattern  similar 
to  that  described  here,  although  others 
from  the  same  locality  show  Pattern 
Type  A.  Ameiva  festiva  from  Barro 
Colorado  Island  and  southeastern  Pana- 
ma are  somewhat  intermediate  between 
Pattern  Types  A  and  B,  but  because  the 
population  more  closely  resembles  the 
latter,  I  have  included  them  here. 

Pattern  Type  C:  Ameiva  festiva 
from  the  Choco  of  Colombia  (including 
Sample  20)  have  a  pattern  similar  to 
Type  A,  but  the  vertebral  stripe  is  ex- 
tremely straight  and  well  defined  even 
in  large  specimens  ( Fig.  12C ) .  The 
dorsolateral  and  ventrolateral  light 
stripes  are  also  well  defined,  may  be 
entire  or  dotted  (rarely  dashed),  and 
are  not  lost  in  large  individuals.  Pre- 
served specimens  are,  with  the  excep- 


tions of  the  light  stripes  and  vertebral 
stripe,  very  dark. 

Pattern  Type  D:  Adult  Ameiva  fes- 
tiva from  Isla  Escudo  de  Veraguas,  off 
the  northwest  coast  of  Panama,  are 
melanistic.  That  this  is  so  is  supported 
by  color  photographs  of  freshly  col- 
lected specimens  provided  by  Charles 
W.  Myers.  Very  little  pattern  is  ap- 
parent on  specimens  I  have  examined 
and,  although  examination  of  juvenile 
specimens  (unavailable  to  me)  might 
prove  them  to  belong  to  Pattern  Type 
A  or  B,  I  prefer  to  assign  them  to  a 
separate  pattern  type  for  the  present. 
Baskin  and  Williams  (1966)  summa- 
rized information  on  insular  populations 
of  melanistic  Ameiva  in  the  Caribbean, 
and  Mayr  (1963)  discussed  similar 
populations  of  Lacerta  in  the  Mediter- 
ranean. Such  melanistic  island  popula- 
tions are  usually  found  on  small,  diy, 
barren  islands  (  Mayr,  1963 ) ;  Heatwole 
and  Torres  ( 1966 )  attributed  the  phe- 
nomenon in  Ameiva  to  local  diftcrentia- 
tion,  those  species  of  Ameiva  exhibiting 
melanism    not    being    closely    related. 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


33 


Mayr  ( 1963 )  suggested  that  the  dark 
pigmentation  might  function  in  heat 
regulation.  The  sample  from  Isla  Es- 
cudo  de  Veraguas  docs  not  fit  the  pat- 
tern in  temis  of  habitat  as  outlined 
above.  According  to  C.  W.  Myers  (per- 
sonal communication),  one  of  the  few 
biologists  to  have  visited  the  island,  Isla 
Escudo  de  Veraguas  ".  .  .  is  covered 
with  humid  forest,  including  swamp 
forest,  except  for  a  well  developed  line 
of  strand  vegetation  at  some  points." 
Specimens  of  festiva  from  the  Bocas  del 
Toro  Archipelago  to  the  west  of  Isla 
Escudo  de  Veraguas  are  not  notably 
mclanistic  and  conform  to  Pattern  Type 
A,  although,  according  to  Myers,  habi- 
tats on  all  of  these  islands  are  similar. 
That  the  melanism  of  festiva  on  Isla 
Escudo  de  Veraguas  is  the  result  of 
local  differentiation  can  hardly  be 
denied,  but  the  adaptive  significance  of 
such   pigmentation   remains    a   mystery. 

Ecological  Notes. — In  the  northern 
part  of  its  range,  Ameiva  festiva  in- 
habits heavy  forest  (Stuart,  1943;  Smith 
and  Laufe,  1946;  Echternacht,  1968) 
when  sympatric  with  other  macroteiids. 
If  other  large  teiids  are  not  present,  the 
niche  of  the  species  is  expanded  to  in- 
clude more  open  habitats  (Echternacht, 
1968).  In  Mexico,  Guatemala,  and 
Honduras,  festiva  is  partly  sympatric 
with  undulata  and  with  various  species 
of  Cnemidophorus.  In  Costa  Rica  and 
Panama,  festiva  and  either  leptophnjs 
or  quadrilineata,  or  all  three  species 
may  occur  together.  In  the  vicinity  of 
Almirante  and  on  the  islands  of  the 
Bocas  del  Toro  Archipelago,  Bocas  del 
Toro  Pro  v.,  Panama,  festiva  and  quad- 
rilineata are  sympatric,  as  they  are  on 
Isla  Escudo  de  Veraguas.  Ameiva  fes- 
tiva typically  occupies  the  more  shaded 
parts  of  the  habitat,  whereas  quadriline- 
ata occurs  in  the  open  (C.  W.  Myers, 
personal  communication).  In  southeast- 
ern Costa  Rica  (vicinity  of  San  Isidro 
del  General,  San  Jose  Prov.;  vicinity  of 
Palmar  Sur  and  on  the  Oso  Peninsula, 
Puntarenas   Prov.)    festiva  is   sympatric 


with  both  quadrilineata  and  leptophnjs. 
Ameiva  festiva  occupies  a  habitat  in- 
termediate to  those  of  the  other  two 
species:  leptophnjs  occupies  more 
shaded  areas  and  quadrilineata,  the 
open  areas.  This  has  been  noted  by 
Hillman  (1969)  and  myself,  but  Dunn 
( 1940a )  reported  the  habitats  of  lepto- 
phnjs and  festiva  reversed  on  Barro 
Colorado  Island  in  Panama;  there  fes- 
tiva occupies  the  less  shaded  areas. 
Hillman  ( 1969 )  found  hatchling  A.  lep- 
tophnjs sympatric  with  adult  festiva  on 
the  Osa  Peninsula  and  concluded  that 
the  difference  in  size  between  the  two 
minimizes  competition.  The  reproduc- 
tive biology  of  festiva  has  been  investi- 
gated by  R.  E.  Smith  (1968a,  b)  in  the 
only  such  study  of  Ameiva  that  has  been 
undertaken.  Smith's  results  suggest  that 
festiva  is  reproductively  active  through- 
out the  year,  and  that  clutch  sizes  aver- 
age between  2.22  and  2.29  eggs. 

Nothing  has  been  reported  on  the 
ecology  of  A.  festiva  in  South  America. 

Remarks. — Ameiva  festiva  was  de- 
scribed by  Lichtenstein  and  von  Mar- 
tens ( 1856 )  on  the  basis  of  two  speci- 
mens, which  are  catalogued  under  one 
number  (ZMB  881)  in  the  Zoological 
Museum  of  Berlin.  The  description  is 
based  on  a  specimen  "Long,  a  rostro  ad 
caud.  bas.  3  1/2",  caud.  ultra  6""  (88.9 
mm).  The  syntypes  presently  have 
snout- vent  lengths  of  86  and  58  mm;  the 
tail  of  the  larger  is  broken.  I  am  herein 
designating  the  larger  syntype  as  the 
lectotype;  shrinkage  resulting  from  over 
one  hundred  years  in  preservative  would 
account  for  the  discrepancy  in  snout- 
vent  length  between  that  given  in  the 
description  and  my  measurement.  The 
smaller  syntype  becomes  a  paralecto- 
tyi^e. 

Dunn  (1943)  described  Ameiva  fes- 
tiva nicefori  from  Saisima,  Depto.  Cun- 
dinamarca,  Colombia,  but  I  have  pre- 
sented evidence  elsewhere  ( Echter- 
nacht, 1970)  that  the  population  de- 
serves specific  recognition.  Dunn  (1940a) 
and  Villa  (1968)   allied  the  population 


34 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


Fig.  13.  Map  showing  locality  records  of  Ameiva  quadrilineata.  To  avoid  crowding  of  symbols, 
some  localities  have  been  omitted.  Samples  used  in  statistical  analyses  were  ( 1 )  Southeastern  Nica- 
ragua, (2)  Tortuguero,  Limon  Prov.,  Costa  Rica,  (3)  Limon  and  vicinity,  Limon  Prov.,  Costa  Rica, 
(4)  Bocas  del  Toro  Prov.,  Panama,  (5)  Isla  Escudo  de  Veraguas,  Panama,  (6)  San  Isidro  del 
General,  San  Jose  Prov.,  Costa  Rica,  (7)  Palmar  Sur  and  vicinity,  Puntarenas  Prov.,  Costa  Rica, 
(8)  Vicinity  of  Rincon,  Osa  Peninsula,  Puntarenas  Prov.,  Costa  Rica,  (9)  Progreso,  Puerto  Ar- 
muelles  and  vicinity,  Chiriqui  Prov.,  Panama,    (10)   Boquete,  Chiriqui  Prov.,   Panama. 


of  Ameiva  on  the  Islas  del  Maiz,  Nica- 
ragua with  undulata  rather  than  festiva 
as  originally  described  (Barbour  and 
Loveridge,  1929a).  My  investigations 
support  this  conclusion,  and  the  popu- 
lation is  discussed  with  undulata  in  the 
present  work. 

Ameiva  quadrilineata  (Hallowell) 

Cnemidophorus  quadrilineatus  Hallowell,  Proc. 
Acad.  Nat.  Sci.  Philadelphia,  1860,  p.  483, 
1860  [Syntypes:  USNM  6053A,  6053B. 
Type  Locality:  "Nicaragua,"  restricted  to 
Greytown,  Depto.  Rio  San  Juan,  by  Tay- 
lor, Univ.  Kansas  Sci.  Bull.,  38(1):271, 
1956]. 


Ameiva  quadrilineata — Cope,  Proc.  Acad.  Nat. 
Sci.  Philadelphia,  1862,  p.  61,  1862.  Bo- 
court,  Mission  Scientifique  au  Mexique  et 
dans  TAmerique  Centrale,  fitudes  sur  les 
Reptiles,  livr.  4,  p.  259,  pi.  20A,  Fig.  9, 
1874.  Giinther,  Biologia  Centrali-Ameri- 
cana,  Reptilia  and  Batrachia,  p.  24,  1885. 
Dunn,  Proc.  Acad.  Nat.  Sci.  Philadelphia, 
92:113,  1940.  Taylor,  Univ.  Kansas  Sci. 
Bull.,  38(1):271,  1956. 

Ameiva  <^ahhiana  Cope,  J.  Acad.  Nat.  Sci. 
Philadelphia,  ser.  2,  8:33,  pi.  28,  Fig.  3, 
1875  [Syntypes:  USNM  32614-16.  Type 
Locality:  Old  Harbor  (=  Puerto  Viejo), 
Limon  Prov.,  Costa  Rica.  Collector:  W.  M. 
Gabb].  Giinther,  Biologia  Centrali-Ameri- 
cana,   Reptilia  and  Batrachia,  p.  23,   1885. 

Ameiva  undulata — Boulenger,  Catalogue  of 
the   Lizards   in  the  British   Museum    (Na- 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


35 


Fig.  14.  Head  scutellation  of  Ameiva  quad- 
lilineata:  (A)  Lateral  view  of  head,  (B)  Dor- 
sal view  of  head,  (C)  Ventral  view  of  head; 
KU  95593  from  1.5  km  E  Palmar  Norte, 
Puntarenas  Prov.,  Costa  Rica.  X  2.  (D)  Dor- 
sal view  of  head;  KU  96849  from  Cayo 
Zapatillo  Grande,  Bocas  del  Toro  Prov., 
Panama,    X   2.7. 


tiiral    History),    2nd    Ed. 
1885   (part). 


vol.    2,    p.    348, 


Distribution. — Atlantic  coastal  low- 
lands from  extreme  southeastern  Nica- 
ragua to  northwestern  Panama;  Pacific 
slopes  of  Costa  Rica  from  Quepos,  Pun- 
tarenas Prov.  (H.  S.  Fitch,  personal 
communication)  to  the  Azuero  Penin- 
sula, Panama;  the  Bocas  del  Toro  Archi- 
pelago, Isla  Escudo  de  Veraguas  and 
Isla  de  Cebaco  of  Panama  and  Isla  del 
Cafia  of  Costa  Rica.  Specimens  have 
been  collected  at  elevations  of  1100- 
1650  m  above  Paso  Ancho  and  on  Cerro 
Hornito,  Chiriqui  Prov.,  Panama  (Fig. 
13).   _ 

Diagnosis. — A  small  lizard  which 
can  be  distinguished  from  other  Middle 
American  Ameiva  by  the  following  com- 
bination of  characters:  Central  gular 
scales    moderately    enlarged,    irregular, 


hJ^^S 


Fig.  15.  Preanal  region  of  Ameiva  quadrili- 
ncata:  (A)  KU  95636  from  6  km  W  Palmar 
Norte,  Puntarenas  Prov.,  Costa  Rica,  X  2. 
(B)  KU  95593  from  1.5  km  E  Palmar  Norte, 
Puntarenas  Prov.,  Costa  Rica,   X  2. 


scales  of  posterior  one-half  of  gular  re- 
gion abruptly  smaller  than  scales  of 
anterior  one-half  (Figs.  14-15);  moder- 
ate number  of  femoral  pores  (mean 
number  for  males,  34.2;  females,  27.8); 
a  transverse  row  of  abruptly  enlarged 
mesoptychial  scales;  eight  longitudinal 
rows  of  ventral  scutes;  narrow  middor- 
sal  field  (mean  PV:  26.1)  which  does 
not  become  darker  laterally;  continu- 
ous, well  defined  dorsolateral  stripe  and 
continuous  to  interrupted  ventrolateral 
stripe,  both  usually  retained  throughout 
life.  The  middorsal  field  of  Ameiva  un- 
diilata  is  broad  (mean  PV:  males,  46.8; 
females,  45.6)  and  the  ventrolateral 
light  stripe  is  usually  interrupted. 
Ameiva  chaitzami  also  has  a  relatively 
broad  middorsal  field  (mean  PV:  38.1) 
and  interrupted  ventrolateral  light 
stripes  and,  in  addition,  lacks  a  uni- 
formly colored  middorsal  field. 

Description. — Maximum  observed 
SVL  88  mm  for  males,  82  mm  for  fe- 
males; supralabials  11-16  (13.7);  infra- 
labials  10-13  (10.1);  supraoculars  6-9 
(6.7);  COF  values  2-6  (3.0);  COP 
values  4-12  (9.1);  loreals  2-4;  nostril  in 
prenasal-postnasal  suture;  prefrontals  in 
contact  with  postnasals;  GAB  116-177 
(140.5);  PV  16-36  (26.1);  PV/GAB 
0.09-0.25  (0.19);  GOR  185-258  (218.6); 
GAB/GOR  0.41-0.78  (0.64)  for  males, 
0.56-0.75  (0.64)  for  females;  SAT  17-22 
(19.8)  for  males,  17-21  (19.5)  for  fe- 
males; total  preanal  scales  4-10  (7.1) 
for  males,  4-9  (6.8)  for  females;  termi- 
nal preanal  scales  1-3  (1.4)  for  males, 


36 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


37 


1-3  (1.3)  for  females;  prebrachials  en- 
larged, in  a  single  row;  postbrachials 
moderately  enlarged,  in  one  row  or  ir- 
regular; preantebrachials  in  three  rows 
proximally,  one  or  two  rows  distally; 
three  rows  of  enlarged  prefemoral 
scales;  infratibials  enlarged,  in  two 
rows;  femoral  pores  22-42  (34.2)  for 
males,  18-39  (27.8)  for  females;  sub- 
digital  lamellae  25-34  (29.5)  for  males, 
25-34  (28.8)  for  females;  longitudinal 
rows  of  ventral  scutes  28-33  (29.8); 
transverse  rows  of  ventral  scutes  4-10 
(8.0). 

Color  and  Pattern  in  Alcohol. — Juve- 
nile males:  Top  of  head  light  brown  to 
gray-brown;  side  of  head  gray  with 
black  wash  above  grading  to  gray  on 
lower  half  of  postnasals,  loreals  and 
supralabials;  middorsal  field  dark  brown 
to  dark  gray-brown,  narrow,  bounded 
laterally  by  narrow,  white  dorsolateral 
light  stripes,  which  originate  above  the 
eyes  and  extend  to  the  base  of  the  tail 
where  they  coalesce  and  form  a  verte- 
bral stripe  extending  well  onto  the  tail; 
dorsolateral  dark  fields  black;  ventro- 
lateral light  stripe  narrow,  white,  orig- 
inating beneath  eye  and  extending  to 
the  groin,  thence  onto  anterior  surface 
of  femur  where  it  runs  between  the  dor- 
sal granular  scales  and  the  ventral 
plates;  short,  narrow,  interrupted  white 


<1 

Fig.  16.  Ontogenetic  change  and  variation  in 
pattern  of  male  Ameiva  quadrilineata.  A-C; 
Pacific  population.  D-G;  Atlantic  population. 
Females  do  not  show  marked  ontogenetic 
changes  in  color  or  pattern  and  resemble 
juvenile  males  as  to  these  characters.  (A)  KU 
95623,  SVL  31  mm,  from  the  Rio  Zapote,  8 
km  E  Palmar  Norte,  Puntarenas  Prov.,  Costa 
Rica;  (B)  KU  95583,  68  mm,  from  8  km  SE 
San  Isidro  del  General,  San  Jose  Prov.,  Costa 
Rica;  (C)  KU  93984,  84  mm,  from  2-5  km 
ESE  Piedras  Blancas,  Puntarenas  Prov.,  Costa 
Rica;  (D)  KU  96837,  40  mm,  from  Isla  Basti- 
mentos,  Bocas  del  Toro  Prov.,  Panama;  (E) 
KU  108282,  62  mm,  from  E  end  Isla  Escudo 
de  Veraguas,  Panama;  (F)  KU  96845,  79 
mm,  from  Cayo  Zapatillo,  Bocas  del  Toro 
Prov.,  Panama;  (G)  Ventral  view  of  KU  96845 
to  show  dark  pigmentation. 


stripe  from  beneath  ear  to  shoulder; 
ventrolateral  dark  field  black  with 
brown  mottling  below;  dorsal  surfaces 
of  forelimbs  mottled  brown  and  black, 
an  interrupted  white  line  extending 
along  the  back  of  the  arm  from  shoulder 
to  wrist;  dorsal  surface  of  hind  limbs 
mottled  black  and  brown  with  white 
spots  on  shank  and  narrow,  white  post- 
femoral  stripe  extending  onto  tail  for 
ca.  10-12  scale  whorls;  ventral  surfaces 
immaculate  except  for  faint  gray  mark- 
ings in  the  gular  region  and  a  faint, 
transverse  gray  line  across  the  gular  re- 
gion at  the  juncture  of  the  large  anterior 
and  small  posterior  scales. 

Through  ontogeny  (Fig.  16)  black 
mottling  appears  in  the  middorsal  area, 
but  never  enough  to  disrupt  the  visual 
impression  of  a  unicolored  field.  Gray 
or  blue-gray  reticulations  appear  in  the 
ventrolateral  dark  field,  this  pattern  ex- 
tending onto  the  lateral-most  enlarged 
ventral  scutes  and  the  enlarged  ventral 
femoral  scales.  The  dorsolateral  dark 
field  remains  black  above,  but  brown 
vertical  bars  or  reticulations  appears  in 
the  lower  half.  The  white  spots  on  the 
hind  limbs  and  the  short  stripe  between 
ear  and  shoulder  disappear,  the  side  of 
the  neck  becoming  uniformly  gray- 
brown  or  black  with  gray  reticulations. 
Ventral  surfaces  may  be  pale  blue  over- 
all or  the  chin,  gular  region  and  throat 
may  be  salmon  colored,  the  remainder 
of  the  venter  pale  blue.  There  may  be 
black  flecks  in  the  midventral  region 
(see  discussion  of  geographic  varia- 
tion ) . 

Females:  Juvenile  females  resemble 
juvenile  males.  Adult  females  tend  to 
retain  the  juvenile  pattern  except  that 
black  flecking  appears  in  the  middorsal 
field  and  small,  brown,  longitudinally 
oriented  flecks  and  blotches  may  appear 
in  the  dorsolateral  dark  field  (Fig.  16). 

There  is  some  geographic  variation 
in  color  and  pattern  and  the  foregoing 
description  applies  to  specimens  from 
the  Pacific  lowlands  of  Costa  Rica. 

Color  and  Pattern  in  Life. — ^The  fol- 


38 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


lowing  description  is  extracted  from  the 
field  notes  of  C.  W.  Myers  and  refers  to 
a  series  of  Ameiva   quadriUneata    (KU 
96853-96865)  collected  13  km  N  Puerto 
Armuelles,   Chiriqui  Prov.,   Panama,   on 
22    April    1965.     Ameiva    quadriUneata 
from  this  locality  do  not  differ  in  color 
and  pattern  from  individuals   found  in 
the  Pacific  lowlands  of  Costa  Rica,  the 
latter  having  been  used  for  the  descrip- 
tion  of   color   and   pattern   in    alcohol. 
The  following  account   (modified  from 
Myers'  field  notes)   sei-ves  to  illustrate 
the  variability  of  ventral  pigmentation. 
Dorsolateral  and  lateral  stripes  vivid 
yellow  in  small   individuals   and   some- 
what duller  in  larger  ones;  dorsolateral 
stripes    nearly    disappearing    in    largest 
males.     Middorsal   region   light   yellow- 
brown  or  gray-brown,  with  or  without 
irregular  black  spotting,  and  turning  to 
a    whitish-gray    or   pale    brown    dorsal 
stripe  on  basal  one-third  of  tail.    Venter 
of  tail  base  copper,  but  all  but  basal 
part  of  tail  is  uniformly  brown  or  gray 
above   and   below    (in   a  juvenile   male 
this  tail  is  light  blue-gray).    Dorsolat- 
eral   area    of    body     (between    yellow 
stripes)    bright    rust,    variably    suffused 
with  black    (in  some  nearly   uniformly 
black    with    rust    spots).     Flanks    light 
brown     or     light     gray-brown,     heavily 
spotted  with  black.    Ventral  color  vari- 
able, as  follows:    KU  96853-5—3  juve- 
niles ( at  least  two  are  males ) :    Ventral 
surfaces    pale   copper.     KU   96856-9 — 4 
females:    Ventral  surfaces  pale  copper, 
perhaps  with  a  yellow  tinge  on  throat. 
KU    96860-1—2    males:     Underside    of 
head  pale  blue;  rest  of  venter  light  blue, 
except  tail  base  and  thighs  behind  fem- 
oral pores,  which  are  light  copper.    KU 
96862-3—2   males:     Underside   of   head 
light    coppery    orange;    rest    of    venter 
light  blue,  except  for  copper  thighs  be- 
hind femoral  pores,  and  light  blue  and 
copper-mottled  tail  base.    KU  96864-5 — 
2  males:    Underside  of  head  red-orange 
(pale  in  smaller  specimen  and  deep  in 
larger)  with  an  occasional  speck  or  spot 
of  blue  showing  through;  rest  of  venter 


light  blue,   except  rear   of  thighs,   and 
tail  base  (copper). 

In  addition  to  the  ventral  colors 
given  above,  both  males  and  females  of 
all  sizes  may  have  immaculate  or  pearly- 
white  ventral  surfaces.  The  chin,  gular 
region  and  throat  may  be  various  shades 
of  yellow. 

Geographic  Variation. — The  range  of 
Ameiva  quadriUneata  is  divided  by  the 
high    mountains    of    the    Cordillera    de 
Talamanca  of  Costa  Rica  and  Panama. 
Although  the  resulting  Atlantic  and  Pa- 
cific  populations   do   not   show   statisti- 
cally significant  differences  for  any  char- 
acters,   it   is    apparent   that    divergence 
has  taken  place.    Mean  values  for  num- 
ber   of   femoral    pores    of   females    and 
number    of   terminal    preanals    of   both 
sexes  are  higher  and  lower,  respectively, 
on  the  Atlantic  slope  than  on  the  Pacif- 
ic.   The  mean  number  of  terminal  pre- 
anal    scales    of   females    from    Boquete, 
Chiriqui  Prov.,  Panama  (Sample  10),  is 
statistically  different  from  mean  values 
of  all  other  samples  and   constitutes   a 
separate   subset.     Several   other   charac- 
ters show  similar  trends,  but  with  one 
sample    running    counter    to    the    trend 
established   by   the   others.     Characters 
having  lower  values  along  the  Atlantic 
slope    are    GAB,    GAB/GOR    for    both 
sexes,    and    total    number    of    preanal 
scales  for  both  sexes.    The  sample  from 
Boquete,  which  is  intermediate  in  geo- 
graphic   position    to    Puerto    Armuelles 
(Pacific  slope)  and  Almirante  (Atlantic 
slope)     in     Panama,     has    intermediate 
mean  values  to  those  of  Puerto  Armu- 
elles and  Almirante  for  some  characters 
and    more    closely    resembles    samples 
from  the  Atlantic  slope  than  those  from 
the   Pacific   slope   for   other   characters. 
Boquete  lies  on  the  Pacific  slope  of  the 
Cordillera    de    Talamanca    but    a    few 
kilometers   from   the   continental   divide 
in  western  Panama.    Sample  10  has  in- 
termediate values  for  number  of  supra- 
labials,  PV,  and  SAT  of  males.    Mean 
values     for     number     of     supraoculars, 
GAB,  GAB/GOR  of  males  and  females, 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


39 


and  total  number  of  preanals  of  both 
sexes  are  nearer  mean  values  for  Atlan- 
tic samples  than  for  Pacific  samples. 
Occasionally,  other  samples  on  the 
Pacific  slope  resemble  the  aggregate  of 
Atlantic  slope  samples  in  terms  of  mean 
values  of  one  character  or  another,  but 
none  with  such  noticeable  frequency  as 
does  Sample  10. 

Samples  8,  9  and  10  show  a  statis- 
tical relationship  to  one  another  for  sev- 
eral characters  and  constitute  a  separate 
subset  for  COF  and  COP  values.  Sam- 
ples 8  and  9  on  a  separate  subset  for 
GAB,  having  very  high  means  for  this 
character.  For  GAB,  Sample  10  has  the 
lowest  observed  mean  value  and  is  simi- 
lar to  Atlantic  slope  samples  in  this. 
Sample  6  has  the  lowest  observed  mean 
PV  and  constitutes  a  separate  subset. 

General  clinal  tendencies  are  noted 
for  some  characters.  The  total  number 
of  preanal  scales  decreases  from  north 
to  south  on  the  Atlantic  slopes,  but  in- 
creases from  north  to  south  in  samples 
from  the  Pacific  slopes  (except  Sample 
10,  which  has  a  very  low  mean);  COP 
values  decrease  from  north  to  south  on 
the  Pacific  side;  COF  values  increase 
from  north  to  south  on  the  Pacific  slope; 
the  number  of  subdigital  lamellae  of 
females  decreases  from  north  to  south 
on  the  Atlantic  slope.  Inclusion  of  more 
samples  in  the  analysis  is  necessary  in 
order  to  substantiate  these  trends. 

Mean  values  for  total  number  of  in- 
fralabials  and  for  numbers  of  rows  of 
longitudinal  and  transverse  ventral 
scutes  show  no  significant  differences 
among  the  ten  samples. 

Some  geographic  variation  is  appar- 
ent in  the  color  and  pattern  of  Ameiva 
quadrilineata.  Although  specimens 
from  Atlantic  and  Pacific  slope  popula- 
tions do  not  differ  markedly  among 
themselves,  there  are  differences  be- 
tween the  two  populations.  Adult  males 
from  the  Atlantic  side  are  never  as 
ornate  in  terms  of  dorsal  and  lateral 
color  and  pattern  as  those  from  the 
Pacific   slopes    (Fig.    16).     Males   from 


the  Atlantic  slopes,  especially  those 
from  northwestern  Panama  (including 
Samples  4  and  5)  have  all  ventral  sur- 
faces variably  flecked  with  black  (Fig. 
16);  the  dorsolateral  light  stripes  of 
specimens  from  this  area  may  be  scal- 
loped over  the  posterior  third  of  the 
body  and  may  become  diffuse,  blending 
with  middorsal  coloration,  in  large  in- 
dividuals; the  lateral  light  stripe  may 
be  interrupted  and  the  dorsal  caudal 
stripe  is  less  evident;  the  lateral  ground 
color  of  some  adults  is  black  mottled 
with  brown  above,  gray  below.  In  gen- 
eral, it  seems  that  the  chin,  gular  region 
and  throat  of  adult  male  quadrilineata 
from  the  Atlantic  slopes  becomes  yel- 
low when  brightly  pigmented,  but  that 
such  lizards  from  the  Pacific  slopes 
show  red  or  orange  coloration.  I  know 
of  only  one  exception  to  this:  KU  95600 
collected  at  Quebrada  Boruca  at  the 
Rio  Puente  Nuevo,  22  km  E  Palmar 
Norte,  Puntarenas  Pro  v.,  Costa  Rica,  on 
15  August,  1965,  was  yellow  on  the 
ventral  surface  of  the  head.  This  gener- 
ality needs  to  be  substantiated;  I  have 
had  no  color-in-life  notes  on  Atlantic 
slope  specimens  from  west  of  the  Bocas 
del  Toro  region  of  Panama. 

Ameiva  quadrilineata  from  Isla  Es- 
cudo  de  Veraguas  of  Panama  and  from 
the  Osa  Peninsula  of  Costa  Rica  do  not 
appear  to  be  any  darker  in  coloration 
than  those  from  the  adjacent  mainlands. 

Ecological  Remarks. — Ameiva  quad- 
rilineata is  ecologically  the  best  known 
member  of  the  genus  ( Hirth,  1963,  1965; 
R.  E.  Smith,  1968a,  b).  Both  Hirth  and 
Smith  worked  with  populations  of  quad- 
rilineata on  the  Atlantic  lowlands,  Costa 
Rica.  Hirth  (1963)  studied  all  aspects 
of  the  ecology  of  A.  quadrilineata  at 
Tortuguero,  Limon  Prov.,  Costa  Rica. 
At  Tortuguero  these  lizards  were  abun- 
dant on  the  beaches,  which  they  shared 
with  Basiliscus  vittatus.  Ameiva  quad- 
rilineata is  also  characteristic  of  the 
beach  fauna  on  the  islands  of  the  Bocas 
del  Toro  Archipelago  and  Isla  Escudo 
de  Veraguas  of  Panama  (C.  W.  Myers, 


40 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


personal  communication)  and  near  the 
northern  (western)  Hmit  of  its  range 
on  the  Pacific  coast  of  Costa  Rica  ( H.  S. 
Fitch,  personal  communication).  I  have 
collected  it  in  open  areas  (clearings  in 
forested  areas,  along  roadsides,  etc.) 
near  San  Isidro  del  General  and  Palmar 
Norte  in  southeastern  Costa  Rica.  Dvmn 
( 1940a )  reported  the  species  to  be  com- 
mon in  open  areas  along  large  rivers  on 
the  Atlantic  side  of  Costa  Rica.  Hill- 
man  (1969:477),  who  studied  quadrili- 
neata  on  the  Osa  Peninsula  of  Costa 
Rica,  found  them  foraging  "in  low  vege- 
tation adjacent  to  an  exposed  surface 
such  as  a  road."  The  species  is  defi- 
nitely more  heliophilic  than  either  fes- 
tiva  or  leptophrys,  with  which  it  is 
sympatric  in  some  areas.  It  is  similar  in 
habitat  preference  to  ameiva.  The  rela- 
tively small  size  of  adult  qiiadrilineata 
undoubtedly  helps  to  reduce  competi- 
tion between  it  and  its  larger,  sympatric 
congeners.  Competition  may  be  rela- 
tively more  intense  between  juvenile 
qiiadrilineata  and  juvenile  leptophrys 
or  festiva.  I  have  no  experience  with 
festiva  on  this  point,  but  I  found  juve- 
niles of  the  other  two  species  common 
in  tree-fall  clearings  in  a  swamp  forest 
near  Palmar  Norte  in  Costa  Rica.  The 
clearings  were  always  occupied  by  one 
of  the  two,  but  not  both  species. 

R.  E.  Smith  (1968a,  1968b)  studied 
Ameiva  qiiadrilineata  and  festiva  at 
Pandora,  Limon  Pro  v.,  Costa  Rica.  His 
investigations  indicate  that  both  species 
may  be  reproductively  active  through- 
out the  year.  The  mean  clutch  size  for 
qiiadrilineata  is  2.05-2.07  eggs,  depend- 
ing upon  method  of  counting,  and  ovi- 
position  occurs  between  17  and  21  days 
following  ovulation.  Smith  also  dis- 
cussed the  adaptive  significance  of  fat 
bodies  and  provided  experimental  evi- 
dence for  a  relationship  between  fat 
body  development  and  gonadal  cycles. 

Ameiva  are  usually  diurnal,  but 
W.  E.  Duellman  (personal  communica- 
tion) found  an  adult  male  quadrilineata 
active  at  night   on  a  road  in   western 


Panama.  There  had  been  a  heavy  rain 
earlier  in  the  day,  and  possibly  the  liz- 
ard was  flooded  from  its  burrow. 

Henry  S.  Fitch  (personal  communi- 
cation) is  studying  a  population  of 
quadrilineata  near  Quepos,  Puntarenas 
Pro  v.,  Costa  Rica.  The  nearest  locality 
from  which  specimens  of  undulata  are 
available  is  near  Parrita,  about  25  km  to 
the  northwest,  but  Fitch  has  observed 
both  species  at  Quepos.  There  quad- 
rilineata occurs  only  on  the  beach,  and 
undulata  is  inland  in  plantations  or  pas- 
tures where  shade  and  ground  cover  are 
abundant.  Quepos  is  the  northern-most 
locality  for  quadrilineata  on  the  Pacific 
coast,  and  the  southern-most  locality  for 
undulata. 

Remarks. — Barbour  and  Noble 
( 1915 )  apparently  confused  Ameiva 
quadrilineata  and  A.  undulata  pulchra. 
Both  were  described  by  Hallowell 
( 1860 )  from  "Nicaragua,"  but  quad- 
rilineata is  restricted  to  the  Atlantic 
coast  of  Nicaragua  and  pulchra  to  the 
Pacific  coast;  Barbour  and  Noble  had 
before  them  specimens  from  "Chinan- 
dega,  Nicaragua,"  which  is  on  the  Pacif- 
ic side.  Moreover,  the  color  description 
given  by  Barbour  and  Noble  fits  that  of 
undulata,  not  quadrilineata. 

Ameiva  undulata  (Wiegmann) 

Cnetnidophorus  undulatus  Wiegmann,  Herpe- 
tologica  Mexicana,  pp.  27-28,  1834  [Syn- 
types:  ZMB  867-869,  869  missing.  Type 
Locality:  "Mexico,"  restricted  to  Tehiian- 
tepec,  Oaxaca,  Mexico  by  Hobart  M.  Smith, 
Proc.  Biol.  Soc.  Washington,  53:56,  1940. 
Collector:     Unknown]. 

Ameiva  undulatus — Gray,  Catalogue  of  the 
lizards  in  the  British  Museum  ( Natural 
History),  p.  20,   1845. 

Ameiva  undulata — Cope,  Proc.  Acad.  Nat.  Sci. 
Philadelphia,  14:62-63,  1862.  Bocourt, 
Mission  Scientifique  an  Mexique  et  dans 
TAmerique  Centrale;  fitudes  sur  les  rep- 
tiles, livr.  3,  pp.  254-259,  pi.  20A,  Fig.  7a- 
7e,  pi.  20B,  Fig.  1.  Gunther,  Biologia  Cen- 
trali-Americana;  Reptilia  and  Batrachia, 
pp.  23-24,  1885.  Boulenger,  Catalogue  of 
the  lizards  in  the  British  Museum  ( Natural 
History),  vol.  2,  pp.  347-348,  1885.  Hart- 
weg  and  Oliver,  Occas.  Papers  Mus.  Zool., 
Univ.     Michigan,     359:7-8,     1937.      Dunn, 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


41 


» 


Proc.  Acad.  Nat.  Sci.  Philadelphia,  92:114- 
115,  1940. 

Ameiva  pulchra  Hallowell,  Proc.  Acad.  Nat. 
Sci.  Philadelphia,  p.  483,  1860  [Syntypes: 
ANSP  9133-9134.  Type  Locality:  "Nica- 
ragua."   Collector:     Unknown]. 

Cnemidophorus  amivoides  Cope,  Proc.  Acad. 
Nat.  Sci.  Philadelphia,  pp.  198-199,  1894 
[Holotype:  AMNH  16316.  Type  Locality: 
La  Carpintera,  San  Jose  Prov.,  Costa  Rica. 
Collector:     A.  Alfaro]. 

Ameiva  undulaia  undulata — Barbour  and 
Noble,  Bull.  Mus.  Comp.  Zool,  59(6):474- 
475,  1915.  Stuart,  Proc.  Biol.  Soc.  Wash- 
ington, 55:145,  1942.  Smith  and  Laufe, 
Univ.  Kansas  Sci.  Bull.,  31(2):62-64,  Fig. 
2b,  1946.  Smith  and  Taylor,  U.  S.  Natl. 
Mus.  Bull.,  199:174,  1950. 

Ameiva  undulata  parva  Barbour  and  Noble, 
Bull.  Mus.  Comp.  Zool.,  59(6)  :476-477, 
1915  [Holotype:  MCZ  5831.  Type  Locali- 
ty: "Guatemala,"  restricted  to  Mazatenan- 
go,  Depto.  Suchitepequez  by  Smith  and 
Laufe,  Univ.  Kansas  Sci.  Bull.,  31(2)  :51, 
1946.  Collector:  Unknown].  Stuart,  Proc. 
Biol.  Soc.  Washington,  55:145,  1942. 
Smith  and  Laufe,  LTniv.  Kansas  Sci.  Bull., 
31(2):51-54,  Fig.  la.  Pi.  la,  1946.  Smith 
and  Taylor,  U.  S.  Natl.  Mus.  Bull.,  199: 
173,  1950.  Taylor,  Univ.  Kansas  Sci.  Bull., 
38(1):26S-271,  1956.  Stuart,  Misc.  Publ. 
Mus.  Zool.,  Univ.  Michigan,   122:77,   1963. 

Ameiva  undulata  pulchra — Dunn,  Proc.  Acad. 
Nat.  Sci.  Philadelphia,  92:115,  1940. 
Stuart,  Proc.  Biol.  Soc.  Washington,  55: 
146,  1942. 

Aineiva  undulata  haitwegi  Smith,  Proc.  Biol. 
Soc.  Washington,  53:55,  1940  [Holotype: 
USNM  108600.  Type  Locality:  Across  the 
Rio  Usumacinta  from  Piedras  Negras, 
Depto.  El  Peten,  Guatemala.  Collector: 
H.  M.  Smith].  Stuart,  Proc.  Biol.  Soc. 
Washington,  55:145,  1942.  Smith  and 
Laufe,  Univ.  Kansas  Sci.  Bull.,  31(2):35- 
36,  Pi.  lib,  1946.  Smith  and  Taylor,  U.  S. 
Natl.  Mus.  Bull.,  199:171-172,  1950. 
Stuart,  Misc.  Publ.  Mus.  Zool.,  Univ.  Mich- 
igan, 122:77,  1963. 

Ameiva  undulata  stuarti  Smith,  Proc.  Biol.  Soc. 
Washington,  53:55-56,  1940  [Holotype: 
USNM  108601.  Type  Locality:  Palenque, 
Chiapas,  Mexico.  Collector:  H.  M.  Smith]. 
Stuart,  Proc.  Biol.  Soc.  Washington,  55: 
146,  1942.  Smith  and  Laufe,  Univ.  Kansas 
Sci.  Bull.,  31(2):50-51,  Fig.  lb,  Pi.  lb, 
1946.  Smith  and  Tavlor,  U.  S.  Natl.  Mus. 
Bull.,  199:173,  1950. 

Ameiva  undulata  amphigramma  Smith  and 
Laufe,  Trans.  Kansas  Acad.  Sci.,  48(3): 
338-344,  1945  [Holotype:  FMNH  100020. 
Type  Locality:  San  Andres  Tuxtla,  Vera- 
cruz, Mexico.  Collector:  H.  M.  Smith]. 
Smith  and  Laufe,  Univ.  Kansas  Sci.  Bull., 


31(2):43-47,  PI.  Ic-Id,  1946.  Smith  and 
Tavlor,  U.  S.  Natl.  Mus.  Bull.,  199-172, 
19.50. 

Ameiva  undulata  gaigeae  Smith  and  Laufe, 
Univ.  Kansas  Sci.  Bull.,  31(2):37-39,  Fig. 
Ic,  Pi.  lie,  1946  [Holotype:  FMNH 
100030.  Type  Locality:  Progreso,  Yucu- 
tan,  Mexico.  Collector:  H.  M.  Smith]. 
Smith  and  Taylor,  U.  S.  Natl.  Mus.  Bull., 
199:172,  1950.  Stuart,  Misc.  Publ.  Mus. 
Zool.,  Univ.  Michigan,   122:77,   1963. 

Ameiva  undulata  podarga  Smith  and  Laufe, 
Univ.  Kansas  Sci.  Bull.,  31(2):40-43,  Fig. 
Id,  2a,  1946  [Holotype:  FMNH  100050. 
Type  Locality:  7  mi.  W  Victoria,  Tamau- 
lipas,  Mexico.  Collectors:  H.  M.  Smith 
and  D.  H.  Dimkle].  Smith  and  Taylor, 
U.  S.  Natl.  Mus.  Bull.,  199:172,  1950. 

Ameiva  imdidata  dextra  Smith  and  Laufe, 
Univ.  Kansas  Sci.  Bull.,  31(2)  :54-,59,  Fig. 
2c,  1946  [Holotype:  FMNH  100080.  Type 
Locality:  nr.  Rincon,  Guerrero,  Mexico. 
Collectors:  E.  H.  Taylor  and  H.  M. 
Smith].  Smith  and  Taylor,  U.  S.  Natl. 
Mus.  Bull.,  199:173,  1950. 

Ameiva  undulata  sinistra  Smith  and  Laufe, 
Univ.  Kansas  Sci.  Bull.,  31(2):59-62,  1946 
[Holotype:  FMNH  100054.  Type  Locali- 
ty: Manzanillo,  Colima,  Mexico.  Collec- 
tor: H.  M.  Smith].  Smith  and  Taylor, 
U.  S.  Natl.  Mus.  Bull.,   199:174,   19.50. 

Ameiva  festiva  miadis  Loveridge  and  Barbour, 
Bull.  Mus.  Comp.  Zool.,  69(7)  :141,  1929 
[Holotype:  MCZ  26970.  Type  Locality: 
Isla  del  Maiz  Grande,  Depto.  Zelaya,  Nica- 
ragua.   Collector:    James  L.  Peters]. 

Ameiva  undulata  miadis — Dunn,  Proc.  Acad. 
Nat.  Sci.  Philadelphia,  92:115,  1940.  Villa, 
Rev.  Biol.  Trop.,  15(1):119,  1968.  Echter- 
nacht,  Breviora,  354:5,  1970. 

Distribution. — The  most  widely  clis- 
tiibuted  of  the  Middle  American  Amei- 
va (Fig.  17),  Ameiva  umJidata  occm-s 
from  near  Las  Varas,  Nayarit,  Mexico, 
to  the  vicinity  of  Quepos,  Puntarenas 
Prov.,  Costa  Rica,  on  the  Pacific  slopes. 
On  the  Atlantic  slopes,  the  species 
ranges  from  Rancho  Santa  Ana  (12.8 
km  SE  Podilla),  Tamaulipas,  Mexico,  to 
Puerto  Lempira,  Cabo  Gracias  a  Dios, 
Honduras,  and  on  more  westerly  Atlan- 
tic slopes  of  Nicaragua  from  Corozo, 
Depto.  Nuevo  Segovia  to  Tuli  Creek, 
Depto.  Rio  San  Juan.  Four  specimens 
(ANSP  1543S-41)  from  Huaunta,  Depto. 
Zelaya,  Nicaragua,  seemingly  document 
the  presence  of  Ameiva  undulata  on  the 
Caribbean   coast   of   that   country    (see 


42 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


24^ 


2 


8^ 


5° 


2' 


9' 


£>. 


-24^ 


21 


27-.a^^S^°  •34,  ,'     W 


0 


200      400 


KILOMETERS 


105^ 


99^ 


Fig.  17.  Map  showing  locality  records  of  Ameiva  undulata.  To  avoid  crowding  of  symbols,  some 
localities  have  been  omitted.  Samples  used  in  statistical  analyses  were  ( 1 )  Gomez  Farias  and  vicin- 
ity, Tamanlipas,  Mexico,  (2)  Tamazunchale  and  vicinity,  San  Luis  Potosi,  Mexico,  (3)  Tierra 
Colorado,  Veracruz,  Mexico,  (4)  Cuautlapam,  Veracruz,  Mexico,  (5)  San  Andres  Tuxtla  and 
vicinity,  Veracruz,  Mexico,   (6)   Vicinity  of  Jesus  Carran?a,    Veracruz,    Mexico,    (7)    Frontera    and 

(8)    Tres   Brazos   and   Balchacaj,  Campeche,  Me.xico,    (9)    Dzilbalchen, 

Piste,    Yucatan,    Mexico,  (11)  El  Cayo  and  vicinity,  Cayo  Dist.,  British 

and  Tikal,  Depto.  El   Peten,    Guatemala,    (13)    Piedras    Negras,    Depto. 

Canihor,   Depto.   Alta  Verapaz,  Guatemala,   (15)  El  Rancho  and  Finca 


vicinity.  Tabasco,  Mexico, 
Campeche,  Mexico,  ( 10 ) 
Honduras,  ( 12 )  Uaxactiin 
El   Peten,  Guatemala,    (14) 


Bucural,  Depto.  Progreso,  Guatemala,  (16)  Copan,  Depto.  Colon,  Honduras,  (17)  La  Ceiba, 
Depto.  Atlantida,  Honduras,  (18)  Coyoles  Central,  Depto.  Yoro,  Honduras,  (19)  Trujillo,  Depto. 
Colon,  Honduras,  (20)  Catacamas,  Depto.  Olancho,  Honduras,  (21 )  Isla  del  Maiz  Grande,  Depto. 
Zelaya,  Nicaragua,  (22)  Manzanillo-Pacific  Coast  Localities,  Colima,  Mexico,  (23)  Colima  and 
vicinity,  Colima,  Mexico,  (24)  Acapulco  and  vicinity,  Guerrero,  Mexico,  (25)  Chilpancingo  and 
vicinity,  Guerrero,  Mexico,  (26)  Chacalapa  and  vicinity,  Oaxaca,  Me.xico,  (27)  Tres  Cruces, 
Oaxaca,  Mexico,  (28)  Tolocito,  Oaxaca,  Mexico,  (29)  Tapanatepec,  Oaxaca,  Mexico,  (30)  Vicinity 
of  Cintalapa,  Chiapas,  Mexico,  (31)  Las  Tazas,  Chiapas,  Mexico  (51  km  E  Altamirano),  (33) 
Esquintla,  Chiapas,  Mexico,  (34)  Sabana  de  San  Quintin,  Chiapas,  Mexico,  (35)  Panajachel, 
Depto.  Solola,  Guatemala,  (36)  Finca  La  Trinidad,  Depto.  Jutiapa,  Guatemala,  (37)  JuHapa 
and  Finca  Mongoy,  Depto.  Jutiapa,  Guatemala,  (38)  Vicinity  of  La  Libertad,  Depto.  La  Liber- 
tad,  El  Salvador,  (39)  Chinandega,  Depto.  Chinandega,  Nicaragua,  (40)  Matagalpa,  Depto. 
Malagalpa,  Nicaragua,  (41)  Hacienda  La  Norma  and  vicinity,  Guanacaste  Prov.,  Costa  Rica,  (42) 
Canas,  Guanacaste  Prov.,  Costa  Rica,  (43)  Tilaran,  Guanacaste  Prov.,  Costa  Rica,  (44)  El  Zamo- 
rano  and  vicinity,  Depto.  Francisco-Morazan,  Hondmas,  (46)  Isla  Mujeres,  Quintana  Roo,  Mex- 
ico,   (48)   Apatzingan,   Michoacan,   Mexico.    Open   symbols  indicate  literature  records  of  specimen-S 

not  examined  for  the  present  study. 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


43 


Fig.  18.  Head  and  preanal  scutellation  of 
Ameiva  nndulata  (KU  94112;  Sabana  San 
Quintin,  Chiapas,  Mexico):  (A)  Lateral  view 
of  head,  (B)  Dorsal  view  of  head,  (C)  Ventral 
view  of  head,  X  1.1-  (D)  Preanal  region, 
X   1.5. 


Remarks).  The  species  is  known  from 
Isla  Mujeres,  east  of  Quintana  Roo, 
Mexico,  and  from  the  Islas  del  Maiz 
east  of  Nicaragua.  Gene  flow  between 
populations  on  the  Atlantic  and  Pacific 
slopes  occurs  across  the  Isthmus  of  Te- 
huantepec,  in  the  lowlands  of  eastern 
Guatemala,  and  in  Honduras  and  Nica- 
ragua south  of  the  highlands  that  mark 
Nuclear  Central  America.  Ameiva  tin- 
chiJata  is  generally  found  at  elevations 
below  1500  m,  although  there  are  ex- 
ceptions (e.g.,  at  Panajachel,  Depto. 
Solola,  Guatemala). 

Diagnosis. — Ameiva  umJulata  can  be 
distinguished  from  other  Middle  Ameri- 
can Ameiva  by  the  following  combina- 
tion of  characters:  Gentral  gular  scales 
slightly  to  greatly  enlarged,  regular  or 
irregular  in  arrangement;  gradual  re- 
duction in  size  of  scales  radiating  out- 


ward from  central  gular  scales;  prefron- 
tal scales  in  contact  with  postnasal 
scales;  three  parietal  scales  (four  if  me- 
dian parietal  divided);  a  transverse  row 
of  abruptly  enlarged  mesoptychial 
scales;  usually  eight  rows  of  ventral 
scutes;  a  relatively  broad  middorsal 
stripe  bounded  by  narrow  light  stripes 
which  may  be  lost  in  adults;  no  narrow 
vertebral  stripe;  lateral  pattern  empha- 
sizes longitudinal  stripes  or  vertical  bars 
in  adults,  never  spots. 

Description. — Maximum  observed 
SVL  129  mm  for  males,  111  mm  for 
females;  supralabials  12-18  (14.1)  for 
males,  12-18  (14.1)  for  females;  infra- 
labials  7-14  (10.2);  supraoculars  6-10 
(6.4);  COF  values  2-6  (2.2);  GOP 
values  2-14  (10.6);  SO-SG  values  2-10 
(3.8);  loreals  2-8   (4.0);  nostril  in  pre- 


nasal-postnasal     suture; 
contact    with    postnasal; 


prefrontal     in 
median    gular 


Fig.  19.  Head  and  preanal  scntellation  of 
Ameiva  unchdata  (KU  26963;  25  km  SE  Jesus 
Carranza,  Veracruz,  Mexico):  (A)  Lateral 
view  of  head,  (B)  Dorsal  view  of  head,  (C) 
Ventral  view  of  head,  X  1.1.  (D)  Preanal 
region,  X   1-5. 


44 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


Fig.  20.  Preanal  Region  of  Ameiva  undulata 
to  illustrate  two  of  the  pattern  types:  (A) 
KU  87417;  East  of  San  Andreas  de  la  Cruz, 
Guerrero,  Mexico;  Preanal  scales  in  a  single 
row  except  for  paired  terminal  scales,  X  2.7. 
(B)  KU  38232;  Chacalapa,  Oaxaca,  Mexico; 
Preanal  scales  in  a  single  row  including  single 
terminal  preanal,  X   1.5. 

scales  variable  in  size  and  orientation; 
GAB  108-202  (149.5);  PV  31-67  (47.4) 
for  males,  33-65  (46.4)  for  females; 
PV/GAB  0.21-0.40  (0.32)  for  males, 
0.23-0.43  (0.31)  for  females;  GOR  186- 
315  (246.0);  GAB/GOR  0.44-0.77 
(0.61)  for  males,  0.49-0.74  (0.60)  for 
females;  SAT  18-26  (22.3)  for  males, 
18-27  (22.2)  for  females;  total  preanal 
scales  4-13  (7.6)  for  males,  5-13  (7.5) 
for  females;  terminal  preanal  scales  1-3 
(2.0);  one  (usually)  or  two  rows  of  en- 
larged prebrachials;  postbrachials  en- 
larged, in  one  row;  preantebrachials  in 
two  rows,  one  row  or  two  rows  prox- 
imally  and  one  row  distally;  prefemoral 
scales  enlarged,  three  to  five  rows  prox- 
imally  reducing  to  two  (usually)  or 
three  rows  distally;  infratibials  enlarged, 
in  two  rows,  femoral  pores  29-48  (37.4) 
for  males,  22-45  (35.0)  for  females;  sub- 
digital  lamellae  24-38  (30.2);  longitu- 
dinal rows  of  ventral  scutes  26-34 
(30.1);  transverse  rows  of  ventral  scutes 
8-12  (8.2).  For  details  of  scutellation 
see  figures  18-20. 

Color  and  Pattern  in  Alcohol. — 
Throughout  its  broad  range,  Ameiva 
undulata  shows  geographic  variation  in 
color  and  pattern  unmatched  by  other 
Middle  American  species  of  Ameiva 
and  ontogenetic  variation,  especially  of 
males,  greater  than  all,  except  possibly 
A.  ameiva.  The  following  description 
of  color  and  pattern  in  alcohol,  as  well 
as  the  description  of  color  and  pattern 


in  life,  is  based  on  specimens  of  undu- 
lata from  the  vicinities  of  La  Ceiba  and 
Trujillo  along  the  north  coast  of  Hon- 
duras  (Localities  17  and  19,  Fig.  21). 

Juvenile  Males:  Top  of  head  brown; 
side  of  head  black  above,  grading 
through  brown  to  gray  on  supralabials; 
a  white  patch  beneath  and  slightly  be- 
hind eye;  ear  surrounded  by  white 
markings;  middorsal  field  yellowish- 
brown  or  greenish-brown  with  black 
flecking,  bordered  laterally  by  fine  white 
cream-colored  dorsolateral  stripes  which 
originate  above  eye  and  extend  well 
onto  tail;  dorsolateral  dark  stripe  ex- 
tending from  behind  eye  to  base  of  tail, 
velvet  black  with  no  light  bars  or 
blotches,  bounded  below  by  an  inter- 
rupted white  ventrolateral  light  line 
originating  behind  the  ear  and  extend- 
ing to  the  groin,  thence  as  an  uninter- 
rupted white  line  well  onto  tail,  ventro- 
lateral dark  field  gray-brown;  dorsal 
surfaces  of  limbs  mottled  black  and 
brown,  hind  limbs  with  white  spots 
above;  white  postfemoral  stripe  joining 
caudal  extension  of  ventrolateral  light 
stripe;  tail  brown  with  black  flecks 
above;  ventral  surfaces  immaculate  or 
blue-gray. 

Through  ontogeny  (Fig.  21)  the 
dorsolateral  light  stripe  disappears  or  is 
retained  only  on  the  anterior  half  of  the 
body.  The  ventrolateral  light  stripe  is 
usually  lost  entirely.  Light  blue  vertical 
bars  appear  on  the  flanks,  extending 
from  the  border  of  the  middorsal  field 
or  near  it  to  or  nearly  to  the  enlarged 
ventral  scutes.  Shorter  bars  and  blotches 
may  appear  between  the  primary  bars. 
The  light  spots  around  the  ear  are  lost 
and  the  venter  may  become  light  blue 


Fig.  21.  Ontogenetic  change  in  pattern  of 
Avieiva  undulata  from  northern,  coastal  Hon- 
duras. Males:  (A)  KU  101247,  SVL  71  mm; 
(B)  KU  101238,  92  mm;  (C)  KU  101237, 
110  mm.  Females:  (D)  KU  101242,  39  mm; 
(E)  JRM  2318,  88  mm;  (F)  JRM  2554,  109 
mm.  Small  males  resemble  small  females  as 
exemplified  by  KU  101242   (D  above). 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


45 


46 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


overall  except  for  black  patches  in  the 
lateral-most  scales  and  on  the  enlarged 
plates  on  the  anterior  surface  of  the 
femur.  Traces  of  red  pigment  are  visi- 
ble in  the  gular  region  of  some  large 
males. 

Females:  Juvenile  females  are  simi- 
lar to  juvenile  males.  Through  ontog- 
eny, females  generally  retain  the  juve- 
nile pattern,  except  that  the  dorsolateral 
light  stripe  may  be  lost  on  the  pos- 
terior one-half  of  the  body  and  regu- 
larly spaced  light  brown  blotches  ap- 
pear in  the  dorsolateral  dark  field  (Fig. 
21). 

Color  and  Pattern  in  Life. — The  fol- 
lowing is  based  on  field  notes  and  color 
photographs  of  Ameiva  tindulata  col- 
lected during  July,  1967,  near  La  Ceiba 
and  Trujillo  on  the  north  coast  of  Hon- 
duras. 

Adult  Males:  Top  of  head  gray- 
brown  to  light  brown;  middorsal  field 
gray-green  on  neck  becoming  bright 
leaf-green  on  shoulder  and  extending 
one-half  to  two-thirds  length  of  trunk, 
then  becoming  golden-brown  on  pos- 
terior part  of  body;  entire  middorsal 
field  irridescent  in  sunlight;  dorsal  sur- 
face of  tail  gray-brown  with  black  fleck- 
ing; middorsal  field  from  shoulder  to 
tail  lightly  to  moderately  blotched  with 
black  which  sometimes  appear  paired; 
remnant  of  dorsolateral  light  stripe 
cream,  if  present,  and  only  visible  on 
anterior  two-thirds  of  body;  lateral  pat- 
tern of  seven  to  ten  vertical  blue-white 
or  greenish-white  bars  which  may  touch 
middorsal  field  but  which  usually  are 
separated  from  it  and  which  taper  so 
that  they  are  wider  than  or  equal  to  the 
interspaces  dorsally  but  much  narrower 
than  interspaces  ventrally;  short  blue 
bars  extending  upward  between  the 
prominent  lateral  bars,  sometimes  con- 
necting with  them;  lateral  ground  color 
black  above  (in  area  of  dorsolateral 
dark  field  of  juveniles),  reddish-brown 
below.  Some  individuals  retain  a  rem- 
nant of  the  cream-colored  ventrolateral 
light  stripe  so  that  the  dorsolateral  dark 


field  may  be  defined.  Dorsal  surfaces  of 
limbs  brown  with  indistinct  black  mot- 
tling; ventral  coloration  of  limbs,  body 
and  tail  blue-white  with  black  flecking 
and  blotches,  especially  on  the  hind 
limbs  and  laterally  on  the  body;  chin, 
gular  region  and  neck  range  in  color 
from  white  to  brick  red  with  no  indica- 
tion of  yellow  pigmentation. 

Adult  Females:  Dorsal  pattern  simi- 
lar to  that  of  males  except  that  cream- 
colored  dorsolateral  light  stripes  usually 
are  retained;  dorsal  coloration  similar  to 
that  of  males  but  several  shades  darker 
overall;  ventrolateral  light  stripe  cream, 
continuous,  broken  or  broken  anteriorly, 
entire  posteriorly;  dorsolateral  dark  field 
black  with  brown  or  cream  blotches, 
each  wider  than  or  equal  to  width  of 
interspaces;  ventrolateral  field  brown  or 
gray-brown  with  darker  mottling;  dor- 
sal surfaces  of  limbs  as  for  males;  ven- 
tral surfaces  immaculate  or  with  slight 
bluish  tint  laterally  and  on  hind  legs. 
Females  lack  the  bright  vertical  bars  on 
the  flanks,  never  show  reddish  pigment 
on  the  ventral  surfaces  of  the  head  and 
neck  and  never  develop  blue  ventral 
coloration  as  markedly  as  do  males. 

Geographic  Variation. — The  follow- 
ing samples  were  removed  during  STP 
analysis  of  sexually  dimorphic  charac- 
ters because  of  inadequate  sample  sizes: 
For  characters  of  males,  samples  3,  9 
and  48;  for  characters  of  females,  sam- 
ples 7,  9,  23,  25,  26,  34,  38,  46  and  48. 
In  addition,  sample  3  was  removed  in 
the  analysis  of  number  of  loreals  of 
females  and  both  3  and  22  were  re- 
moved in  the  analysis  of  SO-SC  values 
of  both  sexes,  again  for  reasons  of  inade- 
quate sample  size. 

The  most  obvious  patterns  of  geo- 
graphic variation  for  meristic  characters 
of  Ameiva  undulata  are  clinal  trends 
along  the  northeast  and  northwest  coasts 
of  Mexico  and  along  the  Pacific  coast  of 
Middle  America  from  the  Isthmus  of 
Tehuantepec  to  Nicaragua  and  Costa 
Rica  ( Fig.  22 ) .  In  some  cases  one  sam- 
ple along  such  a  cline  is  "out  of  phase," 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


47 


interrupting  what  otherwise  is  a  rather 
smooth  trend.  Such  samples  are  indi- 
cated in  parentheses  in  the  following 
account. 

Those  characters  showing  an  in- 
crease in  sample  means  from  Tamaulipas 
to  the  Isthmus  of  Tehuantepec  include 
number  of  supraoculars  (2),  number  of 
paravertebral  granules  of  males,  num- 
ber of  paravertebrals  of  females  (3), 
total  preanal  scales  of  males,  total  pre- 
and  scales  of  females  (4)  and  number 
of  femoral  pores  of  males.  Shorter 
clines  in  the  same  direction,  but  involv- 
ing only  samples  1  through  3  or  4  are 
apparent  for  PV/GAB  of  males  and 
SAT  of  males  (the  size  of  sample  3  is 
too  small  for  inclusion  in  the  analysis 
for  both  characters),  as  well  as  number 
of  femoral  pores  of  females.  The  cline 
for  total  preanal  scales  of  males  and  of 
females  extends  southward  across  the 
isthmus  and  thence  eastward  along  the 
Pacific  coast  as  far  as  locality  33  for 
males  and  locality  36  for  females.  Two 
characters,  COF  values  and  number 
loreals,  show  clinal  increases  from  north 
to  south  across  the  isthmus  and  along 
the  coast  to  varying  degrees.  For  both 
characters,  lowest  mean  values  are  for 
sample  5.  COF  values  increase  through 
samples  28  and  29  and  through  samples 
6,  30  and  33  to  36.  The  number  of 
loreals  increases  through  samples  28,  27 
and  26  and  through  samples  28,  29  and 
30.  Mean  numbers  of  supraoculars 
show  the  same  trans-isthmian  pattern 
as  COF  values,  but  in  the  opposite  di- 
rection and  with  the  lowest  value  at 
sample  35,  not  36.  CAB  increases  from 
locality  5  through  28  and  29  or  6  and 
30  to  33  and  36. 

Decrease  in  mean  values  from  Ta- 
maulipas to  the  Isthmus  of  Tehuantepec 
is  notable  for  the  number  of  supralab- 
ials  of  females.  Shorter  clines  in  the 
same  direction  but  involving  only  sam- 
ples 1  through  3  or  4  are  apparent  for 
CAB  and  CAB /COR  of  males  (sample 
3  is  too  small  for  inclusion  in  the  analy- 
sis of  CAB /COR  of  males)  and  number 


Fig.  22.  Major  patterns  of  geographic  varia- 
tion in  Ameiva  ttndulata.  Arrows  represent 
geographical  trends  involving  several  charac- 
ters. Samples  on  one  side  of  the  heavy,  dashed 
line  differ  from  adjacent  samples  on  the  other 
side  of  the  line  in  a  statistically  significant 
manner  for  five  or  more  characters  but  do  not 
differ  in  such  a  way  from  adjacent  samples  on 
the  same  side  of  tlie  line.  See  text  for  further 
elal)oration. 


of  subdigital  lamellae.  Both  COP  and 
SO-SC  values  increase  from  south  to 
north  across  the  isthmus  and  a  west  to 
east  and  northeast  increase  is  noted  for 
SO-SC  values. 

Northwest  to  southeast  clines  are 
apparent  along  the  south  coast  of  Mex- 
ico also.  Characters  showing  increased 
sample  means  from  Colima  to  the  Isth- 
mus of  Tehuantepec  or  at  least  to  the 
vicinity  of  Chacalapa,  Oaxaca  (Sample 
26)  include  number  of  paravertebral 
granules  of  males  and  of  females,  PV/ 
CAB  of  males  and  of  females,  total  pre- 
anal scales  of  females  and  SAT  of  males 
(inadequate  sample  size  precludes  in- 
clusion of  sample  48  in  analysis  of  these 
characters).  Although  the  clinal  trend 
terminates  at  the  isthmus  for  the  other 
characters  mentioned,  that  for  total  pre- 
anals  of  females  continues  along  the 
Pacific  coast  to  locality  36  in  Cuatemala. 
A  short  trend  in  the  same  direction  is 
noted  for  number  of  transverse  rows  of 
ventral  scutes  from  localities  22  and  23 
through  48  to  24  and  25. 

Trends  along  the  same  coast,  but 
showing  a  decrease  in  sample  means 
from  Colima  to  the  Isthmus  of  Tehuan- 


48 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


tepee  are  noted  for  number  of  siipra- 
labials  of  males  (sample  48  excluded 
from  the  analysis)  and  females  (sam- 
ples 26  and  48  excluded),  number  of 
infralabials,  GAB/GOR  of  females, 
number  of  tenninal  preanals  and  num- 
ber of  subdigital  lamellae.  These  trends 
generally  involve  only  coastal  samples 
(samples  22,  24,  26,  27  and  29),  but  in- 
clude inland  samples  (samples  23,  25 
and  48)  for  supralabials  of  males  and 
total  number  of  infralabials. 

A  number  of  characters  show  clinal 
tendencies  for  increase  or  decrease  in 
sample  means  between  Nicaragua  or 
Costa  Rica  and  localities  33,  37  and 
especially  36  to  the  northwest.  There  is 
a  marked  tendency  for  clines  to  either 
begin  or  end  in  the  area  of  southeastern 
Mexico  and  southern  Guatemala  with 
no  clines  running  through  to  the  south- 
east or  northwest  along  the  coast.  Num- 
bers of  supraoculars  show  an  increase 
from  sample  40  (.x  =  6.13)  in  Nicara- 
gua to  sample  36  (x  =  9.97)  in  Guate- 
mala, at  which  point  there  is  an  abrupt 
decrease  in  sample  mean  and  a  clinal 
increase  from  sample  33  (x  =  6.28)  to 
sample  5  ( x  =  6.70 )  on  the  Gulf  side  of 
the  Isthmus  of  Tehuantepec  in  Mexico. 
Samples  34  (Ghiapas)  and  35  (Guate- 
mala) are  significantly  different  from 
the  nearest  samples  to  the  east  in  Guate- 
mala for  this  character,  but  sample  36 
(Guatemala)  is  not  significantly  differ- 
ent from  sample  33  (Chiapas).  A  simi- 
lar trend  for  increase  is  noted  between 
sample  39  (x  =  2.07)  in  Nicaragua  and 
sample  37  (x  =  2.29)  in  Guatemala  for 
COF  values.  Sample  36  (x  =  3.77), 
located  just  west  of  sample  37,  is  signifi- 
cantly different  from  it  and  is  the  end  of 
a  cline  beginning  at  sample  5  (x  = 
2.13)  and  which  progresses  across  the 
Isthmus  of  Tehuantepec  through  sam- 
ples 28  and  29  or  samples  6,  30  and  33. 
Samples  33  and  36  have  very  high  mean 
values  for  COF  and  are  statistically  iso- 
lated from  samples  to  the  north.  Fem- 
oral pore  means  for  both  males  and 
females  also  show  increases  from  sam- 


ple 40  to  sample  37,  but  the  trend  may 
be  traced  from  samples  41,  42  and  43  in 
Costa  Rica  as  well.  For  this  character 
in  females  the  cline  continues  to  sample 
15  and  may  also  be  traced  from  the 
Costa  Rican  samples  or  sample  40  in 
Nicaragua  through  samples  39  and  44 
to  sample  16  in  western  Honduras.  The 
number  of  supralabials  of  males  in- 
creases from  sample  37  north  and  west- 
ward to  sample  29,  and  GAB/GOR  for 
females  increases  from  sample  37  to 
sample  33  and  from  sample  15  to  sam- 
ple 33,  both  trends  running  to  the  west 
or  northwest.  Mean  numbers  of  sub- 
digital  lamellae  increase  from  samples 
20  and  44  through  samples  38  and  37  to 
sample  36  on  the  coastal  plain  of  Guate- 
mala and  to  sample  33  by  way  of  sam- 
ples 15  and  35.  Total  number  of  pre- 
anal  scales  of  females  follow  the  same 
trend.  These  two  characters  are  the 
only  ones  involved  in  clines  running 
through  the  area  of  samples  36  and  37. 
Mean  numbers  of  supralabials  of  fe- 
males increase  from  west  to  east,  be- 
ginning at  sample  30  and  terminating  at 
samples  14  and  37  by  way  of  samples 
33,  35  and  15.  Sample  36  (x  =  13.5), 
with  a  mean  for  this  character  lower 
than  that  of  sample  30  (x  ^  13.6)  is  not 
part  of  the  trend. 

Mean  numbers  of  paravertebral 
granules  of  males  and  of  females  and 
PV/GAB  of  females  show  trends  for  in- 
crease from  sample  40  in  Nicaragua  or 
sample  20  in  Honduras  southwestward 
and  along  the  Pacific  coast  to  sample  36 
in  Guatemala.  With  respect  to  number 
of  paravertebrals  of  females,  sample  36 
is  significantly  different  from  sample  37. 
For  both  number  of  paravertebrals  of 
females  and  PV/GAB  of  females,  the 
cline  may  be  traced  to  either  sample  36 
or  sample  15  in  Guatemala. 

Mean  SO-SC  values  and  mean  num- 
ber of  loreals  show  increased  values 
from  north  to  south  along  a  line  be- 
tween sample  9  on  the  Yucatan  Penin- 
sula and  samples  37  and  38  in  south- 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


49 


eastern    Guatemala    and    El    Salvador, 
respectively. 

There  seems  to  be  a  trend  toward 
increase  in  mean  numbers  of  paraverte- 
bral granules  of  females,  GAB/GOR  of 
females  and  total  number  of  preanal 
scales  of  males  from  sample  8  in  south- 
western Campeche  through  sample  10 
in  Yucatan  to  sample  46  on  Isla  Mujeres. 
There  is  a  similar  trend  beginning  at 
sample  7  in  Tabasco  and  terminating  at 
sample  10  for  mean  numbers  of  femoral 
pores  of  males.  For  all  of  these  charac- 
ters, however,  sample  9  in  central  Cam- 
peche ( Dzilbalchen )  is  too  small  for 
meaningful  comparison  (n  =  3)  where 
females  are  concerned  and  such  trends 
might  owe  their  existences  to  lack  of 
data  from  this  locality.  A  clinal  de- 
crease from  sample  8  through  sample  9 
and  10  to  sample  46  is  apparent  for 
mean  number  of  rows  of  transverse  ven- 
tral scutes.  An  increase  along  the  same 
line  is  noted  for  mean  number  of 
loreals. 

Most  samples  differ  from  adjacent 
samples  in  a  statistically  significant 
manner  for  at  least  one  character.  For 
purposes  of  this  discussion,  a  character 
judged  sexually  dimorphic  would  count 
as  two  characters  because  separate  con- 
sideration is  given  to  males  and  females. 
If,  for  example,  two  samples  were  sig- 
nificantly different  with  respect  to  num- 
ber of  femoral  pores  of  males  and  fe- 
males, they  would  be  said  to  differ  sig- 
nificantly for  two  characters,  not  one. 
Viewing  all  characters  at  once  gives  one 
the  impression  that  each  sample  is 
more-or-less  statistically  isolated  from 
its  neighbors.  Although  this  may  be 
true,  it  renders  analysis  and  formation 
of  meaningful  generalizations  extremely 
difficult.  However,  if  only  pairs  of  sam- 
ples differing  statistically  by  (arbi- 
trarily) five  or  more  characters  are 
examined,  some  interesting  patterns 
emerge.  These  are  discussed  below.  It 
must  be  emphasized  that  comparisons 
are,  in  most  cases,  being  made  only  be- 
tween adjacent  samples.    Two  samples 


adjacent  to  one  another  in  Honduras 
may  be  statistically  different  from  one 
another  on  the  basis  of  one  or  more 
characters,  but  neither  may  differ  from 
a  sample  in,  for  example,  northern 
Mexico.  Of  course,  the  opposite  is  also 
possible. 

The  population  of  Ameiva  iindulata 
on  Isla  del  Maiz  Grande  (sample  21) 
differs  from  samples  42,  43  and  44  for 
five  characters  each,  from  samples  39 
and  40  six  characters  each  and  from 
sample  20  for  ten  characters.  With  the 
exceptions  of  samples  40  and  41,  which 
differ  from  each  other  for  five  charac- 
ters, none  of  these  mainland  samples 
differ  from  adjacent  samples  among 
those  mentioned  by  more  than  four 
characters.  This  strongly  emphasizes 
the  divergence  of  the  Ameiva  on  Isla 
del  Maiz  Grande  from  the  nearest  main- 
land populations,  or  at  least  from  the 
available  samples  of  those  populations, 
and  is  indicated  by  the  heavy,  dashed 
line  in  figure  22.  Although  sample  20 
differs  from  sample  21  for  twice  or  more 
the  number  of  characters  separating 
sample  21  from  other  mainland  samples, 
there  is  still  little  basis  for  concluding 
that  A.  undulata  on  the  Islas  del  Maiz 
were  derived  from  elsewhere  because 
the  Honduranian  population  from  which 
sample  20  was  drawn  may  have  been 
modified  itself  through  introgression  of 
characteristics  of  populations  to  the 
north  and  west  (samples  17,  18  and  19). 

The  latter  three  samples  represent  a 
population  quite  distinct  from  neigh- 
boring samples  on  the  basis  that  none 
of  the  three  differ  among  themselves  by 
more  than  one  character  ( 17  and  18  dif- 
fer from  19  for  SO-SG  values;  other 
characters  show  non-significant  differ- 
ences among  sample  means),  but  all 
differ  from  other  nearby  samples  by 
more  than  five  characters  (see  Fig.  22). 
Samples  16,  20  and  44,  which  encircle 
samples  17,  18  and  19,  do  not  differ 
among  themselves,  but  all  show  con- 
siderable divergence  from  samples  17, 
18  and  19  for  meristic  and,  as  will  be 


50 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


discussed  later,  color  and  pattern  char- 
acters. Samples  17,  18  and  19  differ 
from  sample  11  (El  Cayo  and  vicinity, 
British  Honduras)  for  only  three,  one 
and  one  characters,  respectively.  Sam- 
ple 16,  on  the  other  hand,  differs  from 
sample  11  for  four  characters,  none  of 
which  are  among  those  few  which  dis- 
tinguish sample  11  from  samples  17,  18 
and  19. 

Samples  9-16,  31,  34,  35,  37,  38  and 
46  all  show  statistically  significant  dif- 
ferences involving  fewer  than  five  char- 
acters when  compared  among  them- 
selves with  respect  to  adjacent  samples. 
However,  sample  8  differs  from  samples 
12,  13,  31,  33  and  34  for  ten,  ten,  eight, 
ten  and  seven  characters,  respectively. 
In  addition  to  sample  8,  sample  31  dif- 
fers from  samples  6,  7,  30,  33  and  35  for 
six,  seven,  nine,  ten  and  nine  characters 
respectively.  Sample  33,  in  addition  to 
differing  from  samples  8  and  31  by 
more  than  five  characters,  differs  from 
samples  34  and  35  by  six  and  seven 
characters,  respectively.  Sample  36  dif- 
fers from  samples  15,  35  and  37  for 
seven,  eight  and  seven  characters,  re- 
spectively. Samples  8,  7,  6,  30,  33  and 
36  constitute  a  series  of  localities  be- 
ginning in  the  Tres  Brazos-Balchacaj 
area  of  Campeche,  progressing  west 
along  the  Gulf  coast  and  thence  south 
across  the  Isthmus  of  Tehuantepec  and 
eastward  along  the  Pacific  slopes  to  the 
vicinity  of  Finca  La  Trinidad,  Depto. 
Jutiapa,  Guatemala.  These  samples,  as 
indicated  above,  all  differ  from  adjacent 
more  inland  samples  to  the  south,  east 
or  north  by  five  or  more  characters.  If 
a  line  is  drawn  to  emphasize  the  diver- 
gence of  these  "peripheral"  samples 
from  the  other,  more  inland,  samples 
(Fig.  22),  none  of  the  samples  falling 
within  the  curvature  of  the  line  differ 
from  adjacent  samples  by  as  many  as 
five  characters  with  the  exception  of 
sample  15,  which  differs  from  sample  35 
for  six  characters. 

Sample  localities  from  the  northern 
Gulf  region  of  Mexico  (samples  1-5)  are 


isolated  from  sample  localities  along  the 
southwest  Pacific  coast  (samples  22-27, 
48)  by  the  Mexican  Plateau.  In  gen- 
eral, however,  samples  on  the  one  coast 
differ  for  five  or  more  characters  only 
from  the  most  distant  samples  on  the 
other  coast.  For  example,  sample  1 
(Tamaulipas)  differs  from  sample  27 
(south-central  Oaxaca)  for  seven  char- 
acters, but  differs  from  sample  22  (Co- 
lima)  for  only  three  characters.  Sample 
5  ( San  Andres  Tuxtla  and  vicinity, 
Veracruz)  differs  from  sample  27  for 
onlv  four  characters,  but  differs  from 
sample  22  for  nine.  Samples  1-5  and 
22-27,  48  do  not  differ  from  adjacent 
samples  on  the  same  coast  by  more  than 
two  characters,  with  the  exception  of 
samples  25  and  26,  which  differ  for  four 
characters. 

Sample  27  (Tres  Cruces,  Oaxaca), 
however,  seems  to  be  strongly  diver- 
gent from  adjacent  samples  to  the  east. 
Sample  27  differs  from  samples  6,  28 
and  29  for  ten,  six  and  eight  characters, 
respectively.  This,  coupled  with  the 
isolation  afforded  by  the  Mexican  Pla- 
teau, is  the  basis  for  separating  off  the 
samples  from  the  Pacific  slopes  of  Mex- 
ico northwest  of  the  Isthmus  of  Tehuan- 
tepec as  shown  in  figure  22. 

The  groupings  of  samples  set  apart 
according  to  the  method  just  described 
fits  well  with  the  general  pattern  of 
clinal  trends  discussed  earlier.  There  is 
also  a  generally  good  fit  by  these  group- 
ings with  the  placement  of  highland 
regions  with  elevations  above  1500 
meters,  an  elevation  above  which  Amei- 
va  rarely  occur  in  Middle  America. 

Using  the  14  samples  (8,  15-21,  27, 
31,  33,  35,  37  and  44)  which  were  found 
to  differ  from  adjacent  samples  by  five 
or  more  characters,  it  is  possible  to  ar- 
rive at  some  idea  of  which  characters 
are  most  discriminatory  in  distinguish- 
ing adjacent  samples.  The  14  samples 
were  compared  with  adjacent  samples 
to  give  68  different  comparisons.  It  was 
found  that  in  53  per  cent  of  these  com- 
parisons   (36  out  of  the  possible  68), 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


51 


mean  GAB  differed  in  a  statistically 
significant  manner.  GAB  is  apparently 
the  primary  meristic  indicator  of  sam- 
ple differences,  followed  by  GOR  (43 
per  cent  of  the  comparisons),  SO-SC 
values  (31%),  number  of  subdigital 
lamellae  (31%)  and  COP  values  (29%). 
All  other  characters  were  involved  in 
26  per  cent  or  less  of  the  comparisons 
each.  Two  characters,  number  of  su- 
pralabials  of  males  and  number  of  trans- 
verse rows  of  ventral  scutes,  were  found 
not  to  distinguish  any  of  the  68  pairs  of 
samples  compared.  All  five  of  the  char- 
acters which  seem  most  important  in 
this  respect  do  not  appear  together  in 
distinguishing  two  adjacent  samples, 
but  they  do  appear  in  groups  of  2,  3  or 
4  in  various  comparisons.  GAB  and 
GOR,  as  might  be  expected,  occur  to- 
gether most  often  in  distinguishing  pairs 
of  adjacent  samples,  appearing  in  26  of 
the  68  comparisons  (38%).  GAB  and 
number  of  subdigital  lamellae  occur  to- 
gether in  distinguishing  15  pairs  among 
the  68  (22%),  whereas  GAB  and  SO-SC 
values,  and  GAB  and  COP  values  both 
occur  together  in  14  of  68  comparisons 
(20%).  Other  combinations  of  the  five 
characters  taken  two  at  a  time  occur 
less  frequently. 

The  condition  of  the  preanal  scales 
of  Ameiva  tindtilata  can  be  analyzed  in 
a  number  of  ways.  Two  of  these,  the 
total  number  of  preanals  and  the  num- 
ber of  terminal  preanal  scales  have 
been  defined  and  discussed  with  respect 
to  STP  analysis.  These  characters,  along 
with  a  third  which  combines  the  two, 
warrant  further  discussion. 

For  STP  analysis,  the  number  of 
terminal  preanal  scales  was  coded  1,  2 
or  3,  indicating  one,  two  or  three  scales 
in  the  terminal  position.  An  inherent 
weakness  in  this  system  is  that  a  sample 
mean  or  range  gives  no  indication  of 
the  actual  frequency  distribution  of 
single,  double  and  triple  terminal  pre- 
anals in  the  sample.  A  sample  of  two 
individuals,  one  having  one  terminal 
preanal,  the  other  having  three,  would 


have  the  same  mean  number  of  terminal 
preanals  as  another  sample  of  two  in- 
dividuals where  each  animal  had  two 
terminal  preanal  scales.  This  is  an  over- 
simplification complicated  by  larger 
sample  sizes.  Even  statistics,  such  as 
standard  deviation,  are  of  little  help  in 
visualizing  the  frequency  distribution. 
For  this  reason,  the  number  of  terminal 
preanal  scales  was  coded  so  that  three 
terminal  preanal  scales  received  the 
same  value  as  one.  This  system  has 
some  validity  in  that  only  scales  in  the 
midline  are  being  compared.  Individ- 
uals having  two  terminal  preanals  con- 
tinue to  receive  a  code  value  of  two.  If 
this  is  done,  most  samples  still  have  a 
mean  number  of  tenninal  preanals  near 
or  approaching  2.00,  but  certain  sam- 
ples have  much  lower  values.  The  low- 
est of  these  are  samples  1  (x  =  1-67), 
8  (1.60),  17  (1.67),  22  (1.42),  23 
(1.61),  25  (1.76),  26  (1.66),  27  (1.33) 
and  48  (1.78).  Means  of  other  samples 
ranged  from  1.80  (sample  9)  to  2.00 
(many  samples).  It  will  be  noted  that 
most  of  the  low  sample  means  are  en- 
countered in  samples  22-27  and  48  be- 
tween Colima  and  Tres  Cruces,  Oaxaca, 
in  Mexico.  Sample  24,  consisting  of  14 
individuals  each  with  two  terminal  pre- 
anal scales,  does  not  fit  the  pattern. 

Coupled  with  the  number  of  termi- 
nal preanal  scales  is  the  pattern  of  these 
scales.  They  may  be  (A)  mostly 
(>50%)  single  (Fig.  20B),  (B)  mostly 
paired,  (C)  single  except  the  terminal 
scales  paired  (Fig.  20A),  or  (D)  mixed. 
The  latter  category  includes  individuals 
not  fitting  into  any  other  category  if  the 
single  or  paired  condition  is  based  on 
three  or  more  scales  or  pairs  of  scales  in 
a  row.  Most  individuals  of  Ameiva  tin- 
dtilata fall  into  class  C  (71.6%;  n  r= 
1003),  whereas  fewer  are  referable  to 
classes  A  (18.2%),  B  (4.5%)  and  D 
(5.7%).  Two  series  of  samples,  how- 
ever, show  marked  deviation  from  this 
distribution.  If  samples  22-27  and  48, 
from  the  Pacific  slopes  of  Mexico  north 
and   west   of   the   Isthmus    of   Tehuan- 


52 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


tepee,  are  eombined,  there  are  about 
equal  numbers  of  individuals  referable 
to  classes  A  (33.9%),  B  (27.3%)  and  D 
(30.6%),  but  only  8.3  percent  are  refer- 
able to  class  C  (n  =  121).  Individual 
lizards  in  samples  41-43  from  Costa 
Rica  are  distributed  as  follows:  class 
A  (44.2%);  class  B  (3.9%);  class  C 
(35.1%);  class  D  (16.9%)  (n  =  77). 
On  the  basis  of  this  character  alone 
samples  41-43  differ  markedly  from 
samples  to  the  north  (samples  39-40  in 
Nicaragua),  but  the  divergence  of  sam- 
ples 22-27  and  48  parallels  that  noted 
for  number  of  terminal  preanal  scales 
and  appears  to  reinforce  the  evidence 
for  a  break  in  gene  flow  between  popu- 
lations of  iindiilata  east  and  west  of  a 
line  drawn  between  sample  27  and 
samples  28  and  29.  If  samples  21-27, 
41-43  and  48  are  removed,  the  distribu- 
tion of  individuals  in  the  remaining 
samples  becomes:  class  A  (13.4%); 
class  B  (1.1%);  class  C  (84.6%);  class 
D  (0.9%)   (n  =  805). 

Another  character  not  analyzed  by 
STP,  but  warranting  further  mention  is 
the  pattern  of  the  median  gular  scales. 
Median  gular  scales  may  be  either  ir- 
regular in  arrangement  or  oriented 
longitudinally  in  a  more-or-less  regular 
row.  If  irregular,  they  are  usually  at 
least  slightly"  enlarged  (39.0%;  404  of 
1026  individuals),  although  a  few  in- 
dividuals have  midgular  scales  no  larger 
than  surrounding  scales  (0.4%).  If 
longitudinally  oriented  and  enlarged, 
they  may  be  paired  (2.1%),  in  a  row  of 
single  scales  (34.8%)  or  mixed  paired 
and  single  (23.1%;  at  least  three  scales 
or  pairs  of  scales  in  a  row  were  deemed 
necessary  to  classify  a  lizard  as  having 
longitudinally  oriented  scales).  Sam- 
ples 3-8,  15,  22-30,  33,  and  35-36  are 
made  up  of  individuals  more  than  70 
percent  of  which  have  enlarged,  longi- 
tudinally oriented  gulars.  Samples  5,  7, 
8  and  28  contain  only  such  individuals. 
These  sample  localities  are  distributed 
along  the  Gulf  coast  of  Mexico  east  and 
west    of   the    Isthmus    of   Tehuantepec 


from  northwestern  Veracruz  to  the  Tres 
Brazos-Balchacaj  region  of  Campeche, 
across  the  Isthmus  and  along  the  Pacific 
coast  from  Colima  to  southeastern 
Guatemala  (Fig.  17).  Samples  9-14,  17- 
19,  31  and  34  from  the  Yucatan  Penin- 
sula, eastern  Mexico,  northern  Guate- 
mala and  northern  Honduras  are  com- 
posed of  individuals  more  than  70  per- 
cent of  which  have  enlarged,  irregularly 
arranged  gulars.  Samples  13  and  34 
consist  of  only  such  individuals. 

These  patterns  are  generally  re- 
flected in  the  lines  between  adjacent 
samples  depicting  significant  differences 
involving  five  or  more  characters  (Fig. 
22).  Gular  pattern  tends  to  place  sam- 
ples 17-19  from  northern  Honduras  with 
samples  from  northern  Guatemala  and 
Yucatan  Peninsula.  The  line  separating 
samples  6-8,  30  and  33  from  samples  12- 
13,  31  and  34  is  represented,  but  sam- 
ples 15  and  35  are  not  differentiated 
with  respect  to  gular  scale  pattern  and 
these  samples  do  not  differ  from  sample 
36.  Samples  15,  35  and  36  differ  from 
one  another  for  at  least  five  of  the  char- 
acters analyzed  by  STP.  With  respect 
to  gular  pattern,  sample  36  differs 
sharply  from  sample  37  and  most  other 
samples  to  the  south  and  east.  Sample 
36  contains  only  one  individual  out  of 
30  (3.3%)  with  irregularly  arranged 
gulars,  whereas  sample  37  contains  13 
of  31  (41.9%)  individuals  with  irregular 
gular  scales. 


Fig.  23.  Selected  geographic  variants  of  color 
pattern  of  adult  Ameiva  undulata.  (A)  KU 
101300,  male,  SVL  126  mm,  (B)  KU  101301, 
female,  109  mm,  both  from  Isla  del  Maiz 
Grande,  Depto.  Zelaya,  Nicaragua;  (C)  KU 
87409,  male,  93  mm,  from  5  km  E  Vista  Her- 
mosa,  Oaxaca,  Mexico;  (D)  UMMZ  41435, 
female,  99  mm,  from  Cuautlapam,  Veracruz, 
Mexico.  (E)  KU  95645,  male,  92  mm,  from 
3  km  E  Mapastepec,  Chiapas,  Mexico;  (F)  KU 
95643,  female,  82  mm,  from  4.5  km  E  Mapa- 
stepec, Chiapas,  Mexico.  (G)  KU  101260, 
male,  93  mm,  (H)  KU  101267,  female,  90 
mm,  both  from  El  Zamorano,  Depto.  Fran- 
cisco-Morazan,  Honduras. 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


53 


54 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


Samples  16  ( 19%  with  irregularly 
arranged  gvilars;  n  =  16),  40  (20%;  n 
=  15)  and  2  (73%;  n  =  15)  stand  out 
as  having  unusually  high  or  low  num- 
bers of  individuals  with  irregularly 
arranged  gular  scales  relative  to  adja- 
cent samples.  Sample  2  would  seem  to 
belong  with  samples  from  the  Yucatan 
Peninsula,  northern  Guatemala  and 
northern  Honduras,  whereas  samples  16 
and  40  are  more  like  samples  involved 
in  the  Isthmus  of  Tehuantepec  "X." 

In  samples  other  than  those  spe- 
cifically mentioned  above,  the  percent- 
age of  irregularly  arranged  gular  scales 
ranged  from  33  to  63. 

Discussion  of  geographic  variation 
in  color  and  pattern  over  the  broad 
range  of  Ameiva  undulata  is  compli- 
cated by  the  presence  of  both  onto- 
genetic variation  (especially  among 
males)  and  sexual  dimorphism  (Stuart, 
1942).  The  magnitude  of  ontogenetic 
change  encountered  is  apparent  in  fig- 
ure 21,  and  sexual  dimorphism  in  pat- 
terns of  adults  from  selected  popula- 
tions is  represented  in  figure  23.  That 
geographic  variation  is  considerable  is 
also  apparent  from  figure  23.  There  are 
two  basic  pattern  types  (Stuart,  1942). 
The  first  of  these  is  characterized  by 
an  emphasis  on  longitudinal  stripes 
rather  than  blotches  or  bars  on  the 
flanks,  by  the  small  amount  of  sexual 
dimorphism  evident,  and  by  its  re- 
stricted geographic  range.  This  pattern 
type  occurs  only  in  samples  7  and  8  of 
the  present  study,  but  it  apparently 
grades  into  other  pattern  types  to  the 
west  and  south.  Smith  and  Laufe  ( 1946, 
PI.  IB)  have  illustrated  an  individual 
exhibiting  this  color  pattern  from  Teno- 
sique.  Tabasco,  Mexico.  These  authors 
cite  specimens  having  this  pattern  from 
Palenque  and  San  Ricardo,  Chiapas; 
Frontera,  Teapa  and  Tenosique,  Ta- 
basco; Balchacaj  and  Tres  Brazos,  Cam- 
peche.  I  have  examined  two  specimens 
from  San  Ricardo,  Chiapas  (FMNH 
106715,  male;  UIMNH  26157,  female). 
The  male  has  10  light-colored  blotches 


in  the  dorsolateral  dark  field,  a  pattern 
not  characteristic  of  the  striped  forms 
of  undulata.  Nine  males  (FMNH 
106715;  UIMNH  8437,  8439,  30229, 
39233-4,  39237-40)  from  Finca  San  Bar- 
tola,  ca  12  mi  SW  of  Cintalapa  and 
near  San  Ricardo,  also  show  prominent 
bars  and/or  blotches  on  the  flanks.  The 
specimens  from  Finca  San  Bartola  and 
San  Ricardo  comprise  sample  30  of  this 
study;  on  the  basis  of  both  meristic  and 
pattern  characters,  this  sample  is  dis- 
tinct from  those  to  the  north.  The 
striped  pattern  type  seems  to  be  re- 
stricted to  the  lowlands  of  Tabasco,  ex- 
treme western  Campeche  and  northern 
Chiapas.  The  pattern  is  not  unlike  that 
of  quadrilineata  (Fig.  16)  of  Nicaragua, 
Costa  Rica  and  Panama  or  chaitzami 
(Fig.  27)  of  eastern  Chiapas,  Mexico 
and  Guatemala.  Some  undulata  having 
the  striped  pattern  share  with  chaitzami 
and,  to  a  much  lesser  extent,  quadrili- 
neata, the  presence  of  a  secondary  dark 
stripe  in  the  middorsal  field  medial  to 
the  dorsolateral  light  stripes. 

The  second  basic  pattern  type  is 
characterized  by  a  highly  variable  pat- 
tern of  bars  and /or  blotches  on  the 
flanks  of  adult  males  often  accompanied 
by  loss  of  the  ventrolateral  light  stripe 
or  its  reduction  to  a  series  of  dashes  or 
spots.  However,  some  adult  females,  as 
well  as  juveniles  of  both  sexes,  retain  a 
pattern  of  longitudinal  stripes  ( Figs. 
21,  23).  Specimens  of  Ameiva  undidata 
included  in  samples  4,  6  and  28  possess 
this  pattern  type  (Fig.  23  C  and  D; 
Smith  and  Laufe,  1946,  Plate  I  C  and 
D).  The  dorsolateral  dark  field  is  re- 
tained in  both  sexes,  but  the  ventro- 
lateral light  stripe  may  be  completely  or 
partially  lost  in  adult  males  such  that 
the  ventral  border  of  the  dark  field  is 
indistinct.  The  ventrolateral  light  stripe 
is  usually  retained  in  females.  The  dor- 
solateral light  stripe,  which  is  usually 
retained  in  females  may  be  completely 
lost  in  males  or  retained  only  anteriorly. 
In  many  specimens  the  middorsal  field 
contains   dark  patches  which  lie  para- 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


55 


vertebrally  iind  may  be  paired.  This  is 
especially  notable  in  specimens  from 
sample  6.  A  characteristic  of  individuals 
from  these  samples  is  a  light  stripe  with 
indistinct  borders  lying  in  the  dorso- 
lateral dark  field  and  extending  from 
the  ear  to  the  groin  (Fig.  23  C  and  D). 
This  stripe  is  usually  continuous  but,  if 
broken,  the  interspaces  are  much  nar- 
rower than  the  remnants  of  the  stripe. 
The  stripe  is  white  or  blue-white  in 
males,  pale  brown  in  females  and  is  nar- 
rower and  less  distinct  in  females.  The 
ventrolateral  field  contains  vertically 
oriented  bars  or  blotches  on  a  dark 
background  in  adult  males.  Some  males 
in  sample  5  show  a  tendency  for  the 
blotches  in  the  dorsolateral  field  to  be 
continuous  with  those  in  the  ventrolat- 
eral field.  The  ventrolateral  field  of 
females  is  mottled.  Females  in  samples 
6  and  28  occasionally  have  a  secondary 
dark  stripe  between  the  dorsolateral 
light  stripes  and  the  lighter  brown  area 
of  the  middorsal  field,  the  same  as  de- 
scribed for  individuals  from  samples  7 
and  8. 

Adult  male  A.  wulnhta  in  samples 
1  and  2  are  similar  in  pattern  to  those 
just  described,  but  the  dorsolateral  dark 
field  contains  a  series  of  large,  quad- 
rangular, white  or  blue-white  blotches 
which  are  two  or  three  times  as  wide  as 
the  interspaces.  These  blotches  are 
never  fused  into  a  solid  stripe.  The 
dorsolateral  dark  field  of  females  con- 
tains similar,  but  less  prominent 
blotches.  The  dorsolateral  fight  stripe, 
which  is  retained  in  its  entirety  in  some 
adult  females,  terminates  at  one-half  to 
two-thirds  of  the  distance  from  occiput 
to  groin  in  juvenile  males  and  may  be 
present  only  on  the  neck  in  adults. 

Adult  males  from  samples  22,  23  and 
48  are  similar  to  those  of  samples  1  and 
2,  except  that  the  blotches  in  the  dorso- 
lateral dark  field  may  be  subequal  in 
width  to  the  interspaces.  Samples  24 
and  25  contain  adult  males  with  dorso- 
lateral blotching  as  in  samples  22  and 
23,    but    also    contain    some    males    in 


which  the  blotches  are  fused  into  a  con- 
tinuous stripe  as  described  for  samples 
4,  6  and  28.  In  addition,  there  is  a 
greater  tendency  for  the  dorsolateral 
light  stripe  to  be  retained  in  males  from 
samples  24  and  25  than  in  males  from 
samples  22  and  23. 

With  few  exceptions,  other  samples 
of  Ameiva  undulata  from  Pacific  drain- 
age localities  in  Mexico  present  a  variety 
of  lateral  patterns  ranging  from  pres- 
ence of  blotches  in  the  dorsolateral  dark 
field  and  spots,  blotches  or  bars  in  the 
ventrolateral  field  to  having  the  entire 
flanks  barred  or  a  combination  of  both 
patterns.  No  one  pattern  type  prevails 
among  males  of  any  one  sample.  Adult 
males  in  sample  44  (El  Zamorano  and 
vicinity,  Depto.  Francisco-Morazan, 
Honduras)  are  unusual  in  that  both 
males  and  females  possess  numerous 
(12-13  per  side)  small  blotches  in  the 
dorsolateral  dark  field.  These  blotches 
are  tear-drop  shaped,  and  some  extend 
ventrally  (Fig.  23  G  and  H).  This  pat- 
tern is  not  apparent  elsewhere. 

Samples  9-14,  16-19,  21,  34,  39-43 
and  46  are  characterized  by  a  lateral 
pattern  of  white  or  blue-white  bars 
which  extend  from  the  dorsolateral  light 
stripe  or  just  beneath  it  to  or  almost  to 
the  enlarged  ventral  scutes  ( Figs.  21  C 
and  23  A;  Taylor,  1956,  Fig.  67).  The 
bars  show  little  or  no  differentiation 
into  dorsolateral  and  ventrolateral  com- 
ponents. The  dorsolateral  dark  field  is 
usually  present  in  adult  males,  but  it  is 
difficult  to  define  because  it  grades  grad- 
uallv  into  the  ventrolateral  field.  In 
such  males  the  dorsolateral  and  ventro- 
lateral light  stripes  are  retained  only 
anteriorly  and  the  ventrolateral  light 
stripe  may  be  completely  lacking  in 
large  adults.  Sexual  dimorphism  is 
marked.  With  the  exception  of  those 
from  Isla  del  Maiz  Grande  (sample  21), 
adult  females  retain  a  juvenile  pattern 
with  the  addition  of  blotches  in  the 
dorsolateral  dark  field  (Fig.  21  D-F). 
Adult  females  from  Isla  del  Maiz 
Grande  resemble  adult  males  in  pattern 


56 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


TABLE    2. — Total   number   of   lateral   bars 
for   some   samples   of   male   Ameiva   undulata. 
Arranged    in    descending    order    according    to 
sample  mean. 


Sample 
Number 

X 

S.D. 

N 

Range 

46 

29.7 

1.11 

7 

28-31 

12 

25.4 

3.47 

9 

20-31 

9 

24.6 

3.58 

5 

20-28 

10 

24.3 

3.20 

14 

19-29 

34 

21.6 

1.67 

5 

20-24 

42 

21.0 

0 

4 

21 

21 

20.0 

1.95 

20 

16-23 

41 

20.0 

1.81 

12 

16-22 

43 

19.8 

1.34 

7 

18-22 

18 

19.6 

2.25 

16 

16-23 

14 

19.5 

1.77 

8 

17-22 

13 

19.4 

1.81 

9 

16-21 

39 

18.6 

1.78 

10 

17-22 

40 

18.3 

1.97 

6 

16-21 

17 

17.8 

1.28 

8 

16-20 

19 

16.8 

1.92 

5 

15-20 

(Fig.  23  A-B),  but  not  in  color.  The 
total  number  (sum  of  number  on  both 
sides)  of  vertical  bars  of  males  varies 
considerably  among  these  samples 
(Table  2).  The  largest  sample  means 
are  those  of  samples  on  the  Yucatan 
Peninsula  and  Isla  Mujeres.  The  sample 
from  Isla  Mujeres  (sample  46)  has  a 
mean  of  29.7,  the  largest  recorded. 
Lowest  sample  means  occur  in  the  two 
northern  coastal  samples  from  Hon- 
duras (samples  17  and  19),  with  means 
of  17.8  and  16.8,  respectively.  Males 
from  samples  9-14,  17-19,  34  and  46  may 
be  differentiated  from  barred  males 
from  other  samples  in  that  they  have 
little  or  no  secondary  spotting  or 
blotching  between  the  bars  in  the  ven- 
trolateral field.  Taken  as  a  whole, 
barred  males  from  other  samples  have 
such  markings  to  a  greater  degree,  al- 
though individuals  from  these  samples 
may  show  few  secondary  markings. 

The  general  ontogenetic  changes  ex- 
hibited by  the  barred  populations  of 
Ameiva  undulata  are  illustrated  in  Fig- 
ure 21.  Neill  (1961,  1965),  reported  that 
young   undulata   from   parts   of   British 


Honduras  lack  lateral  light  stripes,  the 
ontogenetic  progression  being  from  un- 
striped  hatchlings  through  light-striped 
subadults  to  barred  adults.  Presumably 
this  refers  to  ontogeny  of  males  only. 
Neill  ( 1965 )  also  called  attention  to 
two  small  specimens  (MCZ  71623-24) 
from  Augustine,  Cayo  Dist.,  British 
Honduras.  MCZ  71623  has  four  light 
lines  on  each  side,  whereas  MCZ  71624 
has  three.  Augustine  is  in  the  ecolog- 
ically distinctive  Mountain  Pine  region 
of  British  Honduras. 

Among  samples  of  Ameiva  undulata 
there  is  considerable  geographic  varia- 
tion in  amount  of  dark  pigment  scat- 
tered as  flecks  or  blotches  in  the  mid- 
dorsal  area.  These  dark  markings  usual- 
ly take  the  form  or  more-or-less  paired 
or  alternating,  paravertebral  spots,  al- 
though in  heavily  pigmented  specimens 
a  reticulate  pattern  may  be  evident. 
The  latter  is  especially  evident  in  speci- 
mens from  Isla  del  Maiz  Grande  (Fig. 
23  A-B).  Both  sexes  may  exhibit  such 
markings.  Although  they  are  sometimes 
evenly  distributed,  the  markings  are  of- 
ten restricted  to  or  heaviest  on  the  pos- 
terior one-half  or  two-thirds  of  the 
back.  I  have  not  attempted  to  quantify 
this  aspect  of  pattern  but  feel  justified 
in  making  the  generalization  that  the 
samples  along  the  Pacific  slopes  from 
southeastern  Chiapas,  Mexico  through 
El  Salvador  (samples  33,  35-38)  and 
northward  into  Honduras  south  of  the 
northern  coastal  ranges  (samples  16,  20 
and  44)  are  the  most  heavily  marked  of 
the  mainland  populations  sampled. 
Specimens  from  Isla  del  Maiz  Grande 
(sample  21)  have  very  heavy  middorsal 
marking,  as  noted  above,  but  this 
marking  takes  a  different  form,  appear- 
ing to  be  a  continuation  of  the  lateral 
pattern.  Samples  39  and  40  are  variable 
with  respect  to  this  character  and  sam- 
ples 41-43  show  very  light  markings.  A 
few  specimens  of  Ameiva  undulata  are 
available  from  the  Meseta  Centrale  of 
Costa  Rica.  This  population  was  named 
by    Cope     (1894)     as    Cnemidophorus 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


57 


Fig.  24.  Aiijeiva  undulata  (Holotype  of 
Cnemidopliorus  amivoides  Cope;  AMNH 
16316)  from  La  Caipintera,  Cartago  Prov.?, 
Costa  Rica,  illustrating  the  heavy  blotching  in 
the  middorsal  area  and  the  divided  inter- 
parietal; a  male,  SVL  64  mm. 

amivoides  (see  Remarks).  These  speci- 
mens all  show  a  high  degree  of  marbling 
or  blotching  in  the  middorsal  region 
(Fig.  24),  a  pattern  sharply  divergent 
from  that  of  other  populations  of  undu- 
lata at  the  southern  end  of  its  range. 
Further  study  of  this  population  is  de- 
sirable. Samples  other  than  those  spe- 
cifically mentioned  above  show  variable 
markings  from  light  to  moderate  in 
extent. 

There  is  little  geographic  variation 
with  respect  to  color  other  than  that  di- 
rectly associated  with  pattern.  Studies 
of  such  variation  are  hampered  by  a 
paucity  of  good  color-in-life  descrip- 
tions. Such  descriptions  are  available 
for  samples  from  Michoacan,  Mexico 
(Duellman,  1961),  the  Yucatan  Penin- 
sula (Maslin,  1963;  Duellman,  1965a), 
British  Honduras  (Neill,  1965)  and 
Chiapas  (Alvarez  del  Toro,  1960). 
Other  descriptions  have  been  gleaned 
from  my  own  field  notes  and  those  of 
William  E.  Duellman. 

The  most  striking  aspect  of  geo- 
graphic variation  in  color  has  to  do  with 
color  in  the  gular  region  of  adult  males. 
As  noted  for  Ameiva  festiva  and  quad- 
rilineata,  the  gular  coloration  of  males 
may  be  associated  with  reproductive 
condition.  In  many  specimens  through- 
out the  range,  gular  coloration  (and 
ventral  coloration  in  general)  is  white 
or  cream.    In  other  specimens,  the  en- 


tire venter  is  pale  blue.    A  number  of 
individuals,    however,    show    either   red 
or  yellow  pigment  in  the  gular  region 
and  over  the  anterior  part  of  the  chest 
and   ventral   surfaces   of  the   forelimbs. 
The  red  coloration  ranges  from  orange 
or  brick-red  to  scarlet.    The  yellow  pig- 
ment is  a  bright  lemon  yellow.   Whether 
one  color  precedes  the  other  in  a  dy- 
namic system  of  change  is  not  known, 
but  I  have  observed  no  specimens  in- 
termediate in  coloration.   The  specimens 
upon    which    this    discussion    is    based 
were    collected    during    June,    July    or 
August  between   1959   and    1966  by   a 
number   of  collectors.    Yellow  pigment 
has  been  observed  to  the  exclusion  of 
red  at  only  three  localities  in   Mexico: 
along  the  Rio  Frio,  Tamaulipas;  Finca 
Orizaba,     Chiapas;     and    Tapanatepec, 
Oaxaca.    At  some  localities,  individuals 
may  have  either  red  or  yellow  pigment 
in    the    gular   region.     Thus,    Duellman 
(1965a)  reported  that  about  half  of  his 
sample  from  Piste,  Yucatan  had  orange 
throats,  whereas  half  had  yellow  throats, 
corroborating  an  observation  by  Maslin 
(1963).     Other    localities    from    which 
both    colors    have    been    reported    are 
Felipe    Carillo    Puerto,    Quintana    Roo 
(Duellman,    1965a),    British    Honduras 
( Neill,  1965 )  and  the  Tepalcatepec  Val- 
ley   of    Michoacan    (Duellman,    1961). 
Specimens    from    a    number    of    other 
localities    in   Veracruz,    Tabasco,    Cam- 
peche,  Yucatan,  Quintana  Roo,  Chiapas, 
Jalisco,    Honduras    and    Isla    del    Maiz 
Grande,      Nicaragua     had      only      red- 
throated  individuals. 

There  seems  to  be  some  geographic 
variation  in  coloration  of  the  middorsal 
area.  This  area  is  iridescent  moss- 
green  anteriorly  grading  to  yellowish- 
brown  posteriorly  in  specimens  col- 
lected by  me  at  La  Ceiba,  Coyoles  and 
Trujillo,  Honduras  (samples  17-19,  re- 
spectively). It  is  reported  as  "olive" 
for  specimens  from  British  Honduras 
(Neill,  1965)  and  Chiapas  (Alvarez  del 
Toro,  1960).  Specimens  from  the  Te- 
palcatepec  Valley,    Michoacan    (Duell- 


58 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


man,  1961);  Piste,  Yucatan  (Maslin, 
1963);  along  the  Rio  Frio,  Tamaulipas; 
16  km.  SW  Frontera,  Tabasco;  Mapas- 
tepec,  Chiapas;  vicinity  of  Melaque, 
JaHsco;  El  Zamorano,  Depto.  Francisco- 
Morazan,  Honduras  (personal  observa- 
tion) have  brown  middorsal  pigmenta- 
tion. Few  descriptions  of  lateral  ground 
color  of  living  lizards  are  available,  but 
there  may  be  considerable  geographic 
variation  for  this  character  as  well. 

Ecological  Notes. — Despite  its  broad 
range  and  relative  abundance  in  many 
parts  of  that  range,  there  have  been  no 
thorough  ecological  investigations  of 
Ameiva  undulata.  Literature  on  this 
subject  is  limited  to  miscellaneous  ob- 
servations. Thus,  Stuart  (1951)  found 
undulata  altitudinally  distributed  to 
about  2000  m  in  the  "moderately  hu- 
mid" region  near  Panajachel,  Depto. 
Solola,  Guatemala,  but  commented  that 
in  more  moist  areas  the  altitudinal  limit 
is  reached  about  400  m  lower.  Else- 
where, the  species  seems  to  occur  below 
1500  m,  and  its  vertical  distribution  may 
be  limited  to  below  the  level  of  occa- 
sional frosts  (1200-1500  m,  depending 
upon  humidity),  a  factor  considered  by 
Stuart  (1966)  to  be  of  possible  bio- 
logical significance. 

Throughout  most  of  its  range, 
Ameiva  undulata  is  found  in  forest  or 
forest-edge  situations  (Alvarez  del 
Toro,  1966;  Duellman,  1965a;  Echter- 
nacht,  1968;  Rand,  1957;  Stuart,  1935, 
1950,  1954b,  1958)  or,  in  drier  regions, 
dense  thickets  and  scrub  (Maslin,  1963; 
Schmidt  and  Stuart,  1941;  Stuart,  1948). 

Stuart  (1950),  Duellman  (1966), 
and  Echternacht  (1968)  commented  on 
the  sciophilic  nature  of  Ameiva  iindu- 
lata,  although  I  pointed  out  that  even 
the  more  shade-tolerant  species  of 
Am^eiva  may  utilize  open  areas  in  the 
absence  of  competition  from  more  he- 
liophilic  macroteiids  like  Cnemidopho- 
rus.  Stuart  (1950)  reported  undulata 
to  be  abundant  in  the  grassland  and  ri- 
parian habitats  of  Depto.  Alta  Verapaz, 
Guatemala,     and     Echternacht     ( 1968 ) 


found  the  species  to  be  common  in  open 
situations  near  Panajachel,  Depto.  So- 
lola, Guatemala.  Although  the  lizards 
shun  the  open  beach,  I  found  the 
species  occupying  clearings,  paths  and 
other  open  areas  as  well  as  thickets  and 
piles  of  debris  in  the  coconut  groves  of 
Isla  del  Maiz  Grande,  Depto.  Zelaya, 
Nicaragua.  In  northern  Honduras, 
western  Nicaragua,  northwestern  Costa 
Rica,  and  in  those  parts  of  Mexico  with 
which  I  am  familiar,  undulata  occurs  in 
shaded  areas;  the  more  open  parts  of 
the  habitat  are  occupied  by  various 
species  of  Cnemidophorus. 

Virtually  nothing  has  been  reported 
on  the  reproduction  of  this  species  of 
Ameiva.  As  I  have  remarked  elsewhere, 
research  on  this  topic  may  reveal  the 
sequence  and  basis  for  changes  in  gular 
pigmentation  of  adult  males. 

Stuart  (1958)  collected  56  juveniles 
and  only  four  adults  of  Ameiva  undu- 
lata at  Tikal,  El  Peten,  Guatemala,  be- 
tween February  and  May,  1956,  but  he 
cited  a  collection  made  by  Gosner  at 
the  same  locality  at  unspecified  dates  in 
1947  and  1949  that  contained  a  "fine 
series  of  adults."  This,  along  with  my 
own  observations,  suggests  that  adult 
and  subadult  Ameiva  may  have  differ- 
ent seasonal  activity  periods.  However, 
there  have  been  no  observations  of  a 
quantitative  nature  to  substantiate  this. 

Remarks. — I  have  examined  four 
specimens  of  Ameiva  undulata  (ANSP 
15438-15441)  from  "Wounta"  (=  Hua- 
unta)  Haulover,  Depto.  Zelaya,  Nica- 
ragua. This  locality  is  at  least  290  kilo- 
meters from  other  known  localities  for 
the  species.  Duellman  (1958)  exam- 
ined three  specimens  (ANSP  15445- 
15447)  of  the  snake  Leptodeira  annu- 
lata  rhomhifera  Giinther  from  the  same 
locality,  far  removed  from  the  re- 
mainder of  the  range  of  the  species.  Al- 
though it  is  possible  that  a  snake  might 
have  been  carried  to  Huaunta  by  man, 
it  is  unlikely  that  a  moderately  large, 
active  lizard  could  be  so  transported. 
Ameiva  undulata  shuns  deep  forest,  and 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


59 


it  is  unlikely  to  have  arrived  at  Huaunta 
along  rivers  from  the  west,  but  it  is  pos- 
sible that  the  species  exists  in  isolated 
or  semi-isolated  populations  along  the 
Atlantic  coast  of  Nicaragua  northward. 
An  alternative  hypothesis  would  be  that 
the  A7newa  and  Leptodeira  populations 
at  Huaunta  are  relict  and  that  a  more 
careful  survey  of  the  region  would  un- 
cover other  such  species  as  well.  This 
part  of  Central  America  certainly  de- 
serves more  attention  from  biologists 
than  it  has  received. 

Such  an  explanation  does  not  apply 
for  records  of  Ameiva  undidata  from 
Panama  (Brennan  and  Yunker,  1966). 
All  specimens  so  reported  are  lepto- 
phrys. 

Taylor  (1956)  placed  Cnemidopho- 
rus  amivoides  Cope  from  the  Meseta 
Central  of  Costa  Rica  in  the  synonymy 
of  Ameiva  undidata,  and  I  see  no  rea- 
son to  dispute  this  judgment.  I  have 
called  attention  to  the  unusual  color 
pattern  of  specimens  from  the  Meseta 
Central  (Fig.  24),  but  so  few  are  avail- 
able that  further  comment  is  unjustified 
at  this  time. 

Dunn  (1940a)  and  Villa  (1968)  in- 
cluded the  population  of  Ameiva  on  the 
Islas  del  Maiz,  Depto.  Zelaya,  Nica- 
ragua (sample  21)  in  the  species  undu- 
lata.  The  population  was  described  as 
a  subspecies  of  festiva  by  Barbour  and 
Loveridge  (1929a).  I  have  substan- 
tiated the  assignment  of  this  population 
to  undulata  ( Echternacht,  1970). 

Ameiva  chaitzami  Stuart 

Ameiva  chaitzami  Stuart,  Proc.  Biol.  Soc. 
Washington,  55:143-145,  1942  [Holotype: 
UMMZ  90638.  Type  Locality:  "Along 
Cahabon-Languin  trail  about  2  km  north 
of  Finca  Canihor  .  .  .  ,  Alta  Verapaz, 
Guatemala."  Collector:  L.  C.  Stuart]. 
Stuart,  Misc.  Publ.  Mus.  Zool.,  Univ.  Mich- 
igan No.  122,  p.  77,  1963.  Echternacht, 
Breviora,  354:  7,  1970. 

Ameiva  undidata  stuarti — Stuart,  Occas.  Papers 
Mus.  Zool,  Univ.  Michigan  No.  471,  p.  21, 
1943. 

Ameiva  undulata  thomasi  Smith  and  Laufe, 
Univ.  Kansas  Sci.  Bull.  31(2):47-50,  Pi. 
lA,  1946  [Holotype:    FMNH  100006.   Type 


92» 


'     "\  — 


5° 


I 


',• 


s 


10  0  50 

H 1 

KILOMETERS 


92° 


90° 


89° 


88° 


Fig.  25.  Map  showing  locality  records  of 
Ameiva  chaitzami.  Samples  used  in  statistical 
analyses  were  ( 1 )  Comitan,  Chiapas,  Mexico, 
(2)  Vicinity  of  San  Antonio  Huista,  Depto. 
Huehuetenango,  Guatemala,  (3)  combined 
sample  of  those  specimens  originally  desig- 
nated by  Stuart  (1942)  as  holotype  and  para- 
types  from  the  Languin-Cahabon  road  near 
Finca  Canihor,  Depto.  Alta  Verapaz,  Guate- 
mala, and  specimens  from  the  vicinity  of 
Poptun,  Depto.  El  Peten,  Guatemala,  later 
collected  by  Stuart  and  identified  as  A. 
chaitzami. 


Locality:  La  Libertad,  Chiapas,  Mexico, 
near  Rio  Cuilco  where  it  crosses  Guate- 
malan border.  Collector:  H.  D.  Thomas]. 
Smith  and  Taylor,  U.  S.  Natl.  Mus.  Bull. 
199,  p.  173,  1950. 

Distribution. — Known  from  the  val- 
leys of  the  upper  tributaries  of  the  Rio 
Grijalva  in  Chiapas,  Mexico  and  west- 
central  Guatemala,  from  along  the  Lan- 
guin-Cahabon trail  near  Finca  Canihor, 
Alta  Verapaz,  Guatemala,  and  near 
Poptun,  El  Peten,  Guatemala  (Fig.  25). 

Diaanosis. — Ameiva  chaitzami  can 
be  distinguished  from  other  Middle 
American  congeners  by  the  following 
combination  of  characters:  Small  size 
(maximum  observed  SVL  of  males  85 
mm,  of  females  75  mm);  central  gular 
scales  enlarged,  in  longitudinal  arrange- 
ment; gradual  reduction  in  size  of  scales 
radiating  outward  from  central  gular 
scales;  prefrontal  scales  in  contact  with 


60 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


postnasal  scales;  three  parietal  scales 
(four  if  median  parietal  scale  divided); 
a  transverse  row  of  abruptly  enlarged 
mesoptychial  scales;  usually  eight  trans- 
verse rows  of  ventral  scutes;  relatively 
narrow  middorsal  stripe  (mean  PV  =: 
37.1);  no  narrow,  well-defined  vertebral 
stripe;  usually  a  dark  secondary  stripe 
medial  to  the  dorsolateral  light  stripes; 
dorsolateral  blotches  of  males  fused  to 
the  dorsolateral  light  stripe  such  that 
the  latter  has  a  well-defined  dorsal  bor- 
der and  an  irregular  ventral  border. 

Adult  male  Ameiva  chaitzami  can 
be  distinguished  from  other  Ameiva 
readily  on  the  basis  of  pattern.  Juvenile 
individuals  and  females  are  difficult  to 
differentiate  from  some  undidata,  espe- 
cially those  from  the  coastal  lowlands 
of  eastern  Veracruz,  Tabasco,  Chiapas 
and  extreme  western  Campeche,  Mex- 
ico. However,  there  are  no  signs  of 
intergradation  or  hybridization  of  chait- 
zami with  other  species  of  Ameiva.  The 
only  Ameiva  with  which  chaitzami 
comes  in  contact  is  the  large  form  of 
undulata  having  a  color  pattern  as 
shown  in  figure  21,  and  there  is  no  dif- 
ficulty in  distinguishing  the  two  species. 

Description. — Maximum  observed 
SVL  85  mm  for  males,  75  mm  for  fe- 
males; supralabials  13-16  (14.2);  infra- 
labials  9-13  (10.1);  supraoculars  6-8 
(6.2);  COF  value  2;  COP  value  10-13 
(11.6);  SO-SC  value  2-8  (2.3);  loreals 
2-4  (3.7);  nostril  in  prenasal-postnasal 
suture;  prefrontals  in  contact  with  post- 
nasals; central  gular  scales  enlarged, 
longitudinal  in  arrangement;  GAB  83- 
134  (108.6);  PV  26-51  (37.1);  PV/GAB 
0.27-0.42  (0.34);  GOR  174-243  (203.6) 
for  males,  171-255  (205.1)  for  females; 
GAB/GOR  0.44-0.65  (0.54);  SAT  20-24 
(21.2);  total  preanal  scales  5-9  (6.8); 
terminal  preanal  scales  1-3  (2.0);  one 
row  of  enlarged  prebrachials;  one  or 
two  rows  of  enlarged  postbrachials;  en- 
larged preantebrachials  in  two  rows  or 
two  rows  proximally  and  one  row  dis- 
tally;  prefemoral  scales  enlarged,  four 
(usually)   or  three  rows  proximally  re- 


FiG.  26.  Head  and  preanal  scutellation  of 
Ameiva  chaitzami  (UMMZ  90643,  paratype, 
from  along  the  Cahabon-Langiiin  trail  about 
2  km  N  Finca  Canihor ) :  ( A )  Lateral  view  of 
head,  ( B )  Dorsal  view  of  head,  ( C )  Ventral 
view  of  head,  X  2.  (D)  Preanal  region, 
X  2.7. 

ducing  to  three  (usually)  or  two  rows 
distally;  infratibials  enlarged,  in  two 
rows;  femoral  pores  30-41  (35.0)  for 
males,  26-36  (32.7)  for  females;  sub- 
digital  lamellae  24-33  (28.6);  longitudi- 
nal rows  of  ventral  scutes  27-32  (30.0); 
transverse  rows  of  ventral  scutes  8-10 
(8.1).  For  details  of  scutellation  see 
figure  26. 

Color  and  Pattern  in  Alcohol. — De- 
scriptions of  specimens  in  preservative 
have  been  presented  by  Stuart  ( 1942 ) 
and  Smith  and  Laufe  ( 1946 ) .  Juvenile 
males  possess  a  broad  middorsal  field 
bounded  laterally  by  narrow,  white  dor- 
solateral stripes  which  originate  just  be- 
hind the  eye  and  extend  well  onto  the 
tail.  Between  the  blue-gray  or  gray- 
brown  middorsal  pigment  and  the  dor- 
solateral light  stripes  there  are  often 
secondary  stripes,  velvet  black  in  color, 
which  extend  from  the  occiput  to  the 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


61 


base  of  the  tail.  The  medial  border  of 
the  secondary  stripe  is  irregular.  The 
niiddorsal  field  may  be  flecked  witli 
black  or  be  nearly  uniform  in  colora- 
tion. A  white  ventrolateral  light  stripe, 
which  may  be  complete  or  interrupted, 
extends  from  the  posterodorsal  corner 
of  the  ear  to  the  groin,  beginning  again 
lieliind  the  leg  and  terminating  on  the 
tail.  A  white  postfemoral  stripe  is  con- 
tinuous with  the  ventrolateral  light 
stripe  behind  the  leg.  The  black  dorso- 
lateral dark  field  is  prominent  and  con- 
tains faint,  light-colored  blotches  or 
bars.  The  ventrolateral  field  contains 
wliitc  mottling  or  bars  on  a  black  back- 
ground. The  same  pattern  is  evident  on 
dorsal  surfaces  of  the  limbs,  but  the 
mottling  is  more  pronounced.  Ventral 
surfaces  are  cream  or  white  in  color. 

Juvenile  females  resemble  juvenile 
males  except  that  there  are  no  light- 
colored  blotches  or  bars  in  the  dorso- 
lateral dark  field  and  the  ventrolateral 
and  limb  mottling  is  less  well  de- 
veloped. 

Through    ontogeny     (Fig.    27)     the 
light  ventrolateral  and  dorsal  limb  mot- 
tling   or   barring    becomes    much    more 
pronounced  and  the  ventrolateral  light 
stripe  becomes  interrupted  such  that  it 
merges  with  the  ventrolateral  mottling. 
The  dorsolateral  blotches  become  white 
or  blue-white  in  color  and  fuse  with  the 
dorsolateral  light  stripe  in  males,  giving 
that  stripe  an  irregular  ventral  border. 
Adult    females    resemble    juveniles     of 
both    sexes    in    color    and    pattern,    ap- 
parent ontogenetic  changes  being  slight. 
Color   and   Pattern   in   Life. — There 
are    no    descriptions    of    the    coloration 
of    living    Ameiva    chaitzami,    although 
Stuart's  (1942)   description  of  the  holo- 
type  may  have  been  written  before  ex- 
tensive fading  had  occurred  in  preserva- 
tive.   He  stated  that  the  middorsal  field 
and  top  of  the  head  are  olive-brown  and 
that  the  legs  and  arms  are  olive-brown 
above     with     black     and     bluish-white 
mottling.    The  anterior  surfaces  of  the 
thighs   are  said  to  be  black  with  blue 


spots,  the  venter  l:)hiisli  and  the  head 
blue  mottled  with  black.  There  is  no 
mention  of  yellow  or  red  pigment  in  tlie 
gular  region  of  this  species,  the  only 
Middle  American  Ameiva  with  which  I 
have  had  no  field  experience. 

Geographic  Variation. — The  follow- 
ing characters  show  no  significant  dif- 
ferences among  sample  means:  Femoral 
pores  of  males,  number  of  supralabials, 
number  of  infralabials,  number  of  ter- 
minal preanals,  number  of  transverse 
rows  of  ventral  scutes,  SAT  and  COF, 
COP  and  SO-SC  values. 

The  three  samples  of  Ameiva  chait- 
zami (Fig.  25 )  were  collected  in  eastern 
Chiapas,    Mexico    (Sample    1),   western 
Guatemala    (2)    and  central  to   eastern 
Guatemala   (3).    Samples   1  and  2  are 
much  closer  geographically  than  either 
is  to  sample  3  and  sample  1  lies  slightly 
northwest  of  sample  2.    This  geographic 
arrangement  is   reflected  in  the  results 
of    STP    analysis    of    characters    which 
show      significant      differences      among 
means.    Five  characters  show  samples  1 
and  2  as  constituting  one  subset,  sample 
3  a  second  subset  significantly  different 
from  the  first.    These  five  characters  are 
numbers  of  loreals  and  subdigital  lamel- 
lae,  GAB /COR,   numbers   of  longitudi- 
nal rows  of  ventrals  and  total  preanal 
scales.     For   two    characters,    GAB    and 
PV/GAB,  samples  2  and  3  form  a  sub- 
set  distinct   from   that   containing   only 
sample   1.    For  both  PV  and  GOR  of 
females,  all  three  sample  means  fall  into 
separate,    significantly    distinct    subsets. 
This  is  true  of  samples  1  and  2  for  num- 
ber   of    femoral    pores    of   females    and 
GOR  of  males;  sample  3  is  too  small  for 
inclusion  in  STP  analysis.    Only  for  one 
character,    number    of    supraoculars,    is 
there  overlap  of  sample  means:    there  is 
a  gradient  from  sample  3  through  sam- 
ple 2  to  sample  1,  samples  3  and  2  con- 
stituting one  subset  and  samples  2  and 
1  another. 

Because  of  the  inadequate  number 
of  samples  and  the  geographic  localities 
from   which   the   samples   were   drawn, 


62 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


63 


detection  of  clinal  trends  must  await 
accumulation  of  additional  large  sam- 
ples as  the  range  of  the  species  becomes 
more  fully  known. 

There  seems  to  be  no  notable  geo- 
graphic variation  in  color,  pattern  or 
arrangement  of  gular  scales  among  the 
three  available  samples.  A  comparison 
of  color  patterns  of  equal-sized  (68 
mm)  males  from  samples  1  and  3  may 
be  seen  in  Figure  27  (B  and  G,  re- 
spectively ) . 

Ecological  Notes.— Stuart  ( 1942; 
personal  communication)  provided  the 
only  insight  into  the  ecology  of  Ameiva 
chaitzami.  He  is  of  the  opinion  that  the 
species  is  restricted  to  pine  savannas  on 
the  lowlands  of  central  Guatemala 
where  its  population  density  may  be 
kept  at  a  low  level  by  regular  burning- 
off  of  ground  cover.  The  species  is 
found  with  Ameivo  tmdulata,  but  never 
with  Cnemidophorus.  Other  Middle 
American  species  of  Ameiva  are  sym- 
patric  with  species  of  Cnemidophorus 
over  at  least  part  of  their  range.  Stuart 
(1942)    mentioned  a  chaitzami  with  a 


SVL  of  66  mm  having  "well  formed 
eggs." 

Smith  and  Laufe  (1946)  stated  that 
these  lizards  ".  .  .  probably  occur  in  all 
the  dry,  hot  valleys  of  the  upper  tribu- 
taries of  the  Rio  Grijalva  in  the  interior 
of  Chiapas  and  of  western  central 
Guatemala." 

Remarks. — Less  is  known  about  the 
biology  of  Ameiva  chaitzami  than  of 
any  other  Middle  American  Ameiva. 
The  overall  range  of  the  species  has  yet 
to  be  defined. 

The  divided  interparietal  (median 
parietal),  which  was  used  by  Stuart 
( 1942 )  to  diagnose  Ameiva  chaitzami, 
is  a  variable  character  found  through- 
out the  genus  in  Middle  America  with 
varying  frequency  ( Echternacht,  1970). 
Likewise,  I  have  found  the  lateral  en- 
largement of  the  gular  scales  cited  by 
Smith  and  Laufe  ( 1946 )  to  be  an  unre- 
hable  character.  Color  pattern  and  size 
remain  the  most  useful  characters  for 
distinguishing  this  probable  sibling  spe- 
cies from  its  closest  relative,  undulata. 


DISCUSSION 


INTRASPECIFIC  RELATIONSHIPS 

In  my  opinion  the  Ameiva  of  Mid- 
dle America  belong  to  three  natural 
groups  the  relationships  of  which  will 
remain  obscure  until  their  affinities  with 
South  American  representatives  of  the 
genus  are  known.  The  three  groups 
contain  one,  two  and  three  species  ( Fig. 
28). 


Fig.  27.  Ontogenetic  change  in  pattern  of 
Ameiva  chaitzami.  A-D  and  F,  UMMZ  94905, 
E,  UMMZ  94901,  all  from  the  vicinity  of 
Comitan,  Chiapas,  Mexico.  Field  Tag  Num- 
bers and  SVL  are:  Males,  (A)  473,  45  mm, 
(B)  482,68  mm,  (C)  461,  73  mm.  Females, 
(D)  483,  55  mm,  (E)  365,  68  mm,  (F)  501, 
75  mm.  (G)  For  comparison,  UMMZ  90640, 
a  male  paratype  68  mm  SVL  from  along  the 
Cahabon-Languin  trail  about  2  km  N  Finca 
Canihor,  Alta  Verapaz,  Guatemala. 


Ameiva  ameiva,  the  only  species  in 
Group  I,  can  be  distinguished  from  the 
others  on  the  basis  of  a  number  of 
scale  and  pattern  characteristics.  Chief 
among  these  are  the  presence  of  five 
parietal  scales,  10-12  enlarged  ventral 
scutes,  a  large  number  of  scales  around 
the  tail,  high  GAB  and  GOR  counts  and 
a  pattern  which  emphasizes  spots  in- 
stead of  stripes  in  adults.  Additionally, 
ameiva  is  the  only  species  included  in 
the  present  study  having  the  bulk  of  its 
range  in  South  America.  The  Pana- 
manian portion  of  that  range  is  but  a 
minor  and  disjunct  part  of  the  whole. 
Group  I  is  farther  removed  phylogenet- 
ically  from  Groups  II  and  III  than  these 
are  from  each  other. 

Group  II  includes  Ameiva  festiva 
and  leptophrijs.  Both  species  are  essen- 
tially   lower    Central   American   in    dis- 


64 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


Fig.  28.  Relationships  of  the  Middle  American 
species  of  Ameiva.  Group  I:  A,  A.  ameiva. 
Group  II:  F,  A.  f estiva;  L,  A.  leptophrys. 
Group  III:  Q,  A.  quadrilineata;  C,  A.  chait- 
zami;  U,  A.  undulata.  Lengths  of  the  lines 
and  positions  of  branches  do  not  reflect  a 
time  scale. 

tribution.  Ameiva  festiva  occurs  in  Co- 
lombia, and  leptophrys  is  expected 
there.  The  group  is  characterized  by 
the  relatively  high  number  of  femoral 
pores  and  a  tendency  for  the  enlarged, 
posterior  scales  of  the  head  to  become 
disrupted  (Fig.  7).  Disruption  of  head 
scales  is  more  noticeable  in  leptophrys 
than  in  festiva  but  is  a  more  commonly 
encountered  feature  of  these  species 
than  of  other  Middle  American  species 
of  Ameiva. 

Group  III  consists  of  Ameiva  quad- 
rilineata, chaitzami  and  undulata.  The 
group  is  entirely  Middle  American  in 
distribution,  and  is  characterized  by 
similarities  in  pattern  and  scale  charac- 
ters. The  pattern  emphasizes  stripes, 
rather  than  spots,  and  features  a  broad, 
middorsal  field  without  a  vertebral 
stripe.  Prefrontal  and  postnasal  scales 
are  in  contact.  Except  for  an  occasional 
longitudinal  division  of  the  interparietal 
scale,  there  is  little  disruption  of  pos- 
terior scales  of  the  head.  Ameiva  chait- 
zami is  very  closely  related  to  undulata, 
from  which  it  is  distinguishable  pri- 
marily on  the  basis  of  the  color  pattern 
of  males.  However,  the  two  species  are 
sympatric  in  central  Guatemala.    Clari- 


fication of  the  relationships  of  chait- 
zami, its  range  and  biology  remain 
problems  of  major  importance. 

Elucidation  of  the  relationships  of 
the  Middle  American  species  of  Ameiva 
with  those  of  South  America  may  show 
that  Groups  II  and  III  are  much  more 
closely  related  than  I  have  indicated. 
These  may  constitute  a  single  group.  I 
should  emphasize  that  these  three 
groups  of  Middle  American  Ameiva  are 
not  the  equivalents  of  the  species 
groups  of  Cnemidophorus  established 
by  Burt  ( 1931 ) .  Nevertheless,  it  seems 
apparent  that  my  Group  I  would  belong 
to  a  difterent  species  group  (sensu 
Burt)  than  the  other  species  of  Middle 
American  Ameiva,  regardless  of  whether 
these  other  species  are  found  to  belong 
to  one  or  several  species  groups.  The 
relationships  indicated  without  substan- 
tiating comment  by  Barbour  and  Noble 
(1915)  are  suspect.  The  taxonomic  im- 
portance of  extensive  division  of  head 
scales  in  Ameiva  bridgesi  Cope,  orcesi 
Peters  and  septemlineata  Dumeril  and 
Dumeril  (Peters,  1964)  and  the  rela- 
tionships of  these  species  to  leptophrys 
and  festiva  require  further  study.  The 
relationships  of  niceforoi  also  need  clari- 
fication ( Echternacht,  1970). 

INTERSPECIFIC  PATTERNS  IN 
GEOGRAPHIC  VARIATION 

Assessing  interspecific  trends  in  vari- 
ation is  considerably  more  difficult  than 
determining  intraspecific  trends.  This  is 
so  for  two  reasons:  1)  The  narrow  lin- 
earity of  lowland  Middle  America  vir- 
tually dictates  that  clines  will  run  paral- 
lel to  the  coastlines  along  a  northwest 
to  southeast  axis  and  2)  samples  of  two 
or  more  species  from  the  same  locality 
and  of  sufficient  size  for  statistical  anal- 
ysis are  rarely  available.  Samples  from 
only  ten  localities  where  sympatric  oc- 
currence of  two  or  more  species  was 
found  were  large  enough  for  analysis 
(Table  3),  and  only  three  of  the  six 
Middle  American  species  of  Ameiva 
were  present  in  more  than  one  sample. 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


65 


TABLE  3. — Localities  from  which  samples  of  two  or  more  species  were  available  for  statistical 

analysis. 


Locality 


Species 


Costa  Rica:    Limon:    Limon,  Los  Diamantes  and  vicinity 

Costa  Rica:    Limon:    Tortuguero 

Costa  Rica:     Puntarenas:     Golfito  Region 

Costa  Rica:    Puntarenas:    Osa  Peninsula 

Guatemala:    El  Peten:    Piedras  Negras 

Honduras:    Northern  Coastal  Localities 

Panama:    Bocas  del  Toro:    Almirante  and  vicinity 

Panama:    Canal  Zone 

Panama:    Isla  Escudo  de  Veraguas 

Panama:    Eastern  Darien 


festiva. 

quadrilineata 

festiva. 

quadrilineata 

festiva. 

quadrilineata,    leptophrys 

festiva. 

(juadiilineata,    leptophrys 

festiva. 

tmdidata 

festiva. 

itudulata 

festiva. 

quadrilineata 

ameiva 

festiva,   leptophrys 

festiva. 

quadrilineata 

festiva. 

leptophrys 

Ameiva  undtdata  was  present  with  fes- 
tiva in  but  two  samples,  and  festiva  was 
found  with  quadrdineata  in  two.  Al- 
though no  rigorous  statistical  analysis 
was  attempted,  some  parallel  trends  are 
evident.  For  seven  characters  (infra- 
labials,  supraoculars,  COP,  GAB,  GOR, 
subdigital  lamellae  and  scales  around 
the  tail)  an  increase  or  decrease  in 
sample  mean  for  festiva  was  paralleled 
by  a  like  shift  in  mean  for  the  same 
character  for  tindulata  when  the  two 
localities  of  sympatry  (Piedras  Negras, 
El  Peten,  Guatemala,  and  northern  Hon- 
duras) were  compared.  Similar  trends 
were  noted  for  festiva  and  quadrdineata 
for  ten  characters  (GAB,  GOR,  GAB/ 
GOR,  longitudinal  rows  of  ventral 
scutes,  transverse  rows  of  ventral  scutes, 
femoral  pores,  subdigital  lamellae,  total 
preanal  scales,  temiinal  preanal  scales, 
and  scales  around  the  tail)  when  step- 
wise pair  comparisons  of  samples  were 
made  between  localities  beginning  with 
Tortuguero  in  Costa  Rica  and  running 
through  the  Canal  Zone  to  Darien, 
Panama,  and  thence  to  the  Golfito  and 
Osa  Peninsula  regions  of  Costa  Rica. 
Perfect  agreement  was  noted  only  for 
GOR,  but  all  other  characters  men- 
tioned showed  parallel  mean-shifts  for 
at  least  two-thirds  of  the  possible  pair 
comparisons.  Similar  parallel  changes 
were  noted  for  festiva  and  leptophrys 
( supralabials,  infralabials,  COF,  COP, 
transverse  rows   of  ventrals  and  scales 


around  the  tail  with  COP  and  COF 
showing  perfect  agreement)  and  lepto- 
phrys and  quadrilineata  (infralabials 
and  supraoculars).  Such  parallel  shifts 
in  sample  means  would  tend  to  indicate 
that  these  characters  are  responding  in 
similar  ways  to  environmental  differ- 
ences between  sample  localities.  Con- 
siderable additional  study  will  be  neces- 
sary before  any  light  will  be  shed  on  the 
question  of  which  environmental  factors 
are  most  important  in  determining  the 
direction  of  shift  of  sample  mean  for  a 
character  from  one  sample  locality  to 
another.  Because  only  increase  or  de- 
crease in  the  value  of  the  mean  were 
noted  from  sample  to  sample,  and  not 
the  magnitude  of  the  increase  or  de- 
crease, it  is  probable  that  some  shifts 
are  only  the  result  of  small  sample  size 
and  that  additional  data  would  show 
that  there  has  been,  in  fact,  no  shift 
at  all. 

An  important,  although  likewise  un- 
explained, trend  is  that  of  insular  popu- 
lations of  Ameiva  to  have  higher  mean 
GAB  and/ or  GOR  counts  than  main- 
land populations  (Table  4).  Samples 
are  available  from  Isla  Mujeres,  Quin- 
tana  Roo,  Mexico  (undidata),  Isla  del 
Maiz  Grande,  Zelaya,  Nicaragua  (undu- 
lata),  Isla  Escudo  de  Veraguas,  Panama 
(festiva,  quadrilineata)  and  Isla  San 
Jose,  Archipelago  de  las  Perlas,  Panama 
(ameiva).  Samples  from  the  Osa  Penin- 
sula, Puntarenas,  Costa  Rica  of  festiva, 


66 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


TABLE  4. — Relati\'e  positions  of  island  samples    with    respect    to    mean    Granules    Around    the 
Body  (GAB)  and  Granules  Occiput  to  Rump   (GOR)  when  sample  means  are  ordered  from  largest 

to   smallest. 


Total  Number 

Relative 

Position 

Species 

of  Samples 

GAE 

; 

GOR 

ameiva 

7 

1 

2*' "          1" 

festiva:    Isla  Escudo  de  Veraguas 

20 

1" 

1" 

V             1" 

fcstica:    Osa  Peninsula 

20 

4" 

4" 

15"           16" 

k'pto))hnjs 

4 

2 

2 

quadrilineata:    Isla  Escudo  de  Veraguas 

10 

7 

5 

quadrilineata:    Osa  Peninsula 

10 

1 

1 

imdulata:    Isla  Mujeres 

45 

2 

10 

uudulata:    Isla  del  Maiz  Grande 

45 

1 

2 

"  Males  where  sexual  dimorphism  is  evident. 

"  Females  where  sexual  dimorphism  is  evident. 

''  Tied  with  sample  3  (See  Appendix  B)  for  this  position. 


leptophrijs,  and  quadrilineata  also  have 
relatively  high  dorsal  granule  counts; 
the  peninsula  may  be  effectively  an  is- 
land as  far  as  Ameiva  are  concerned 
inasmuch  as  it  is  separated  from  the 
mainland  by  a  swampy  area.  It  will  be 
obvious  from  an  examination  of  table  4 
that  not  all  insular  samples  have  high 
doi-sal  granule  counts,  but  the  tendency 
is  evident  in  a  remarkable  number  of 
instances.  There  is  no  clear  relationship 
between  high  GAB  and  GOR  counts  on 
the  one  hand  and  the  ecology  of  the 
islands  upon  which  the  lizards  are  living 
on  the  other.  It  would  seem  that  Amei- 
va adapted  to  life  on  dry  islands  would 
have  fewer,  but  larger,  dorsal  granules 
than  Ameiva  of  the  same  size  living  on 
wet  islands  as  a  selective  strategy  for 
prevention  of  excess  water  loss.  Fewer 
and  larger  granules  would  mean  less  ex- 
posed skin  between  the  granules 
through  which  water  might  be  lost  than 
more  and  fewer  scales  on  animals  of 
equal  size.  Such  an  hypothesis  is  con- 
traindicated  by  the  observation  that  the 
islands  from  which  samples  of  Ameiva 
are  available  range  from  very  dry  (Isla 
Mujeres)  to  very  wet  (Isla  Escudo  de 
Veraguas  and  the  Osa  Peninsula).  It 
would  be  instructive  to  have  samples  of 
one  species  of  Ameiva  from  a  number 
of  localities  for  which  weather  records 
are    available    and    ranging    from    rela- 


tively dry  to  relatively  wet.  Such  sam- 
ples are  not  presently  available  and, 
until  they  are,  this  problem  is  likely  to 
remain  unsolved. 

Sample  mean  snout-vent  length 
(SVL)  data  are  not  presented  in  this 
report  because  collections  are  usually 
strongly  biased  in  favor  of  large,  bright- 
ly colored  males.  The  relatively  drab 
juvenile  and  adult  female  Ameiva  are 
often  overlooked  by  collectors  who  ob- 
tain Ameiva  incidental  to  those  genera 
they  themselves  are  working  on.  A  sea- 
sonal bias  also  acts  to  reduce  the  num- 
bers of  hatchling  Ameiva  in  collections 
in  that,  in  areas  having  pronounced  wet 
and  dry  seasons,  reproduction  may  be 
temporally  restricted  with  hatchlings 
present  only  at  certain  times  of  the  year. 
Collections  made  at  other  times  do  not 
include  hatchlings.  All  of  the  insular 
samples  of  Ameiva  available  to  me  were 
obtained  in  June,  July,  or  August,  and 
all  are  the  result  of  but  one  or  two  field 
trips.  With  the  exception  of  my  own 
collection  from  Isla  del  Maiz  Grande, 
all  of  these  collections  were  made  by 
individuals  primarily  interested  in  other 
genera.  For  these  reasons,  I  have  little 
faith  in  the  mean  SVL  of  samples  used 
in  this  study  as  an  estimate  of  the  true 
population  means.  Nevertheless,  it  is 
apparent  that  the  insular  samples  have 
generally  higher  mean  SVL  than  main- 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


67 


TABLE    5. — Correlation   between   snout-vent    length    (SVL)     and    granules    around    the    body 
(GAB)  or  granules  from  occiput  to  nunp  (GOR)  for  five  species  of  Middle  American  Ameiva.    Cor- 
relation coefficient  followed  by  sample  size  in  parentheses. 


ametva 


festiva 


leptophrys      quadrilineata  undulata 


SVL-GAB  Males 
SVL-GAB  Females 
SVL-GOR  Males 
SVL-GOR  Females 


0.1101(112)  0.0897(243)      0.1345  ( 101 ) -0.0289  ( 180)  0.1606(409) 

-0.0403(133)  0.1531  (233) -0.1040(98)    -0.0950(114)  0.2445(561) 

0.0737(114)  0.1857(268)  0.2262(188)  0.1629(507) 

-0.0888(138)  0.0682(264)  -0.2283(126)  0.1550(448) 


land  samples,  making  it  necessary  to  ask 
the  question  whether  GAB  and  GOR 
counts  are  in  any  way  correlated  with 
SVL.  Correlation  analysis  was  carried 
out  on  a  species-by-species  basis  (Table 
5).  There  is  no  indication  that  either  of 
the  tsvo  dorsal  granule  counts  are  highly 
correlated  with  SVL  and  the  high  mean 
GAB  and  GOR  values  of  island  samples 
is  not  an  artifact  resulting  from  biased 
estimates  of  sample  mean  SVL. 

HISTORY  OF  AMEIVA  IN 
MIDDLE  AMERICA 

There  is  no  fossil  record  of  macro- 
teiids  in  Middle  America.  Therefore, 
any  discussion  of  the  history  of  Ameiva 
in  that  region  must  lean  heavily  on 
what  is  known  of  present  distribution 
patterns  and  on  geological  and  faunal 
summaries  relating  to  Middle  America 
and  northern  South  America.  Geological 
theories  concerning  Middle  America 
have  undergone  an  evolution  of  their 
own  over  the  past  50  years;  geological 
data  were  summarized  by  Lloyd  ( 1963 ) , 
Vinson  and  Brineman  (1963),  and  Mal- 
donado-Koerdell  ( 1964 ) .  Duellman 
(1966),  Savage  (1960,  1966)  and  Stuart 
(1935,  1951,  1954a,  1954b,  1957,  1964, 
1966)  provided  thoroughly  documented 
discussions  of  the  ecology,  origins,  and 
history  of  the  herpetofauna  of  Middle 
America,  whereas  Haffer  ( 1967,  1970 ) 
presented  an  important  assessment  of 
the  geologic-climatic  history  of  the  criti- 
cal Gulf  of  Uraba-northwestern  Colom- 
bia region.  Although  the  following  dis- 
cussion is  based  in  large  part  on  these 
works,   it   is   a   highly   speculative   and 


personal  view  and  should  be  read  with 
that  in  mind. 

Dunn  ( 1931 )  considered  the  family 
Teiidae  to  be  a  part  of  his  South  Amer- 
ican element,  having  its  origins  in  South 
America  with  some  genera  invading 
Middle  America  and  one,  Cnemido- 
phorus,  reaching  North  America.  The 
South  American  element  itself  was  de- 
rived from  a  generalized  tropical  Amer- 
ican herpetofauna  which  ranged  over 
much  of  North  and  South  America  dur- 
ing the  early  Tertiary  (  Savage,  1960 ) . 
Subsequent  authors  have  followed  this 
arrangement  (Duellman,  1965b;  Savage, 
1960,  1966;  Schmidt,  1943;  Stuart,  1957, 
1964),  although  Savage  (1966)  sug- 
gested that  the  ancestral  stock  of  Cnemi- 
dophorus  was  isolated  north  of  the 
Isthmus  of  Panama  and  that  of  Ameiva 
south  of  the  Isthmus  when  the  Pana- 
manian portal  was  open  throughout 
most  of  the  Tertiary.  Northward  move- 
ments of  the  South  American  element 
would  have  been  possible  after  the 
closure  of  the  portal  in  the  late  Pliocene 
or  early  Pleistocene  (Lloyd,  1963;  Mal- 
donado-Koerdell,  1964).  As  outlined  by 
Stuart  (1957)  and  Savage  (1966,  Fig. 
23),  Ameiva,  with  the  exception  of  A. 
ameiva,  probably  followed  humid,  low- 
land routes  from  South  America  into 
Middle  America.  Movement  through 
the  Pacific  lowlands  progressed  only  as 
far  as  the  Golfito-Palmar-San  Isidro  del 
General  region  of  Costa  Rica.  An  At- 
lantic route  was  followed  northward  in 
Mexico,  dispersal  to  the  Pacific  lowlands 
occurring  at  several  points  north  of 
what    is    now    Lake    Nicaragua.     Some 


68 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


southward  movement  along  the  Pacific 
lowlands  of  Nicaragua  and  Costa  Rica 
at  a  later  time  is  indicated  by  the  pres- 
ent distribution  of  A.  tinchilota  ( Fig. 
17).  These  routes  may  have  been 
broader  than  at  present  because  of  low- 
ered sea  levels,  especially  in  areas 
where  the  continental  shelf  is  extensive 
(Bennett,  1968). 

Of  all  of  the  Ameiva  found  in  Mid- 
dle America,  A.  ameiva  has  made  the 
smallest  inroads  there.  Heatwole  (1966) 
placed  its  arrival  in  the  Upper  Pliocene 
at  the  earliest.  As  an  alternative  hy- 
pothesis, Bennett  (1968)  suggested 
that  the  species  dispersed  into  Panama 
during  the  period  of  human  occupation 
of  Darien.  The  A.  ameiva  of  Panama 
resemble  those  of  western  Venezuela; 
Bennett  suggested  than  an  open  corri- 
dor between  the  two  areas  for  the 
length  of  time  suggested  by  Heatwole 
( over  one  million  years )  v^^ould  have  re- 
sulted in  formation  of  clines.  Subse- 
quent closure  of  the  corridor  would 
have  isolated  opposite  ends  of  such 
clines,  resulting  in  morphologically  di- 
vergent populations.  Other  than  a  study 
emphasizing  color  pattern  ( Miiller, 
1929),  no  detailed  study  of  the  geo- 
graphic variation  of  South  American  A. 
ameiva  has  been  reported.  Donoso- 
Barros  ( 1968 )  characterized  the  west- 
ern population  of  the  species  in  Vene- 
zuela as  having  a  light-colored  vertebral 
stripe.  Such  a  stripe  is  evident  in  those 
A.  ameiva  from  eastern  Panama,  but 
lacking  in  those  from  the  western  part 
of  that  country  (Fig.  4).  With  the 
examination  of  additional  samples,  rem- 
nants of  a  cline  as  postulated  by  Ben- 
nett may  be  found  for  this  and  other 
characters  (but  see  Remarks,  p.  19).  If 
remnants  of  a  cline  are  found,  it  would 
indicate  that  entry  of  the  species  into 
Middle  America  occurred  at  a  time  in- 
termediate to  the  extremes  suggested  by 
Heatwole  and  Bennett.  Recent  evi- 
dence of  alternating  dry  interglacial  and 
humid  glacial  periods  during  the  Pleis- 
tocene   so    convincingly    presented    by 


Haffer  (1967,  1970)  would  allow  for 
several  intervals  during  which  suitable 
conditions  prevailed  for  the  northward 
movements  of  A.  ameiva.  Haffer  (1970) 
suggested  that  the  humid  forests  of  gla- 
cial times  were  forced  southward  dur- 
ing interglacials  and  during  post-Pleis- 
tocene dry  periods  and  that  central  and 
eastern  Panama,  as  well  as  the  Gulf  of 
Uraba-lower  Rio  Atrato  region  of  Co- 
lombia, were  not  heavily  forested.  Thus, 
it  would  have  been  possible  for  A. 
ameiva  to  have  moved  into  Panama 
prior  to  the  period  of  human  occupancy 
of  the  Darien  and  yet  not  long  enough 
for  establishment  of  clines.  Indeed,  if 
Haffer's  scheme  is  followed,  the  corridor 
between  central  Panama  and  Colombia 
would  have  been  interrupted  and  re- 
established several  times,  mediating 
against  formation  of  clinal  variation.  In 
any  case,  it  is  certain  that  the  Pana- 
manian A.  ameiva  originated  from  a 
population  of  South  American  Ameiva 
similar  to  the  present  population  of  the 
species  in  western  Venezuela.  Dispersal 
into  Middle  America  was  contained  by 
the  availability  of  non-forested  habitats. 
The  species  probably  reached  the  is- 
lands of  the  Archipelago  de  las  Perlas 
at  a  time  when  a  land  connection  existed 
between  these  islands  and  the  mainland 
during  a  period  of  lowered  sea  level 
(Bennett,  1968). 

With  the  exception  of  Ameiva  amei- 
va already  discussed,  none  of  the  species 
of  Ameiva  presently  inhabiting  Middle 
America  would  have  required  a  non- 
forested  corridor  for  northward  disper- 
sal. Although  the  species  show  a  graded 
dependence  upon  forested  habitat,  all 
are  found  in  or  at  the  edge  of  forested 
areas.  Ameiva  f estiva  and  leptoplinjs 
may,  upon  completion  of  studies  now  in 
progress,  be  found  to  be  closely  allied  to 
the  A.  septemlineata — hridgesi — orcesi 
assemblage  of  northwestern  South 
America.  Ameiva  leptophrys,  with  its 
present  range  limited  to  eastern  Pana- 
ma and  the  Pacific  slopes  of  western 
Panama    and    southeastern    Costa    Rica 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


69 


( Fig.  5 )  may  have  evolved  ( along  with 
several  other  species  of  reptiles  and  am- 
phibians) on  insular  isolates  during  the 
Middle   Tertiary    (Savage,    1966).    This 
seems  possible  in  view  of  both  the  pres- 
ent distribution  of  the  species   and  its 
broad  sympatry  with  festiva.    The   ap- 
parent absence  of  populations  of  Jepto- 
phrys  on  islands  at  present  is  a  minor 
set-back  to  a  theory  of  insular  origin  in 
that   suitable   habitat   may   have   disap- 
peared from  the  islands  in  the  relatively 
recent   past,    or   that   further   collecting 
will  reveal  the  presence  of  the  species. 
Ameiva  festiva  occurs  in  both  Mid- 
dle and  South  America  ( Fig.  9 ) ,  as  does 
A.  ameiva,  but  festiva  is   continuously 
distributed   over   its    continental    range. 
Populations  of  festiva  on  the  west  and 
east  ( or  north )  slopes  of  the  Andes  have 
diverged  markedly  in  color  pattern  ( Fig. 
12 ) .   On  this  basis,  the  population  found 
in  the  Choco  of  Colombia  seems  to  have 
diverged   relatively   early  from   the   an- 
cestral stock.   It  may  possibly  have  been 
isolated  in  the  Choco  region  during  a 
glacial    period    during    the    Pleistocene. 
Haffer   ( 1967 )    documented  similar  iso- 
lation  in   birds   and   fomially   proposed 
the  name  "Choco-Refuge"  for  the  area. 
I  have  seen  no  specimens  intermediate 
in  pattern  between  those  of  the  Choco 
(Pattern  Type  C;   see  page  31   for  de- 
scriptions of  pattern  types)    and  speci- 
mens from  populations  to  the  north  and 
east.    Because  of  similarity  in  color  pat- 
tern of  festiva  from  eastern  Colombia 
and  those  of  the  San  Bias  and  Golfito- 
Puerto  Armuelles  regions  of  lower  Mid- 
dle  America,   it   seems   likely   that   the 
latter    are    derived    from    the    former, 
rather  than  from  the  Chocoan  popula- 
tion.    The    species    moved    northward, 
through   lowland   forests   along  the  At- 
lantic slopes.    The  evolution  of  Pattern 
Type  A  probably  occurred  in  situ  as  a 
result   of   environmental  selection  pres- 
sures.   The  present  distribution  of  fes- 
tiva  in    Middle   America   is    limited    to 
lowland  forests.    Expansion  of  its  range 
in  northern  parts  of  the  region  has  been 


prevented  by  lack  of  suitable  liabitat 
and  possibly  by  competition  from  Amei- 
va undtdata  and  Cnemidophorus.  Such 
expansion  in  South  America  has  prob- 
ably been  limited  by  the  lack  of  suit- 
able habitat  in  Venezuela  and  eastern 
Colombia  and  possibly  by  competition 
from  Ameiva  hridgesi  in  western  Co- 
lombia. 

The  early  history  of  the  Ameiva  un- 
didata-q  uadrdineata-chaitzami  complex 
in  Middle  America  may  have  been  simi- 
lar to  that  of  A.  festiva.  Having  origi- 
nated from  an  unknown  South  Ameri- 
can ancestral  stock,  the  forerunners  of 
the  extant  species  moved  into  Middle 
America  and  along  both  Atlantic  and 
Pacific  routes.  Extensive  northward 
movement  along  the  Pacific  lowlands 
was  probably  prohibited  by  ecological 
barriers,  possil^ly  the  climatic  filter  bar- 
rier illustrated  by  Savage  ( 1966,  Fig. 
23).  Northward  migration  along  the 
Atlantic  slopes  progressed  to  Nuclear 
Central  America  and  thence,  along  two 
routes,  on  into  what  is  now  Mexico. 
Presumably,  ecological  divergence  in 
Costa  Rica  and  adjacent  Panama,  asso- 
ciated with  exploitation  of  beach  and 
open-ground  habitats,  resulted  in  the 
southern  segment  of  the  basal  stock  be- 
coming A.  quadrdineata.  The  range  of 
this  species  is  now  disjunct  (Fig.  13), 
but  in  the  past  it  probably  extended 
along  the  Atlantic  and  Pacific  slopes  of 
the  Western  Sierra  ( terminology  of  Ben- 
nett, 1968)  in  Panama  and  was  joined 
in  the  lowlands  of  the  isthmus.  This 
contention  is  supported  by  the  simi- 
larity of  samples  from  opposite  slopes 
in  the  eastern  part  of  the  ]3resent  range 
of  the  species,  a  similarity  not  found  be- 
tween samples  from  opposite  slopes 
farther  west.  Populations  of  quadrdi- 
neata on  Isla  del  Cano,  Costa  Rica,  and 
Isla  de  Cebaco  and  Isla  Escudo  de 
Veraguas,  Panama,  apparently  owe  their 
presence  to  jiast  land  connections  with 
the  mainland  during  the  Pleistocene 
(Bennett,  1968). 

Ameiva   widulata,   which   continued 


70 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


to  inhabit  more  forested  areas  than 
quadrilineata,  migrated  northward  to 
fill  its  present  range  (Fig.  17).  Smith 
and  Laufe  (1946),  having  to  rely  on 
Schnchert  ( 1935 )  for  geological  back- 
ground, discussed  the  subsequent  his- 
tory of  the  species.  They  postulated  a 
broad  distribution  of  tinchilata  in  Mid- 
dle America  during  the  Miocene,  with 
four  secondary  centers  of  dispersal,  two 
on  either  side  of  the  Isthmus  of  Tehuan- 
tepec  ( Smith  and  Laufe,  1946,  Fig.  4A ) . 
Further  differentiation  was  promoted  by 
the  opening  of  the  Tehuantepec  portal 
during  the  Upper  Miocene  and  Lower 
Pliocene  and  by  subsequent  migration, 
isolation  or  ecological  divergence  of 
populations.  If  we  accept  the  South 
American  origin  of  the  family  Teiidae 
(Dunn,  1931),  and  the  timing  of  the 
reconnection  of  Middle  and  South 
America  at  the  Isthmus  of  Panama  as 
Early  Pliocene  (Lloyd,  1963;  Maldo- 
nado-Koerdell,  1964),  we  must  reject 
the  hypothesis  of  Smith  and  Laufe  and 
can  eliminate  from  consideration  the 
presence  or  absence  of  an  open  portal  at 
the  Isthmus  of  Tehuantepec.  It  seems 
probable  that  one  line  of  undulata  stock 
moved  through  Nuclear  Central  Amer- 
ica along  the  southern  lowlands,  cross- 
ing to  the  Atlantic  lowlands  through  the 
Isthmus  of  Tehuantepec  depression. 
Movement  along  the  Pacific  slopes  was 
limited  by  xeric  conditions  to  the  north. 
Similar  conditions  in  northeastern  Mex- 
ico prevented  further  northward  migra- 
tion on  the  Atlantic  slopes.  Eastward 
migration  along  the  Atlantic  lowlands 
from  the  Isthmus  of  Tehuantepec  may 
have  been  blocked  by  unsuitable  eco- 
logical conditions  in  the  form  of 
swamps,  or  by  competition  from  a  sec- 
ond line  of  undulata  stock  that  had 
migrated  along  the  Atlantic  lowlands 
of  Nuclear  Central  America  to  reach 
Guatemala  and  the  Yucatan  Peninsula. 
This  second  line  would  have  given  rise 
to  the  extant  populations  of  undulata  on 
Isla  de  Cozumel  off  the  Yucatan  Penin- 
sula and  on  Islas  del  Maiz  east  of  Nica- 


ragua when  these  islands  were  con- 
nected with  the  mainland.  Migration  of 
the  ancestral  stock  southward  from  Nu- 
clear Central  America  into  what  is  now 
Costa  Rica  would  account  for  the  pres- 
ent populations  along  the  southern 
Pacific  slopes  and  on  the  Meseta  Cen- 
tral of  Costa  Rica.  I  have  postulated 
that  the  first  of  the  two  lines  gave  rise 
to  those  populations  having  individuals 
with  color  patterns  emphasizing  stripes 
or  combinations  of  stripes  and  lateral 
bars  or  blotches.  The  second  line  gave 
rise  to  those  subpopulations  character- 
ized by  having  barred  flanks. 

Ameiva  chaitzami  apparently  evolved 
rather  recently  from  a  striped  fonn  of 
undulata,  possibly  because  of  ecological 
isolation  in  the  arid  valley  of  the  Rio 
Grijalva  and  its  tributaries.  If  further 
samples  linking  populations  of  chait- 
zami with  undulata  are  found,  it  is  pos- 
sible that  study  will  show  that  chaitzami 
is  a  form  of  undulata,  as  inadvertently 
proposed  by  Smith  and  Laufe  ( 1946 ) 
and  that  the  sympatry  of  the  two  seen 
today  in  Guatemala  is  the  result  of  sec- 
ondary contact. 

Although  I  think  it  unlikely,  studies 
of  South  American  Ameiva  may  show 
that  I  have  underestimated,  rather  than 
overestimated,  the  amount  of  diver- 
gence of  the  undulata  complex  from 
other  species.  If  so,  this  complex,  like 
Cnemidophorus,  may  be  found  to  have 
its  center  of  radiation  in  northern  Mid- 
dle America  and  may  be  considered  a 
component  of  the  Middle  American 
herpetofaunal  element  ( =  "Hanging 
Relicts,"  Schmidt,  1943;  "Autochthonous 
Middle  American  Element,"  Stuart, 
1950 )  proposed  by  Savage  ( 1966 ) .  Such 
an  arrangement  would  mean  that  the 
complex  was  isolated  in  Middle  Amer- 
ica with  the  opening  of  the  Panamanian 
portal  and,  with  some  adjustments  in 
timing,  would  support  the  hypothesis  of 
Smith  and  Laufe  ( 1946 )  concerning 
evolution  of  Ameiva  undulata  itself  in 
Middle  America. 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


71 


SUMMARY 


The  genus  Ameiva  is  broadly  dis- 
tributed in  Middle  and  South  America, 
as  well  as  on  islands  throughout  the 
Caribbean.  The  genus  is  represented  in 
Middle  America  by  six  species:  Aiiwiva 
ameiva  (Linnaeus),  leptophrys  (Cope), 
f  estiva  ( Lichtenstein  and  Von  Mar- 
tens), quadrilineata  ( Hallowell ) ,  tin- 
dulata  (Wiegmann)  and  chaitzami 
Stuart.  These  species  are  variously  dis- 
tributed throughout  the  lowland  tropics 
of  Middle  America  from  Nayarit  and 
central  Tamaulipas,  Mexico,  through 
Panama.  Ameiva  ameiva  and  f estiva, 
and  possibly  leptophrys  occur  in  South 
America  as  well.  Insular  populations 
are  known  of  all  species,  except  lepto- 
phrys and  chaitzami. 

Geographic  variation  is  extensive  in 
scutellation  and,  in  some  species,  color 
pattern.  Sexual  dimoiphism  and  onto- 
genetic change  in  color  pattern  is  pro- 
nounced in  some  species.  Certain  char- 
acters, such  as  total  number  of  supra- 
labials,  number  of  dorsal  granules  from 
occiput  to  rump  and  scales  around  the 
tail,  are  sexually  dimorphic  in  some 
species,  not  in  others.  On  the  other 
hand,  several  characters  show  no  sexual 
dimorphism  at  all.  Ontogenetic  changes 
are  most  striking  among  males.  Males 
of  some  species  undergo  marked 
changes  in  ventral  coloration,  especially 
in  the  gular  region;  these  may  be  asso- 
ciated with  reproductive  state.  Clines 
are  apparent  in  a  great  many  characters 
and,  as  might  be  expected  in  the  context 
of  the  essential  linearity  of  the  Middle 
American  land  mass,  most  clines  nm 
parallel  to  the  coastlines.  This  is  best 
illustrated  by  undulata,  the  species 
having  the  greatest  range  in  Middle 
America  and  for  which  the  most  samples 
were  available.  In  this  species,  the 
number  of  granules  around  the  body 
and  the  number  from  occiput  to  rump 
proved  the  most  sensitive  indicators  of 
differences  between  adjacent  samples. 
These    two     characters     are    relatively 


highly  correlated.  The  numbers  and 
arrangement  of  posterior  head  scales 
and  of  supraoculars  were  found  to  be 
unstable.  The  former  is  especially  true 
of  leptophrys  and,  to  a  lesser  extent, 
festiva.  The  condition  of  a  longitudinal- 
ly divided  interparietal,  occasionally 
used  to  diagnose  some  species  of  Amei- 
va, was  found  to  be  of  wide  occurrence 
in  several  species. 

In  Middle  America,  the  species  of 
Ameiva  run  an  ecological  gamut  from 
the  savanna-dwelling  ameiva  to  festiva, 
which  inhabits  lowland  tropical  rain 
forest.  In  the  presence  of  Cnemido- 
phorus,  Ameiva  tend  toward  less  open 
habitats,  although  the  same  species  may 
show  more  latitude  in  habitat  choice  if 
Cnemidophorus  is  absent.  As  many  as 
three  species  of  Ameiva  occur  sym- 
patrically  in  lower  Central  America. 
Competition  among  them  is  partially 
averted  by  partitioning  of  the  available 
habitat.  Typically,  one  species  is  a 
forest-dweller,  another  inhabits  open 
areas  and  the  third  is  intermediate.  Size 
differences  are  also  important  in  this 
context,  and  partitioning  by  subadults 
may  be  different  than  that  by  adults. 
Little  is  known  of  the  demography  of 
the  Middle  American  Ameiva.  All  are 
bisexvial. 

The  species  of  Ameiva  of  Middle 
America  have  been  assigned  to  three 
groups.  These  groups  cannot  be  given 
formal  taxonomic  recognition  until  their 
relationships  with  South  American  spe- 
cies of  Ameiva  have  been  ascertained. 
The  composition  of  the  groups  may 
have  to  be  modified  when  these  rela- 
tionships are  better  understood.  Group 
I  contains  only  Ameiva  ameiva.  This 
group  has  diverged  considerably  from 
Groups  II  and  III,  and  can  be  distin- 
guished on  the  basis  of  a  large  number 
of  characters  of  scutellation  and  color 
pattern.  The  group  is  primarily  South 
American  in  distribution.  Group  II  con- 
sists of  festiva  and  leptophrys,  both  of 


72 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


which  have  ranges  centered  in  lower 
Central  America.  These  two  species 
have  in  common  such  characters  as  a 
high  number  of  femoral  pores  and  a 
tendency  for  the  posterior  head  scales 
to  be  disrupted.  Group  III  is  made  up 
of  unchdata,  quadrilineota  and  chait- 
zami  and  is  entirely  Middle  American 
in  distribution.  The  group  is  character- 
ized by  certain  details  of  scutellation 
and,  especially,  color  pattern. 

The  genus  Ameiva  is  thought  to 
have  originated  in  South  America  and 
to  have  invaded  Middle  America  fol- 
lowing closure  of  the  Panamanian  por- 
tal in  Early  Pliocene.  Of  the  Middle 
American  species,  Ameiva  ameiva  now 
occupies  a  disjunct  range  in  central  and 
western  Panama  and  in  South  America 
east  of  the  Andes.  Ameiva  festiva  oc- 
curs on  eastern  and  western  sides  of  the 
Andes;  it  is  thought  that  the  eastern 
population  gave  rise  to  the  Middle 
American  population.  The  species  pres- 
ently has  a  continuous  range  along  the 
Atlantic  lowlands  to  Tabasco,  Mexico, 
and  along  Pacific  slopes  to  southeastern 
Costa  Rica.  Based  on  present  distribu- 
tion, leptophrys  may  have  evolved  as  an 
insular  isolate  in  the  Panamanian  portal. 
Ameiva  quadriUneata  is  thought  to  be  a 
lower  Central  American  derivative  of 
early  iimhdata  stock,  and  chaitzami  a 
recently  evolved  offshoot  of  a  striped 
form  of  umlulata.  Forerunners  of  un- 
dulata  itself  are  thought  to  have  mi- 
grated northward  on  either  side  of  Nu- 
clear Central  America.  Those  moving 
along  the  Pacific  routes  gave  rise  to  the 
populations  in  which  striped,  barred 
and  blotched  patterns  are  emphasized. 
Those  moving  through  the  Atlantic  low- 
lands have  given  rise  to  those  popula- 
tions in  which  lateral  bars  are  empha- 
sized. Populations  of  tindulata  from  the 
Atlantic    lowlands    of    Mexico    between 


central  Tamaulipas  and  Campeche  are 
derivatives  of  the  line  which  moved 
along  the  Pacific  routes  and  crossed  to 
the  Atlantic  slopes  through  the  depres- 
sion at  the  Isthmus  of  Tehuantepec. 
Without  exception,  insular  populations 
of  Ameiva  in  Middle  America  occur  on 
islands  of  the  continental  shelf.  These 
populations  were  isolated  from  main- 
land populations  when  rising  sea  levels 
severed  connections  with  the  mainland 
during  the  Pleistocene. 

The  genus  Ameiva  is  the  most  wide- 
ranging  of  the  exclusively  terrestrial 
Neotropical  lizards.  Over  its  entire 
range,  the  genus  has  exploited  a  broad 
spectrum  of  available  habitats.  For 
these  reasons,  a  better  understanding  of 
the  ecological  relationships  of  Ameiva 
may  lead  to  a  better  understanding  of 
tropical  vertebrates  in  general.  To  date, 
very  little  ecological  research  has  dealt 
with  Ameiva.  Interesting  problems  in- 
clude the  relationships  of  characters  of 
scutellation  and  color  pattern  to  envi- 
ronmental parameters,  the  cause(s)  and 
significance  of  color  changes  in  adult 
males,  and  the  ecological  relationships 
of  sympatric  species  of  Ameiva  and 
Cnemidophonis,  regardless  of  whether 
or  not  these  two  genera  are  shown  to  be 
congeners.  Taxonomic  problems  of  im- 
portance to  complete  understanding  of 
the  Middle  American  Ameiva  include 
elucidation  of  their  relationships  with 
South  American  species  (and  the  rela- 
tionships of  the  South  American  species 
to  one  another),  the  relationships  of 
chaitzami  to  undtdata,  and  the  status  of 
the  population  of  undulata  on  the  Me- 
seta  Central  of  Costa  Rica.  These  are 
but  a  few  suggestions  for  future  lines  of 
research.  There  seem  to  be  sufficient 
problems  to  keep  a  number  of  investi- 
gators occupied  for  years  to  come. 


RESUMEN 


El  genero  Ameiva  esta  ampliamente 
distribuido  tanto  por  America  Media  y 
Sur    America,    como    por    las    islas    del 


Caribe.  El  genero  esta  representado  en 
America  Media  por  seis  especies:  Am- 
eiva    ameiva     (Linnaeus),     leptophrys 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


73 


(Cope),  f estiva  ( Lichtenstein  y  Von 
Martens),  qiiodrilineata  ( Hallowell ) ,  y 
chaitzami  Stuart.  Estas  especies  se  en- 
cuentran  variablemente  distribuidas  al 
traves  de  las  tierras  bajas  tropicales  de 
America  Media,  desde  Nayarit  y  Ta- 
maulipas  central,  Mexico,  hasta  Pana- 
ma. Ameiva  amewa  y  f estiva,  y  posible- 
mente  leptophrys  ocurren  tambien  en 
Sur  America.  Se  conocen  poblaciones 
insulares  de  todas  las  especies  excepto 
de  leptophrys  y  chaitzami. 

La  variacion  geografica  de  la  escu- 
telacion  y,  en  algunas  especies,  el  patron 
de  coloracion  es  amplia.  El  dimorfismo 
sexual  y  el  cambio  ontogenetico  del 
patron  de  coloracion  es  pronunciado  en 
algunas  especies.  Ciertos  caracteres, 
como  el  mimero  total  de  supralabiales, 
numero  de  granulos  dorsales,  desde  el 
occipito  a  la  anca,  y  las  escamas  alrede- 
dor  de  la  cola,  son  sexualmente  dimor- 
ficos  en  algunas  especies  pero  no  en 
otras.  Por  otro  lado,  algunos  caracteres 
no  presentan  dimorfismo  sexual  del 
todo.  Los  cambios  ontogeneticos  son 
mas  sorprendentes  en  los  machos.  Los 
machos  de  algunas  especies  sufren  cam- 
bios marcados  en  la  coloracion  ventral, 
aspecialmente  en  la  region  gular,  y  estos 
se  pueden  asociar  con  el  estado  de  re- 
produccion.  Gradaciones  son  aparentes 
en  muchos  caracteres  y  en  su  mayoria, 
corren  paralelas  a  las  lineas  costera, 
como  se  puede  esperar  del  contexto  del 
arreglo  linear  de  la  masa  terrestre  de  la 
America  Media.  Esto  puede  ser  ilus- 
trado  mejor  con  undulata,  la  especie 
que  tiene  la  extension  mas  larga  por 
America  Media  y  de  la  cual  se  tienen 
el  mayor  numero  de  muestras.  En  esta 
especie,  el  numero  de  granulos  alrede- 
dor  del  cuerpo,  y  el  numero  desde  el 
occipito  a  la  anca  probaron  ser  los  mas 
sensibles  indicadores  de  diferencias  en- 
tre  muestras  adjacentes.  Estos  dos  car- 
acteres estan  relativemente  muy  corre- 
lacionados.  El  numero  y  arreglo  de  las 
escamas  supraoculares  y  las  posteriores 
a  la  cabeza  es  inestable.  Lo  primero  es 
especialmente    cierto    en    leptophrys   y, 


en  menor  grado,  festiva.  La  condicion 
de  poseer  la  escama  interparietal  divi- 
dida  longitudinalmente,  empleada  oca- 
cionalmente  en  las  diagnosis  de  algunas 
especies  de  Ameiva,  fue  encontrado 
siendo  de  gran  ocurrencia  en  varias 
especies. 

En  la  America  Media,  las  especies  de 
Ameiva  pasan  por  una  gama  ecologica 
desde  la  ameiva  que  vive  en  las  sabanas 
hasta  la  festiva  que  habita  en  las  tierras 
bajas  del  bosque  tropical  lluvioso.  En 
presencia  de  CnemidopJiorus,  Ameiva 
tiende  a  vivir  en  habitats  menos  abier- 
tos,  aunque  la  misma  especie  puede 
presentar  mayor  latitud  en  la  escogen- 
cia  del  habitat  si  Cnemidophorus  esta 
ausente.  Hasta  tres  especies  de  Ameiva 
ocurren  simpatricamente  en  la  baja  Cen- 
tro  America.  La  competencia  entre  ellas 
se  previene  por  medio  de  repartimiento 
del  habitat  utilizable.  Tipicamente,  una 
especie  habita  el  bosque,  otra  habita  las 
areas/  abiertas  y  la  tercera  es  inter- 
media. Las  diferencias  en  tamano  tam- 
bien son  importantes  en  este  contexto, 
y  la  reparticion  del  habitat  por  los  sub- 
adultos  puede  ser  diferente  de  la  de  los 
adultos.  Se  sabe  muy  poco  sob  re  la 
demografia  de  Ameiva  de  America  Me- 
dia.   Todas  son  bisexuales. 

Las  especies  de  Ameiva  de  America 
Media  han  sido  asignadas  a  tres  grupos. 
A  estos  grupos  no  se  les  puede  dar  re- 
conocimiento  taxonomico  formal  hasta 
que  sus  relaciones  con  las  especies  de 
Ameiva  de  Sur  America  se  determinen. 
La  composicion  de  los  grupos  podra 
tener  que  modificarse  cuando  estas  rela- 
ciones se  comprendan  mejor.  El  grupo 
I  contiene  solo  a  Ameiva  ameiva.  Este 
grupo  se  ha  apartado  considerablemente 
de  los  grupos  II  y  III,  ye  se  puede  dis- 
tinguir  en  base  al  gran  numero  de 
caracteristicas  en  la  escutelacion  y  pa- 
tron de  coloracion.  Este  grupo  es  prin- 
cipalmente  suramericano  en  distribu- 
cion.  El  grupo  II  consiste  de  festiva  y 
leptophrys,  ambas  con  distribuciones 
concentradas  en  la  baja  Centro  Amer- 
ica.  Estas  dos  especies  tienen  en  comun 


74 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


caracteres  como  un  numero  grande  de 
poros  femorales  y  la  tendencia  a  poseer 
escamas  rotas  en  la  parte  posterior  de 
la  cabeza.  El  grupo  III  se  compone  de 
undulata,  qiiadrilineata  y  chaitzami  y 
esta  distribuido  enteramente  por  Amer- 
ica Media.  Este  grupo  esta  caracteri- 
zado  por  ciertos  detalles  de  escutelacion 
y,   especialmente,   el  patron   de  colora- 


cion. 


Se  cree  que  el  genero  Ameiva  se 
origino  en  Sur  America  y  que  invadio 
la  Ameria  Media  despues  del  cierre  del 
paso  panamefio  en  el  Plioceno  tempra- 
no.  Ameiva  ameiva  posee  una  distribu- 
cion  discontinua  en  Panama  central  y 
oeste,  y  en  Sur  America  al  este  de  los 
Andes.  Ameiva  f estiva  ocurre  en  los 
lados  este  y  oeste  do  los  Andes;  se  cree 
que  la  poblacion  del  oeste  dio  origen  a 
la  poblacion  de  America  Media.  Esta 
especie  ahora  tiene  una  distribucion 
continua  a  lo  largo  de  las  tierras  bajas 
del  Atlantico  hasta  Tabasco,  Mexico,  y 
a  lo  largo  del  Pacifico  hasta  el  sureste 
de  Costa  Rica.  Basado  en  la  distribu- 
cion presente,  leptophrys  pudo  haber 
evolucionado  de  un  aislamiento  insular 
en  el  paso  panamefio.  Se  cree  que  quad- 
rilineata  se  derivo,  en  la  parte  baja  de 
Centro  America,  de  una  linea  temprana 
de  undulata  y  chaitzami  evoliciono 
recientemente  de  una  forma  de  undu- 
lata listada.  Se  cree  que  los  precursores 
de  undulata  misma  emigraron  hacia  al 
norte  por  ambos  lados  de  Centro  Amer- 
ica unclear.  Esos  que  tomaron  las  rutas 
del  lado  del  Pacifico  dieron  origen  a  las 
poblaciones  en  las  cuales  las  listas, 
barras,  y  manchas  son  enfatizadas.  Las 
que  se  desplazaron  por  las  tierras  bajas 
del  Atlantico  dieron  origen  a  las  pobla- 
ciones en  las  cuales  se  enfatizan  las 
barras  laterales.  Las  poblaciones  de 
undulata  de  las  tierras  bajas  del  Atlan- 
tico de  Mexico,  entre  Tamaulipas  cen- 
tral y  Campeche,  son  derivadas  de  la 
linea  que  se  movio  a  lo  largo  de  las 


rutas  del  Pacifico  y  que  crusaron  la  de- 
presion  del  istmo  de  Tehuantepec  para 
dirigirse  a  la  region  Atlantica.  Sin  ex- 
cepcion,  las  poblaciones  insulares  de 
Ameiva  en  America  Media  ocurren  en 
islas  de  la  plataforma  continental.  Estas 
poblaciones  fueron  aisladas  de  las  pob- 
laciones del  continente  cuando  los  ni- 
veles  del  mar  subieron  durante  el  Pleis- 
toceno  y  separaron  las  conecciones  con 
tierra  firme. 

El  genero  Ameiva  es  el  mas  amplia- 
mente  distribuido  de  las  lagartijas  Neo- 
tropicales  terrestres.  Sobre  su  total  dis- 
tribucion, el  genero  ha  expotado  un 
gran  espectro  de  habitats  utilizables. 
Por  estas  razones,  una  mejor  compren- 
sion  de  las  relaciones  ecologicas  en 
Ameiva  puede  llevar  a  una  mejor  com- 
prension  de  los  vertebrados  tropicales 
en  general.  Hasta  esta  fecha,  muy  poca 
investigacion  ecologia  a  tratado  sobre 
Ameiva.  Problemas  interresantes  inclu- 
yen  las  relaciones  de  los  caracteres  de 
escutelacion  y  el  patron  de  coloracion 
con  parametros  ambientales,  la  causa(s) 
y  significado  de  los  combios  de  colora- 
cion en  machos  adultos,  y  las  relaciones 
ecologicas  entre  especies  simpatricas  de 
Ameiva  y  Cnemidophorus,  indiferente- 
mente  de  presentar  o  no  a  los  dos  ge- 
neros  como  si  fuesen  congenericos. 
Problemas  taxonomicos  de  importancia 
para  completar  la  comprension  de  Am,- 
eiva  de  America  Media  incluyen  la 
aclaracion  de  sus  relaciones  con  las 
especies  de  Sur  America  (y  las  rela- 
ciones de  las  especies  suramericanas 
entre  si),  las  relaciones  entre  chaitzami 
y  undulata,  y  la  posicion  relativa  o 
status  de  undulata  en  la  Meseta  Central 
de  Costa  Rica.  Estas  son  no  mas  que 
algunas  sugerencias  para  futuras  lineas 
de  investigacion.  Parece  haber  sufici- 
entes  problemas  para  mantener  ocupa- 
dos  a  un  numero  de  investigadores 
durante  varios  aiios. 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


75 


APPENDIX  A 

An  example  of  summarized  data  as  used  in  this  report,  along  with  results  of 
Gabriel's  Simultaneous  Test  Procedure  analysis  for  number  of  femoral  pores  of  Pana- 
manian Ameiva  amewa.  For  a  complete  set  of  the  statistical  results  of  this  study, 
order  NAPS  Document  #01660  from  CCM  Information  Corporation,  866  3rd  Ave- 
nue, New  York,  New  York  10022;  remitting  $2.00  for  microfiche  or  $7.00  for  photo- 
copies. 


Ameiva  amewa:   Femoral  Pores 


Sample 
Number 

X 

Standard 
Deviation 

Sample 
Size 

Range 

STP 

6 

35.0 

2.76 

23 

29-40 

5 

34.1 

1.64 

11 

31-36 

7 

32.4 

1.85 

20 

29-37 

3 

32.3 

2.07 

24 

28-36 

2 

31.8 

1.93 

21 

29-35 

1 

31.7 

2.61 

40 

25-38 

4 

30.1 

1.28 

13 

27-32 

All 

32.4 

2.69 

270 

25-41 

76 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


APPENDIX  B 

Specimens  Examined 


Ameiva  ameiva 

Sample  1.  PANAMA:  Canal  Zone:  An- 
con,  MCZ  19805-11,  UMMZ  98415-16,  98418; 
Balboa,  AMNH  37129,  FMNH  13441,  UMMZ 
98411-14;  Bruja  Point,  FMNH  13445;  nr 
Corozal,  MCZ  19813;  Diablo  Heights,  UMMZ 
98417;  6.4  km  E  Lacone,  KU  62701;  Mira- 
flores,  FMNH  16584-5.  Panama:  nr  City  of 
Panama,  MCZ  9924-27,  9929-30,  9933-41; 
Panama  City,  El  Cangrejo,  KU  96742-43; 
Tapia,  AMNH  28005-6,  28008. 

Sample  2.  PANAMA:  Canal  Zone:  Gatiin, 
FMNH  16555-57,  16572-74,  16580,  16582-83; 
Cristobal,  FMNH  34163-64.  Colon:  Achiote, 
KU  76109-18,  96737. 

Sam^ple  3.  PANAMA:  Panama:  Bejuco, 
KU  107546;  Nuevo  Gorgona,  AMNH  89925- 
46;  ca  12  km  NW  San  Carlos  on  road  to  El 
Valle,  KU  107547. 

Sample  4.  PANAMA:  Panama:  Archi- 
pelago de  las  Perlas,  Isla  San  Jose,  AMNH 
66491,  USNM  120611-23. 

Sample  5.  PANAMA:  Herrera:  Las  Minas, 
ANSP  22452;  Parita,  USNM  127291-94, 
127296;  8  km  SE  Parita,  KU  107538-39;  Pese, 
KU  107540-45. 

Sample  6.  PANAMA:  Los  Santos:  Gua- 
nico,  KU  107521-35;  Playa  Guanico,  near 
mouth  of  Rio  Guanico,  KU  107909;  Las  Pal- 
mitas,  USNM  148207-11,  KU  110731-34. 

Sample  7.  PANAMA:  Panama:  S  Slope 
Cerro  Campana,  KU  76094-106,   107548-54. 

Additional  Specimens  Not  Used  In  STP 
Analysis.  COSTA  RICA:  No  definite  locality, 
BMNH  80.6.21.1-2.  PANAMA:  Canal  Zone: 
Alajuela,  UMMZ  76104(4);  Ancon,  KU 
110740;  Bas  Obispo,  USNM  54129;  Camp 
Chagres,  KU  76107-8;  Chagres,  USNM  5519; 
Culebra,  FMNH  16559-61;  Culebra  Cut, 
FMNH  16554;  Fort  Kobbe,  KU  110739;  Fort 
Kobbe,  Venado  River,  KU  110742-43;  El 
Vigia,  UMMZ  76012;  Frijoles,  UMMZ  63760; 
Gamboa,  AMNH  32818;  Juan  Mina,  Chagres 
River,  KU  108258-61;  Lower  Chagres,  AMNH 
37073-4,  37077;  Madden  Dam,  UMMZ 
76015(2);  Madden  Forest  Preserve,  KU 
96740-41;  Rio  Abajo,  FMNH  130685;  Rio 
Frijoles,  USNM  8394;  Road  K9,  FMNH 
130972;  Rosseau,  KU  62702;  Summit,  FMNH 
130683,  131309;  nr  Summit  Experimental 
Gardens,  UMMZ  95484;  Tabernilla,  FMNH 
16585-87,  USNM  54130.  Chiriqui:  2.5  km  E 
La  Concepcion,  KU  96736;  Remedios,  AMNH 
28390;  3  km  W  Rio  Chorcha,  KU  95540-44; 
16  km  W  Rio  Chorcha,  KU  95545;  Rio  Salado, 
KU  95546-47.  Cocle:  Bank  of  Rio  Chico  de 
Nata,  KU  95548-52;  El  Valle,  AMNH  89924; 
16  km  S,  9  km  W  Penonome,  KU  107536;  19 
km  S,  9  km  W  Penonome,  Puerto  Posoda,  KU 


107537.  Colon:  Portobello,  USNM  65122; 
Quebrada  Bonita,  KU  96738-39;  Santa  Rosa, 
AMNH  71705.  Herrera:  Tres  Puntas,  ANSP 
22470-74.  Los  Santos:  Jobero,  KU  110735- 
36;  Pocri,  KU  107520;  Punta  Mala,  AMNH 
71704.  Panama:  Cafiitas,  UMMZ  124148-50; 
Cerro  Azul,  AMNH  89947-48;  Chorrera, 
AMNH  71703,  FMNH  57597-98;  Flora  de 
Lagunas  (Rio  Caiiitas),  UMMZ  124146-47; 
Gatuncillo,  UMMZ  76013;  Isla  San  Miguel, 
MCZ  9942-47;  Isla  Taboga,  FMNH  16576, 
USNM  54197,  54205,  102754-56;  Isla  Tabo- 
guilla,  USNM  102809-10;  Juan  Diaz,  KU 
110737-38;  Nuevo  Emperador,  FMNH  130937, 
130939-45,  KU  110471;  Pacora,  AMNH  32817, 
FMNH  120956;  Rio  Bayano,  USNM  53980; 
Rio  Mamoni,  FMNH  16566-68;  Rio  Mamoni, 
5  km  E  Chepo,  KU  76119-25;  Rio  Tapia,  24 
km  E  Panama  City,  AMNH  42953.  Veragita^-: 
26  km  W  Santiago,  KU  95553-55;  Sona, 
USNM  51972-73. 

Ameiva  chaitzami 

Sample  1.  MEXICO:  Chiapas:  Comitan, 
UMMZ  94905(24),  94906(5),  94907. 

Sample  2.  GUATEMALA:  Htiehuete- 
nango:  San  Antonio  Huista,  UMMZ  120192; 
1  km  N  San  Antonio  Huista,  UMMZ  120189, 
120190(4),  120191(2);  1  km  W  San  Antonio 
Huista,  UMMZ  120195;  1-2  km  W  San  An- 
tonio Huista,  UMMZ  120193(18);  1-2  km  E 
San  Antonio  Huista,  UMMZ  120194(4). 

Sainple  3.  GUATEMALA:  Alta  Verapaz: 
Languin-Cahabon  road,  MCZ  52170;  On  Lan- 
guin-Cahal^on  road  ca  2  km  N  Finca  Canihor, 
UMMZ  90638-41,  90643.  El  Peten:  1  km  N 
Popti'm,  UMMZ  124355;  2  km  SW  Poptun, 
UMMZ  124356;  2  km  NE  Poptun  nr  source 
Rio  San  Juan,  UMMZ  124357. 

Additional  Specimens  Not  Used  In  STP 
Analysis.  MEXICO:  Chiapas:  La  Libertad, 
MCZ  56007,  USNM  137216;  La  Libertad,  nr 
Rio  Cuilco  where  it  crosses  the  Guatemalan 
border,  FMNH  100006,  100712-15;  unknown, 
but  pro1ia]:)ly  near  La  Libertad,  FMNH 
100716. 

Ameiva  festiva 

Sample  1.  GUATEMALA:  El  Peten: 
Piedras  Negras,  FMNH  115429-43,  USNM 
133826-36,  UIMNH  11349-62. 

Sample  2.  GUATEMALA:  Alta  Verapaz: 
Chinaja,  KU  55839,  55849-50,  59566;  ca  3  km 
S  Chinaja,  KU  55841-48,  55857.  El  Peten:  6 
km  NNW  Chinaja,  KU  55836;  11  km  NNW 
Chinaja,  KU  55837-38;  ca  15  km  NW  Chinaja, 
KU  55851-56,  59567-73;  Toocog,  15  km  SE 
La  Libertad,  KU  55840. 

Sample  3.    GUATEMALA:    Alta  Verapaz: 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


77 


Finca  Chama,  UMMZ  91319,  91321(2), 
91322(2),  91323(3),  91324(2),  91325(2), 
91326(4),  91327,  91328(3),  91329(2),  91330, 
91331(8),  91332. 

Saiyiple  4.  HONDURAS:  Atlantida:  ca 
15  km  E  La  Ceiba,  mtns  above  Corozal,  JRM 
2299,  2418-21,  2439-40,  2466-69;  Along  Rio 
Cangrejal  ca  12  km  SSE  La  Ceiba,  KU 
101195-97;  Tela,  Lancetilla,  UMMZ  70324(3). 

Sample  5.  NICARAGUA:  Zelaya:  Bo- 
nanza, KU  84870-72,  86039-40,  101201-12;  6 
km  E  Bonanza,  KU  101198;  On  banks  of  Rio 
Tunkey  4  km  E  Bonanza,  KU  101199-200; 
Eden  Mine,  ANSP  2133-35. 

Sample  6.  NICARAGUA:  Zelaya:  Cukra, 
AMNH  16587-96;  Kanawa,  AMNH  16597, 
16605-16;  Between  Cukra  and  Kanawa, 
AMNH  16600,  16602-4. 

Sample  7.  NICARAGUA:  Zelaya:  El 
Recreo,  16  km  W  Rama  on  Rio  Mico,  UCLA 
9769-70;  El  Recreo,  S  side  Rio  Mico,  KU 
101874;  16  km  above  El  Recreo  on  Rio  Mico, 
UMMZ  79838(12);  11.2  km  above  Rama, 
UMMZ  79840(5),  79841(2);  Junctions  Rio 
Mico  and  Rio  Siquia,  UMMZ  79839(4). 

Satnple  8.  NICARAGUA:  Rio  San  Juan: 
Tuli  Creek,  AMNH  16666-74,  16676-700. 

Sample  9.  COSTA  RICA:  Heredia:  Puer- 
to Viejo,  KU  67298-309,  91787,  104081-82; 
UMMZ  125505;  1.5  km  N  Puerto  Viejo,  KU 
67310-11;  2  km  NE  Puerto  Viejo,  KU  95558- 
68;  3  km  S  Puerto  Viejo,  KU  95556-57;  ca 
12.8  km  NE  Puerto  Viejo  along  Rio  Sarapiqui, 
UMMZ  125506(2);  Rio  Sarapiqui  at  Puerto 
Viejo,  UMMZ  123582-83;  Finca  La  Selva  on 
Rio  Puerto  Viejo,  KU  100683-85. 

Sample  10.  COSTA  RICA:  Cartago:  Tur- 
rialba,  FMNH  101906,  103107,  MCZ  55091- 
92,  55094,  USNM  37737,  KU  25633-34, 
25636-39,  34783-89,  34826-27,  40552,  UMMZ 
117492;  5  km  SE  Turrialba,  KU  25174-75; 
6.4  km  SE  Turrialba,  KU  34778-79;  ca  4  km 
SW  Moravia  de  Turrialba,  KU  67312;  4.8  km 
W  Turrialba,  Hacienda  Florencia,  KU  34790- 
91. 

Sajnple  11.  COSTA  RICA:  Limon:  Vi- 
cinity of  mouth  of  Rio  Tortuguero,  AMNH 
89189-203,  89205-15. 

Sample  12.  COSTA  RICA:  Limon:  Gua- 
piles,  MCZ  15335-36;  La  Lola,  KU  34798-801, 
34807-13,  UMMZ  117493(9);  Los  Diamantes, 
FMNH  101250,  101252-54,  101256-59, 
101261-63,  101904-5,  101907,  KU  25581-86, 
25619-32,  34772-76,  34802-6;  Puerto  Limon, 
KU  34797;  Zent,  MCZ  15376-81. 

Sample  13.  PANAMA:  Bocas  del  Tow: 
Almirante,  FMNH  154445-48,  154451,  MCZ 
19785-87,  KU  80207-11;  3.2  km  NW  Almi- 
rante, KU  96745-46;  4.8  km  NW  Almirante, 
KU  96747;  5.6  km  NW  Almirante,  UU  5162; 
9.6  km  NW  Almirante,  KU  96748;  12.8  km 
NW  Almirante,  KU  96749-52. 

Satnple   14.     PANAMA:     Bocas   del    Tow: 


Isla  Escudo  de  Veraguas,  KU  108262-71,  UU 
5153-54. 

Sample  15.  PANAMA:  Canal  Zone:  Bar- 
ro  Colorado  Island,  AMNH  6.3401,  89949, 
FMNH  13342-43,  MCZ  20612-13,  28066-67, 
KU  76126-27,  UMMZ  63746-55,  63766,  63779, 
124156. 

Sample  16.  PANAMA:  Darien:  Rio 
Mono,  ca  5  km  above  junction  with  Rio  Tuira, 
KU  96781-84;  Rio  Tuira  at  Rio  Mono,  KU 
96785-804. 

Sample  17.  COSTA  RICA:  Puntarenas: 
Agua  Buena,  Caiias  Gordas,  KU  40545-48, 
40553-58;  Golfito,  KU  67313;  2.5  km  ESE 
Piedras  Blancas,  KU  93978;  Rio  Zapote,  8  km 
E  Palmar  Norte,  KU  93977,  95569. 

Sample  18.  COSTA  RICA:  Puntarenas: 
3  km  NW  Rincon  de  Osa,  KU  101487-88;  7.2 
km  SE  Rincon  de  Osa,  KU  101489-90;  9.6 
km  NW  Rincon  de  Osa,  KU  101483-86;  10  km 
SSE  Rincon  de  Osa,  KU  101473-80;  11.7  km 
SSE  Rincon  de  Osa,  KU  101481-82. 

Sample  19.  COLOMBIA:  Cordoba:  Cafio 
Guarmal,  FMNH  165260-66;  Upper  Rio  Ure, 
FMNH  165225,  165227-30,  165748-67, 
165769. 

Sainple  20.  COLOMBIA:  Choco:  An- 
dagoya,  USNM  124246;  Andagoya-Condoto 
Area,  UMMZ  121463;  Condoto,  UMMZ 
121465(13);  Tado,  ANSP  26265-66. 

Additional  Specimens  Not  Used  In  STP 
Analysis.  BRITISH  HONDURAS:  Cayo: 
Valentin,  UMMZ  92372;  Stann  Creek:  Mid- 
dlesex, FMNH  4460;  Toledo:  1.6  km  E  Swa- 
zey  Branch  of  Monkey  River,  MCZ  71606-7; 
District  Unknown:  Silk  Grass  Creek  State  For- 
est, FMNH  4459;  Bobowina,  FMNH  49313-14. 
COLOMBIA:  Boyaca:  Muzo,  MCZ  56233; 
Choco:  Boca  de  la  Raspadura,  AMNH  18266- 
68;  El  Valle,  Bahia  Solano,  Mutis,  USNM 
151512;  El  Valle,  Pizarro,  USNM  151513-14; 
Pizarro,  FMNH  43816-28;  Rio  San  Pado,  1.6  hn 
below  Boca  de  la  Raspadura,  AMNH  18274;  Si- 
erra Bauda,  ANSP  25526;  Sierra  de  Darien, 
ANSP  25527-28;  Upper  Rio  San  Juan,  FMNH 
15652-53;  Santander:  El  Centro  Santander, 
ANSP  25075,  25427;  Departamento  Unknown: 
E  base  Cordillera  de  Bogota  at  extreme  limit 
of  llanos,  ANSP  24165;  "Tambo  S.  Monica, 
Atlantic  Slope,"  AMNH  18265;  Region  of 
Truando,  ANSP  9071.  COSTA  RICA:  Ala- 
juela:  3.2  km  NE  Muelle  Canal,  UCR  23-26; 
Isla  Bonita,  FMNH  103103-6;  Cartago:  nr 
Peralta,  Lake  Bonilla,  KU  43886-8;  nr  Peralta, 
Tunnel  Camp,  KU  34828,  38825;  Reventazon, 
USNM  37492;  Rio  Pacuare,  nr  Pacuare,  KU 
34777;  Suiza,  FMNH  101251,  101255,  KU 
25635,  34792;  Turrialba,  FMNH  101909;  6.4- 
8.0  km  S  Turrialba,  FMNH  101260;  Guana- 
caste:  Laguna  de  Arenal,  UCR  232-33,  268- 
69;  Tenorio,  KU  34814-24;  Tilaran,  ANSP 
24561;  5  km  ENE  Tilaran,  KU  40551;  Tron- 
cadero,  nr  Lake  Arenal,  KU  40539-40;  Here- 


78 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


dia:    Barro  Colorado,  KU  100679-80;  Behveen 
Puerto    Viejo   and    Cariblanco,    KU    40537-38; 
Limon:    Bataan,    KU   34780-82;    Pandora,    KU 
86573-76,    100681-82;    Suretka,    MCZ    19777, 
KU    40541-44,    40559-63;    Puntarenas:     Agua 
Buena,  Caiias  Gordas,  KU  40549-50;  San  Jose: 
San     Isidro     del     General,     FMNH     101908, 
101910,  KU  34793-94;  8  km  E  San  Isidro  del 
General,  FMNH   120236;  24-32  km   NW   San 
Isidro  del  General,  San  Juan  Mtns.,  KU  34795- 
96;     Province     Unknown:      Guayabo,     FMNH 
6167;    Rio    Frio,    USNM     19534-36;    Locality 
Unknown,  USNM  30558,  32612-13.    GUATE- 
MALA:    Alfa    Verapaz:     Cubiliquitz,    UMMZ 
91334,    91335(2),    91336(2);    Finca    Chama, 
MCZ    28452-54,     UMMZ    91320(2),     91329, 
91331;      Panzos,      UMMZ     9137;      Samazana, 
UMMZ  91333(4);  El  Quiche:    Finca  Tesoro, 
UMMZ    89197(4);    Izabdh     "Babos    Pi.,    Mt. 
Playita,"    FMNH    20081,    20109-11;    Quiriqua, 
FMNH  20487-88;  El  Peten:    Pacomon,  USNM 
71394;    Poptim-San    Luis    Road,    ca    15    km    S 
Poptim,    UMMZ    124359;    12    km    S    PopUm, 
UMMZ   124360;    10  km  NW  Poptun,  UMMZ 
124.361(5);     Ramate,    USNM    71406,    71817, 
71823,    71826-27,    71829-31,    71836;    Localiiij 
Unknown:    MNHN  5480.    HONDURAS:     Co- 
pan:    19.2  km  ENE  Copan,  JRM  2895,  2897; 
Cortes:    Hacienda  Santa  Ana,  San  Pedro  Sula, 
FMNH  5067-71;  Mtns  W  of  San  Pedro  Sula, 
FMNH    5074;    Garcias   a    Dios:     Rio    Segovia, 
USNM  24527-28;  Santa  Barbara:   Jaral,  FMNH 
5073;     Yoro:      Mataderos     Mountains,     FMNH 
21781,    MCZ    38924;    Portillo    Grande,    MCZ 
38925;     Subirana    Valley,    MCZ    38926;     De- 
partamento      Unknown:       Carmelina,      USNM 
62972;   Lake  Ticamaya,  FMNH   5072.     MEX- 
ICO:   Chiapas:    Palenque,   USNM    133818-25, 
UIMNH  11343,  KU  94107-8;  Tabasco:    Teapa, 
USNM    47454-56.     NICARAGUA:      Rio    San 
Juan:     Greytown,    USNM    19533-34,    19640-1; 
Zelatfa:    3-4  km  W  Bluefields,  KU   101213-15; 
Cupitna,  AMNH   16633-51;  Rio  Escondido  70 
km    from    Bluefields,    USNM     19735,     19871. 
PANAMA:     Boca,s-  del  Toro:     11   km   NW  Al- 
mirante,    FMNH    68171;    Bastimentos,    USNM 
150021;   Boca  del   Drago,    USNM    142332-33; 
Cayo   Agua,    USNM    150014-15;    Rio   Cahuita, 
ca   5    km    above    mouth,    KU    108278-79;    Rio 
Changena,    FMNH     130971,    KU     110744-45; 
Canal    Zone:      Cerro    Azul,     FMNH     130684; 
Cristobal,     FMNH    34165;     Gamboa,     FMNH 
131308;     Gatun,     FMNH     16654-56,     USNM 
54134;   Chiriqui:     Boquete,   UMMZ   58296;   El 
Volcan,  USNM  129924;  Finca  Palosanto  7  km 
NW  El  Volcan,  Rio  Chiriqui  Viejo,  KU  96808- 
11;  Finca  Santa  Clara,  KU  110753-54;  Volcan 
Chiriqui,    UMMZ    101787;    Code:     El    Valle, 
FMNH    47456;    Finca    El    Valle    de    Anton, 
FMNH  60174;  CoUn:    Achiote,  KU  76129-30; 
Candelaria      Hydrographic      Station,      FMNH 
68172-73;  Portobello,  FMNH  16652-53,  USNM 
54072-73,    80044;    3.5    km    SE    Puerto    Pilon, 


KU  110755;  Rio  Candelaria  nr  Hydrographic 
Station,  FMNH  68174-76;  Darien:  Cafia, 
USNM  50134,  50136,  50138-39,  ,50142-43;  El 
Real,  KU  80529;  Laguna,  KU  76131;  Rio 
Cupe,  ca  12  km  SSW  Boca  de  Cupe,  KU 
96805;  Rio  Esnape,  Sambu  Valley,  MCZ 
17224-29;  Rio  Jaque,  1.5  km  above  Rio 
Imamado,  KU  110751-52;  Rio  Jesuito, 
Sapo  Mtns,  MCZ  17179-80,  17189;  Rio 
San  Antonio  nr  base  of  La  Jarcia,  KU  110750; 
Tacarcuna,  KU  76132-39;  Tacarcuna  Village, 
USNM  141819-20;  Panama:  S  Slope  Cerro 
Campana,  KU  76128;  Cerro  Tefe,  KU  96806-7; 
E  Slope  Cerro  Jefe,  KU  80530-37;  NW  Slope 
Cerro  Prominente,  KU  80538-39;  Chepo, 
FMNH  16656;  El  Valle  de  Anton,  trail  to  La 
Mesa,  FMNH  68177-79;  Rio  Abajo,  FMNH 
57599;  Rio  Siluganti,  UMMZ  124151-55;  San 
Bias:  Camp  Sasardi,  KU  110748-49;  Camp 
Summit,  KU  110746-47;  Veraguas:  Mouth  of 
Rio  Concepcion,  KU  108272-77;  Province  Uti- 
known:  Cabima,  USNM  48504;  Chico,  UMMZ 
76011;  Nombre  de  Dios,  MCZ  26944;  Punta 
de  Pena,  USNM  38693-98;  "Veragua,"  ZMB 
881a,  881b. 

Ameiva  leptophrys 

Sample  1.  COSTA  RICA;  Puntarenas: 
Camp  Seattle,  6.4  km  inland  from  Rincon  de 
Osa,  USC(CRE)  752;  Vicinity  of  Rincon  de 
Osa,  KU  101491-528,  101825. 

Sample  2.  COSTA  RICA:  Puntarenas: 
Goto,  Colfo  Dulce,  UMMZ  71999-72002;  Es- 
quinas  Forest  Preserve  between  Golfito  and 
Palmar,  KU  34766-70;  3.2-4.8  km  W  Palmar 
Norte  on  road  to  Puerto  Cortez,  USC(CRE) 
7101c-7101f;  6  km  W  Palmar  Norte,  KU 
93982-83,  95570-82;  6  km  SE  Palmar  Norte, 
KU  67314-16;  Rio  Zapote,  8  km  E  Palmar 
Norte,  KU  93980-81;  Villa  Neilly,  nr  town  of 
Corredor,  USC(CRE)  179a;  12.3  km  WNW 
\'il]a  Neilly,  KU  67317. 

Sample  3.  PANAMA:  Canal  Zone:  Barro 
Colorado  Lsland,  AMNH  47018-19,  63403(2), 
89953,  ANSP  23021-22,  FMNH  13307,  1.3344- 
47,  13375-76,  MCZ  28063-65,  29386,  KU 
76140-45,  80540,  UMMZ  63756-59,  63780;  nr 
Coro/al,  MCZ  19814;  Corozal  Hill,  USNM 
102743. 

Sample  4.  PANAMA:  Darien:  Rio  Chu- 
cunaque,  ca  7  km  above  Rio  Morti,  KU 
107561-63;  Rio  Cupe,  ca  12  km  SSW  Boca  de 
Cupe,  KU  96815;  Rio  Tuira  at  Rio  Mono,  KU 
96816-25;  Rio  Ucurganti,  ca  7  km  above 
mouth,  KU  107564-65;  Tacarcuna,  KU  76158- 
61;  Tacarcuna  Village,  USNM  141817-18; 
Three-Falls  Creek,  AMNH  37870. 

Additional  Specimens  Not  Used  In  STP 
Anahisis.  COSTA  RIGA:  Puntarenas:  Par- 
rita.  La  Tulieta,  Unca  La  Ligia,  USC(CRE) 
8256,  8260]>8260c;  2.5  km  ESE  Piedras  Blan- 
cas,  KU  93979;  Rincon  de  Osa,  RDS  1122-23, 
1170,  1177;  San  Jose:  San  Isidro  del  General, 
FMNH   103111,   103117,  KU  25640-42;  Prov- 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


79 


ince  Unknown:  Pozo  Azul  de  Parris,  ANSP 
21470;  Santo  Domingo  de  San  Mateo,  USNM 
37754.  PANAMA:  Canal  Zone:  Alajuela, 
UMMZ  76009;  Camp  Chagres,  KU  76146-48; 
Corozal,  AMNH  67073;  Gatim,  USNM  54142; 
0.5  km  S  Juan  Mina,  FMNH  68159;  Madden 
Dam,  UMMZ  76010;  Miraflores,  FMNH  16650; 
Rio  Cardenas,  Corozal,  USNM  53868;  Chiri- 
qui:  Progreso,  MCZ  19865;  13  km  N  Puerto 
Armuelles,  KU  96814;  Code:  El  Valle,  AMNH 
89950-52,  USNM  129906,  KU  76162-63; 
Colon:  Achiote,  KU  76149-51;  Portobello, 
USNM  92593;  Quebrado  Bonita  nr  Buenavista, 
KU  96812;  1  km  W  Rio  Piedras,  KU  96813; 
San  Bias  Point,  AMNH  71706;  Darien:  Cana, 
USNM  50135,  50137,  50140-41;  Canclones, 
UMMZ  124975;  nr  mouth  of  Rio  Canclones, 
UMMZ  124974;  Taque,  KU  110767-68;  La- 
guna,  KU  76156-57;  Mt.  Sapo,  ANSP  22963, 
MCZ  17201-3;  Rio  Jaque,  1.5  km  above  Rio 
Imamodo,  KU  110769-70;  Rio  Jesuito,  Sapo 
Mtns,  MCZ  17196-97,  17199;  Herrera:  Las 
Minas,  ANSP  22450;  Los  Santos:  N  Slope 
Cerro  Cambutal,  KU  107555-56;  Guanico 
Abajo,  KU  110771;  Panama:  Cerro  Azul, 
USNM  54136,  54147;  Cliffs  W  of  Cerro  Cam- 
pana,  FMNH  60178-79;  S  Slope  Cerro  Cam- 
pana,  KU  76152-53,  107558-59;  Chepo,  FMNH 
16645;  Las  Cumbres,  KU  96827;  Nuevo  Em- 
perador,  FMNH  130938;  nr  Panama  City, 
MCZ  10925,  10927;  2.8  km  ENE  Panama 
City,  Rio  Pacora,  KU  96826;  9  km  NNE  Pa- 
cora,  Rio  Pacora,  KU  107557;  Rio  Mamoni,  5 
km  E  Chepo,  KU  76154-55;  Rio  Siluganti, 
UMMZ  124144-45;  Rio  Tocumen,  MCZ  19816; 
San  Bias:  Camp  Sasardi,  KU  110756-65;  Camp 
Summit,  KU  110766;  Veragtias:  Mangillo, 
ANSP  22450. 

Ameiva  quadrilineata 

Sample  1.  NICARAGUA:  Rio  San  Juan: 
Greytown,  USNM  19537-45,  20741-42,  32230; 
Rio  San  Juan,  USNM  24984. 

Sample  2.  COSTA  RICA:  Limon:  Colo- 
rado Bar,  AMNH  16726-28;  Tortuguero, 
USC(CRE)  2653(2);  Vicinity  of  mouth  of 
Rio  Tortuguero,  AMNH  89217-46. 

Sample  3.  COSTA  RICA:  Liinon:  Los 
Diamantes,  UMMZ  117491;  La  Lola,  USC 
(CRE)  128(3),  8063,  8070(3);  Zent,  MCZ 
15382-85. 

Sample  4.  PANAMA:  Bocas  del  Tow: 
Almirante,  FMNH  60171-72;  Careening  Cay, 
KU  96840-44;  Cayo  Zapatillo  Grande,  KU 
96845-51;  Isla  Bastimentos,  Bastimentos,  KU 
96831-39;  Isla  Bastimentos,  Punta  Vieja, 
USNM  150016-20,  150022-23;  Isla  Colon, 
road  from  Bocas  to  la  Gruta,  KU  96829-30. 

Sample  5.  PANAMA:  Bocas-  del  Tow: 
Isla  Escudo  de  Veraguas,  KU  108280-85,  UU 
5166. 

Sample  6.  COSTA  RICA:  San  Jose:  San 
Isidro  del  General,  FMNH  103115,  KU  25643- 


50,  34685-89,  USC(CRE)  2742;  4.8  km  NE 
San  Isidro  del  General,  USC  (CRE)  2843;  5.4 
km  SW  San  Isidro  del  General,  UMMZ 
117490(2);  3.2  km  W  San  Isidro  del  Gen- 
eral, KU  25651;  8  km  SE  San  Isidro  del 
General,  KU  95583-90. 

Sample  7.  COSTA  RICA:  Ptmtarenas: 
1.5  km  E  Palmar  Norte,  KU  95592-93;  6  km 
W  Palmar  Norte,  KU  95633-40;  3.2-4.8  km  W 
Palmar  Norte  on  road  to  Puerto  Cortez,  USC 
(CRE)  7101a-7101b;  5  km  SE  Palmar  Sur, 
KU  67318-19;  6  hn  SE  Palmar  Sur,  KU 
67320-25;  Rio  Zapote,  8  km  E  Palmar  Norte, 
KU  95607-20,  95622-26,  104083-85. 

Sample  8.  COSTA  RICA:  Puntarenas: 
Rincon  de  Osa,  RDS  1136-38,  1144,  1147, 
1152,  1154-63,  KU  101463-69,  USC  (CRE) 
7239(14);  3  km  NW  Rincon  de  Osa,  KU 
101470-72;  11.7  km  SSE  Rincon  de  Osa,  KU 
101459-62. 

Sample  9.  PANAMA:  Chiriqui:  Comarca 
del  Baru,  Puerto  Annuelles,  FMNH  68166-67, 
Progreso,  UMMZ  58186-202;  8  km  S  Progreso, 
KU  96852;  Puerto  Annuelles,  MCZ  45668-71; 
13  km  N  Puerto  Annuelles,  KU  96853-65. 

Sample  10.  PANAMA:  Chiriqui:  Bo- 
quete,  ANSP  21941-43,  UMMZ  58183-85, 
58209-18. 

Additional  Specimens  Not  Used  In  STP 
Analysis.  COSTA  RICA:  Li7n6n:  Colorado 
Bar,  AMNH  16729-75;  La  ForUina,  AMNH 
99679;  Old  Harbor,  USNM  32614-16;  Suretka, 
KU  40578;  Puntarenas:  0.8-1.6  km  E  Bani 
along  N  bank  of  Rio  Baru,  USC  (CRE)  7096; 
Buenos  Aires,  FMNH  2519;  Isla  del  Cano, 
UMMZ  71194;  Goto,  UMMZ  72005-6;  El  Gen- 
eral (Viejo)  between  Palmar  Sur  and  Golfito, 
KU  34690;  Esquinas,  KU  34691;  Golfito, 
USC(CRE)  7111(5),  7233(3);  Km  47  on 
railway  to  Golfito,  USC  (CRE)  176(8),  177 
(2),  178;  Finca  El  Helechales,  15  km  NE 
Potrero  Grande,  USC  (CRE)  8268,  8271;  27 
km  E  Palmar  Norte,  Rio  Punta  Nuevo,  KU 
95596-606;  30  km  E  Palmar  Norte,  Rio  La 
Vieja,  KU  95627-32;  2-5  km  ESE  Piedras 
Blancas,  KU  93984-86;  Puerto  U\dla,  UMMZ 
72004;  Rincon  de  Osa,  RDS  1139-40,  1142, 
1145-46,  1148-51,  1153;  San  Isidro  del  Gen- 
eral, FMNH  103116;  22  km  SE  San  Isidro  del 
General,  USC  (CRE)  2648;  57  km  SE  San 
Isidro  del  General,  KU  95591;  Villa  Neilly, 
USC(CRE)  179;  1.6  k-m  E  Volcan  de  Buenos 
Aires,  Cone  Finca,  UMMZ  117573;  San  Jose: 
San  Isidro  del  General,  3.2  km  W  Intabee, 
FMNH  103102.  NICARAGUA:  Rio  San 
Juan:  Greytown,  USNM  6053(2).  PANAMA: 
Chiriqui:  SW  Slope  Cerro  Homito,  KU  96867; 
11  km  NE  Gualaco  Road  to  Valle  Homito,  KU 
96866;  El  Hato,  edge  of  lava  flow,  FMNH 
130686;  Llano  del  Volcan  above  Paso  Ancho, 
FMNH  60175-77,  68168-70;  Herrera:  Las 
Minas,  ANSP  22454-55;  Veraguas:  8  km  SW 
El  Maria,  KU  107566;  Isla  Cebaco,  KU  96828. 


80 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


Ameiva  undulata 

Sample  1.  MEXICO:  Tamaulipas:  Go- 
mez Farias,  UMMZ  110802,  110839;  nr  Gomez 
Farias,  UMMZ  111145;  8  km  NE  Gomez 
Farias  along  Rio  Sabinas,  UMMZ  101476-78, 
101479(6),  101480-82,  101515-16,  104059 
(7),  105489,  111142-44;  Pano  Ayiictle,  8  km 
NE  Gomez  Farias,  UMMZ  98982. 

Sample  2.  MfiXICO:  San  Luis  Potosi: 
Axtla,  AMNH  67341;  Hiiichihuayan,  USNM 
133859;  Tamazimchale,  AMNH  66065-7, 
FMNH  38611,  UIMNH  27360,  51207;  3.2  km 
NE    Tamazunchale,    TCWC    4090-92,    UMMZ 

119819,  UIMNH  16804;  Xilitla  Region,  KU 
24056-7. 

Sample  3.  MEXICO:  Veracruz:  Tierra 
Colorado,  FMNH  75793,  126933-41,  MCZ 
56004. 

Sample  4.  MfiXICO:  Veracruz:  Cuaut- 
lapam,  KU  105823,  UMMZ  41423-32,  41434- 
42,  41444-51,  41453-54. 

Sample  5.  MfiXICO:  Veracruz:  Cate- 
maco,  UMMZ  118727;  0.8  km  W  Laguna  de 
Catemaco,  UMMZ  126412;  4  km  NE  Cate- 
maco,  UMMZ  126409;  Goyame,  UIMNH 
36877,  39217-18,  UMMZ  111456,  114794;  1 
km  S  Dos  Amates,  between  Sontecomapan 
and  Catemaco,  UMMZ  126413;  Laguna  En- 
cantada,  3  km  NE  San  Andres  Tuxtla,  UMMZ 

119820,  126407;  Rancho  El  Tular,  ca  8  km  N 
San  Andres  Tuxtla,  UIMNH  39212;  San  An- 
dres Tuxtla,  USNM  46899-904,  UIMNH 
24691,  27347,  28054,  28056;  nr  San  Andres 
Tuxtla,  FMNH  126709;  2  km  NE  San  Andres 
Tuxtla,  UMMZ  121156;  ca  4  km  N  San  An- 
dres Tuxtla,  UIMNH  39212;  San  Martin 
Mountains,  UIMNH  35460;  Sontecomapan, 
UMMZ  114795,  126408. 

Sample  6.  MfiXICO:  Veracruz:  Hacienda 
La  Oaxaquena,  AMNH  62331-33;  20  km  S 
Jesus  Carranza,  KU  23953,  24186;  25  km  SE 
Jesus  Carranza,  KU  26956-57,  26959,  26962- 
63,  26965-71;  35  km  SE  Jesus  Carranza,  KU 
23950;  20  km  ENE  Jesus  Carranza,  KU  26955, 
26961. 

Sample  7.  MfiXICO:  Tabasco:  Frontera, 
USNM  25091,  46659,  47453;  16  km  SW  Fron- 
tera, KU  95669-78. 

Sample  8.  MfiXICO:  Campeche:  Bal- 
chacaj,  FMNH  106716-18,  106721,  UIMNH 
26167-70;  Tres  Brazos,  FMNH  106719, 
UIMNH  26172-75,  UMMZ  81924. 

Sample  9.  MfiXICO:  Campeche:  Dzil- 
balchen,  KU  75589-98. 

Sample  10.  MfiXICO:  Yucatan:  Piste, 
KU  70598-600,  70604-5,  70609,  70611-15, 
70617,  70620-26,  70628-29,  70631-32,  70634- 
35,  70637-41. 

Sample  11.  BRITISH  HONDURAS: 
Cayo:  Central  Farm,  MCZ  71608-9;  El  Cayo, 
UMMZ  75012;  1.6  km  NW  El  Cayo,  USNM 
71372-73;  Xunantimich  Ruins,  MCZ  71619-21. 

Sample    12.      GUATEMALA:      El    Peten: 


Tikal,  UMMZ  117875(8),  117876(7);  Uaxac- 
tim,  AMNH  68507-16,  70938-42,  UMMZ 
70415-19. 

Sample  13.  GUATEMALA:  El  Peten: 
Piedras  Negras,  MCZ  66965,  UIMNH  11363- 
93. 

Sample  14.  GUATEMALA:  Alta  Verapaz: 
Canihor,  UMMZ  91305(21),  91306(3),  91307 
(4),  91308;  Finales,  UMMZ  91316. 

Sample  15.  GUATEMALA:  Progreso:  El 
Rancho,  UMMZ  106994(11),  106995(6); 
Finca  Bucural,  UMMZ  107000(9),  107001(4). 

Sample  16.  HONDURAS:  Copdn:  Go- 
pan,  UMMZ  83035(16). 

Sample  17.  HONDURAS:  Atlantkla:  1 
km  W  La  Ceiba,  KU  101227;  2  km  SE  La 
Ceiba  on  Rio  Cangrejal,  KU  101228-38;  8  km 
SE  La  Ceiba  on  Rio  Cangrejal,  KU  101239-42; 
12  km  SSE  La  Ceiba  on  Rio  Cangrejal,  KU 
101252;  ca  15  km  E  La  Ceiba  in  mountains 
above  Corozal,  JRM  2318,  2441-42,  2460-61, 
2470. 

Sample  18.  HONDURAS:  Yoro:  0.5  km 
N  Coyoles,  JRM  2004-7;  2  km  S  Coyoles 
along  Rio  Aguan,  KU  101216-26,  101254-59, 
107911-14,  109973-74;  5  km  E  Coyoles,  JRM 
2082-89;  25  km  WSW  Coyoles,  Rancho  San 
Lorenzo,  JRM  2116. 

Sample  19.  HONDURAS:  Colon:  0.5  km 
SW  Trujillo,  JRM  2554;  1  km  SSW  Trujillo, 
KU  101249-51;  0.5-1.5  km  W  Trujillo,  JRM 
2511;  1-3  km  W  Trujillo,  KU  101243-47;  2 
km  E  Trujillo,  jRM  2533-36;  2  km  W  Tru- 
jillo, KU  101248. 

Sample  20.  HONDURAS:  Olancho:  0.5- 
1.0  km  WNW  Catacamas,  JRM  1433-37;  1 
km  NW  Catacamas,  JRM  1551;  1.5  km  NW 
Catacamas,  JRM  1554-55;  2-3  km  NW  Cata- 
camas, JRM  1526-29;  4.5  km  SE  Catacamas, 
Esquela  Agricola  Nacional  de  Agricultura, 
JRM  1586-87,  1622,  1628-30;  6.5  km  SE  Cata- 
camas, JRM  1638-39. 

Sample  21.  NICARAGUA:  Zelat/a:  Isla 
del  Maiz  Grande,  AMNH  97045-67,  97637, 
MCZ  26970,  26972,  26974,  USNM  94053, 
KU  85988,  85997,  85999,  101268,  101273-74, 
101276-77,  101281-86,  101292,  101299-301, 
101305-6,  101309-10. 

Sample  22.  MfiXICO:  Colima:  Hacienda 
Paso  del  Rio,  FMNH  75794,  126436,  126438- 
41,  126443-47,  126449-51,  167438,  UMMZ 
80118(2),  80119;  Manzanillo,  FMNH  100054; 
1.6  km  W  Pascuales,  UMMZ  80113(3);  0.4 
km  E  Pascuales,  UMMZ  80114;  6  km  SW 
Tecuman,  UMMZ  80110;  8  km  SW  Tecuman, 
UMMZ  80111(3),  80112(5),  80120. 

Sample  23.  MfiXICO:  Colima:  Colima, 
AMNH  5185,  15448-49;  NW  of  Colima, 
AMNH  15838-42;  0.8  km  SW  Colima,  AMNH 
12634-35;  2.4  km  SW  Colima,  AMNH  12643; 
6.4  km  SW  Colima,  KU  29529;  near  Salva- 
dor, MCZ  52172. 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


81 


Sample  24.  MfiXICO:  Guerrero:  Mtns 
N  of  Acapulco,  FMNH  126351,  126355, 
UIMNH  27344;  8  km  E  Coyuca,  USNM 
133849-53;  Between  km  431-432,  camp  nr  El 
Triente,  FMNH  126353;  2  km  N  El  Triente, 
UMMZ  119564;  1.6  km  N  Organos,  S  of  El 
Triente,  UIMNH  27345;  E  San  Andreas  de  la 
Cruz,  KU  87417. 

Sample  25.  MEXICO:  Guerrero:  Aca- 
huizotla,  TCWC  7780,  9587-88,  11320, 
UMMZ  119563;  Agua  del  Obispo,  UMMZ 
119565,  TCWC  7575,  KU  87413-16;  nr  Agua 
del  Obispo  between  Rincon  and  Cajones, 
FMNH  126352,  126354,  MCZ  56005;  Mtns 
near  Agua  del  OlMspo,  TCWC  7576;  8  km  SW 
Chilpancingo,  TCWC  9590;  1.6  km  SW  Cot- 
otlipa,  TCWC  9591. 

Sample  26.  MEXICO:  Oaxaca:  6.4  km  S 
Candelaria,  KU  38234;  Chacalapa,  KU  38230- 
33;  3.2  km  W  El  Soledad,  UIMNH  8421-27. 

San^ple  27.  MEXICO:  Oaxaca:  Tres 
Cruces,  32  km  SW  Tehuantepec,  UMMZ 
81897(2),  81898(3),  81899(2),  81900(3), 
81901(5),  81902(4),  81903(5),  81904(6). 

Sample  28.  MEXICO:  Oaxaca:  3.2  km 
E  Tollocito,  KU  39722-32,  39734,  44656-57. 

Sample  29.  MEXICO:  Oaxaca:  Tapa- 
natepec,  AMNH  80002-3,  MCZ  27283-300; 
2.4  km  S,  7.2  km  E  Tapanatepec,  TCWC 
17047-48;  4  km  W  Tapanatepec,  KU  59577- 
80;  Rio  Novillero,  4.3  km  W  Tapanatepec, 
UIMNH  39204-8. 

Sample  30.  MEXICO:  Chiapas:  Finca 
San  Bartola,  ca  19  km  SW  Cintalapa,  UIMNH 
8437-41,  39229-41;  nr  San  Ricardo,  FMNH 
106715,  UIMNH  26157. 

Sample  31.  MEXICO:  Chiapas:  Abel- 
lanal,  2  km  SE  Florida,  TCWC  19764-65; 
Florida,  50  km  E  Altimirano,  TCWC  19758- 
63,  19766-86;  Las  Tazas,  51  km  E  Altimirano, 
TCWC  19757. 

Sample  33.  MEXICO:  Chiapas:  Esquint- 
la,  LACM  9527-28;  Distrito  Soconusco,  6  km 
NE  Esquintla,  UMMZ  86862-71,  86873-88. 

Sample  34.  MEXICO:  Chiapas:  Sabana 
de  San  Quintin,  KU  94109-17,  94119. 

Sample  35.  GUATEMALA:  Solold:  Fin- 
ca Santa  Buenaventura,  1  km  NW  Panajachel, 
UMMZ  98215(2);  Panajachel,  MCZ  27515-20, 
KU  59581,  95653,  UMMZ  98201(3),  98214 
(3);  1  km  E  Panajachel,  KU  95657;  3  km  E 
Panajachel,  KU  95654-56;  3  km  NE  Pana- 
jachel, UMMZ  98202;  1  km  NW  Panajachel, 
UMMZ  98203(2),  98204;  2  km  NE  Pana- 
jachel, UMMZ  98212;  4  km  NE  Panajachel, 
UMMZ  98213;  Solola,  UMMZ  98205(2);  On 
road  to  Solola,  1  km  from  Panajachel,  UMMZ 
98206. 

Sample  36.  GUATEMALA:  Jutiapa:  Fin- 
ca La  Trinidad,  UMMZ  107421,  107422(8), 
107423(5),  107424(3),  107425(6),  107427 
(3),  107428,  107429(3). 

Sample  37.    GUATEMALA:    Jutiapa:    Ha- 


cienda Mongoy,  UMMZ  107002(6),  107003, 
107004(3);  jutiapa,  UMMZ  106997(4), 
106998,  106999(16). 

Sample  38.  EL  SALVADOR:  Libertad: 
2  km  SE  Colon,  KU  62608-70;  Laguna  de 
Chanmico,  UMMZ  117486(2);  0.8  km  N,  16 
km  W  La  Libertad,  TCWC  17059;  Rancho 
Belmar,  nr  La  Libertad,  LACM  9370.  San 
Salvador:  Instituto  Tropical,  San  Salvador, 
KU  62066-67;  0.4  km  NW  Instituto  Tropical, 
UMMZ  117487;  0.8  km  NW  Instituto  Tropical, 
UMMZ  117488;  1.6  km  NW  San  Salvador, 
KU  42262. 

Sample  39.  NICARAGUA:  Chinamle^a: 
4  km  N,  2  km  W  Chichigalpa,  KU  86036; 
Chinandega,  MCZ  9540,  9546;  Finca  San 
Isidro,  10  km  S  Chinandega,  KU  86030-31; 
Foothills,  N  Slope  Volcan  San  Cristobal,  KU 
86032-35;  Hacienda  Bellavista,  Volcan  Casita, 
KU  101875-77;  San  Antonio,  KU  86008-26. 

Sample  40.  NICARAGUA:  Matagalpa: 
Guasquali,  UMMZ  116420(3);  Matagalpa, 
UMMZ  116414,  116418(5);  0.8  km  E  Mata- 
galpa, UMMZ  116416;  1.6  km  W  Matagalpa, 
UMMZ  116417,  116419;  2.4  km  E  Matagalpa, 
UMMZ  116415(2). 

Sample  41.  COSTA  RICA:  Guanacasie: 
Hacienda  Coyolar,  4.8  km  N,  4.0  km  W  Li- 
beria, USC(CRE)  8206(15),  8207(2);  Haci- 
enda La  Norma,  5  km  N,  4.5  km  W  Liberia 
on  Rio  Colorado,  USC(CRE)  100(2),  103 
(3),  104,  105(3),  106(8),  107(4),  111,  251. 

Sample  42.  COSTA  RICA:  Guanacaste: 
Canas,  USNM  80894;  1.5  km  E  Canas,  USC 
(CRE)  208-9;  1  km  S  Canas,  USC(CRE) 
201;  7  km  N,  3  km  E  Cai^as,  USC  (CRE)  270- 
77;  Finca  Taboga,  11  km  S  Caiias,  USC 
(CRE)  631,  7166(2);  Rio  Javillo,  2  km  S 
Las  Cafias,  USC  (CRE),  202-3. 

Sample  43.  COSTA  RICA:  Guanacaste: 
On  first  river  N  Santa  Rosa,  KU  40531-36; 
Road  from  Silencio  to  Tilaran,  1.8  km  from 
Tilariin,  USC  (CRE)  6240;  Road  from  Silen- 
cio to  Tilaran,  2.6  km  from  Tilaran,  USC 
(CRE)  6239(2);  Tilaran,  ANSP  24566-67; 
Rio  Santa  Rosa  drainage  1.2  km  by  road  SW 
Tilaran,  USC  (CRE)  7161;  1.6  km  SW  Tila- 
ran, USC(CRE)  6270;  2  km  W  Tilaran,  USC 
(CRE)  732;  4.3  km  by  road  NNE  Tilaran, 
USC(CRE)  8082;  4  km  ESE  Los  Angeles  de 
Tilaran,  KU  40527-28;  5.8  km  from  Tilaran 
on  road  to  Los  Angeles,  USC  (CRE)  6267; 
Troncadero,  Laguna  de  Arenal,  KU  40529-30. 

Sample  44.  HONDURAS:  Francisco- 
Morazdn:  El  Zamorano,  Esquela  Agricola 
Panamericana,  AMNH  70368,  MCZ  49765-67, 
10001-3,  KU  101260-67;  Rio  Yeguare,  AMNH 
70369-79;  Valley  of  Rio  Yeguare  S  of  Teguci- 
galpa, AMNH  70470-73. 

Sample  46.  MEXICO:  Quintana  Roo: 
Isla  Mujeres,  USNM  47568-70,  47651,  KU 
70585-88. 


82 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


Sample  48.  MEXICO:  Michoacdn:  Apat- 
zingan,  FMNH  38977-85. 

Additional  Specimens  Not  Used  In  STP 
Analysis.  BRITISH  HONDURAS:  Belize: 
Aml:!ergris  Cay,  3.2  km  N  San  Pedro,  MCZ 
71611;  Belize,  USNM  51879-80,  58373-78; 
Caijo:  Augustine,  Mountain  Pine  Ridge,  MCZ 
71622-24;  Corozal:  0.3  km  N  Corozal,  MCZ 
71610;  Orange  Walk:  Gallon  Jug,  MCZ 
71612-18;  Stann  Creek:  Mango  Creek,  MCZ 
71627-28;  8  km  SW  Mango  Creek,  MCZ 
71626;  6.4  km  S  Waha  Leaf  Creek,  MCZ 
71625;  Toledo:  1.6  km  W  Svvazey  Bridge, 
Monkey  River,  MCZ  71629.  COSTA  RICA: 
Alajuela:  San  Mateo,  USNM  37491;  "Auto- 
pista-AJajuela,"  UCR  uncatalogued;  Cartage: 
La  Carpintera,  AMNH  16314,  16316;  Guana- 
caste:  Bebedero,  USC(CRE)  7162;  1.6  km 
N  Guayabo  de  Bagaces,  USC(CRE)  7024; 
Playa  Samara,  outer  coast  of  Nicoya  Peninsula, 
USC(CRE)  8252;  Parrita,  La  Julieta,  Finca 
La  Ligia,  USC(CRE)  8255(4),  8260,  8262; 
1.6  km  S  Santa  Cruz,  USC(CRE)  8218;  1.8 
km  N  Santa  Cruz,  USC(CRE)  8217(2); 
Puntarenas:  4.3  km  E  Esparta,  UCR  547;  San 
Jose:  Barrio  Mexico,  San  Jose,  KU  40526. 
EL  SALVADOR:  La  Paz:  3  km  E  San  Ra- 
fael Obrajuela,  KU  62065;  San  Miguel:  San 
Pedro,  MCZ  57081.  GUATEMALA:  Guate- 
mala: El  Rosario,  FMNH  68708-10;  Izabal: 
1.6  km  NE  York,  KU  59574-75;  San  Marcos: 
Finca  Carolina,  MCZ  22144-48,  27301-5;  San- 
to Rosa:  Finca  Las  Vinas,  UMMZ  107073-74; 
Departamento  Unknotvn:  MCZ  5831,  99840- 
41.  HONDURAS:  Colon:  Belfate,  AMNH 
58607-9;  Comaijagua:  La  Libertad,  MCZ 
38923;  Copdn:  Hugito,  ANSP  22195-98;  Cor- 
tes: E  side  Lago  de  Yojoa,  KU  67332;  Rancho 
Agua  Azul,  Lago  de  Yojoa,  MCZ  49964;  3.2 
km  W  San  Pedro  Sula,  MCZ  29387-88;  El 
Paraiso:  Arenal,  25  km  E  Jalapa  ( Nicaragua ) , 
UCLA  14759;  Valle  de  Jamastran,  AMNH 
70333-35;  Francisco-Morazdn:  Cantarranas. 
ANSP  22199-212;  Gracias  a  Dios:  Tancin,  ca 
15  km  NW  Puerto  Lempira,  JRM  1732,  1752. 
MEXICO:  Campeche:  3.2  km  E  Barro  San 
Pedro  Y  Pablo,  AMNH  88877;  Champoton, 
UMMZ  72956-57;  Chiapas:    3.2  km  E  El  Real, 


KU  43663-64;  Palenque,  MCZ  66966;  Across 
Rio  Usimiacinta  from  Piedras  Negras  (Guate- 
mala), USNM  108600;  San  Jnanita,  Palenque, 
USNM  108601;  Colima:  Queseria,  UMMZ 
80109;  nr  Queseria,  FMNH  126437,  126442, 
126448;  Guerrero:  nr  Rincon,  FMNH  100080; 
Oaxaca:  La  Concepcion,  nr  Tehuantepec, 
AMNH  66932;  Cosolapa,  USNM  133920;  Ju- 
quila  Mixes,  AMNH  89761-65;  Mitla,  AMNH 
91023-24;  9.6  km  N  Putla,  UIMNH  .52952-53; 
Rio  Cacahuatepec,  UIMNH  529,50;  Rio  Canoa, 
26  km  W  Pinotepec  Nacional,  UIMNH  .52951; 
Sta.  Lucia,  Tehuantepec,  AMNH  65077-78, 
65085;  Tehuantepec,  USNM  30163;  Zacatepec, 
UIMNH  52948-49;  Puehla:  Necapa,  Rio  Ne- 
capa,  AMNH  76441-43;  nr  Vegas  de  Xuchil, 
AMNH  58217;  Quintana  Roo:  8  km  inland 
from  Vigia,  Ascension  Bay,  UMMZ  78586; 
San  Luis  Potosi:  Ciudad  Maiz,  FMNH  126942; 
nr  Cuidad  Maiz,  FMNH  126946;  Palictla, 
AMNH  67358;  8  km  S  Valles,  FMNH  126947; 
Tabasco:  La  Venta,  USNM  117350;  Tamau- 
lipas-:  S  of  Antiguo  Morelos,  FMNH  126944- 
45;  Hacienda  La  Clementina,  FMNH  126943, 
126948,  MCZ  56006,  USNM  106141-42; 
Rancho  Santa  Ana,  12.8  km  E  Padilla,  MCZ 
43634;  Rio  Guayalejo,  nr  Magiscatzin,  MCZ 
45557,  UMMZ  88232;  11  km  W  Victoria, 
FMNH  100050;  Veracruz:  Jalapa,  FMNH 
126706;  Orizaba,  USNM  133839;  Panuco, 
MCZ  19255-59;  4  km  E  Papantla,  KU  24189, 
24194;  San  Andres  Tuxtla,  FMNH  100020;  16 
km  WNW  Temapache,  KU  61791-92;  Yuca- 
tan: Chichen  Itza,  MCZ  52171;  Culebra  Cay, 
Ascencion  Bay,  UMMZ  78587;  Progreso, 
FMNH  100030;  Estado  Unktwwn:  "Mexico," 
( Restricted  to  Tehuantepec,  Oaxaca,  by  Smith, 
1940),  ZMB  868.  NICARAGUA:  Esteli: 
Finca  Daraili,  5  km  N,  14  km  E  Condega,  KU 
85984;  Madriz:  1.6  km  SE  Yalaguina,  TCWC 
17051;  Managua:  12  km  E  Managua,  UCLA 
14787;  6.4  km  E  San  Lorenzo,  KU  42249; 
Rio  San  Juan:  Tuli  Creek,  AMNH  16675; 
Rivas:  within  3  km  of  Moyogalpa,  Isla  Ome- 
tepe,  KU  86029;  Zelaija:  Huaunta  Haulover, 
ANSP  15438-41;  Departamento  Unknown: 
ANSP  9133-34. 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


83 


LITERATURE  CITED 


Alvarez  Del  Toro,  M. 

1960.  Los  Reptiles  De  Chiapas.  Institute 
Zoologico  del  Estado,  Tuxtla  Gutier- 
rez, Chiapas,  Mexico,  204  pp. 

Barbour,  T.  and  Loveridge,  A. 

1929a.  Vertebrates  from  the  Corn  Islands. 
Reptiles  and  Amphibians.  Bull.  Mus. 
Comp.  Zool.,  69:138-146. 

1929b.  Typical      reptiles      and      amphibians. 

Bull.  Mus.  Comp.  Zool.,  69:205-360. 
Barbour,  T.  and  Noble,  G.  K. 

1915.    A  revision  of  the  lizards  of  the  genus 

Ameiva.      Bull.     Mus.     Comp.     Zool., 

59:417-479. 

BaSKIN,    J.    N.    and   WlLLL«iMS,   E.    E. 

1966.  The  Lesser  Antillean  Ameiva  (Sau- 
ria,  Teiidae).  Re-evaluation,  Zoo- 
geography and  the  Effects  of  Preda- 
tion.  Studies  Fauna  Curacao  and 
other  Carib.  Isls.,  23:144-176. 

Bennett,  C.  F. 

1968.    Human    influences    on    the    zoogeog- 
raphy   of    Panama.     Ibero-Americana, 
Univ.  California  Press,  No.  51,  vii  + 
112  pp. 
Boulenger,  G.  a. 

1885.    Catalogue  of  the  lizards  in  the  British 
Museum    (Natural    History),    vol.    II. 
London,  xiii  -f  497  pp. 
Brennan,  J-  M-  AND  Yunker,  C.  E. 

1966.  The  chiggers  of  Panama  (Acarina: 
Trombiculidae ) .  In:  Ectoparasites 
of  Panama  ( R.  L.  Wenzel  and  V.  J. 
Tipton,  Eds.).  Field  Mus.  Nat. 
Hist.,  Cliicago.    pp.  221-266. 

Burt,  C.  E. 

1931.  A  study  of  the  teiid  lizards  of  the 
genus  Cnemidophorus  with  special 
reference  to  their  phylogenetic  rela- 
tionships. Bull.  U.  S.  Natl.  Mus., 
154:1-286. 
Cochran,  D.  M. 

1946.    Notes  on  the  herpetology  of  the  Pearl 
Islands,    Panama.     Smithsonian    Misc. 
Coll.,  106:1-8. 
Cope,  E.  D. 

1875.  On  the  Batrachia  and  Reptilia  of 
Costa  Rica.  ].  Acad.  Nat.  Sci.,  Phila- 
delphia, ser.  2,  8:93-157. 

1894.    Third   addition    to   the   knowledge   of 
the   Batrachia   and   Reptilia   of  Costa 
Rica.     Proc.    Acad.    Nat.    Sci.    Phila- 
delphia,  1894:194-206. 
Donoso-Barros,  R. 

1968.  The  lizards  of  Venezuela  (check  list 
and  key).  Carribean  J.  Sci.,  8:105- 
122. 


DUELLMAN,    W.    E. 

1958.    A  monographic  study  of  the  colubrid 

snake  genus  Leptodcira.    Bull.  Amer- 
ican Mus.  Nat.  Hist.,  114:1-152.    pis. 

1-31. 
1961.    The      amphibians      and      reptiles      of 

Michoacan,     Mexico.      Univ.     Kansas 

Publ.  Mus.  Nat.  Hist.,   15:1-148. 
1965a.  Amphibians     and     reptiles    from     the 

Yucatan     Peninsula,     Mexico.      Univ. 

Kansas    Publ.    Mus.    Nat.    Hist.,    15: 

577-614. 
1965b.  A     biogeographic     account     of     the 

herpetofauna   of    Michoacan,    Mexico. 

Univ.   Kansas   Publ.   Mus.   Nat.   Hist., 

15:627-709. 
1966.    The  Central  American  herpetofauna: 

An    ecological    perspective.      Copeia, 

1966:700-719. 
Dunn,  E.  R. 

1931.    The     herpetological     fauna     of     the 

Americas.    Copeia,    1931:106-119. 
1940a.  New    and    noteworthy    herpetological 

material    from    Panama.     Proc.    Acad. 

Nat.  Sci.  Philadelphia,  92:105-122. 
1940b.  Some  aspects  of  heipetology  in  lower 

Central    America.     Trans.    New    York 

Acad.  Sci.,  2:156-158. 
1943.    A  new  race  of  Arneiva  festiva  from 

Colombia.     Notulae    Naturae    of    the 

Acad.  Nat.  Sci.  Pliiladelphia,  no.  126, 

pp.  1-2. 

EcHTERNACHT,    A.    C. 

1968.  Distributional  and  ecological  notes  on 
some  reptiles  from  northern  Hon- 
duras.    Herpetologica,    24:151-158. 

1970.    Taxonomic    and    ecological    notes    on 
some    Middle    and    South    American 
lizards    of    the    genus    Ameiva    (Teii- 
dae).   Breviora,  354:1-9. 
EsTES,   R. 

1969.  Relationships  of  t\vo  Cretaceous  liz- 
ards (Sauria,  Teiidae).  Breviora,  no. 
317:1-8. 

Gabriel,  K.  R. 

1964.  A  procedure  for  testing  the  homoge- 
neity of  all  sets  of  means  in  analysis 
of  variance.    Biometrica,   20:459-477. 

Gabriel,  K.  R.  and  Sokal,  R.  R. 

1969.  A  new  statistical  approach  to  geo- 
graphic variation  analysis.  Syst. 
Zool.,  18:259-278. 

Gorman,  G.  C. 

1970.  Chromosomes  and  the  systematics  of 
the  family  Teiidae  (Sauria,  Reptilia). 
Copeia,   1970:230-245. 

Haffer,  J. 

1967.  Speciation  in  Colombian  forest  birds 
west  of  the  Andes.  Amer.  Mus. 
Novitates,  2294:1-57. 


84 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


1970.  Geologic  climatic  history  and  zoo- 
geographic  significance  of  the  Uraba 
region  of  northwestern  Colombia. 
Caldasia,  10:603-636. 

Hallowell,  E. 

1860.  Report  upon  the  Reptilia  of  the  north 
Pacific  exploring  expedition,  under 
command  of  Capt.  ]ohn  Rogers, 
U.S.N.  Proc.  Acad.  Nat.  Sci.  Phila- 
delphia, 1860:480-510. 

Heatwole,  H. 

1966.  The  eff^ect  of  man  on  distribution  of 
some  reptiles  and  amphibians  in 
eastern  Panama.  Herpetologica,  22: 
55-59. 

Heatwole,  H.  and  Torres,  F. 

1967.  Distribution  and  geographic  variation 
of  the  Ameivas  of  Puerto  Rico  and 
the  Virgin  Islands.  Studies  Fauna 
Curagao  and  other  Carib.  Isls.,  24: 
63-111. 

HiLLMAN,   P.   E. 

1969.  Habitat  specificity  in  three  sympatric 
species  of  Ameiva  (Reptilia:  Teii- 
dae).    Ecology,  50:476-481. 

HiRTH,    H.    F. 

1963.  The  ecology  of  two  lizards  on  a 
tropical  beach.  Ecol.  Monogr.,  33: 
83-112. 

1965.  Temperature  preferences  of  five 
species  of  Neotropical  lizards.  Her- 
petologica, 20:273-276. 

LiCHTENSTEIN,    H.    AND    VoX    MaRTENS,    E. 

1856.    Nomenclator    Reptilium    et    Amphibi- 
orum     Musei     Zoologici     Berolinensis 
.  .  .  Berlin,  Konigl.  Akad.  Wissensch., 
iv  -f  48  pp. 
Lloyd,  J.  J. 

1963.  Tectonic  history  of  the  south  Central 
American  orogen.  Mem.  American 
Assoc.  Petrol.  Geol.  2:88-100. 

Maldoxado-Koerdell,  M. 

1964.  Geohistory  and  paleogeography  of 
Middle  America.  In:  Handbook  of 
Middle  American  Indians,  vol.  1  ( R. 
Wauchope  and  R.  C.  West,  Eds.). 
Univ.  Texas  Press,  Austin,  Texas, 
pp.  3-32. 

Maslin,  T.  p. 

1963.  Notes  on  a  collection  of  herpetozoa 
from  the  Yucatan  Peninsula  of  Mex- 
ico. Univ.  Colorado  Studies,  Ser. 
Biol.,  no.  9,  pp.   1-20. 

Mayh,  E. 

1963.  Animal  Species  and  Evolution.  Bel- 
knap Press,  Cambridge,  Massachu- 
setts,   xiv  -j-  797  pp. 

MiJLLER,    V.    L. 

1929.  Uber  einige  Rassen  der  Ameiva 
ameiva  aus  Venezuela.  Zool.  Anz., 
83:97-112,  193-211. 


Neill,  W.  T. 

1965.  New  and  noteworthy  amphibians  and 
reptiles  from  British  Honduras.  Bull. 
Florida  State  Mus.,  9:77-130. 

Neill,  W.  T.  axd  Allex,  R. 

1961.  Further  studies  on  the  herpetology 
of  British  Honduras.  Herpetologica, 
17:37-52. 

Peters,  J.  A. 

1964.  The  lizard  genus  Ameiva  in  Ecuador. 
Bull.  So.  California  Acad.  Sci.,  63: 
113-127. 

Peters,  J.  A.,  Burt,  W.  H.,  Sibley,  C.  C, 
Bogert,  C.  M.,  Hubbell,  T.  H., 
Clexch,  W.  J.,  AND  Rogers,  J.  S. 

1954.    Symposium:     Subspecies    and    CUnes. 
Syst.  Zool.,  3:97-125. 
Power,  D.  M. 

1970.    Geographic    variation    of    red- winged 
blackbirds  in  central  North  America. 
Univ.   Kansas   Publ.   Mus.   Nat.   Hist., 
19:1-83. 
Raxd,  a.  S. 

1957.  Notes  on  amphibians  and  reptiles 
from  El  Salvador.  Fieldiana:  Zool- 
ogy, 34:505-534. 

Savage,  J.  M. 

1960.  Evolution  of  a  peninsular  herpeto- 
fauna.    Syst.  Zool.,  9:184-212. 

1966.  The  origins  and  history  of  the  Cen- 
tral American  herpetofauna.  Copeia, 
1966:719-766. 

Savage,  J.  M.  and  Heyer,  W.  R. 

1967.  Variation  and  distribution  in  the 
tree-frog  genus  PhijUomedusa.  Bei- 
trage  zur  Neotropischen  Fauna,  5: 
111-131. 

Schmidt,  K.  P. 

1943.  Corollary  and  commentary  for  "Cli- 
mate and  Evolution."  Amer.  Midi. 
Nat.,  30:241-253. 

Schmidt,  K.  P.  and  Stuart,  L.  C. 

1941.  The  herpetological  fauna  of  the  Sa- 
lama  Basin,  Baja  Verapaz,  Guate- 
mala. Zool.  Ser.  Field  Mus.  Nat. 
Hist.,  24:233-247. 

Schneider,  C.  R. 

1965.  Besnoitia  panaviensis,  sp.  n.  (Proto- 
zoa: Toxoplasmatidae )  from  Pana- 
manian lizards.  J.  Parasit.,  51:340- 
344. 

Schuchert,  C. 

1935.    Historical    geology    of    the    Antillean- 
Caribbean    region.     John    Wiley    and 
Sons,  New  York,  xxvi   +  811  pp. 
Schwartz,  A. 

1965.  The  Ameiva  (ReptiHa:  Teiidae)  of 
Hispaniola  I.  Ameiva  lineolata  Du- 
meril  and  Bibron.  Carib.  J.  Sci.,  5: 
45-57. 


ECHTERNACHT:    MIDDLE  AMERICAN  LIZARDS 


85 


1967.  The  Aineiva  (Lacertilia,  Teiidae)  of 
Hispaniola.  III.  Ameiva  iaenixira 
Cope.  Bull.  Mus.  Comp.  Zool.,  135: 
345-375. 

1968.  Two  new  subspecies  of  Ameiva 
( Lacertilia,  Teiidae )  from  Hispan- 
iola.   Herpetologica,  24:21-28. 

Schwartz,  A.  and  Klinikowski,  R.  F. 

1966.  The  Ameiva  (LacertiUa,  Teiidae)  of 
Hispaniola.  II.  Geographic  variation 
in  Ameiva  chnjsolaema  Cope.  Bull. 
Mus.  Comp.  Zool.,  133:425-487. 

Schwartz,  A.  and  McCoy,  C.  J. 

1970.  A  systematic  review  of  Ameiva 
aiiheri  Cocteau  (Reptilia,  Teiidae) 
in  Cuba  and  the  Bahamas.  An. 
Carnegie   Mus.,  41(4)  :45-168. 

Smith,  H.  M. 

1940.    Descriptions      of     new     lizards      and 

snakes   from   Mexico   and   Guatemala. 

Proc.    Biol.    Soc.    Washington,   53:55- 

64. 
1946.    Handbook      of      Lizards.       Comstock 

Publ.   Co.,   Ithaca,   New  York,   xxi   + 

557  pp. 

1967.  The  perspective  of  subspecies  in  ani- 
mal evolution.  The  Biologist,  49:43- 
51. 

Smith,  H.  M.  and  Laufe,  L.  E. 

1946.    A  summary  of  Mexican  lizards  of  the 
genus     Ameiva.      Univ.     Kansas     Sci. 
Bull.,  31:7-73. 
Smith,  H.  M.  and  White,  F.  N. 

1956.  A  case  for  the  trinomen.  Syst.  Zool., 
5:183-190. 

Smith,  R.  E. 

1968a.  Experimental  evidence  for  a  gonadal- 
fat  body  relationship  in  two  teiid 
lizards  (Ameiva  festiva,  Ameiva 
qiiadrilineata).  Biol.  Bull.,  134:325- 
331. 

1968b.  Studies  on  reproduction  in  Costa 
Rican  Ameiva  festiva  and  Ameiva 
qiiadrilineata  (Sauria:  Teiidae).  Co- 
peia,   1968:236-239. 

Stuart,  L.  C. 

1935.  A  contribution  to  a  knowledge  of 
the  herpetology  of  a  portion  of  the 
savanna  region  of  central  Peten, 
Guatemala.  Misc.  Publ.  Mus.  Zool. 
Univ.   Michigan,  29:1-56,  4  Pis. 

1942.  Comments  on  the  imdidata  group  of 
Ameiva  (Sauria).  Proc.  Biol.  Soc. 
Washington,  55:143-150. 

1943.  Comments  on  the  herpetofauna  of 
the  Sierra  de  los  Cuchumatanes  of 
Guatemala.  Occas.  Papers  Mus. 
Zool.  Univ.  Michigan,  471:1-28,  1  Pi. 

1948.  The  amphibians  and  reptiles  of  Alta 
Verapaz,  Guatemala.  Misc.  Publ. 
Mus.  Zool.  Univ.  Michigan,  69:1-109. 


1950.  A  geographic  study  of  the  herpeto- 
fauna of  Alta  Verapaz,  Guatemala. 
Contrib.  Lab.  Vert.  Biol.  Univ. 
Michigan,  45:1-77,  9  Pis. 

1951.  The  herpetofauna  of  the  Guatemalan 
Plateau,  with  special  reference  to  its 
distribution  on  the  southwestern 
highlands.  Contrib.  Lab.  Vert.  Biol. 
Univ.  Michigan,  49:1-71,  7  Pis. 

1954a.  A  description  of  a  subhumid  corridor 
across  northern  Central  America, 
with  comments  on  its  herpetofaunal 
indicators.  Contrib.  Lab.  Vert.  Biol. 
Univ.  Michigan,  65:1-26. 

1954b.  Herpetofauna  of  the  southeastern 
highlands  of  Guatemala.  Contrib. 
Lab.  Vert.  Biol.  Univ.  Michigan,  68: 
1-65,  4  Pis. 

1955.  A  brief  review  of  the  Guatemalan 
lizards  of  the  genus  Anolis.  Misc. 
Publ.  Mus.  Zool.  Univ.  Michigan,  91: 
1-31. 

1957.  Herpetofaimal  dispersal  routes 
through  northern  Central  America. 
Copeia,  1957:89-94. 

1958.  A  study  of  the  herpetofauna  of  the 
Uaxactun-Tikal  area  of  northern 
El  Peten,  Guatemala.  Contrib.  Lab. 
Vert.   Biol.   Univ.   Michigan,   75:1-30. 

1964.  Fauna  of  Middle  America.  In: 
Handbook  of  Middle  American  In- 
dians, vol.  1  ( R.  Wauchope  and  R. 
C.  West,  Eds.).  Univ.  Texas  Press, 
Austin,  Texas,    pp.  316-362. 

1966.    The     environment     of     the     Central 
American       cold-blooded      vertebrate 
fauna.    Copeia,  1966:684-699. 
Taylor,  E.  H. 

1956.  A  review  of  the  lizards  of  Costa  Rica. 
Univ.  Kansas  Sci.  Bull.,  38:1-322. 

Terry,  R.  A. 

1956.  A  geological  reconnaissance  of  Pan- 
ama. Occas.  Papers  California  Acad. 
Sci.,  No.  23.    91  pp. 

Vanzolini,  p.  E.  and  Valencia,  J. 

1965.  The  genus  Dracaena,  with  a  brief 
consideration  of  macroteiid  relation- 
ships (Sauria,  Teiidae).  Arq.  Zool., 
13:7-45. 

Villa,  J.  D. 

1968.    A  new  colubrid  snake  from  the  Corn 
Islands,  Nicaragua.    Rev.  Biol.  Trop., 
15:117-121. 
Vinson,  G.  L.  and  Brineman,  J.  H. 

1963.    Nuclear  Central  America,  hub  of  the 
Antillean      Transverse      Belt.       Mem. 
American  As.soc.  Petrol.  Geol.   2:101- 
112. 
Wilson,  E.  O.  and  Brown,  W.  L. 

1953.  The  subspecies  concept  and  its  taxo- 
nomic  application.  Syst.  Zool,  2:97- 
111. 


86 


MISCELLANEOUS  PUBLICATION  MUSEUM  OF  NATURAL  HISTORY 


YUNKER,   C.    E.    AND   RaDOVSKY,    F.    J. 

1966.  The  Dermanyssid  mites  of  Panama. 
In:  Ectoparasites  of  Panama  (R.  L. 
Wenzel  and  V.  J.  Tipton,  Eds.). 
Field  Mus.  Nat.  Hist.,  Chicago,  pp. 
83-103. 

ZWEIFEL,    R.    G. 

1959.    Variation    in    and   distribution   of   liz- 


ards of  western  Mexico  related  to 
CnemidopJiorus  sacki.  Bull.  Amer- 
ican Mus.  Nat.  Hist.,  117:57-116. 

The  manuscript  for  this  publication  was 
completed  in  March,  1970,  and  submitted  as  a 
dissertation  in  partial  fulfillment  of  the  re- 
quirements for  the  PhD  at  the  University  of 
Kansas. 


3  2044  093  361    616 


Date  Due 


BOUND 


1972