1. Herpetologica, 64(1), 2008, 92–108
E 2008 by The Herpetologists’ League, Inc.
TWO NEW SPECIES OF ANOLES FORMERLY REFERRED TO AS
ANOLIS LIMIFRONS (SQUAMATA: POLYCHROTIDAE)
GUNTHER KOHLER1,3
¨ AND JAVIER SUNYER1,2
1
Forschungsinstitut und Naturmuseum Senckenberg, Senckenberganlage 25, 60325 Frankfurt a.M., Germany
2
´ ´ ´n,
Gabinete de Ecologıa y Medio Ambiente, Departamento de Biologıa, UNAN-Leo Nicaragua
ABSTRACT: We describe two new species of anoles (genus Anolis) from Panama formerly referred to as
Anolis (or Norops) limifrons. Both new species differ from A. limifrons by having a large bilobed hemipenis
(small and unilobed in A. limifrons). The new species differ from each other in male dewlap size and
coloration. We provide an identification key and standardized descriptions of A. limifrons and the two new
species described herein.
´ ´
RESUMEN: Describimos dos especies nuevas de anoli (genero Anolis) de Panama anteriormente referidas
como Anolis (o Norops) limifrons. Ambas nuevas especies difieren de A. limifrons en tener un hemipene
˜ ´
grande y bilobulado (pequeno y unilobulado en A. limifrons). Las dos especies nuevas difieren entre sı en el
˜ ´ ´
tamano y coloracion de la papera gular de los machos adultos. Se proporciona una clave dicotomica de
´
identificacion y descripciones estandar de A. limifrons y de las otras dos nuevas especies.
Key words: Anolis; New species; Panama; Polychrotidae; Reptilia; Squamata
ANOLES are a diverse and taxonomically an adult male specimen (now ANSP 7890).
poorly understood group of lizards that are According to Savage (1973:36), the holotype
distributed widely across the tropical and of A. bransfordii was ‘‘collected by Bransford
subtropical portions of North, Central and at Machuco (5 Machuca) on the Rıo San ´
¨
South America (Kohler, 2003; Savage, 2002). ´
Juan, Departamento Rıo San Juan, Nicara-
Several new species have been described in gua.’’ In 1882, Thominot described Anolis
recent years indicating that more field work rivieri based on a juvenile specimen (now
and study of museum material is needed to MNHN 1884.221) from ‘‘Panama.’’ Boulenger
document the real diversity of anoles. The (1885) described Anolis godmani based on
comparative study of hemipenial variation in four specimens from ‘‘Guatemala’’ and three
anoles revealed substantial cryptic diversity ´
specimens from ‘‘Irazu, Costa Rica.’’ Stuart
¨ ¨
(Kohler and Kreutz, 1999; Kohler et al., 2003). (1955:30) presumed that in ‘‘Anolis godmani
In 1862, Cope described the new species Boulenger … [there] may have been a mixup
Anolis (Dracontura) limifrons based on two in locality data in the Godman-Salvin collec-
syntypes (now ANSP 7900–01) from ‘‘Vera- tions, one of the cotypes having been listed as
gua.’’ According to Savage (1970:279), ‘‘today of Guatemala whereas it probably came from
the old Veragua comprises the Provincias of Costa Rica,’’ and he also stated that ‘‘the
´
Veraguas, Chiriquı and Bocas del Toro.’’ material [including the holotype of A. god-
Barbour (1934:139) described the type locality mani] may have been received by the British
of A. limifrons as ‘‘Cucuyos, Veragua Prov., Museum somewhat after the main bulk of the
Panama [an abandoned mine on the Rıo ´ earlier parts of the collection had been turned
Santiago].’’ In 1871, Cope named Anolis over to the Museum.’’ In 1956, Taylor
trochilus based on an adult male specimen described the new species Anolis biscutiger
(now ANSP 7804) from ‘‘San Jose, Costa ´ based on an adult male (KU 40771) from
Rica.’’ Peters (1873) added another nominal ‘‘Golfito, Puntarenas Province, Costa Rica.’’
species, Anolis pulchripes, based on a speci- Dunn (1930) regarded rivieri, trochilus and
´
men (now ZMB 7827) from ‘‘Chiriquı.’’ A year bransfordii as synonyms of limifrons. Barbour
later, Cope (1874) described the new species (1934:140) stated that ‘‘Dunn has seen the
Anolis bransfordii from ‘‘Nicaragua’’ based on types of limifrons, rivieri, pulchripes and
rodriguezii and declares them all the same
3
CORRESPONDENCE: e-mail, gkoehler@senckenberg.de species.’’ The names trochilus, pulchripes,
92

2. March 2008] HERPETOLOGICA 93
bransfordii and rivieri have remained in the in order to document any variation. We
synonymy of A. limifrons Cope whereas god- measured scale sizes using the ocular micro-
mani has been retained as a valid species until meter of a stereo microscope (Leica MZ 12)
recently (Peters and Donoso-Barros, 1970; and rounded the values to the nearest
Savage and Villa, 1986; Taylor, 1956; Villa et 0.01 mm. For all other measurements we used
¨
al., 1988). Savage (2002) and Kohler (2003) precision calipers and rounded the values to
recognized a northern species, rodriguezii, the nearest 0.1 mm. We measured head length
and a southern one, limifrons, with biscutiger from tip of snout to the anterior margin of the
and godmani as synonyms of limifrons. ear opening. We measured snout length from
Here we report the results of our study of tip of snout to the anterior border of the orbit.
the variation in hemipenial and scalation We determined head width as the distance
morphology as well as morphometrics of the between rictuses. We counted dorsal and
small anoles occurring from eastern Honduras ventral scales at midbody along the midline.
to eastern Panama commonly referred to as We measured tail height and width at the point
Anolis (or Norops) limifrons. reached by the heel of the extended hind leg.
We counted subdigital lamellae on phalanges ii
MATERIALS AND METHODS to iv of the 4th toe. We considered the scale
directly anterior to the circumnasal to be a
For this study, we examined 1428 specimens
prenasal. The capitalized colors and color
of Anolis limifrons. We provide a list of the
codes (the latter in parentheses) are those of
comparative specimens examined in the Ap-
Smithe (1975–1981). To measure dewlap area,
pendix. Abbreviations for museum collections
we took photographs of males with their
follow those of Leviton et al., (1985) except
dewlaps artificially extended using small fore-
´
MHCH (Museo Herpetologico de Chiriquı, ´
ceps. The head portion was magnified and
´
David, Panama), and JS field numbers, which
printed, then superimposed on millimetric
refer to specimens that will be deposited in the
paper, and the total number of 1-mm squares
Museo de Ciencias Naturales de la Universidad
contained in the extended dewlap counted. A
Centroamericana (UCA), Managua, Nicara-
straight line was drawn between both the
gua. In the course of this study, G.K. had the
anterior and posterior insertions of the dewlap.
privilege of examining all extant primary types
We also determined the HL on the printout.
of nominal species regarded as synonyms of
We used the following equation to convert the
¨
Anolis limifrons by previous authors (Kohler,
magnified dewlap area to the real size: X 5
2003; Peters and Donoso-Barros, 1970; Savage,
[(!Y / A) ? B]2; where: X is the real area of the
2002). For the synonymy lists, we included only
dewlap in mm2; Y is the total area (mm2) of the
those works that cite actual specimens. No-
dewlap at a magnified scale; A is the HL
menclature of scale characters follows that of
measure (mm) of the anole at a magnified
¨
Kohler (2003). Terminology for hemipenial
scale; and B is the HL measure (mm) of the
morphology follows that of Myers et al. (1993)
anole at the real size. We executed statistical
and Savage (1997). We everted the hemipenes
techniques using the computer program Sta-
in the field after euthanasia and before
tistica version 6.1. Juveniles were not included
preserving the lizards in 70% ethanol by
in the statistical analyses. We used discrimant
manually applying pressure to the ventral base
function analysis to evaluate the phenetic
of tail. Once the hemipenes had been everted
distinctness of a priori groups. Discriminant
to some extent, we injected 70% ethanol in the
scores (DC) were calculated by multiplying
area of the hemipenial pockets, usually
selected variables by their associated canonical
resulting in complete eversion of both hemi-
coefficients. Each specimen was then plotted
penes. With the needle still attached to the
along the axae providing maximal separation of
lizard’s tail, we placed the lizard for 1–2 min
the a priori groups.
into the preservative to allow fixation of the
everted hemipenis. We did not tie off the
hemipenes at the base because this might cause RESULTS
damage to these fragile organs. We intended to Two distinctly different hemipenial mor-
evert the hemipenes in all available adult males photypes are evident in the specimens we

3. 94 HERPETOLOGICA [Vol. 64, No. 1
examined. In Type A (n 5 188 adult males (Fig. 1) that correctly classified 72.7% of the
with everted hemipenes), the hemipenis is a specimens. The first function is DS 5 0.471297
relatively large bilobed organ; both the (number of medial ventral scales in one head
truncus and the lobes have a strongly calycu- length) +0.673627 (number of medial dorsal
late surface. The sulcus spermaticus bifurcates scales in one head length) 20.392336 (sub-
at the base of the apex and the branches digital lamellae) +0.176923 (total number of
continue to the tip of the lobes. In Type B (n loreal scales) +0.286056 (scales between su-
5 85 adult males with everted hemipenes), praorbital semicircles) 20.651674 (scales be-
the hemipenis is much smaller relative to body tween posterior canthals) +0.025958 (scales
size as compared to the Type A hemipenis. around midbody). The second function is DS
Also, it is unilobed and the truncus and lobes 5 0.284551 (number of medial ventral scales in
are not calyculate. The sulcus spermaticus one head length) 20.173192 (number of
opens at the base of the apex. While these two medial dorsal scales in one head length)
hemipenial types show a broadly sympatric 20.610593 (subdigital lamellae) 20.171498
geographical distribution pattern in western (total number of loreal scales) 20.293526
Panama, variation in hemipenial morphology (scales between supraorbital semicircles)
within these discrete types is negligible both +0.054329 (scales between posterior canthals)
within populations and in a geographical 20.603442 (scales around midbody).
context. Within Type A, two distinct types Based on our data, we recognize three
can be distinguished in respect of the relative species of this complex: Species A: Hemipenis
size and the coloration of the male dewlap. unilobed; male dewlap small (smaller than
Males with bilobed hemipenes from the 100 mm2), dull white with a small basal
Province of Bocas del Toro, Panama, and orange blotch; distributed from eastern Hon-
adjacent southeastern Costa Rica have a small duras to central Panama west of the Canal
dewlap (smaller than 100 mm2) that is dull Zone (Fig. 2). Species B: Hemipenis bilobed;
white with a small basal orange blotch. Males male dewlap small (smaller than 100 mm2),
with bilobed hemipenes from central and dull white with a small basal orange blotch;
eastern Panama have a large dewlap (larger distributed in the western and central portions
than 150 mm2) that is almost uniformly of the Province Bocas del Toro, Panama, and
orange. All Type B males have a small dewlap adjacent southeastern Costa Rica (Fig. 2).
(smaller than 100 mm2) that is dull white with Species C: Hemipenis bilobed; male dewlap
a small basal orange blotch. large (larger than 150 mm2), almost uniformly
In external morphology, there is great orange; distributed in central and eastern
overlap in the ranges of all examined charac- Panama; expected in adjacent northwestern
ters of scalation and morphometrics (Table 1). Colombia (Fig. 2). In western Panama (Prov-
We conducted a discriminant function analy- ´
inces of Chiriquı, Bocas del Toro and
sis (DFA) based on seven pholidotic charac- Veraguas), the bilobed form (our Species B)
ters (number of medial ventral scales in one is restricted to the Caribbean versant; whereas
head length; number of medial dorsal scales in the unilobed form (our Species A) occurs on
one head length; subdigital lamellae; total both versants. Although no cases of actual
number of loreal scales; scales between syntopy of these two forms have been
supraorbital semicircles; scales between pos- documented, they show a broadly sympatric
terior canthals; scales around midbody) and distribution pattern on the Caribbean versant
did a priori assignments to groups based on of western Panama.
our hemipenis and male dewlap findings Unfortunately, none of the type specimens
(Group 1 5 Type B hemipenis, male dewlap mentioned in the introduction are males with
small, dull white with a small basal orange their hemipenes everted. Also, data on dewlap
blotch; Group 2 5 Type A hemipenis, male coloration in life is not available for these type
dewlap small, dull white with a small basal specimens. However, the type locality data of
orange blotch; Group 3 5 Type A hemipenis, most of these nominal species allow for a
male dewlap large, more or less uniformly reliable allocation to one of the three species
orange). This DFA yielded a scatter diagram that we distinguish. Thus, on geographic

4. TABLE 1.—Selected measurements, proportions and scale characters of Anolis limifrons, A. apletophallus, and A. cryptolimifrons (specimens with * listed in Appendix). Range is
March 2008]
followed by mean value and one standard deviation in parentheses. Morphomentric data were only taken from adults.
A. limifrons A. apletophallus A. cryptolimifrons
= 50 = 25 = 20
R 21 R 24 R 10
Maximum SVL = 41.5 47.0 45.0
R 43.5 46.5 42.0
Tail length / SVL = 1.53–2.52 (2.21 6 0.19) 1.52–2.29 (2.03 6 0.17) 1.83–2.19 (2.07 6 0.10)
R 1.60–2.30 (2.05 6 0.22) 1.82–2.32 (2.06 6 0.13) 1.87–2.08 (1.98 6 0.08)
Tail diameter vertical / horizontal = 1.00–1.30 (1.16 6 0.07) 1.03–1.27 (1.14 6 0.08) 1.00–1.25 (1.15 6 0.06)
R 1.00–1.27 (1.14 6 0.07) 1.00–1.25 (1.11 6 0.07) 1.00–1.17 (1.21 6 0.06)
Head length / SVL = 0.25–0.28 (0.27 6 0.01) 0.25–0.29 (0.27 6 0.01) 0.25–0.29 (0.27 6 0.01)
R 0.24–0.28 (0.26 6 0.01) 0.25–0.29 (0.27 6 0.01) 0.26–0.28 (0.27 6 0.01)
Head length / head width = 1.44–1.80 (1.67 6 0.07) 1.54–1.84 (1.67 6 0.06) 1.59–1.77 (1.67 6 0.06)
R 1.57–1.77 (1.67 6 0.06) 1.56–1.70 (1.65 6 0.04) 1.62–1.75 (1.66 6 0.04)
Interparietal plate / ear = 0.79–5.20 (2.03 6 0.78) 0.71–4.12 (1.71 6 0.70) 0.54–2.29 (1.47 6 0.46)
R 1.09–4.75 (2.19 6 0.98) 1.01–2.45 (1.69 6 0.44) 0.50–1.68 (1.09 6 0.39)
Shank length / SVL = 0.26–0.33 (0.30 6 0.02) 0.29–0.32 (0.30 6 0.01) 0.26–0.31 (0.29 6 0.02)
R 0.26–0.31 (0.28 6 0.01) 0.26–0.32 (0.30 6 0.02) 0.27–0.29 (0.28 6 0.01)
Axilla–groin distance / SVL = 0.36–0.47 (0.42 6 0.03) 0.34–0.46 (0.39 6 0.03) 0.39–0.46 (0.43 6 0.02)
R 0.41–0.53 (0.45 6 0.02) 0.35–0.48 (0.41 6 0.03) 0.40–0.46 (0.43 6 0.02)
HERPETOLOGICA
Subdigital lamellae of 4th toe 20–27 (23.33 6 1.61) 21–26 (23.59 6 1.12) 23–29 (25.03 6 1.47)
Number of scales between supraorbital semicircles 0–4 (1.78 6 0.84) 1–3 (2.04 6 0.50) 1–3 (2.20 6 0.61)
Number of scales between interparietal plate and supraorbital semicircles 1–5 (2.68 6 0.86) 1–4 (2.72 6 0.68) 2–4 (3.27 6 0.64)
Number of scales between subocular scales and supralabial scales 0 0 0
Number of supralabial scales to level below center of eye 5–8 (6.48 6 0.69) 6–8 (6.76 6 0.56) 5–7 (6.73 6 0.52)
Number of infralabials to level below center of eye 5–8 (6.30 6 0.70) 5–8 (6.37 6 0.57) 6–8 (6.62 6 0.56)
Total number of loreals 24–68 (41.13 6 9.56) 25–74 (45.00 6 8.78) 36–69 (49.07 6 7.67)
Number of horizontal loreal scale rows 5–8 (6.10 6 0.79) 5–8 (6.38 6 0.73) 6–8 (6.70 6 0.70)
Number of postrostrals 5–9 (6.86 6 0.82) 6–8 (7.23 6 0.59) 6–9 (7.17 6 0.53)
Number of postmentals 4–8 (6.45 6 0.91) 6–9 (7.28 6 0.88) 6–8 (7.03 6 0.87)
Number of scales between nasals 7–11 (8.91 6 0.86) 8–11 (9.69 6 0.66) 9–12 (9.77 6 0.77)
Number of scales between 2nd canthals 7–15 (10.59 6 1.69) 9–13 (10.61 6 1.06) 9–16 (12.00 6 1.55)
Number of scales between posterior canthals 10–18 (13.77 6 2.10) 10–18 (13.23 6 1.86) 12–19 (15.21 6 1.76)
Number of medial dorsal scales in one head length 38–70 (54.26 6 5.76) 42–74 (60.65 6 7.10) 46–70 (58.87 6 5.35)
Number of medial ventral scales in one head length 26–58 (40.62 6 6.15) 36–60 (45.78 6 5.44) 34–50 (41.31 6 4.01)
95

5. 96 HERPETOLOGICA [Vol. 64, No. 1
(1873; in part); Dunn (1930; in part);
Barbour (1934; in part); Gaige et al.,
(1937); Brattstrom and Howell (1954);
Taylor (1956; in part); Meyer and Wilson
(1973); Fitch and Seigel (1984); Vences et
al., (1998); Poe (2004; in part).
Anolis limifrons limifrons: Etheridge (1959; in
part).
Anolis pulchripes Peters, 1873:739. Holotype
ZMB 7827 from ‘‘Chiriqui.’’
Anolis trochilus Cope, 1871:215. Holotype
´
ANSP 7804 from ‘‘San Jose, Costa Rica.’’
Cope (1876); Savage (1970).
Norops limifrons: Wilson and McCranie
FIG. 1.—Discriminant function analysis of the Central ¨ ¨
(1994); Kohler (1999); Kohler (2001); Koh- ¨
American anoles formerly referred to as Anolis limifrons. ¨
ler et al., (2001); Kohler and McCranie
See text for details. (2001).
Diagnosis.—A medium-sized species
reasons the following taxa (respective type (snout–vent length [SVL] in largest specimen
localities in parentheses) are clearly referable 43.5 mm) of the genus Anolis (sensu Poe,
to our Species A: Anolis limifrons Cope 2004) that is most similar in external mor-
(Cucuyos, Veragua Province, Panama); A. phology to a cluster of Central American
´,
trochilus Cope (San Jose Costa Rica); A. species that are long-legged (longest toe of
´
pulchripes Peters (Chiriquı, Panama); A. adpressed hindlimb reaches to mid-eye or
bransfordii Cope (Nicaragua); A. godmani: beyond), have a single elongated prenasal
´
Boulenger (Irazu, Costa Rica); and A. biscuti- scale, smooth to slightly keeled ventral scales,
ger Taylor (Golfito, Puntarenas Province, Costa and slender habitus, often delicate (i.e., Anolis
Rica). Because A. limifrons Cope is the oldest dollfusianus, A. ocelloscapularis, A. rodrigue-
available name for this species, our Species A zii, A. yoroensis, A. zeus). Within this cluster
has to be referred to that name and the other of species, A. limifrons can be readily
nominal species remain in the synonymy of A. distinguished by male dewlap coloration (dull
limifrons. The holotype of A. rivieri Thominot white with a small basal orange blotch in A.
is a juvenile with unspecific locality data limifrons vs. uniformly dull white in A. zeus,
(‘‘Panama’’), and its taxonomic identity cannot and almost uniformly orange to orange-yellow
be determined. Therefore, we consider A.
in the remaining species. Additionally, A.
rivieri Thominot to be a nomen dubium.
limifrons differs from the species in this
Interestingly, there is no available scientific
cluster by the following characteristics (con-
name for either of our Species B and C. We
dition for A. limifrons in parentheses): Anolis
therefore describe them as new species below.
dollfusianus: ventrals weakly keeled (smooth).
Anolis limifrons Cope, 1862 Anolis ocelloscapularis: An ocellated shoulder
spot present (absent); ventrals weakly keeled
Anolis biscutiger Taylor, 1956:81. Holotype (smooth); hemipenis bilobed (unilobed). Ano-
KU 40771 from ‘‘Golfito, Puntarenas Prov- lis rodriguezii: hemipenis bilobed (unilobed).
ince, Costa Rica.’’ Anolis yoroensis: ventrals weakly keeled
Anolis bransfordii Cope, 1874:67. Holotype (smooth).
ANSP 7890 from ‘‘Nicaragua.’’ Description (Fig. 3).—Maximum SVL 41.5
Anolis godmani Boulenger, 1885:85. Syntypes mm in males, 43.5 mm in females; ratio tail
(BMNH and MCZ) from ‘‘Guatemala’’ and length/SVL 1.53–2.52 (2.17 6 0.21); tail
‘‘Irazu, Costa Rica.’’ Barbour (1834), Taylor slightly compressed in cross section, ratio tail
(1956). height/tail width 1.00–1.30 (1.16 6 0.07); ratio
Anolis limifrons Cope, 1862:178. Syntypes axilla to groin distance/SVL 0.36–0.53 (0.43 6
ANSP 7900–01 from ‘‘Veragua.’’ Bocourt 0.03); ratio head length/SVL 0.24–0.28 (0.26

6. March 2008] HERPETOLOGICA 97
FIG. 2.—Map indicating known collecting sites mentioned in text of Anolis apletophallus (triangles), A.
cryptolimifrons (squares), and A. limifrons (circles) in Honduras, Nicaragua, Costa Rica and Panama. Each symbol
can represent one or more nearby localities. Areas above 500 and 1000 m are shaded gray. Open symbols: specimens
with no everted hemipenis; Solid symbols: specimens with everted hemipenis.
6 0.01); ratio snout length/head length 0.41– hind limbs; scales on snout varying from
0.52 (0.45 6 0.02); ratio head length/head almost non-keeled to keeled; 5–9 (6.9 6 0.8)
width 1.44–1.80 (1.67 6 0.06); longest toe of postrostrals; 7–11 (8.9 6 0.9) scales between
adpressed hind limb reaching to a point nasals; 1 large elongated prenasal scale in
between anterior to eye and tip of snout; ratio contact with both rostral and first supralabial,
shank length/SVL 0.26–0.33 (0.29 6 0.02); occasionally only in contact with rostral; scales
ratio shank length/head length 0.97–1.24 (1.11 in distinct prefrontal depression generally
6 0.07); longest finger of extended forelimb slightly tuberculate posteriorly, wrinkled an-
reaching to a point between nostrils and tip of teriorly, some of them keeled; supraorbital
snout; longest finger of adpressed forelimb semicircles well developed, separated by 0–4
reaches in between anterior to insertion of (1.8 6 0.8) scales; supraorbital disc composed
hind limbs and slightly beyond to insertion of of 5–14 distinctly enlarged keeled scales;

7. 98 HERPETOLOGICA [Vol. 64, No. 1
to the canthals mostly tuberculated; 4–7
keeled subocular scales arranged in a single
row; 5–8 (6.5 6 0.7) supralabials to level
below center of eye; 2–5 suboculars broadly in
contact with supralabials; ear opening medi-
um-sized, ratio tympanum height/interparietal
scale length 0.58–1.36 (0.86 6 0.16); mental
distinctly wider than long, completely divided
medially, bordered posteriorly by 4–8 (6.5 6
0.9) postmentals; 5–8 (6.3 6 0.7) infralabials
to level below center of eye; sublabials
undifferentiated; keeled granular scales pre-
sent on chin and throat; dewlap extending
from level below oral ricti to axilla, in some
specimens extending 1–2 mm posterior to
axilla; dorsum of body with weakly keeled
granular scales (at least anteriorly) with
rounded posterior margins, 2 medial rows
slightly enlarged, 38–70 (54.3 6 5.8) medial
dorsal scales in one head length; 70–112 (91.5
6 9.0) medial dorsal scales between axilla and
groin; lateral scales homogeneous, ventrals at
midbody smooth, slightly bulging, non–imbri-
cate, 26–58 (40.6 6 6.1) ventral scales in one
head length; 49–79 (64.6 6 5.7) ventral scales
between axilla and groin; 108–157 (132.1 6
10.6) scales around midbody; caudal scales
FIG. 3.—Head of Anolis limifrons (SMF 86900). Scale
strongly keeled; caudal middorsal scales
bars equal 1.0 mm. slightly enlarged, without whorls of enlarged
scales, although an indistinct division in
circumorbital row usually incomplete, there- segments is discernible; a pair of slightly
fore, 0–3 enlarged supraorbitals in contact enlarged postanal scales usually present; no
with supraorbital semicircles; a single large tube-like axillary pocket present; scales on
elongated superciliary; 3–6 rows of small dorsal surface of forelimb keeled, imbricate;
keeled scales extending between enlarged digital pads dilated; distal phalanx narrower
supraorbitals and superciliaries; a very shallow than and raised from dilated pad; 20–27 (23.3
parietal depression present in most speci- 6 1.6) lamellae under phalanges ii–iv of
mens; interparietal scale well developed, fourth toe; 7–11 (8.5 6 0.9) scales under
usually surrounded by scales of moderate size distal phalanx of fourth toe.
anteriorly and by small to moderate size scales Description of completely everted hemipe-
posteriorly; 1–5 (2.7 6 0.9) scales present nis.—Small unilobed organ; sulcus spermati-
between interparietal and supraorbital semi- cus bordered by well developed sulcal lips and
circles; canthal ridge distinct, composed of 6– opens at base of apex; no discernable surface
11 (7.5 6 0.8) canthal scales, with 3–5 (4.0 6 structure on truncus and lobes; no asulcate
0.7) larger posterior scales; 7–15 (10.6 6 1.7) processus present (Fig. 4).
scales present between second canthals; 10–
18 (13.8 6 2.1) scales present between Anolis apletophallus sp. nov.
posterior canthals; 24–68 (41.1 6 9.6) loreal Anolis limifrons: Bocourt (1873; in part);
scales in a maximum of 5–8 (6.1 6 0.8) Dunn (1930; in part); Barbour (1934; in
horizontal rows, with the scales of lower rows part); Breder (1946).
and those adjacent to the canthals mostly Anolis limifrons limifrons: Etheridge (1959; in
keeled, and those of upper rows non-adjacent part).

8. March 2008] HERPETOLOGICA 99
FIG. 4.—Hemipenis of Anolis limifrons (SMF 85246). (a)
sulcate view; (b) asulcate view. Scale bar equals 1.0 mm.
Holotype (Fig. 5).—SMF 85307, an adult
male from Panama City, Metropolitan Na-
tional Park (8u589600N, 79u329460W), 45 m,
´
Panama Province, Panama. Collected 26
¨
January 2006 by Gunther Kohler, Javier
Sunyer, Abel A. Batista R. and Marcos Ponce.
Field tag number GK 1672.
Paratypes.—SMF 85308–19, same collect-
ing data as holotype. SMF 85308–13 are adult
males, SMF 85314–19 are adult females.
Diagnosis.—A medium-sized species (SVL
in largest specimen 47.0 mm) of the genus
Anolis (sensu Poe, 2004) that is most similar in
FIG. 5.—Head of holotype of Anolis apletophallus
external morphology to a cluster of Central (SMF 85307). Scale bars equal 1.0 mm.
American species that are long-legged (lon-
gest toe of adpressed hindlimb reaches to
mid-eye or beyond), have a single elongated slightly imbricate (smooth and non-imbricate).
prenasal scale, smooth to slightly keeled Anolis yoroensis: Ventrals weakly keeled and
ventral scales, and slender habitus, often slightly imbricate (smooth and non-imbricate).
delicate (i.e., Anolis dollfusianus, A. limifrons, Anolis zeus: Hemipenis unilobed (bilobed);
A. ocelloscapularis, A. rodriguezii, A. yoroen- male dewlap uniformly dull white (almost
sis, A. zeus). Within this cluster of species, A. uniformly orange).
apletophallus is most similar to A. limifrons Description of the holotype.—Adult male as
from which it is readily distinguished by indicated by everted hemipenes; SVL
hemipenis morphology: hemipenis small and 44.0 mm; tail length 96.0 mm, tail complete;
unilobed in A. limifrons, large and bilobed in tail slightly compressed in cross section, tail
A. apletophallus (Fig. 6). Anolis apletophallus height 1.75 mm, tail width 1.70 mm; axilla to
differs from the remaining species in this groin distance 17.6 mm; head length 12.0 mm,
cluster by the following characteristics (con- head length/SVL ratio 0.27; snout length
dition for A. apletophallus in parentheses): 5.2 mm; head width 7.1 mm; longest toe of
Anolis dollfusianus: Hemipenis unilobed (bi- adpressed hind limb reaching to a point
lobed); ventrals weakly keeled and slightly between eyes and rostrils; shank length
imbricate (smooth and non-imbricate). Anolis 13.0 mm, shank length/head length ratio 1.08;
ocelloscapularis: Ventrals weakly keeled and longest finger of extended forelimb reaching to
slightly imbricate (smooth and non-imbricate); a point slightly beyond nostrils; longest finger
an ocellated shoulder spot present (absent). of adpressed forelimb just reaches anterior
Anolis rodriguezii: Ventrals weakly keeled and insertion of hind limbs. Most scales on snout

9. 100 HERPETOLOGICA [Vol. 64, No. 1
between posterior canthals; 64 (right)–61 (left)
loreal scales in a maximum of 8 horizontal rows,
with the scales of lower rows mostly keeled,
and those of upper rows mostly tuberculated; 8
(right)–6 (left) keeled subocular scales ar-
ranged in a single row; 7 supralabials to level
below center of eye; 4 (right)–3 (left) subocu-
lars broadly in contact with supralabials; ear
opening 0.90 3 1.40 mm (length 3 height);
mental distinctly wider than long, completely
divided medially, bordered posteriorly by
keeled 6 postmentals (outer pair larger); 7
infralabials to level below center of eye;
sublabials undifferentiated; keeled granular
scales present on chin and throat; dewlap
extending from level below oral ricti to 3 mm
beyond level of axilla; dorsum of body with
weakly keeled granular scales with rounded
posterior margins, 2 medial rows slightly
enlarged, largest dorsal scales about 0.21 3
0.23 mm (length 3 width); about 64 medial
dorsal scales in one head length; about 85
medial dorsal scales between axilla and groin;
lateral scales homogeneous, average size
0.16 mm in diameter; ventrals at midbody
smooth, slightly bulging, non-imbricate, about
0.34 3 0.34 mm (length 3 width); about 42
FIG. 6.—Hemipenis of Anolis apletophallus (SMF ventral scales in one head length; about 66
80719). (a) sulcate view; (b) asulcate view. Scale bar
equals 1.0 mm.
ventral scales between axilla and groin; 142
scales around midbody; caudal scales strongly
keeled; 7 postrostrals; 10 scales between nasals; keeled; caudal middorsal scales slightly en-
1 large elongated prenasal scale in contact with larged, without whorls of enlarged scales,
both rostral and first supralabial; scales in although an indistinct division in segments is
distinct prefrontal depression slightly tubercu- discernible; a pair of enlarged postanal scales
late posteriorly, wrinkled anteriorly; supraor- present, about 0.69 mm wide; no tube-like
bital semicircles well developed, separated by 3 axillary pocket present; scales on dorsal surface
scales; supraorbital disc composed of 13–15 of forelimb keeled, imbricate, about 0.28 3
distinctly enlarged keeled scales; circumorbital 0.25 mm (length 3 width); digital pads dilated;
row incomplete, therefore, one enlarged su- distal phalanx narrower than and raised from
praorbital in contact with supraorbital semicir- dilated pad; 24 lamellae under phalanges ii–iv
cles; a single large elongated superciliary; about of fourth toe; 9 (right)–8 (left) scales under
3 or 4 rows of small keeled scales extending distal phalanx of fourth toe.
between enlarged supraorbitals and super- Description of completely everted hemipe-
ciliaries; a very shallow parietal depression nis.—Medium-sized bilobed organ; sulcus
present; interparietal scale well developed, 1.4 spermaticus bordered by well developed sulcal
3 1.0 mm (length 3 width), surrounded by lips and bifurcating at base of apex; shortly after
scales of moderate size; 3 scales present the bifurcation, branches open into broad
between interparietal and supraorbital semi- concave areas, one on each lobe; asulcate
circles; canthal ridge distinct, composed of 4 surface of apex and distal truncus strongly
large (posterior two largest) and 4 small calyculate, base of truncus with transverse
anterior canthal scales; 10 scales present folds; no asulcate processus present, although
between second canthals; 17 scales present a slightly elevated ridge present (Fig. 6).

10. March 2008] HERPETOLOGICA 101
Coloration in life.—Dorsal ground color
Warm sepia (color 221A); flanks Dark Drab
(119B); dorsal surface of head Dark Drab
(119B); venter dull white suffused with Dark
Drab (119B); dorsal surface of limbs Mars
Brown (223A); tail Drab Gray (119D) with
transverse Sepia (119) (but slightly more
reddish) bands; iris Raw Sienna (136); dewlap
Orange Yellow (18).
Variation.—The paratypes agrees well with
the holotype in general appearance, morpho-
metrics and scalation (see Table 1). In most of
the male paratypes, one pair of slightly
enlarged postanal scales is present. The
female paratypes have no dewlap and no
enlarged postanal scales. The coloration in life
of male paratype’s dewlap (SMF 85308) was
recorded as Trogon Yellow (153).
Etymology.—The name apletophallus is
formed from the Greek words apletos (im-
mense) and phallos (penis) and is used as a
noun in apposition.
Natural history notes.—All type specimens
were collected active during the day in
secondary forest. The lizards were spotted
on tree trunks and fallen branches, from near
ground level to 1.5 m above the ground.
Anolis cryptolimifrons sp. nov. FIG. 7.—Head of holotype of Anolis cryptolimifrons
(SMF 85230). Scale bars equal 1.0 mm.
Anolis limifrons: Poe (2004; in part).
Anolis limifrons limifrons: Etheridge (1959; in
part.). ventral scales, and slender habitus, often
delicate (i.e., A. apletophallus, A. dollfusianus,
Holotype (Fig. 7).—SMF 85230, an adult A. limifrons, A. ocelloscapularis, A. rodrigue-
male from Cerro Brujo (9u11916.40N, zii, A. yoroensis, A. zeus). Within this cluster
82u11925.40W), 10 m, Bocas del Toro Prov- of species, A. cryptolimifrons is most similar to
ince, Panama. Collected 19 January 2006 by A. limifrons and A. apletophallus. Anolis
¨
Gunther Kohler, Javier Sunyer, Abel A. cryptolimifrons is readily distinguished from
Batista R. and Marcos Ponce. Field tag A. limifrons by hemipenis morphology: hemi-
number GK 1502. penis small and unilobed in A. limifrons, large
Paratypes.—SMF 85231–44, same collect- and bilobed in A. cryptolimifrons. Anolis
ing data as holotype. Most of the paratypes are cryptolimifrons differs from A. apletophallus
females, except SMF 85236–37, 85239–40, in male dewlap size and color in life (small and
85242–43 (adult males). dull white with a small basal orange blotch in
Diagnosis.—A medium-sized species (SVL N. cryptolimifrons versus dewlap large and
in largest specimen 45.0 mm) of the genus almost uniformly orange in N. apletophallus).
Anolis (sensu Poe, 2004) that is most similar in Anolis cryptolimifrons differs from the re-
external morphology to a cluster of Central maining species in this cluster by the following
American species that are long-legged (lon- characteristics (condition for A. cryptolimi-
gest toe of adpressed hindlimb reaches to frons in parentheses): Anolis dollfusianus:
mid-eye or beyond), have a single elongated Hemipenis unilobed (bilobed); ventrals weak-
prenasal scale, smooth to slightly keeled ly keeled (smooth); fewer than 80 dorsals

11. 102 HERPETOLOGICA [Vol. 64, No. 1
between levels of axilla and groin (more than of 4 large (posterior two largest) and 4 small
87); male dewlap almost uniformly orange- anterior canthal scales; 16 scales present
yellow (dull white with a small basal orange between second canthals; 19 scales present
blotch). Anolis ocelloscapularis: An ocellated between posterior canthals; 61 loreal scales in
shoulder spot present (absent); ventrals weak- a maximum of 6 horizontal rows, with the
ly keeled (smooth); male dewlap almost scales of lower rows mostly keeled, and those
uniformly orange (dull white with a small of upper rows mostly tuberculated; 6 (right) –
basal orange blotch). Anolis rodriguezii: Male 5 (left) keeled subocular scales arranged in a
dewlap almost uniformly orange (dull white single row; 7 supralabials to level below center
with a small basal orange blotch). Anolis of eye; 2 suboculars broadly in contact with
yoroensis: Ventrals weakly keeled (smooth); supralabials; ear opening 0.5 3 1.3 mm
fewer than 87 dorsals between levels of axilla (length 3 height); mental distinctly wider
and groin (more than 87); male dewlap almost than long, completely divided medially, bor-
uniformly orange (dull white with a small dered posteriorly by 8 keeled postmentals
basal orange blotch). Anolis zeus: Hemipenis (outer pair larger); 7 infralabials to level below
unilobed (bilobed); male dewlap uniformly center of eye; sublabials undifferentiated;
dull white without a basal orange blotch (basal keeled granular scales present on chin and
orange blotch present). throat; dewlap extending from level below oral
Description of the holotype.—Adult male as ricti to about 1 mm anterior to axilla; dorsum
indicated by everted hemipenes; SVL of body with weakly keeled granular scales
41.0 mm; tail length 88.0 mm, tail complete; with rounded posterior margins, 2 medial
tail slightly compressed in cross section, tail rows slightly enlarged, largest dorsal scales
height 1.50 mm, tail width 1.25 mm; axilla to about 0.19 3 0.20 mm (length 3 width);
groin distance 18.0 mm; head length about 54 medial dorsal scales in one head
11.0 mm, head length/SVL ratio 0.27; snout length; about 97 medial dorsal scales between
length 5.2 mm; head width 6.6 mm; longest axilla and groin; lateral scales homogeneous,
toe of adpressed hind limb reaching to average size 0.10 mm in diameter; ventrals at
anterior portion of eye; shank length midbody smooth, slightly bulging, non-imbri-
11.1 mm, shank length/head length ratio cate, about 0.24 3 0.27 mm (length 3 width);
1.01; longest finger of extended forelimb about 46 ventral scales in one head length;
reaching nostrils; longest finger of adpressed about 68 ventral scales between axilla and
forelimb not reaching anterior insertion of groin; 148 scales around midbody; caudal
hind limbs. Most scales on snout keeled, some scales strongly keeled; caudal middorsal scales
wrinkled; 8 postrostrals; 10 scales between slightly enlarged, without whorls of enlarged
nasals; 1 large elongated prenasal scale in scales, although an indistinct division in
contact with both rostral and first supralabial; segments is discernible; a pair of enlarged
scales in distinct prefrontal depression slightly postanal scales present; no tube-like axillary
tuberculate; supraorbital semicircles well de- pocket present; scales on dorsal surface of
veloped, separated by 2 scales; supraorbital forelimb keeled, imbricate, about 0.24 3
disc composed of 9–10 distinctly enlarged 0.26 mm (length 3 width); digital pads
keeled scales; circumorbital row complete, dilated; distal phalanx narrower than and
therefore, no enlarged supraorbitals in contact raised from dilated pad; 24 lamellae under
with supraorbital semicircles; a single large phalanges ii–iv of fourth toe on right foot
elongated superciliary; about 5 rows of small (fourth toe of left foot missing); 7 scales under
keeled scales extending between enlarged distal phalanx of fourth toe.
supraorbitals and superciliaries; a shallow Description of completely everted hemipe-
parietal depression present; interparietal scale nis.—Medium-sized bilobed organ; sulcus
well developed, 1.30 3 0.95 mm (length 3 spermaticus bordered by well developed sulcal
width), surrounded by scales of moderate size lips and bifurcating at base of apex; shortly after
anteriorly and small size posteriorly; 3 scales the bifurcation, branches open into broad
present between interparietal and supraorbital concave areas, one on each lobe; asulcate
semicircles; canthal ridge distinct, composed surface of apex and distal truncus strongly

12. March 2008] HERPETOLOGICA 103
relationship between the new species and its
congener Anolis limifrons Cope.
Natural history notes.—All type specimens
were collected active during the day in a patch
of secondary forest. The lizards were spotted
on branches and leaves of bushes and small
trees, 0.5 to 1.5 m above the ground.
Key to the males of the species formerly
referred to as Anolis limifrons
1a. Male dewlap large, larger than 150 mm2
(Fig. 9a), almost uniformly orange _ _ _ _ _ _ _ _ _ _ _ _
Anolis apletophallus
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
1b. Male dewlap small, smaller than 100 mm2
(Figs. 9b,c), dull white with a small basal
orange-yellow blotch 2 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
2a. Hemipenis unilobed Anolis limifrons _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
2b. Hemipenis bilobed Anolis cryptolimifrons _ _ _ _ _ _ _ _ _
DISCUSSION
Until recently, hemipenial morphology in
the diverse group of anoles has been mostly
ignored. The minority of male anoles housed
in museum collections worldwide have evert-
ed hemipenes. We urge future collectors of
these anoles to attempt to fully evert the
hemipenes of their specimens. Study of these
FIG. 8.—Hemipenis of holotype of Anolis cryptolimi- organs may reveal more cryptic species of
frons (SMF 85230). (a) sulcate view; (b) asulcate view. anoles than originally envisaged by most
Scale bar equals 1.0 mm. workers. This distinction is of great conserva-
tional importance: whereas a large amount of
ecological data is available for Anolis limifrons
calyculate, base of truncus with transverse and A. apletophallus, very little is known from
folds; no asulcate processus, although a slightly A. cryptolimifrons, a species with a relatively
elevated ridge present (Fig. 8). restricted distribution. Although A. limifrons
Coloration in life.—Dorsal ground color and A. cryptolimifrons both occur in the same
Raw Umber (color 123 in Smithe 1975–1981) general area of mainland western Panama
suffused with Vandyke Brown (221) at mid- (Caribbean versant), no actual instance of
dorsum; venter Pale Horn Color (92); iris syntopy has been documented. Both taxa
Cinnamon (123A); dewlap dull white with a appear to use the same habitat and same
small Chamois (123D) basal blotch. perching sites and might exclude each other.
Variation.—The paratypes agrees well with Savage (2002) reported populations of
the holotype in general appearance, morpho- Anolis limifrons-like lizards from the slopes
metrics and scalation (see Table 1). In most of ´
of the volcanoes Irazu and Turrialba with
the male paratypes, one pair of slightly keeled ventral scales but otherwise identical in
enlarged postanal scales is present. The scutellation and coloration. Boulenger (1885)
female paratypes have no dewlap and no based the description of A. godmani on this
enlarged postanal scales. characteristic, and in earlier publications
Etymology.—The species name cryptolimi- Savage (e.g., Savage, 1973; Savage and Villa,
frons is used as a noun in apposition and 1986) recognized this form as a valid species
reflects the similarity and suspected close and used the name A. godmani for it. More

13. 104 HERPETOLOGICA [Vol. 64, No. 1
species) which has distinctly keeled ventral
scales. Also, the scales on head and dorsum
are strongly keeled, much more than in any
other examined A. limifrons. Unfortunately,
no data on hemipenis morpholgy nor on male
dewlap coloration are available for represen-
tatives of this population. Further collecting
and study is needed in order to evaluate the
status of the populations of A. limifrons-like
lizards from the slopes of the vulcanos Irazu´
and Turrialba in Costa Rica.
Acknowledgments.—Collecting and exportation permits
were provided by J. Guevara Sequeira, SINAC Central,
´
Ministerio del Ambiente y Energıa (MINAE), San Jose, ´
´
Costa Rica; A. Barahona, T. Garcıa, A. P. Martinez, E.
˜ ´
Munoz Galeano, and C. Romero, Corporacion Hondur-
˜
ena de Desarollo Forestal (COHDEFOR), Tegucigalpa,
Honduras; M. Fonseca Cuevas, S. Tijerino, B. Quintero, I.
´
Ortega, M. G. Camacho, C. Mejıa, and C. Peres-Roman, ´
Ministerio del Ambiente y los Recursos Naturales
(MARENA), Managua, Nicaragua; Y. Hidalgo, Autoridad
Nacional del Ambiente (ANAM), Panama City, Panama.
A. Batista and M. Ponce (Panama), and A. Hertz, S.
Lotzkat, D. Manzanarez and L. Obando (Nicaragua),
actively helped in the field. M. Dehling, J. Ferrari, G. Paiz
and J. R. McCranie generously donated specimens for this
study. M. Piepenbring, Botanisches Institut J. W. Goethe-
¨
Universitat, Frankfurt, Germany, B. E. Sanjur, Facultad
de Ciencias Naturales y Exactas de la Universidad
´ ´
Autonoma de Chiriquı (UNACHI), David, Panama, and
J. Vegas, Panama City, provided logistical support for our
studies in Panama. This paper is based in part upon work
supported by the Deutscher Akademischer Austausch-
¨
dienst (DAAD) to J. Sunyer and to G. Kohler through the
Partnership Program between the J. W. Goethe-Universi-
¨
tat Frankfurt am Main, Germany, und der Facultad de
Ciencias Naturales y Exactas der Universidad Autonoma ´
´
de Chiriquı (UNACHI), David, Panama. We thank L.
¨
Czupalla and J. Kohler for providing some of the drawings
used in this paper. For the loan of and/or access to
specimens, we thank L. Ford, C. J. Raxworthy and D. R.
Frost, American Museum of Natural History (AMNH),
New York; T. Daeschler and A. Gilmore, Academy of
Natural Sciences (ANSP), Philadelphia; S. P. Rogers,
Carnegie Museum of Natural History (CM), Pittsburgh;
A. Resetar, Field Museum of Natural History (FMNH),
Chicago; G. Lenglet, Institut Royal des Sciences Natur-
FIG. 9.—Specimens in life with extended dewlaps (a) elles de Belgique (IRSNB), Bruxelles; W. E. Duellman
´
Anolis apletophallus male (Panama City, Panama, Panama; and J. E. Simmons, University of Kansas, Natural History
not preserved); (b) A. cryptolimifrons male (Cerro Brujo, Museum (KU), Lawrence; D. Rossman, Museum of
Bocas del Toro, Panama; not preserved); (c) A. limifrons, Natural Science, Louisianna State University (LSUMZ),
´
male (Los Algarrobos, Chiriquı, Panama; not preserved). Baton Rouge; J. Hanken and J. P. Rosado, Museum of
Comparative Zoology, Harvard University (MCZ), Cam-
´
bridge; A. Batista and M. Ponce, Museo Herpetologico de
recently, Savage (2002) considered it to be an ´
Chiriquı (MHCH), David; F. Tiedemann, Naturhistor-
individual variant and therefore a synonym of ˜
isches Museum (NMW), Wien; F. Bolanos, Museo de
A. limifrons. We have examined a single ´
Zoologıa Universidad de Costa Rica (UCR), San Jose; K. ´
L. Krysko and F. W. King, Florida Museum of Natural
female (SMF 86924) from this general area History (UF), Gainesville; C. A. Phillips and J. Petzing,
collected at 1500 m elevation (about 160 m Illinois Natural History Survey, Center for Biodiversity
above the highest Costarican record for this (UIMNH), Champaign; R. A. Nussbaum and G. Schnei-

14. March 2008] HERPETOLOGICA 105
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¨
(UMMZ), Ann Arbor; R. W. McDiarmid and W. R. Herpeton, Offenbach, Germany.
Heyer, National Museum of Natural History (USNM), KOHLER, G. 2003. Reptiles of Central America. Herpeton
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¨
¨
Bohme, Zoologisches Forschungsinstitut und Museum A. (Werner, 1917), a valid species of anole from Guate-
Koenig (ZFMK), Bonn; and R. Gunther, Museum fur
¨ ¨ mala and El Salvador (Squamata: Sauria: Iguanidae).
¨
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¨
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12:15–24. SAVAGE, J. M. 1973. Herpetological collections made by
ETHERIDGE, R. 1959. The relationships of the anoles Dr. John F. Bransford, Assistant Surgeon, U.S.N.
(Reptilia: Sauria: Iguanidae). An interpretation based during the Nicaragua and Panama Canal Surveys
on skeletal morphology. Ph.D. Dissertation, University (1872–1885). Journal of Herpetology 7:35–38.
of Michigan, Ann Arbor, Michigan, U.S.A. SAVAGE, J. M. 1997. On terminology for the description of
FITCH, H. S., AND R. A. SEIGEL. 1984. Ecological and the hemipenes of squamate reptiles. Herpetological
taxonomic notes on Nicaraguan anoles. Milwaukee Public Journal 7:23–25.
Museum Contributions in Biology and Geology 57:1–13. SAVAGE, J. M. 2002. The amphibians and reptiles of Costa
GAIGE, H. T., N. HARTWEG, AND L. C. STUART. 1937. Notes Rica. A herpetofauna between two continents, between
on a collection of amphibians and reptiles from eastern two seas. University of Chicago Press, Chicago, Illinois,
Nicaragua. Occasional Papers Museum of Zoology U.S.A.
University of Michigan 357:1–18. SAVAGE, J. M., AND J. VILLA. 1986. Introduction to the
KOHLER, G. 1999. The amphibians and reptiles of
¨ Herpetofauna of Costa Rica. Society for the Study of
Nicaragua—a distributional checklist with keys. Courier Amphibians and Reptiles, Contributions to Herpetolo-
Forschungsinstut Senckenberg 213:1–212. gy 3:1–207.

15. 106 HERPETOLOGICA [Vol. 64, No. 1
SMITHE, F. B. 1975–1981. Naturalist’s color guide. Part I. 505352–53, 505354–56all H, 505357–60, 505361H, 505362–
Color guide. 182 color swatches. American Museum of 63, 505364H, 505365, 505366H, 505367, 505368–70all H; Rıo ´
Natural History, New York, New York, U.S.A. Agua Salud, Panama Canal: USNM 505432–34, 505435–
both H H H
STUART, L. C. 1955. A brief review of the Guatemalan 36 , 505437, 505438 , 505439–40, 505441 , 505442,
lizards of the genus Anolis. Miscellaneous Publications 505443–45all H, 505446–49, 505450H, 505451, 505452H,
Museum of Zoology University of Michigan 91:1–31. 505453, 505454–55both H, 505456; Rıo Gigantito, Panama
´
TAYLOR, E. H. 1956. A review of the lizards of Costa Rica. Canal: USNM 505515–20, 505521–22both H, 505523,
H
University of Kansas Science Bulletin 38:1–322. ´
505524 , 505525–40; Rıo Mendoza, Panama Canal: USNM
`
THOMINOT, A. 1882. Note sur un Anolis d’espece nouvelle. 505628–42, 505643H, 505644–48, 505649H; Tigre Island,
´ ´
Bulletin de la Societe philomatique de Paris 6:251–253. Panama Canal: USNM 505484–500, 505501–05all H,
VENCES, M., M. FRANZEN, A. FLASCHENDRAGER, R.
¨ ¨ 505506–10, 505511H, 505512–14; Quipo, on Rıo Ciri, W ´
SCHMITT, AND J. REGOS. 1998. Beobachtungen zur
¨ ´ ´n:
side L. Gatun: AMNH 42919; Darie 0.5 hr below junction
Herpetofauna von Nicaragua: Kommentierte Artenliste ´ ´
of Rıo Jaque and Rıo Imamado: USNM 161216; Pirri
der Reptilien. Salamandra 34:17–42. ´ ´
Range, near head of Rıo Limon: USNM 50151; Chalichi-
VILLA, J., L. D. WILSON, AND J. D. JOHNSON. 1988. Middle man’s Creek: AMNH 42916–17; Camp Creek: AMNH
American Herpetology. University of Missouri Press, ´
42920; Rıo Chucunague [Chucunaque]: AMNH 37902–03,
Columbia, Missouri, U.S.A. ´
49214; Rıo Chucunague [Chucunaque], 3 mi W of Camp
WILSON, L. D., AND J. R. MCCRANIE. 1994. Comments on Townsend: AMNH 102557–58; Yavisa, backyard junkpiles:
the occurence of a salamander and three lizard species ´
CM 74030–31; Yavisa, trail along Rıo Chucunague [Chucu-
in Honduras. Amphibia–Reptilia 15:416–421. naque]: CM 74038; 7–11 km SW El Real between Rıo ´
´
Presencia and Rıo Morgentese, 100–350 m: UMMZ
´
155802–03; Rıo Sucubti: AMNH 42909–14; Tapia: AMNH
.Accepted: 9 January 2008 ´
25021, 25023, 25025–26; Panama: Metropolitan National
.Associate Editor: Michael Harvey Park, Panama City (8u589600N, 79u329460W), 45 m: SMF
85307*H, 85308*, 85309–10both*H, 85311–19all*; ‘‘Panama
Prov.’’: AMNH 71727, 71729, 76001–03, 89883; Canal
APPENDIX Zone: AMNH 67078, 67081–82, 71716, 71730–33, 75990H,
Specimens Examined.—*Specimens used in Discrimi- 75991–92, 89880–82, 85605–07, 107432–64, 107465H,
nant Function Analysis 107466, USNM 54325; Las Cascadas, Canal Zone: MCZ
H
Specimens with everted hemipenis 19414*, 19416*, 175185*, 175187*, 175189–90both*,
´
Anolis apletophallus.—Panama: Colon: Cano Saddle, 175194*, 175196–97both*, 175200*, 175203*, 175207*,
Close’s plantation: USNM 69586; Chagres River: USNM 175209*, 175213–16all*, 175223*, 175225*, 175233*; Canal
´ ´
102849; Rıo Indio, near Gatun: USNM 54007; Gatun: ´ Zone, Camp Mary Caretta (5 Camp Santa Margarita):
USNM 54011–12; Bohio Peninsula – East, Panama Canal: USNM 25162–63; Canal Zone, Gamboa: USNM 193351;
USNM 505231–37, 505238–42all H, 505243–51, 505252H, ´
Canal Zone, Rıo Frijoles, 3 mi N Gamboa: UF 124417;
505253, 505254–56all H, 505257, 505258H, 505259, Gamboa, at confluence of Panama Canal and Chagres River
505260H, 505261–69, 505270–73all H, 505274, 505275– (9u069540N, 79u419420W): USNM 297807–09; Gamboa:
76both H, 505277–85, 505286H, 505287–88, 505289–90both H, ´
SMF 84954, 83084–85; Canal Zone, Rıo Medio: USNM
505291–92, 505293–301all H; Buena Vista Peninsula, 102725; Lion Hill: USNM 54172; Old Panama: USNM
Panama Canal: USNM 505199–230; Gigante Ridge, Pana- 50129; Panama, Cabima (Pacific slope): USNM 48500–01;
ma Canal: USNM 505371–82, 505383–84both H, 505385, Panama, Cocoli: USNM 193365, 193371, 523377–78;
505386–88all H, 505389–93, 505394–95both H, 505396–97, Puerta Obaldia, Quebrada Repressa: USNM 150127;
505398–99both H, 505400, 505401H; Juan Gallegos Island – Venado Beach: USNM 193359; Punta de Pena: USNM
East, Panama Canal: USNM 505571–82, 505583–84both H, 38712; Toro Point: USNM 53725; Trinidad River: USNM
505585, 505586H, 505587–89, 505590H, 505591–97, ´
63992–94, 63997; Viento Frıo: USNM 48597; La Joya,
505598–99both H, 505600, 505601H; Juan Gallegos Island – ´
Pacific side: ANSP 25136*; Rıo Tatare, Pacific side: ANSP
North, Panama Canal: USNM 505602–04, 505605–08all H, 25137; Gatun: ANSP 19520–22all*, 19523, 24863–65all*,
´
505609–16, 505617H, 505618–22, 505623H, 505624, 24866–67; Canal Zone, Fort Gulick, Atlantic side: ANSP
505625H, 505626, 505627H; Juan Gallegos Island – South, 25104*; Canal Zone, Empire: ANSP 19545; Chico: CM
Panama Canal: USNM 505541–49, 505550H, 505551–52, 6859; Barro Colorado Island: AMNH 75986, ANSP 24487–
505553H, 505554, 505555–57all H, 505558, 505559H, 92all*, 24493*H, 24494–500, 24559, CM 7664, 7666, 7669,
505560–63, 505564–68all H, 505569, 505570H; Limbo 7671, 7673–75, 7681, 7686–93, 7699, 7705, 7707–10, 7712–
Camp, Panama Canal: USNM 505457–73, 505474H, 19, 7725; Barro Colorado Island, Wheeler trail: UMMZ
505475–79, 505480–81both H, 505482, 505483H; Lion Hill 63688; Lutz Creek below Donats bridge: CM 7659; Rıo ´
Island, Panama Canal: USNM 505302–05, 505306–08all H, Pequeni, head of Madden Lake: ANSP 21694; Juan Mina,
505309, 505310H, 505311–18, 505319–20both H, 505321–31, Madden Lake watershed: CM 74047; near Fort Clayton
505332H, 505333–34; Pena Blanca Peninsula, Panama
˜ Reservation: UIMNH 42184; Cerro Campana: AMNH
Canal: USNM 505402–06, 505407H, 505408–10, 505411– 75999; Cerro Campana, 800–900 m: AMNH 10666; Altos
15 all H
, 505416, 505417 , 505418–22, 505423H, 505424–25,
H
de Maje AMNH 109623H, 109624–35; Serranıa de Maje
´: ´ ´,
505426–27both H, 505428, 505429–30both H, 505431; Poach- ´ ˜ ´ ´
proximities of Union Saldana, Rıo Chiman (8u51959.00N,
ers Peninsula: USNM 505650–59, 505660–61both H, ´
78u35913.60W), 470 m: MHCH 1146; Serranıa de Maje Rıo ´, ´
505662–64, 505665H, 505666–67, 505668–69both H, Ambroya: MHCH 1082H, 1086, 1090; San Blas: Armila: ´
H
505670, 505671–73all H, 505674–76; Puma Island, Panama USNM 150099, 150100 , 150101–108; Armila, Quebrada
Canal: USNM 505335–44, 505345H, 505346–50, 505351H, Venado: USNM 150110–13; Nusagandi, near field station

16. March 2008] HERPETOLOGICA 107
(9u20.509N, 78u59.649W), 300–360 m: SMF 80717H, 80718, ´
mouth of Rıo Tortuguero, ca. 50 mi NW Limon: AMNH ´
80719H , 80720*H; km 14.6 on El Llano – Cartı road, 370 m:
´ 89174H; ca. 5 mi N Limon: AMNH 89177H; Zent
´
´
AMNH 110572; km 12.8 on El Llano – Cartı road, 290 m: [10u01960 N, 83u16960 W, 31 m]: USNM 137767;
AMNH 110573–74. Approximately 17.0 km WSW Puerto Limon between ´
´
Anolis cryptolimifrons.—Costa Rica: Limon: SE side ´ ´
Rıo Blanco and Rıo Toro R12882–88; Siquirres large
Cerro Nimaso: UCR 8477*H; Panama: Bocas del Toro: stream outside of town R12880–81; Motel Matama,
Cerro Brujo (9u11916.40N, 82u11925.40W), 10 m: SMF ´
3.5 km N Limon: AMNH 138604–05; 4 mi SW La
85230*H, 85231–35all*, 85236–37both*H, 85238–41all*, Fortuna: IRSNB 11684; Atalanta Farm, Estrella Valle:
85242–43both*H, 85244*; vicinity of Almirante: ANSP ANSP 21465; Puerto Limon: ANSP 19570, 19571H,
´
34047–50all*H, 34051*, 34052H, 34053–54, USNM 19572–78, IRSNB 13804, ZSM 85/1998, 86/1998; La
´
279062–71, 279130–33; Rıo Changuinola, near Quebrada ´ ´
Castilla, lower Rıo Reventazon: ANSP 23710–37, 24501–
El Guabo, 16 km airline W Almirante, 200–250 m: 04, 34747; Tortuguero: UF 135783–84; Tortuguero, just N
AMNH 119043H, 119044–49; Cayo Agua, Punta Norte: of Caribbean Conservation Commission Camp: USNM
USNM 150005, 150007–09; Cayo Agua, near Punta ´
244861; 2.4 km E Siquirres, along Rıo Pacuare: CM
´
Limon: USNM 338690–92; Isla Bastimentos, Old Point: ´
89566–67; Rte 32, 69 km E Rıo Hondo, 2 km N on dirt
USNM 297888–97; Isla Bastimentos: SMF 85229, road: ANSP 32372, 32374; 2–3 km (air) NW Bribri at Rıo ´
85245*H; Isla Colon, ca. 0.8 mi N of Bocas del Toro
´ ´
Carbon along road to Uatsi: ANSP 32559; RB Hitoy
´
(town): USNM 338214–16; Isla Colon, just N of Bocas del Cerere: SMF 86925*H, 86926*, 86927–28both*H; Estacion ´
Toro (town), along beach at fairgrounds, E side of Biologica Tierra Media, Matina: UCR 12399*H; Quebrada
´
H
´
isthmus: USNM 346901; Isla Colon, La Gruta: USNM Uatsi: UCR 13031* ; Sendero San Mateo, Cerro Uatsi:
313767–78, 313794, 338217, 338218H, 338219–22, UCR 13195*H; Finca Bryan Kubicki, Guayacan: UCR ´
338223H, 338224–26, 338227–29all H, 338230–32; Isla 16914*H; Puntarenas: ‘‘Puntarenas Prov.’’: AMNH 16357;
Cristobal, Bocatorito camp: USNM 348191H, 348192–94,
´ ´
7 mi E Golfito: LSUMZ 30260; Penınsula de Osa, Golfo
348195H, 348196–201, 348202H, 348203–05; NW side of ´ ´
Dulce, Puerto Jimenez, jardin at Jimenez Yacht Club:
Isla Cristobal: USNM 348206H, 348207–10; Isla Pastores,
´ SMF 81512–15; EB San Gerardo, Monteverde: UCR
Ford Point: USNM 313847–48; Isla Popa, 1 km SE of 13652*H; Hotel Sunset, 1 km N Santa Elena, Monteverde
Deer Island channel: USNM 298121–35; Isla Popa, south region (10u19936.90N, 84u49924.10W), 1450–1475 m:
end of, 1 km E of Sumwood Channel: USNM 319213–25, SMF 85549–53all*, 85554*H; San Jose: San Jose: ANSP
´ ´
347260–63, 347264–65both H, 347266–71, 347272–75all H, ´
7804, USNM 80902–05; San Jose, grounds of Hotel Irazu: ´
347276–77, 347278H, 347279, 347280H, 347281–83; Isla UMMZ 143761; 6.0 km N San Isidro de Perez Zeledon: ´
Popa (9u139140N, 82u089280W), 10 m: SMF 85247– UTA R12868; Talamanca: USNM 75956; Moravia de
48both*H, 85249*, 85250*H; Isla Popa, NNE beach Chirripo: UMMZ 128952; Near San Isidro (9u24929.10N,
(09u13924.40N, 82u06936.60W), 10–20 m: SMF 85399H, 83u44906.60W), 880 m: SMF 86933–35; Road from
85400; Laguna de Tierra Oscura, 3.7 km S of Tigre Key: General Viejo to Santa Elena (9u20906.50N,
USNM 313838–51, 348467–70all H, 348471–80; Long Bay 83u39911.40W), 650 m: SMF 86929; Honduras: Colon: ´
Point and Flat Rock Point, between, on E side of island, ´
Quebrada Machın (15u199100N, 85u179300W), 540 m:
ca. 100 yds from beach: USNM 297816–17; midpoint on USNM 536490–91, 541026–29; R. B. Rıo Platano, El ´ ´
W side of Cayo Carenero: USNM 347938; N end of Cayo ´ ´
Ocotillal, Cabeceras de Rıo Platano (15u40.39N,
Roldan: USNM 348043; Isla Solarte, 10 m: SMF 85u17.19W) 370–410 m: SMF 86215H, 86216, 86217H;
85251*H, 85252–53both*; USNM 338552, 338553–55all H, ´
Gracias a Dios: Mocoron R46171–72; confluence of Rıo ´
338556–57, 338558H, 338559–61, 338562–63both H, ´
Wampu and Quebrada Waskista (15u009N, 84u599W),
338564, 338565H, 338566–68. 85 m: USNM 330183–84; confluence of Rıo Wampu and ´ ´
Anolis limifrons.—Costa Rica: ‘‘Costa Rica’’: USNM ´
Rıo Patuca (14u589N, 84u599W), 60 m: USNM 330181–
38334, 70406–10, 81198; Parismina nivel del mar: USNM both
82; Quebrada Waskista, 85 m: SMF 80708–09 *; R. B.
75444–46; Colombiana: USNM 67347–48; Alajuela: Rıo ´ ´ ´
Rıo Platano, Raudal Kiplatara, (15u59.99N, 84u94.89W),
Frıo: USNM 19514; Pizote: UCR 9988–89both*H,
´ 50–255 m: SMF 86172H, 86173–74, 86175H, 86176–82,
10646*H; Pilon, Bijagua: UCR 10504*H; Laguna Lagarto
´ 86183–84both H, 86186; R. B. Rıo Platano, Rıo Cuyamel
´ ´ ´
Lodge, Boca Tapada: UCR 12609*H; Rıo Tapezco: UCR
´ (15u58.29N, 84u99.39W), 115–345 m: SMF 86188,
16515*H; Cartago: 10.0 km NE Turrialba on E bank of 86189H, 86190, 86191H, 86192–96, 86198H, 86204–05;
´ ´
Rıo Reventazon R12873–75; 2.0 km W Pavones de ´ ´
R. B. Rıo Platano, Pomokir (15u49.39N, 84u94.89W), 150–
Turrialba R12878–79; 3.0 km NE Pavones de Turrialba 240 m: SMF 86207, 86208–09both H, 86211, 86213–14both H;
´ ´
at Rıo Chitarıa R12869–72, R12876–77; Tapanti: ZFMK ´ ´
R. B. Rıo Platano, Crique Unawas (15u12.79N, 84u92.39W),
48716; ‘‘Turrialba Prov.’’: AMNH 69707–10; Turrialba: 180–305 m: SMF 86220–21, 86222–3both H, 86224–28,
SMF 77206, USNM 133180, 192586, 523375–76; Esta- 86229H; Quebrada Waskista-Rıo Wampu confluence
´ ´
´ ´
cion Biologica Copal, Tausito, Pejibaye: UCR 16127*; (15u009N, 84u599W), 85 m: SMF 86887; Cabeceras del
1 Km E La Pastora (9u58906.10N, 83u44918.90W), 1500 m: ´
Rıo Rus Rus, 190 m: SMF 86888; Olancho: confluence of
SMF 86924; Heredia: Rara Avis, Catarata (10u16.929N, ´ ´
Rıo Wampu and Quebrada Siksatara (15u039N, 85u029W),
84u02.749W), 700 m: SMF 81814, 81815H; Puerto Viejo: ´
95 m: USNM 330180; confluence of Rıo Aner and Rıo ´
ZFMK 48723–37, USNM 245041; La Selva Biological ´
Wampu (15u049N, 85u069W), 110 m: SMF 80704, USNM
´
Station, 2.6 km SE of Puerto Viejo de Sarapiquı: USNM ´ ´
330176–77; confluence of Rıo Wampu and Rıo Sausa ´
505677–90, 505691H, 505692–95, 505696H, 505697, (15u049N, 85u069W), 100 m: SMF 80705, 80706–07both*,
505698H, 505699–701; Finca Santiago (near La Selva): ´
USNM 330178–79; confluence of Rıo Yanguay and Rıo ´
´ ´
SMF 78433*; Limon: ‘‘Limon Prov.’’: AMNH 89171–73, ´
Wampu (15u039N, 85u089W), 110 m: USNM 330175;
89175–76, 95095, 99671–76, 149611–20; ca 5/4 mi S Parque Nacional Patuca, Matamoros (14u409210N,