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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..
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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
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The manuscript for this publication was
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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