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A CRYPTIC NEW SPECIES OF ANOLE (SQUAMATA: DACTYLOIDAE)
FROM THE LENCA HIGHLANDS OF HONDURAS,
PREVIOUSLY REFERRED TO AS NOROPS CRASSULUS (COPE, 1864)
Erich P. hofmann
Department of Biology, Indiana University of Pennsylvania
975 Oakland Ave., Indiana, Pennsylvania, 15705–1081, USA
Current Address: Department of Biological Sciences, Clemson University
190 Collings St., Clemson, South Carolina 29634, USA
ehofman@clemson.edu
Josiah h. TownsEnd
[Research Associate, Section of Amphibians and Reptiles, Carnegie Museum of Natural History]
Department of Biology, Indiana University of Pennsylvania
975 Oakland Ave., Indiana, Pennsylvania 15705–1081, USA
josiah.townsend@iup.edu
ANNALS OF CARNEGIE MUSEUM
Vol. 85, numbEr 2, PP. 91–111 31 dEcEmbEr 2018
ABSTRACT
We describe a cryptic new species of Norops Wagler, 1830, Norops caceresae, from mixed transitional and broadleaf cloud forest formations in
the Lenca Highlands of southwestern Honduras. This population was previously considered conspecific with Norops crassulus (Cope, 1864) of
Guatemala, El Salvador, and Mexico, despite it being entirely disjunct (> 100 km) from any of those populations. Recent molecular work revealed
consistent, deep mitochondrial and nuclear distinctiveness between this population and all other anoles of the N. crassulus species group, prompting
a thorough morphological investigation of this population. This new species is most similar in external morphology to N. crassulus sensu stricto,
but is readily distinguished by molecular distinctiveness, distribution, and morphology.
KEy words: Chortís Block; lizard; Mesoamerica; morphology; Nuclear Central America; Reptilia; taxonomy
RESUMEN
Describimos una nueva especie críptica de Norops Wagler, 1830, Norops caceresae, de las formaciones mixtas de bosque nuboso transiciónal y
de hoja ancha en las tierras altas de Lenca, en el suroeste de Honduras. Anteriormente, esta población se consideraba conespecíca con Norops
crassulus (Cope, 1864) de Guatemala, El Salvador y México, a pesar de ser completamente disyunto (> 100 km) de cualquiera de esas poblacio-
nes. El trabajo molecular reciente reveló una especicidad mitocondrial y nuclear consistente y profunda entre esta población y todos los demás
Norops del grupo de especies de N. crassulus, lo que provocó una investigación morfológica exhaustiva de esta población. Esta nueva especie
es muy similar en morfología externa a N. crassulus sensu stricto, pero se distingue fácilmente por su distintividad molecular, distribución y
morfología.
Palabras claVE: Bloque Chortís, largatija, Mesoamerica, morfología, Centroamerica Nuclear, Reptilia, taxonomía
INTRODUCTION
The anoles (Squamata: Dactyloidae) of Central America’s
highlands are a ubiquitous component of the region’s di-
verse herpetofauna. Systematic studies based upon mo-
lecular and morphological evidence continue to reveal
underestimated, and often cryptic, diversity, helping to
resolve long-standing taxonomic problems and leading to
the descriptions of numerous new species (e.g., Köhler et
al. 2014a, 2016; Nicholson and Köhler 2014; Poe et al.
2015; Gray et al. 2016, Poe and Ryan 2017).
Norops crassulus (Cope, 1864) has long been a taxo-
nomically confusing highland anole (Stuart 1942, 1955;
Meyer and Wilson 1971). Currently, this taxon is assigned
to populations from a number of disjunct highland areas
across Nuclear Central America, with isolated populations
found at intermediate-to-high elevations (1300–3000 m
above sea level) in Chiapas, Mexico, throughout central
Guatemala, in the Salvadoran Cordillera, and in the Lenca
Highlands (departments of Intibucá, La Paz, and Lempira)
of Honduras (Köhler 2008; McCranie and Köhler 2015).
A fth population, previously referred to as Norops smin-
thus (Dunn and Emlen, 1932) (Nicholson et al. 2012),
N. crassulus (McCranie and Köhler 2015), or Anolis aff.
A. morazani Townsend and Wilson, 2009 (Hofmann and
Townsend 2017), is found in the Sierra de Agalta in De-
partamento de Olancho in eastern Honduras. Despite work
by Stuart (1942, 1955) and Meyer and Wilson (1971), N.
crassulus and the other members of its subgroup (sensu
McCranie and Köhler 2015) remain a complicated and un-
resolved group of anoles.
Hofmann and Townsend (2017) recently presented a
multilocus phylogenetic investigation into the evolution and
biogeography of the Norops crassulus subgroup, revealing
92 annals of carnEgiE musEum Vol. 85
cryptic diversity within the taxon N. crassulus that cor-
responded to disjunct populations following well-known
biogeographic breaks in Nuclear Central America. Most
distinct was the population of N. crassulus sensu lato from
the Lenca Highlands region of Honduras, which was esti-
mated to have diverged from populations in Guatemala, El
Salvador, and Mexico more than 13 million years before
present (YBP) (Fig. 1; Hofmann and Townsend 2017).
Uncorrected p-distances for mitochondrial DNA between
the Lenca population and the other lineages of N. crassu-
lus sensu lato (Table 1) were found to be comparable to
or greater than distances between other recently described
anoles and closely related taxa, and the distinctiveness of
this lineage was well-supported by mitochondrial and nu-
clear gene trees, as well as a coalescent species tree (Hof-
mann and Townsend 2017). This study also revealed that
populations attributed to N. crassulus in previous studies
represented an unnamed sister lineage to Norops anisolepis
(Smith et al., 1968), and was otherwise not closely related
to any of the populations assigned to N. crassulus. The
only other genetic sample assigned to N. crassulus that has
appeared in published phylogenetic studies, SMF 78830
from the Sierra de Agalta in Olancho, Honduras (McCra-
nie and Köhler 2015), was shown to represent a lineage
closely related to N. morazani (Hofmann and Townsend
2017).
In light of the evolutionary distinctiveness of the lineag-
es, we further investigated the taxonomic status of popula-
tions assigned to Norops crassulus sensu lato using analy-
ses of morphological data. Based on the deep evolutionary
history and historical biogeography of this lineage, and the
analyses of morphological variation presented herein, we
provide a formal description of the Lenca Highland popu-
lations of these anoles as a new species.
Fig. 1.—Distribution of Norops crassulus sensu lato used in this study, with inset phylogeny (summarized from Hofmann and Townsend 2017) showing
the divergence times and relationships of the four major lineages.
2018 hofmann and TownsEnd—a cryPTic nEw anolE from honduras 93
MATERIALS AND METHODS
Justication of Nomenclature
In the most comprehensive phylogeny of anoles published
to date, Poe et al. (2017) applied a PhyloCode approach
(Cantino and de Queiroz 2014) to dene clade names under
the crown clade Anolis Cope, 1864. Within the unranked
clade-based taxonomy of anoles of Poe et al. (2017), the
taxa discussed herein are beta anoles in the clade Draconura
Wagler, 1830 (formerly the auratus group of Nicholson et
al. 2012), which is part of the larger Norops clade. Most
recently, Nicholson et al. (2018) showed the concordance
between the clades of Poe et al. (2017) and the rank-based
multi-genera approach to anole classication. In order to
provide a valid taxonomy under the International Commis-
sion on Zoological Nomenclature (ICZN), Nicholson et al.
(2018) converted Poe et al. (2017)’s clade names to a rank-
based, multi-genera classication, which we follow here.
Table 1
Genetic distances (uncorrected pairwise distances) for the four lineages of Norops crassulus sensu lato, summarized from Hof-
mann and Townsend (2017). Intralineage distances for each gene are in the shaded box. Interlineage distances are in the lower-left
triangle. A dash (-) indicates sequence data is only available for a single individual for that gene; therefore, no intraspecic genetic
distances could be calculated.
Gene Lenca
Highlands
Salvadoran
Cordillera Guatemala Mexico
Lenca Highlands
16S 0.005
ND2 0.008
COI 0.007
PRLR 0.002
BDNF 0.000
PTPN12 0.001
Overall 0.002
Salvadoran
Cordillera
16S 0.057 0.001
ND2 0.169 -
COI 0.141 -
PRLR 0.020 -
BDNF 0.005 -
PTPN12 0.010 -
Overall 0.071 -
Guatemala
16S 0.064 0.008 0.001
ND2 0.138 0.084 0.016
COI 0.172 0.054 0.010
PRLR 0.016 0.009 0.008
BDNF 0.005 0.003 0.003
PTPN12 0.009 0.001 0.001
Overall 0.066 0.032 0.007
Mexico
16S 0.071 0.026 0.024 0.000
ND2 0.166 0.119 0.119 0.001
COI 0.138 0.091 0.088 0.001
PRLR 0.022 0.021 0.019 0.000
BDNF 0.003 0.002 0.002 0.000
PTPN12 0.010 0.005 0.004 0.001
Overall 0.069 0.052 0.048 0.001
94 annals of carnEgiE musEum Vol. 85
Taxon Sampling
Institutional abbreviations used here follow Sabaj (2016)
and are as follows: Carnegie Museum of Natural History,
Pittsburgh, Pennsylvania (CM); Museum of Vertebrate
Zoology Herpetology Collection, Berkeley, California
(MVZ:Herp); Natural History Museum of Los Angeles
County, Los Angeles, California (LACM); Senckenberg
Forschungsinstitut und Naturmuseum, Frankfurt am Main,
Germany (SMF); University of Florida, Florida Museum
of Natural History, Gainesville, Florida (UF); University
of Kansas Biodiversity Institute, Lawrence, Kansas (KU).
Specimens from Josiah H. Townsend’s eld series are not-
ed as JHT numbers.
Specimens of N. crassulus representing each of the four
geographically discrete molecular clades were received
from the KU and MVZ:Herp collections, and examined
alongside samples collected by the authors and their col-
laborators (Appendix 1). From 2006 through 2015, JHT
and various collaborators sampled localities in the Hon-
duran departments of Copán, Cortés, Francisco Morazán,
Intibucá, La Paz, Lempira, Ocotepeque, and Santa Bárbara
for N. crassulus and N. crassulus-like anoles. Specimens
were xed in 10% formalin and preserved in 70% etha-
nol, with a tissue sample preserved in SED buffer (20%
DMSO, 0.25 M EDTA, pH 7.5, NaCl saturated; Seutin et
al. 1991; Williams 2007; Mulcahy et al. 2016). Forest for-
mation denitions follow those of Holdridge (1967) and
names of forest types that appear capitalized follow those
dened by Townsend (2014). All GPS coordinates here-
in refer to WGS84 datum. Specimens from the Sierra de
Agalta, Dept. Olancho, Honduras, a population previously
assigned to “N. crassulus,” were not included as part of
this investigation as they are instead most closely related
to N. morazani; a separate study is underway to determine
the taxonomic status of that population.
Morphological Examinations
Morphometric and meristic data were recorded for each
specimen following Köhler (2014). Sex was determined
by the presence of hemipenes and/or a well-developed
dewlap. Female N. crassulus possess a dewlap, but it is
signicantly reduced compared to males (McCranie and
Köhler 2015). Measurements, counts, and descriptions
were recorded following Köhler (2014), though some ab-
breviations were changed for clarity. Köhler (2014) pro-
vided complete denitions and photographic examples
of all morphological characters taken. The rst author
recorded all characters to eliminate any potential inter-ob-
server biases. In total, 103 specimens were examined, of
which data from 100 were used in analyses. The three ex-
cluded specimens were considered juveniles or hatchlings.
Comparative data for other Honduran crassulus subgroup
anoles were taken from McCranie and Köhler (2015) and
each species’ original description.
Fifteen measurements were taken using either a stiff
ruler (for snout-vent length and tail length) or digital
calipers to the nearest 0.01 mm (all other measurements):
snout-vent length (SVL), tail length (TL), head length
(HL), head width (HW), snout length (SL), axilla-groin
distance (AGD), vertical tail diameter (VTD), horizon-
tal tail diameter (HTD), shank length (ShL), longitudinal
diameter of the ear opening (LDE), vertical diameter of
the ear opening (VDE), longitudinal diameter of the pari-
etal scale (LDP), transverse diameter of the parietal scale
(TDP), subdigital pad width (SPW), non-dilated subdigital
pad width (NDPW).
Twenty-one scale counts were recorded: lamellae on
the 4th toe (4TL, taken from both the left and right foot),
ventral scales in one head length (VHL) and dorsal scales
in one head length (DHL), ventral scales between the ax-
illa and groin (VSAG) and dorsal scales between the axilla
and groin (DSAG), scales at midbody (SMB), scales be-
tween the rst (S1C) and second (S2C) canthals, enlarged
supraocular scales (ESO), scales between the supraorbital
semicircles (SOC), scales between the supraorbital semi-
circles and the interparietal plate (SIP), postrostrals (PRS),
internasals (INS), supralabial scales to the level below the
center of the eye (SLS), infralabial scales to the level be-
low the center of the eye (ILS), loreal scales (LS), loreal
scale rows (LSR), subocular scales in contact with supral-
abial scales (both counted and presented as SO/SPL), post-
mentals (PMS), sublabials (SLB), enlarged dorsal scale
rows (EDS).
Twenty-two scale characters were recorded: condition
of the terminal phalanx (CTP), condition of the axillary
region (CAR), condition of the supradigital scales (CSD),
condition of the enlarged supraocular scales (CEO), con-
dition of the supraocular scales (CSO), condition of the
supraorbital semicircles (CSS), condition of the prefron-
tal depression (CFD), condition of the parietal depression
(CPD), relative size of the scales adjacent to the interpa-
rietal plate (SAP), condition of the snout scales (CSN),
condition of the canthal ridge (CR), circumnasal condi-
tion (CNC), prenasal condition (PNC), condition of the
supraciliary scales (SSC), size and condition of the scales
anterior (SAE) and posterior (SPE) of the ear opening,
condition of the outer postmentals (COP), condition of
the ventral (VSC) and dorsal scales (DSC), condition of
the dorsal (TDC), lateral (TLC), and ventral (TVC) caudal
scales. All counts were recorded from the specimens’ right
side; however, where bilateral asymmetry in the character
was apparent, both sides were recorded separately.
In male specimens with extended dewlaps, the number
of gorgetal rows was recorded (GSR), and the dewlap size
was approximated by noting its distal insertion relative
to the eye and proximal insertion relative to the axilla. In
male specimens with fully everted hemipenes, these or-
gans were described largely following Klaver and Böhme
(1986), as well as various descriptions of anole hemipenes
in the literature (e.g., McCranie and Köhler 2015; Köhler
et al. 2016). Following Köhler (2012), color of Honduran
specimens was recorded in life based on photographs and
2018 hofmann and TownsEnd—a cryPTic nEw anolE from honduras 95
also after approximately 30 months in alcohol. Color in
preservative of specimens from other populations was also
recorded.
Analyses
Statistical analyses of morphological data were performed
in R 3.3.1 (R Core Team 2016), and all analyses were
conducted on males and females separately to account for
sexual size dimorphism. In total, 60 males and 40 females
were included in these analyses. TL, VTD, and HTD were
not included in the analyses because many specimens had
incomplete tails, and GSR was not included due to the low
number of male specimens with extended dewlaps. Any
characters exhibiting no variation across specimens were
also left out.
The remaining measurements (except SVL) were re-
gressed against each specimen’s SVL to account for varia-
tion in body size. As in other recent studies of anoles (e.g.,
Köhler et al. 2014a, 2014b, 2016), SL and SHL were also
scaled against HL, and HL was scaled additionally against
HW. Nonparametic Mann-Whitney U tests (MWU, equiv-
alent to the Wilcoxon rank-sum test; Mann and Whitney
1947) were used to determine if any of 15 morphological
measurements and 21 scale counts were signicantly dif-
ferent between populations. Based on the molecular data
presented by Hofmann and Townsend (2017), specimens
from El Salvador and Guatemala were treated as one op-
erational taxonomic unit (OTU), and specimens from the
Lenca Highlands were treated as a second OTU. Because
of the low sample size, specimens from a divergent popula-
tion from Chiapas, Mexico were not included in the MWU
analyses. Principal Component Analyses (PCA) were then
performed separately on the 15 male and female morpho-
logical measurements to determine, in part, if variation in
morphology could be attributed to populations. In these
analyses, samples from the Lenca Highlands were consid-
ered one OTU, while specimens from all other populations
of N. crassulus sensu lato were considered a second OTU.
The data were log transformed prior to analysis and scaled
to their standard deviation in order to normalize their dis-
tribution. PCAs were carried out using ‘prcomp’ and plot-
ted using the ‘autoplot’ command in the package ggfortify
(Tang et al. 2016). According to Kaiser’s criterion, compo-
nents with eigenvalues >1.0 were retained (Kaiser 1960).
RESULTS
Morphological Analyses
Some variation in selected morphological measurements,
proportions, and scale counts was found among N. crassu-
lus populations (Table 2). Thirteen characters in males and
11 in females were signicantly different based on MWU
analyses (Appendix 2). HW/SVL, HL/HW, SHL/SVL,
SHL/HL, SPW/SVL, NDSPW/SVL, ESO, and INS were
signicant in both males and females. Additionally, SVL,
VEO/SVL, LDPS/SVL, DS1H, SMB, S1C, SOSIP, ILS,
and EDS were signicant in either males or females. The
rst ten principal components explained 95% of the cu-
mulative variance in both male and females (Appendix 3).
In males and females, HW/SVL had the strongest positive
loadings in PC1 (0.454 and 0.406, respectively). SL/SVL
(0.536), ShL/SVL (0.461), and SL/HL (0.390) had the
strongest positive loadings in PC2 for males, and HL/HW
(0.448), ShL/SVL (0.395), and VEO/SVL (0.393) were the
strongest in females. The two OTUs formed distinct clus-
ters that overlapped somewhat in morphospace (Fig. 2).
Coloration and pattern were variable across all specimens,
as were dewlap coloration, size, and GSR. No combination
of these traits reliably delimited specimens from any popu-
lation. Based on the examination of seven specimens with
fully everted hemipenes, hemipenial structure also appears
highly conserved between all populations, largely agree-
ing with the description of McCranie and Köhler (2015:
52). However, specimens from Mexico appeared to have
slightly broader lobes than others specimen examined, and
a specimen from El Salvador (KU 184065) appeared to
have a somewhat larger overall organ.
Molecular Evidence and Taxonomic Conclusion
Hofmann and Townsend (2017)’s previous analyses of
mitochondrial and nuclear loci under concatenated and
multispecies coalescent frameworks all unambiguously
recovered the Lenca Highland population of N. crassulus
as a lineage distinct from all other populations assigned
to that name (Fig. 1). The genetic distances between this
population and others assigned to N. crassulus (Table 1)
were notably larger than those separating many recently
described anoles from their closely related congeners for
the mitochondrial loci 16S (5.7–7.1% compared to 3.2–
4.4%; Köhler et al. 2016), ND2 (13.8–16.9% compared
to 7.8–12.5%; Poe et al. 2015; Gray et al. 2016; Grisales-
Martínez et al. 2017; Poe and Ryan 2017), and COI (13.8–
17.2% compared to 7.3%; Poe et al. 2015).
We follow the unied species concept (de Queiroz
2007) in that we dene species as separately evolving
metapopulation lineages, with additional lines of evidence
being relevant to species delimitation as “operational cri-
teria.” It is important that characters, such as morphol-
ogy, distribution, and ecological niche, be considered in
the delimitation of a species, with no one line of evidence
necessarily weighted more heavily than another. Given
the genetic distinctiveness, mid-Miocene divergence, al-
lopatric distributions promoting long-term reproductive
isolation, and subtle morphological differences, we con-
clude that the Lenca Highland population assigned to N.
crassulus represents a species-level lineage distinct from
populations in Guatemala, El Salvador, and Mexico. This
population represents a separately evolving lineage that
has undergone minimal phenotypic change, despite having
diverged from their most recent common ancestor more
96 annals of carnEgiE musEum Vol. 85
Table 2
Selected morphological character comparisons between Norops caceresae, sp. nov., and N. crassulus. Proportion data and averages
± standard deviations of measurements are rounded to the nearest 0.01. Averages ± standard deviations of count data are rounded
to the nearest 0.1. Measurements are in millimeters (mm).
N. caceresae, sp. nov. N. crassulus
LENCA HIGHLANDS SALVADORAN CORDILLERA GUATEMALA MEXICO
Male
(n=12)
Female
(n=9)
Male
(n=10)
Female
(n=6)
Male
(n=33)
Female
(n=24)
Male
(n=5)
Female
(n=1)
Maximum Snout-
Vent Length
(SVL)
57
(47.33 ± 7.20)
54
(49.22 ± 4.09)
49
(44.60 ± 3.03)
52
(45.67 ± 6.09)
55
(41.97 ± 5.39)
54
(44.00 ± 5.96)
47
(43.80 ± 4.66) 48
Complete TL/
SVL11.82–2.02
(1.93 ± 0.09)
1.60–2.00
(1.87 ± 0.14)
1.63–2.39
(2.15 ± 0.34)
2.11–2.26
(2.19 ± 0.06)
1.76–2.53
(2.17 ± 0.22)
1.54–2.44
(2.051 ± 0.27)
2.17–2.57
(2.407 ± 0.16) 1.94
Head Length/SVL 0.26–0.30
(0.28 ± 0.01)
0.25–0.28
(0.27 ± 0.01)
0.25–0.30
(0.27 ± 0.01)
0.25–0.29
(0.27 ± 0.01)
0.27–0.30
(0.29 ± 0.01)
0.24–0.31
(0.28 ± 0.02)
0.27–0.30
(0.28 ± 0.01) 0.27
Head Width/SVL 0.17–0.20
(0.18 ± 0.01)
0.17–0.18
(0.17 ± 0.01)
0.17–0.20
(0.18 ± 0.01)
0.17–0.21
(0.19 ± 0.02)
0.18–0.22
(0.20 ± 0.01)
0.16–0.21
(0.19 ± 0.01)
0.18–0.20
(0.19 ± 0.01) 0.17
Head Length/
Head Width
1.49–1.70
(1.58 ± 0.07)
1.50–1.70
(1.58 ± 0.07)
1.41–1.65
(1.48 ± 0.09)
1.35–1.56
(1.44 ± 0.08)
1.32–1.57
(1.45 ± 0.06)
1.32–1.60
(1.46 ± 0.07)
1.46–1.50
(1.48 ± 0.02) 1.54
Snout Length/SVL 0.13–0.15
(0.14 ± 0.01)
0.11–0.14
(0.13 ± 0.01)
0.12–0.14
(0.13 ± 0.01)
0.12–0.13
(0.12 ± 0.01)
0.12–0.15
(0.13 ± 0.01)
0.11–0.14
(0.13 ± 0.01)
0.12–0.14
(0.13 ± 0.01) 0.12
Snout Length/
Head Length
0.43–0.51
(0.48 ± 0.02)
0.43–0.532
(0.47 ± 0.03)
0.45–0.48
(0.47 ± 0.01)
0.45–0.48
(0.47 ± 0.01)
0.39-0.51
(0.46 ± 0.03)
0.43–0.53
(0.48 ± 0.03)
0.44–0.48
(0.45 ± 0.02) 0.46
Axilla-Groin
Distance/ SVL
0.38–0.45
(0.41 ± 0.03)
0.39–0.50
(0.43 ± 0.03)
0.36–0.47
(0.41 ± 0.04)
0.36–0.455
(0.43 ± 0.04)
0.32–0.45
(0.39 ± 0.03)
0.37–0.47
(0.42 ± 0.03)
0.39–0.43
(0.41 ± 0.02) 0.42
Shank Length/
SVL
0.24–0.29
(0.26 ± 0.01)
0.22–0.27
(0.25 ± 0.02)
0.22–0.25
(0.24 ± 0.01)
0.22–0.25
(0.23 ± 0.01)
0.22–0.27
(0.24 ± 0.01)
0.21–0.25
(0.23–0.01)
0.22–0.26
(0.24 ± 0.01) 0.22
Shank Length/
Head Length
0.84–0.98
(0.93 ± 0.04)
0.79–0.94
(0.90 ± 0.06)
0.83–0.95
(0.87 ± 0.04)
0.79–0.95
(0.87 ± 0.05)
0.74–0.93
(0.84 ± 0.05)
0.74–0.97
(0.84 ± 0.05)
0.82–0.91
(0.86 ± 0.04) 0.82
Subdigital lamel-
lae on Phalanges
II-IV of Toe IV
(Left and Right
side averaged)
22–29
(25.8 ± 1.8)
23–28
(24.9 ± 1.3)
20–26
(23.2 ± 2.0)
20–26
(22 ± 1.6)
20–27
(23.4 ± 2.0)
18–28
(22.6 ± 2.1)
22–26
(23.7 ± 1.2) 20, 23
Subdigital lamel-
lae on distal pha-
lanx of Toe IV
(Left and Right
side averaged)
6–9
(6.9 ± 0.9)
5–8
(6.7 ± 0.8)
5–8
(7.0 ± 0.9)
4–8
(6.3 ± 1.1)
5–8
(6.2 ± 1.0)
4–8
(6.3 ± 0.9)
5–8
(6.5 ± 0.8) 6
Number of ventral
scales in one head
length
18–22
(19.7 ± 1.7)
16–22
(17.6 ± 2.2)
16–32
(20.4 ± 4.7)
18–22
(19.7 ± 2.0)
14–26
(20.55 ± 3.1)
14–22
(17.8 ± 2.5)
20–30
(25.2 ± 4.1) 20
Number of dorsal
scales in one head
length
22–28
(24.5 ± 2.3)
16–24
(20.9 ± 2.5)
20–30
(23.6 ± 3.0)
22–30
(24.3 ± 3.2)
16–32
(25.2 ± 3.4)
10–36
(23.8 ± 6.1)
20–30
(25.2 ± 4.1) 30
Number of ventral
scales between the
levels of
axilla and groin
25–34
(29.3 ± 2.5)
26–33
(30.2 ± 2.5)
24–35
(30.8 ± 3.2)
28–31
(30.3 ± 3.9)
22–36
(29.8 ± 3.5)
23–40
(30.3 ± 4.2)
29–35
(31.8 ± 2.8) 27
Number of dorsal
scales between the
levels of axilla and
groin
34–43
(38.6 ± 2.4)
34–46
(39.6 ± 3.5)
28-49
(39.4 ± 6.0)
36–45
(41.3 ± 4.1)
29-46
(35.5 ± 4.9)
32–63
(42.5 ± 8.8)
32-41
(36.8 ± 3.3) 51
Number of scales
around midbody
88–106
(95.3 ± 6.5)
80–114
(91.6 ± 10.5)
84-106
(89.4 ± 12.7)
80–98
(88.3 ± 6.9)
66-120
(83.4 ± 11.9)
66–116
(84.8 ± 13.6)
60-76
(65.2 ± 6.3) 96
Number of scales
between first can-
thals
7–9
(8.3 ± 0.7)
7–9
(8.2 ± 0.8)
5–9
(7.3 ± 1.3)
6–9
(7.3 ± 1.2)
6–10
(7.5 ± 0.9)
7–9
(7.9 ± 0.7)
7–9
(8.2 ± 0.8) 8
2018 hofmann and TownsEnd—a cryPTic nEw anolE from honduras 97
Table 2
(continued from previous page)
N. caceresae, sp. nov. N. crassulus
LENCA HIGHLANDS SALVADORAN CORDILLERA GUATEMALA MEXICO
Male
(n=12)
Female
(n=9)
Male
(n=10)
Female
(n=6)
Male
(n=33)
Female
(n=24)
Male
(n=5)
Female
(n=1)
Number of
scales
between second
canthals
5–7
(5.9 ± 0.7)
5–6
(5.8 ± 0.4)
5–7
(5.9 ± 0.6)
5–7
(5.7 ± 0.8)
4–7
(5.5 ± 0.7)
5–8
(5.8 ± 0.9)
5–6
(5.4 ± 0.5) 6
Number of
scales between
supraorbital
semicircles
0–1
(0.9 ± 0.3)
1–2
(1.1 ± 0.3)
0–1
(0.8 ± 0.4)
0–1
(0.7 ± 0.5)
0–2
(0.9 ± 0.4)
0–2
(0.9 ± 0.4)
1
(1.0 ± 0.0) 0
Number of
scales between
interparietal
plate and
supraorbital
semicircles
2–3
(2.8 ± 0.4)
2–4
(2.8 ± 0.8)
1–4
(2.6 ± 0.8)
2–3
(2.2 ± 0.4)
1–3
(2.3 ± 0.5)
1–3
(2.3 ± 0.5)
2–3
(2.2 ± 0.4) 2
Number of
Postmentals
4–5
(4.1 ± 0.3)
4
(4.0 ± 0.0)
4–6
(4.3 ± 0.7)
3–6
(4.8 ± 1.3)
4–5
(4.1 ± 0.3)
4–6
(4.1 ± 0.4)
4
(4.0 ± 0.0) 4
Number of
Postrostrals
5–6
(5.4 ± 0.5)
5–7
(5.4 ± 0.7)
5–6
(5.3 ± 0.5)
5–7
(5.8 ± 0.8)
4–10
(5.4 ± 1.0)
4–6
(5.3 ± 0.6)
5–6
(5.2 ± 0.4) 5
Number of
Internasals
6–8
(7.3 ± 0.6)
6–9
(7.1 ± 0.9)
5–8
(6.3 ± 0.8)
6–7
(6.3 ± 0.5)
5–9
(6.5 ± 1.1)
4–8
(6.1 ± 0.9)
6–8
(7.2 ± 0.8) 7
Number of
Supralabials
to level below
center of eye
5–7
(6.0 ± 0.7)
4–6
(5.7 ± 0.5)
6–7
(6.5 ± 0.5)
5–7
(6.0 ± 0.63
5–7
(6.1 ± 0.6)
5–7
(6.0 ± 0.6)
5–6
(5.8 ± 0.4) 6
Number of
Infralabials
to level below
center of eye
5–8
(5.8 ± 0.8)
4–6
(5.2 ± 0.7)
5–7
(6.0 ± 0.7)
5–7
(6.0 ± 0.6)
4–7
(5. 7 ± 0.6)
5–7
(5.8 ± 0.7)
5–6
(5.8 ± 0.4) 7
Number of
enlarged
supraocular
scales
5–10
(7.1 ± 1.4)
5–9
(6.2 ± 1.4)
3–9
(5.2 ± 1.7)
3–6
(4.0 ± 1.3)
3–7
(5.3 ± 1.3)
2–8
(5.0 ± 1.5)
5–6
(5.6 ± 0.5) 6
Number of
Suboculars in
contact with
number of
supralabials
(SO/SL)
2–3/1–3
(2.2 ± 0.4/
2.5 ± 0.7)
1–3/1–3
(2.1 ± 0.6/
2.2 ± 0.7)
1–3/1–3
(2.3 ± 0.7/
2.0 ± 0.1)
2/1–2
(2.0 ± 0.0/
1.5 ± 0.5)
1–4/1–4
(2.5 ± 0.7/
2.3 ± 0.8)
2–3/1–3
(2.5 ± 0.5/
2.2 ± 0.5)
1–3 /1–2
(2.0 ± 1.0/
1.6 ± 0.5)
2/2
Number of
Loreal scales
17–24
(19.5 ± 2.3)
13–27
(18.3 ± 4.1)
17–30
(23.6 ± 4.1)
19–26
(22.2 ± 2.5)
12–36
(19.0 ± 4.9)
13–25
(17.5 ± 3.1)
18–23
(19.4 ± 2.1) 21
Number of
Loreal scale
rows
4–5
(4.1 ± 0.3)
3–5
(4.0 ± 0.9)
4–5
(4.7 ± 0.5)
4–6
(4.8 ± 0.8)
3–6
(4.2 ± 0.6)
3–5
(4.1 ± 0.7)
4–5
(4.6 ± 0.5) 5
Number of
Sublabials
2–4
(2.4 ± 0.7)
2–4
(2.9 ± 0.9)
2–5
(2.9 ± 1.1)
2
(2.0 ± 0.0)
2–5
(2.7 ± 0.9)
2–8
(2.8 ± 1.4)
2–4
(2.8 ± 0.8) 2
Number of
Enlarged
Dorsal Scale
Rows
12–19
(17.2 ± 1.9)
12–20
(16.7 ± 2.9)
12–16
(13.5 ± 1.4)
12–15
(12.5 ± 1.2)
13–19
(15.2 ± 1.4)
12–20
(15.4 ± 2.3)
12–17
(14.4 ± 2.1) 14
Number of
Gorgetal scale
rows2
6–7
(6.3 ± 0.5) - - - 6–8
(6.7 ± 1.0) -5–7
(6.2 ± 0.8) -
Note.1 n = 5 (male), 4 (female) (Salvadoran Cordillera); 24, 16 (Guatemala); 7, 6 (Lenca Highlands); 5, 1 (Mexico). 2 n = 0 (Salvadoran Cordillera);
7 (Guatemala); 11 (Lenca Highlands); 5 (Mexico).
98 annals of carnEgiE musEum Vol. 85
than 13 million years ago (Fig. 1; Hofmann and Townsend
2017). As the type locality of N. crassulus is Cobán, Alta
Verapaz, Guatemala (Cope 1864; Stuart 1942) the name
N. crassulus sensu stricto is restricted to those populations
occurring in the highlands of Guatemala, El Salvador, and
Mexico, with the understanding that further taxonomic in-
vestigation of the populations in Chiapas, Mexico, remains
necessary and may lead to the recognition of additional
taxa. As the Lenca Highland anoles previously assigned
to N. crassulus are not conspecic with that taxon, and no
applicable names are available in synonymy, we describe
this population as a new species below.
SYSTEMATICS
Class Reptilia Laurenti, 1768
Order Squamata Oppel, 1811
Family Dactyloidae Fitzinger, 1843
Subfamily Anolinae Cope, 1864
Genus Norops Wagler, 1830
Norops caceresae, new species
English Common Name: Berta’s Anole
Spanish Common Name: Pichete de Berta
(Figs. 3–5)
Anolis crassulus—Hahn, 1971:111; Meyer and Wilson 1971:106;
1973:16; Wilson 1983:125; Wilson and Townsend 2007:135;
Townsend and Wilson 2009:64; Wilson and Johnson 2010:125 (in
part); Townsend and Wilson 2010:474 (in part).
Norops crassulus—McCranie et al. 1992:208; Wilson and McCra-
nie 1994:148; 2002:91; 2003a:24 (in part); 2003b:37; Köhler and
Obermeier 1998:127 (in part); Köhler 2008:104 (in part); Solís
et al. 2014:129 (in part); Townsend 2014:239 (in part); Johnson et
al. 2015:81 (in part); McCranie 2015:368 (in part); McCranie and
Köhler 2015:49 (in part).
N. crassulus (Honduras)—Köhler et al. 1999:286.
A. crassulus (Honduras)—Hofmann and Townsend 2017:8.
Holotype.—CM 161315 (eld number JHT 3914), Fig-
ures 3 and 4, an adult male, from near Río Agua Negra,
14.459°N, 88.385°W, 1,940 m above sea level, Reserva
Biológica Opalaca, Departamento de Intibucá, Honduras
(Fig. 6), collected on 28 May 2015 by Thomas J. Firneno
Jr., Michael W. Itgen, Josiah H. Townsend, and Kayla D.
Weinfurther.
Paratypes.—(n = 22; 12 males, 10 females); CM
161303–161305, 161308, MVZ:Herp:286121–286122,
KU 348836, KU 348838 [JHT 3739, 3741], (eld num-
bers JHT 3738–39, 3741–43, 3764, 3767, 3781), adult
males, and CM 161301–161302, 161306–161307,
MVZ:Herp:286123–286124, KU 348835, KU 348837
[JHT 3737, 3740] (eld numbers JHT 3735–37, 3740,
3766, 3768–70), adult females, from near El Rodeo,
14.441°N, 88.145°W, 2100 m above sea level, Refugio
de Vida Silvestre Mixcure, Departamento de Intibucá,
Honduras, collected 23 May 2015 by Thomas J. Firneno
Jr., Michael W. Itgen, Josiah H. Townsend, and Kayla D.
Weinfurther; CM 161310, 161312–161314 (eld numbers
JHT 3887, 3906, 3910–11), adult males, CM 161309 (eld
number JHT 3822), an adult female, and CM 161311 (eld
number JHT 3888), a subadult female, with the same col-
lection data as the holotype.
Referred specimens.—(n = 78); KU 219977, from 9.8 km
SW of Siguatepeque, Depto. de Comayagua, Honduras;
UF 166190–91, and KU 194268–70, from Zacate Blanco,
Depto. de Intibucá, Honduras; SMF 86963–67, from near
El Rodeo, Depto. de Intibucá, Honduras; KU 219950–53,
from 18.1 km NW of La Esperanza, Depto. de Intibucá,
Honduras; KU 194271, from 11 km NW of La Esperanza,
Depto. de Intibucá, Honduras; UF 103410–11, from 17 km
NE of La Esperanza, Depto. de Intibucá, Honduras; LACM
47683–85, from 2.4 km ENE of La Esperanza, Depto. de
Intibucá, Honduras; LACM 47686–90, 4.8 km ENE of La
Esperanza, Depto. de Intibucá, Honduras; LACM 47691,
from 12.9 km ENE of La Esperanza, Depto. de Intibucá,
Honduras; UF 121773, about 10 km SE of La Esperanza,
Depto. de Intibucá, Honduras; CM 59118–19 and LSUMZ
38816–24, from La Esperanza, Depto. de Intibucá, Hondu-
ras; KU 219954, from San Pedro La Loma, Depto. de Inti-
bucá, Honduras; FMNH 283624–27, from Santa Catarina,
Depto. de Intibucá, Honduras; UF 166192, from Cerro El
Pelón, Depto. de Intibucá, Honduras; UF 166185–166189
Fig. 2.—Plots of the rst two principal components for males (A) and
females (B). Colors correspond to lineages as in Figure. 1.
2018 hofmann and TownsEnd—a cryPTic nEw anolE from honduras 99
(but see Remarks for UF 166189) from near Guajiquiro,
Depto. de La Paz, Honduras; KU 184084–87, from Cantón
Palo Blanco, Depto. de La Paz, Honduras; KU 1584072–
77, from Cantón El Zancudo, Depto. de La Paz, Honduras;
SMF 78092 from Opatoro, Depto. de La Paz, Honduras;
KU 219955–56, 219978–81 from mountains S of San Pe-
dro de Tutule, Depto. de La Paz, Honduras; KU 194300–
04, from about 5 km S of Santa Elena, Depto. de La Paz,
Honduras; FMNH 236387, KU 209323–26, and 219957–
60, from Narantojos, Depto. de Lempira, Honduras.
Diagnosis.—A medium-sized species (maximum record-
ed snout-vent length 57 mm in males: KU 348838, CM
161310, CM 161315; 54 mm in females: KU 348837,
CM 161307) of beta anole in the genus Norops endemic
to southwestern Honduras. Norops caceresae can be dif-
ferentiated from all other anoles in Honduras, except those
of the N. crassulus species subgroup, by the combination
of having enlarged middorsal scales, strongly keeled ven-
tral scales, two or fewer scales separating the supraorbital
semicircles, four to seven loreal scale rows, suboculars and
supralabials in contact, heterogeneous lateral body scales,
enlarged postcloacal scales in males, and an orange-to-red
male dewlap (in life). Norops caceresae can be diagnosed
from the other species of the Norops crassulus species
subgroup, except N. crassulus sensu stricto, as follows
Fig. 3.—Dorsal and ventral body views of CM 161315, the holotype of Norops caceresae, sp. nov.
100 annals of carnEgiE musEum Vol. 85
(known distributions in parentheses): from N. amplisqua-
mosus McCranie et al., 1993 (northwestern Cortés, Hon-
duras), N. heteropholidotus (Mertens, 1952) (northwestern
El Salvador, southeastern Guatemala, and southwestern
Honduras), N. muralla (Köhler et al., 1999) (northwestern
Olancho, Honduras), N. sminthus (Francisco Morazan and
southern Comayagua, Honduras), and N. wermuthi Köhler
and Obermeier, 1998 (northern Nicaragua and extreme
southeastern Honduras) by its strongly keeled ventral
scales (versus smooth to weakly keeled); from N. anisol-
epis (Chiapas, Mexico) by its larger size (maximum SVL
57 mm in males, 54 mm in females; versus 47 mm in male,
48 mm in female N. anisolepis) and its orange-to-red male
dewlap (versus bright pink male dewlap in N. anisolepis);
from N. haguei (Stuart, 1942) (Alta Verapaz, Guatemala)
by its larger dorsal scales (34–43 between level of axilla
and groin, approximately equal to or slightly smaller in
size than ventrals; versus 41–57, dorsals much smaller
than ventrals in N. haguei); from N. morazani (northern
Francisco Morazán, Honduras) by having a hemipenis
with an undivided asulcate processus (versus divided
asulcate processus in N. morazani), and having 6–9 scales
separating the nasals (2–4 in N. morazani); and from N.
rubribarbaris Köhler et al., 1998 (Santa Bárbara, Hondu-
ras) by having a TL/SVL 1.82–2.02 in males, 1.60–2.00 in
females (versus 2.16–2.54 in males, 2.19–2.21 in females
of N. rubribarbaris), and having 12–20 rows of enlarged
dorsal scales (versus 8–11 in N. rubribarbaris).
Norops caceresae is most similar in external mor-
phology to N. crassulus sensu stricto (highlands of cen-
tral Guatemala, southwestern El Salvador, and Chiapas,
Mexico). Norops caceresae is distinguished from all other
populations assigned to N. crassulus by the following: (1)
a longer head relative to width: HL/HW 1.49–1.70 (1.58 ±
0.07) in males, 1.50–1.70 (1.58 ± 0.07) in females (versus
1.32–1.65 [1.46 ± 0.06] in males, 1.32–1.56 [1.46 ± 0.07]
in females of N. crassulus); (2) a shorter tail relative to
snout–vent length: complete TL/SVL 1.79–2.02 (1.91 ±
0.01) in males, 1.60–2.00 (1.87 ± 0.14) in females (ver-
sus 1.62–2.37 [2.20 ± 0.24] in males, 1.54–2.44 [2.07 ±
0.24] in females of N. crassulus) (3) a longer shank rela-
tive to snout-vent length: ShL/SVL: 0.24–0.29 (0.26 ±
0.01) in males, 0.22–0.27 (0.25 ± 0.02) in females (versus
0.22–0.27 [0.24 ± 0.01] in males, 0.21–0.25 [0.23 ± 0.01]
in females of N. crassulus); (4) a longer shank relative to
head length: ShL/HL: 0.84–1.13 (0.94 ± 0.07) in males,
0.79–0.94 (0.92 ± 0.06) in females (versus 0.74–0.95 [0.83
± 0.12] in males, 0.74–0.97 [0.84 ± 0.05] in females of N.
crassulus); (5) a slightly higher average number of lamel-
lae on Phalanges II-IV of the 4th toe: 25.7 ± 1.8 in males,
24.9 ± 1.3 in females (versus 23.3 ± 1.9 in males, 22.5 ±
2.0 in females of N. crassulus).
Description of the holotype.—Adult male (Figs. 3-4),
medium sized, hemipenes everted, right thigh with skin
and thigh muscle removed; tail complete; SVL 57 mm, tail
length 114 mm, tail slightly laterally compressed, vertical
diameter 3.46 mm, horizontal diameter 2.25 mm, twice as
long as SVL; head length (HL) 15.87 mm, 27.8% of SVL;
head width 9.66 mm, head 1.64 times as long as wide;
snout 7.32 mm, 46.1% of head length, 12.8% of SVL; ax-
illa-groin distance 21.64 mm, 38.0% of SVL; shank length
15.49 mm, 27.2% of SVL, 97.6% of HL; dilated subdigital
pad width 1.01 mm, non-dilated 0.46 mm; when the hind
limb is adpressed, the longest digit reaches approximately
to the middle of the eye.
Scales on dorsal surface of snout large, irregular, keeled;
seven scales between nasals, keeled; one scale separates
nasal from rostral; prenasal divided in vertical series, the
lower in contact with the rostral and rst supralabial and
Fig. 4.—Dorsal, lateral, and ventral views of the head of CM 161315, the
holotype of Norops caceresae, sp. nov.
2018 hofmann and TownsEnd—a cryPTic nEw anolE from honduras 101
the higher in contact with neither; six postrostrals; distinct
frontal depression with scales keeled radially; well-devel-
oped supraorbital semicircles consisting of six enlarged,
irregular scales, three smooth and three with weak keels;
one keeled scale separates supraorbital semicircles at their
closest point; shallow parietal depression, with scales ru-
gose, irregular; interparietal separated from supraorbital
semicircles by three scales; scales on the posterior portion
of the head irregular, rugose to weakly keeled; six moder-
ately enlarged, keeled, supraocular scales; two elongate,
keeled superciliaries; distinct canthal ridge, nine scales be-
tween rst canthals, six between second; 22 weakly keeled
loreal scales in four rows; 6/5 keeled suboculars, two in
contact with the posterior-most supralabial; 6/6 supral-
abials to the level below center of eye; mental wider than
long, completely divided by medial suture, bordered by 4
postmentals with the outer pair greatly enlarged; 6/5 in-
fralabials to the level below center of eye, longest approxi-
mately equal to longest supralabial; chin scales smooth to
weakly keeled, juxtaposed, grading posteriorly into larger,
weakly keeled, imbricate throat scales; middorsal scales of
the neck weakly elevated, forming indistinct crest; scales
anterior to the ear opening keeled and distinctly larger
than those posterior to the ear opening, which are granu-
lar; dorsum of body covered in keeled, subimbricate to im-
bricate scales, with approximately 12 irregular, enlarged
medial dorsal scale rows, largest scales 0.73 x 0.79 mm,
posterior edge rounded or weakly pointed, approximately
24 enlarged dorsal scales in one head length, 37 between
the level of the axilla and groin; dorsals grade into subim-
bricate, heterogeneous lateral scales, then further into
granular, mostly homogeneous scalation; ventrals strongly
keeled, imbricate, pointed, the largest of which are slightly
larger than dorsals (0.9 x 0.9 mm), approximately 20 ven-
trals in one head length, 29 between the axilla and groin;
92 scales at midbody; caudal scales strongly keeled, im-
bricate; dorsal caudal scales slightly enlarged, three scales
per caudal section; postcloacals enlarged; scales on limbs
strongly keeled, imbricate, pointed on forelimbs and upper
half of hindlimbs, ends grading to rounded on the shank of
the hindlimb; scales of ngers and toes distinctly keeled,
multicarinate and parallel; distal phalanx of toes narrower
Fig. 5.—Adult male examples of Norops caceresae, sp. nov., in life. A, adult male from El Rodeo (14.460°N, 88.1585°W), 2,160 m elevation, Depar-
tamento de Intibucá, Honduras, showing the typical coloration of the male dewlap (JHT 2913). B-D, In situ from Yamaranguila (14.364°N, 88.263°W),
2,060 m elevation, Departamento de Intibucá (not collected). Photo A by César Cerrato-Mendoza; photos B–D by Josue Ramos Galdamez.
102 annals of carnEgiE musEum Vol. 85
than and raised above dilated subdigital pad, 29/26 lamel-
lae under phalanges II to IV of fourth toe, 8/9 lamellae
under the distal phalanx; axillary pocket absent; dewlap
large, six gorgetal rows consisting of 6–8 large, keeled gor-
getals, anterior position below the anterior eye opening,
posterior position extends to 7 mm beyond the axilla.
The completely everted hemipenis is a medium-sized,
partially-bilobed organ (lobes connected by thin tissue
through the apex); lobes are somewhat calyculate; an undi-
vided, nub-like asulcal processus is evident approximately
where the two lobes meet, halfway down the apex; truncus
has transverse folds and is approximately equal in length
Fig. 6.—Type locality habitat of Norops caceresae, sp. nov. A, photograph of Reserva Biológica Opalaca, Departamento de Intibucá, Honduras; B,
typical habitat of Norops caceresae. The holotype and some paratypes were collected while sleeping on low vegetation at night along this stream;
photos by Josiah Townsend.
2018 hofmann and TownsEnd—a cryPTic nEw anolE from honduras 103
to the length of each lobe; well-developed sulcal lips sur-
round the sulcus spermaticus, which opens into a broad
area covering the apex between lobes.
After approximately 30 months in preservative, the col-
oration was recorded as follows: dorsal surfaces of head
and body with Hair Brown (277) ground color, with one
pair of well-dened and one pair of poorly-dened Van-
dyke Brown (281) scallops edged in Pratt’s Payne’s Gray
(293), with intervening middorsal area Pratt’s Payne’s
Gray (293); lateral surfaces of head Hair Brown (277) with
supralabial region Dark Pearl Gray (290) and a Dark Pearl
Gray (290) spot directly posterior of eye; infralabial region
speckled Vandyke Brown (281) and Pearl Gray (262); dor-
solateral surfaces with broad Pearl Gray (262) line origi-
nating just anterior of the ear opening and extending to the
level of the adpressed elbow; lateral surfaces of body Glau-
cous (272); ventral surface Pearl Gray (262) with Smoke
Gray (266) suffusions; dorsal surfaces of limbs Glaucous
(272) with Hair Brown (277) suffusions; hind limbs with
paired Sepia (279) crossbands on the medial portions of
the calves; ventral areas of limbs Pearl Gray (262) with
Smoke Gray (266) and Grayish Horn Color (268) suffu-
sions more apparent than on ventral surface of body; dew-
lap extended in preservation, but color completely faded to
Pale Buff (1).
Fig. 7.—Phylogeny of the Norops crassulus species subgroup based on three mitochondrial (16S, ND2, COI) and three nuclear loci (PRLR, BDNF,
PTPN12) showing the phylogenetic placement of Norops caceresae, sp. nov., re-annotated from Hofmann and Townsend (2017: g. S1). Nodes are
labelled with Bayesian posterior probabilities (left) and maximum likelihood bootstrap support values (right).
104 annals of carnEgiE musEum Vol. 85
Variation in paratypes.—Slight variation in meristic
characters is apparent in some paratypes. A median su-
ture only partially divides the mental in CM 161302,
CM 161310, MVZ:Herp:286123, opposed to completely
dividing it in all other paratypes. In most specimens, the
mental is in contact posteriorally with four postmentals,
two greatly enlarged outer and two smaller, longer than
broad inner; in one male MVZ:Herp:286122, one of the
postmentals is divided, creating a fth postmental. Most
paratypes have at most a single scale separating the supra-
orbital semicircles at the narrowest point; in one female,
CM 161307, there are two. Between two and three scales
separate the interparietal plate and the supraorbital semi-
circles; in two females, KU 348835 and CM 161307, there
are four. Most paratypes have between two and four sub-
labials; in one subadult female, CM 161311, and one adult
male, CM 161313, a small scale separates the outer left
postmental from the infralabial, leaving only a single sub-
labial. In two males (CM 161303 and 161305) and two fe-
males (KU 348837, MVZ:Herp:286124), the circumorbital
scales are incomplete, leaving some of the enlarged supra-
oculars in contact with the supraocular semicircles. Varia-
tion in the size of the scales around the midbody is evident,
with females exhibiting wider variation (80–114, 91.6 ±
10.5) than males (86–106, 94.6 ± 6.8). Females exhibited
larger dorsal scales relative to head length, as evidenced
by their count of dorsal scales in one head length (16–24,
20.9 ± 2.5) compared to males (20–28, 24.2 ± 2.5). In six
male paratypes (CM 161310, CM 161313–161314, KU
348838, MVZ:Herp:286121–286122), the dewlap extends
from under approximately the anterior edge of the eye to
5–8 mm past the axilla; in two other male paratypes (CM
161304–161305) it extends 3 mm past the axilla, and in
two additional male paratypes (CM 161303, KU 348836)
it extends from under approximately the middle of the eye
to 2 mm past the axilla.
Variation in color and dorsal pattern are evident in pre-
served male and female paratypes. Adult males typically
exhibited a dorsal coloration and pattern similar to that of
the holotype, with a gray-brown to yellow-brown ground
color and well-dened pale dorsolateral lines extending
from the side of the head onto the body. Dorsal ground
color between these dorsolateral lines is typically darker
than the ground color on the lateral portions of the body.
In most adult male paratypes, this line was more evident
than in the holotype, typically 2–8 dorsal scales wide and
terminating one-third to two-thirds the distance between
the limbs. All male paratypes possess a dorsal pattern con-
sisting of dark brown, paired scallops or chevrons, with
well-dened chevrons in the general shape of equilateral
triangles having the base formed by the dorsolateral stripe
and the vertices contacting the middorsal line. In some in-
dividuals (e.g., CM 161303, CM 161312), these scallops
are relatively pale and only apparent on the anterior half of
the body, while in others (e.g., KU 348836) they are dark,
outlined with pale cream lines, and possess a pale spot in
the center of each chevron. Most female paratypes exhibit
a similar coloration and pattern to that of males, although
in some females (e.g., CM 161306) the dorsolateral stripes
are poorly dened, and pattern is reduced to the outlines
of chevrons on the anterior half of the body. In other fe-
males, the scallops or chevrons only extend from the dor-
solateral stripe to about halfway to the middorsal line (e.g.,
KU 348835). In one female (CM 161311), the dorsolateral
stripes and dorsal scalloping are absent and replaced by a
broad, cream-colored middorsal stripe that is 2–4 enlarged
middorsal scales in width at midbody.
The dewlap coloration in life of two adult males (Fig.
5A, JHT 2913; MVZ:Herp:286122) from El Rodeo, De-
partamento de Intibucá, Honduras, were both Medium
Chrome Orange (75) to Spectrum Orange (9), with Light
Chrome Orange (76) around the margins, and Pale Buff
(1) to Smoky White (261) marginal, gorgetal, and sternal
dewlap scales.
Illustrations.—Figs. 3–5; McCranie and Köhler 2015: 54,
g. 16A (adult male) and 16B (adult male head and dew-
lap) (as Norops crassulus). Note that Köhler 2014: 247,
g. 27(I), 248, g. 28(C), and McCranie and Köhler 2015:
53, g. 15, are of specimens from Olancho, Honduras,
representing N. aff. N. morazani (Hofmann and Townsend
2017).
Distribution.—Norops caceresae occurs at moderate and
intermediate elevations (1,200–2,260 m) in the depart-
ments of Comayagua, Intibucá, La Paz, and Lempira in
southwestern Honduras. Its known distribution is bordered
to the west by the Río Ulúa and the south by Río Lempa.
Natural History.—Norops caceresae inhabits both intact
and disturbed Mixed Transitional Cloud Forest, Broad-
leaf Cloud Forest, and Mixed Cloud Forest between 1,200
and 2,260 m elevation. This diurnal species appears most
abundant near forest edges and along streams and is locally
abundant in the matrix of traditional agriculture and forest
patches that typies much of the Lenca Highlands (Fig. 6).
At night, N. caceresae can be encountered asleep on low
vegetation, typically >1.5 m above the ground. We have
observed this species in sympatry with N. heteropholidotus
around a large stream through relatively intact Mixed
Cloud Forest in the Cordillera de Opalaca in western In-
tibucá, Honduras, and with N. heteropholidotus and N.
laeviventris (Wiegmann, 1834) in and around small Mixed
Cloud Forest fragments at Cerro San Pedro La Loma in
eastern Intibucá.
Phylogenetic relationships.—An updated phylogeny of
the Norops crassulus subgroup is presented in Figure 7.
Norops caceresae is monophyletic in both mtDNA and
nDNA analyses, as well as in concatenated and multi-
species coalescent frameworks (Hofmann and Townsend
2017). Sequences of the mtDNA loci 16S, ND2, and COI,
as well as the nDNA loci PRLR, BDNF, and PTPN12,
for N. caceresae (generated by Hofmann and Townsend
2017) are available on GenBank (Appendix 4). Intraspe-
2018 hofmann and TownsEnd—a cryPTic nEw anolE from honduras 105
cic distances at each of these loci are: 0.5% (16S); 0.8%
(ND2); 0.7% (COI); 0.2% (PRLR); 0.0% (BDNF); 0.1%
(PTPN12). From populations of its closest relative, N.
crassulus sensu stricto, it is 5.7–7.1% divergent at 16S,
13.8–16.9% at ND2, 13.8–17.2% at COI, 1.6–2.2% at
PRLR, 0.3–0.5% at BDNF, and 0.9–1.0% at PTPN12. No-
rops caceresae is a member of the “crassulus” clade (con-
taining N. anisolepis and all N. crassulus populations from
Mexico, El Salvador, and Guatemala; sensu Hofmann and
Townsend 2017) of the Norops crassulus species subgroup
in the genus Norops.
Etymology.—The eponym caceresae is a noun in the gen-
itive case, and is given in honor of Berta Isabel Cáceres
Flores of La Esperanza, Departamento de Intibucá, Hon-
duras (approximately 25 km from the type locality). Berta
Cáceres was a community leader and environmental ac-
tivist who cofounded the Consejo Cívico de Organizacio-
nes Populares e Indígenas de Honduras (COPINH: Civic
Council of Popular and Indigenous Organizations of Hon-
duras), and, in the face of threats against her and her family
and the murders of friends and colleagues, led grassroots
efforts to unite communities against environmentally de-
structive actions and the privatization of native lands in
Honduras. She was awarded the Goldman Prize for Con-
servation in 2015, and continued her work organizing and
ghting for indigenous rights and environmental justice
until she was assassinated in her home in La Esperanza on
3 March 2016. We name this species as a small gesture to
honor Berta and to raise awareness for COPINH and their
work, while continuing to draw attention to the plight of
indigenous and environmental activists in Honduras, doz-
ens of whom have been murdered over the past decade.
Remarks.—Phylogenetic analyses of the Norops crassu-
lus subgroup revealed an unexpected cryptic sister lineage
to N. rubribarbaris (= N. aff. N. rubribarbaris; Hofmann
and Townsend 2017) comprised of two samples from
Dept. La Paz, near Guajiquiro. Of the two samples, only
16S and COI sequences were available, and no viable
tissue remained to sequence additional loci. Only one of
these samples, UF 166189, has been accessioned, and was
part of the series of N. crassulus examined by McCranie
and Köhler (2015); the other sample was deposited but not
accessioned and is presumed lost. UF 166189 is a juvenile
anole with enlarged middorsals, keeled ventrals, and het-
erogeneous lateral scalation, a brown dorsum with a dark
middorsal line, a white venter with some dark mottling,
and mottling on the limbs. These samples suggest a cryptic
lineage potentially exists in sympatry with the newly de-
scribed species, warranting further investigation.
DISCUSSION
With the description of N. caceresae, the Norops crassu-
lus species subgroup now consists of 11 species: N. am-
pliquamosus, N. anisolepis, N. caceresae, N. crassulus, N.
haguei, N. heteropholidotus, N. morazani, N. muralla, N.
rubribarbaris, N. sminthus, and N. wermuthi. Hofmann
and Townsend (2017) showed that this subgroup is mono-
phyletic with respect to other Norops and contains several
cryptic lineages in need of further study.
Norops caceresae is endemic to an area less than 10,000
km2, inhabiting areas above 1,200 m elevation that are as-
sociated with Mixed Transitional Cloud Forest, Broadleaf
Cloud Forest, and Mixed Cloud Forest. This species is
under some degree of threat from anthropogenic habitat
destruction, but appears to be tolerant of anthropogenic
disturbance and able to utilize edges and fence-lines as-
sociated with agricultural clearings. This anole is relatively
abundant where found, and under no direct threat from hu-
man exploitation. Following the IUCN Red List criteria
(2001), we would consider this species to be Least Con-
cern/Near Threatened.
Norops caceresae is notably conserved in external mor-
phology relative to its sister taxon, N. crassulus. Cryptic
species are a critical, yet poorly-reected consideration in
both biodiversity conservation and the exploration of eco-
logical and evolutionary processes (Bickford et al. 2007;
Jörger and Schrödl 2013; Fišer et al. 2018). Many cryptic
species are discovered—often incidentally through broad-
er phylogenetic studies—but left unnamed, masking them
from studies that rely on Linnean binomials as hypotheses
or variables (Fišer et al. 2018; Struck et al. 2018). There-
fore, it is important that such genetically-distinct lineages
are thoroughly examined through integrative taxonomy, in
an effort to better understand natural processes. Recent re-
views by Fišer et al. (2018) and Struck et al. (2018) have
summarized the mechanisms for the formation of cryptic
species: recent divergence, phylogenetic niche conser-
vatism (stasis), convergence, and parallelism. As the age
and relatedness of the externally-similar members of the
Norops crassulus species group belie a recent divergence
or convergent evolution (Hofmann and Townsend 2017),
phylogenetic niche conservatism seems the most likely
means of speciation across these taxa. Allopatric specia-
tion caused by complex geologic history is one of the
dominant drivers of the herpetofaunal biodiversity in the
Chortís Highlands and can lead to morphological stasis, as
is well-documented in Plethodontid salamanders (Rovito
et al. 2012; Rovito 2017; Townsend et al. 2011).
The elevated region that spreads across the departments
of La Paz, Intibucá, and Lempira, geologically linked and
referred to collectively as the Lenca Highlands, has pre-
viously been recognized as being unstudied in terms of
herpetofauna diversity (McCranie 2007). Recent focus on
eldwork in the Lenca Highlands has begun to address this
deciency (McCranie 2014) and has begun to reveal pre-
viously overlooked endemic diversity. In addition to the
description of N. caceresae, Luque-Montes et al. (2018)
recently described a new species of true frog, Rana lenca,
from this region, based on populations previously referred
to as hybrids between the lowland species Rana browno-
rum Sanders, 1973 and R. cf. R. forreri Boulenger, 1883,
106 annals of carnEgiE musEum Vol. 85
and the description of a new species of Bolitoglossa (It-
gen et al. in preparation) endemic to the Lenca Highlands
is also underway. These three taxa will join the salaman-
ders B. celaque McCranie and Wilson, 1993, B. heiroreias
Greenbaum, 2004, B. synoria McCranie and Köhler, 1999,
and Oedipina chortiorum Brodie et al., 2012, the treefrog
Exerodonta catracha (Porras and Wilson, 1987), and the
lizards Abronia montecristoi Hidalgo, 1983, A. salvador-
ensis Hidalgo, 1983, and N. heteropholidotus as part of the
endemic herpetofauna of the Lenca Highlands and adja-
cent areas.
ACKNOWLEDGMENTS
Fieldwork in Honduras was carried out under a series of research permits
issued by the Instituto Nacional de Conservación y Desarrollo Forestal,
Áreas Protegidas y Vida Silvestre [ICF] (Resolución GG-MP-055-2006
and Dictamen DAPVS 0091-2006; Resolución DE-MP-086-2010 and
Dictamen DVS-ICF-045-2010; Resolución DE-MP-095-2014 and Dicta-
men ICF-DVS-112-2014). Invaluable logistical support was provided in
2008, 2009, and 2015 by the Centro Zamorano de Biodiversidad, and we
thank Jorge Iván Restrepo and José Mora (2008–09) and Oliver Komar
and Karla Lara (2015) for facilitating our eldwork. We thank Carol L.
Spencer, Jimmy A. McGuire, and David B. Wake (MVZ) and Rafe M.
Brown, Rich E. Glor, and Luke J. Welton (KU) for specimen and tis-
sue loans from their respective institutions. Cesár Cerrato-Mendoza and
Josue Ramos Galdamez contributed photos in life of N. caceresae, and
Coleman M. Sheehy III (Florida Museum of Natural History) kindly pro-
vided photographs of UF 166189. We are especially grateful to Kaylin
Martin (Carnegie Museum of Natural History [CM]) for her detailed pho-
tographs of the holotype used to prepare Figures 3 and 4, and to her and
Steve Rogers (CM) for cataloging the type series. We are indebted for
the eld assistance provided by Jason M. Butler, César Cerrato, Thomas
J. Firneno, Luis Herrera, Alexander Hess, Michael Itgen, Mariah Ken-
ney, Catherine Krygeris, Ileana Luque-Montes, Melissa Medina-Flores,
Lesster Munguía, Jorge Luis Murillo, Fatima Pereira-Pereira, Sandy
Pereira-Pereira, Josué Vasquez, Hermes Vega, Kayla Weinfurther, and
Larry David Wilson, and the residents of Agua Negra, El Rodeo, Gua-
jiquiro, and Zacate Blanco. Results included in this paper were obtained
with support from the Critical Ecosystem Partnership Fund, Indiana
University of Pennsylvania (IUP) Department of Biology, IUP College
of Natural Sciences and Mathematics, IUP School of Graduate Studies
and Research, IUP Faculty Senate, Commonwealth of Pennsylvania Uni-
versity Biologists (CPUB), and a Pennsylvania State System of Higher
Education Faculty Professional Development Grant. We thank Gunther
Köhler, Kirsten Nicholson, and Larry David Wilson for insightful discus-
sions of current anole taxonomy, as well as two anonymous reviewers for
comments and suggestions that improved this manuscript.
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———. 2010. Conservation of the Honduran herpetofauna: issues
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———. 2003a. The conservation status of the herpetofauna of Honduras.
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———. 2010. The herpetofauna of Mesoamerica: biodiversity signifi-
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108 annals of carnEgiE musEum Vol. 85
aPPEndix 1
Specimens examined. An asterisk (“*”) following a museum voucher number indicates the specimen was also included in the molecular
analyses of Hofmann and Townsend (2017). If an asterisk follows a series (e.g. “A–D”*) or country name, all specimens were used in
molecular analyses.
Norops caceresae, sp. nov. (23 specimens): HONDURAS*: Intibucá: Refugio de Vida Silvestre Mixcure, El Rodeo: CM 161301–
161308; KU 348835–348838, MVZ:Herp:286121–286124; Reserva Biológica Opalaca, Río Agua Negra: CM 161309–161315.
Norops crassulus (81 specimens): EL SALVADOR: Santa Ana: Volcan de Santa Ana, Lago Sur: KU 184063–184071, KU 184078–
184083; Finca El Milagro: KU 289793*. GUATEMALA: Alta Verapaz: Tactic: KU 190610; near Caquipec Mines, 20 km E and 6
km S Coban: MVZ:Herp:143408–143409; 25 km NW (by road) Senahu, Finca Volcan: MVZ:Herp:159923. Baja Verapaz: vicinity
Union Barrios: KU 190619; 1.9 km NNW La Union Barrios: KU 190611–190613; 3 km NW Purulha: KU 190614–190616; 3 km N
Niño Perdido: KU 190623; “156.5 km” [likely in error] ESE of Purulha, Cerro Quisis: MVZ:Herp:160512–160513, 160516–160517;
2.5 mi NE of Chilasco, Finca San Jorge: MVZ:Herp:160518–160523, 160529–160530, 160532, 160534–160538; 5 km ENE of Chi-
lasco, Finca San Jorge: MVZ:Herp:160539–160540; 5 km ESE of Purulha, Finca Mirador: MVZ:Herp:160524–160526; 6.5 km ESE
of Purulha, Finca Mirador: MVZ:Herp:160548. Chimaltenango: Cerro Balamjuyu, 12.5 km SW (by rd) highway from Patzitzia to
Patzún: MVZ:Herp:270039*. El Quiche: 3.3 km NE (by road) of Laj Chimel on road to San Pablo el Baldío: MVZ:Herp:265576*;
3.5 km NE (by road) of Laj Chimel on road to San Pablo el Baldío: MVZ:Herp:265564–265565*, 265577–265578*; 4.9 km NE (by
road) of Laj Chimel on road to San Pablo el Baldío: MVZ:Herp:265579–265580*. Huehuetenango: 3 km NNW of San Mateo Ixtatan:
MVZ:Herp:160563–160566; 2 km W San Mateo Ixtatan: MVZ:Herp:171455–171456, 171554–171557; 4 km W San Mateo Ixtatan:
MVZ:Herp:171553. Jalapa: Cerro Miramundo, ~1 km S of Mataquescuintla-Miramundo: MVZ:Herp:263595*. Zacapa: Sierra de
las Minas, 7.8 km NNW San Lorenzo: KU 190619–190621; 3.4 km WSW San Lorenzo: KU 190622. MEXICO*: Chiapas: Cerro
Boqueron, 0.8 km WSW (by air) of Ejido Boqueron, Municipio Motozintla: MVZ:Herp:256801–256806.
2018 hofmann and TownsEnd—a cryPTic nEw anolE from honduras 109
aPPEndix 2
Mann-Whitney U test summary statistics and P-values. Boldface values are signicant (p<0.05).
Male Female
U p-value U p-value
SVL 175 0.0917 73 0.0394
HL/SVL 232.5 0.6101 143.5 0.7895
HW/SVL 416 0.0013 232 0.0013
HL/HW 38.5 <0.0001 27 0.0003
SL/SVL 199.5 0.2363 155.5 0.5042
SL/HL 210 0.3329 150 0.6286
AG/SVL 196 0.2099 114 0.4943
SHL/SVL 42.5 <0.0001 57 0.0097
SHL/HL 53.5 <0.0001 557 0.0098
LEO/SVL 230 0.5745 146 0.7247
VEO/SVL 236 0.6603 67.5 0.0251
LDPS/SVL 390 0.0072 160 0.4124
TDPS/SVL 260 0.9759 134.5 1.0000
SPW/SVL 407 0.0023 194.5 0.0468
NDSPW/SVL 360 0.0302 194 0.0464
VS1H 295 0.4409 157.5 0.4497
DS1H 285 0.5778 196.5 0.0381
VSAG 292 0.4931 126 0.7760
DSAG 168.5 0.0689 157 0.4722
SMB 104.5 0.0018 89.5 0.1326
S1C 125.5 0.0045 96.5 0.1799
S2C 205 0.2323 127.5 0.7994
ESO 92.5 0.0005 69 0.0260
SOS 243 0.6453 100.5 0.1070
SOSIP 142 0.0080 86 0.0558
PR 233.5 0.5609 140.5 0.8498
IN 133.5 0.0079 60.5 0.0074
SLS 294 0.4167 169.5 0.1627
ILS 265 0.8761 195 0.0283
LS 254 0.9430 144 0.7759
LSR 316 0.1539 157.5 0.4289
SO 328 0.1087 168 0.2021
SL 206.5 0.2620 113 0.3894
PMS 267 0.7486 148.5 0.4551
SLB 308.5 0.2596 103.5 0.2253
EDS 78 0.0002 81.5 0.0745
110 annals of carnEgiE musEum Vol. 85
aPPEndix 3
Principal component scores for the rst ten PCs from male and female morphological measurement analyses.
Male PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PC9 PC10
Standard Deviation 1.940 1.544 1.343 1.177 1.125 0.972 0.900 0.878 0.807 0.752
Proportion of
Variance
0.251 0.159 0.120 0.092 0.084 0.063 0.054 0.051 0.043 0.038
Cumulative
Proportion
0.251 0.410 0.530 0.623 0.707 0.770 0.824 0.875 0.919 0.956
SVL -0.427 -0.043 -0.189 -0.079 0.061 -0.012 0.053 -0.217 0.096 0.123
HL/SVL 0.232 0.290 0.460 -0.101 0.280 -0.028 0.293 -0.059 0.129 0.213
HW/SVL 0.454 0.057 0.075 -0.204 -0.042 0.058 0.334 -0.010 0.041 -0.237
HL/HW -0.361 0.155 0.266 0.231 0.259 -0.085 -0.169 -0.034 0.044 0.449
SL/SVL 0.100 0.536 -0.095 -0.245 0.343 0.136 -0.031 -0.069 -0.106 0.034
SL/HL -0.061 0.390 -0.451 -0.257 0.187 0.172 -0.257 -0.036 -0.208 -0.111
AG/SVL -0.052 0.123 -0.404 0.354 0.287 -0.013 0.289 0.350 0.446 -0.007
SHL/SVL -0.190 0.461 0.210 -0.108 -0.376 -0.084 0.239 -0.011 0.218 -0.020
SHL/HL -0.323 0.285 -0.063 -0.098 -0.532 -0.064 0.062 0.029 0.138 -0.145
LEO/SVL 0.114 0.213 -0.035 0.465 -0.325 0.380 0.097 0.072 -0.492 0.326
VEO/SVL 0.156 0.207 -0.041 0.603 -0.001 0.141 -0.196 -0.166 0.220 -0.430
LDPS/SVL 0.268 -0.047 -0.103 -0.201 -0.214 0.413 -0.319 0.268 0.522 0.430
TDPS/SVL 0.197 0.216 0.165 0.025 -0.087 -0.560 -0.537 0.367 -0.097 -0.054
SPW/SVL 0.215 0.018 -0.426 0.015 -0.120 -0.440 0.307 0.116 -0.188 0.338
NDSPW/SVL 0.277 0.042 -0.174 0.069 -0.141 -0.264 -0.162 -0.752 0.204 0.230
Female PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PC9 PC10
Standard Deviation 2.122 1.485 1.377 1.265 1.069 0.977 0.839 0.801 0.714 0.642
Proportion of
Variance
0.300 0.147 0.126 0.107 0.076 0.064 0.047 0.043 0.034 0.027
Cumulative
Proportion
0.300 0.447 0.574 0.680 0.756 0.820 0.867 0.910 0.944 0.971
SVL -0.392 -0.004 0.191 0.060 -0.213 -0.039 0.043 -0.101 -0.216 0.010
HL/SVL 0.335 0.286 0.218 -0.151 -0.112 0.347 0.230 -0.047 -0.194 -0.021
HW/SVL 0.406 -0.155 -0.093 -0.176 0.018 0.225 0.187 0.234 -0.012 0.223
HL/HW -0.213 0.448 0.314 0.089 -0.119 0.022 -0.034 -0.321 -0.160 -0.304
SL/SVL 0.271 -0.060 0.555 0.035 0.028 0.251 -0.078 -0.076 0.078 0.034
SL/HL 0.031 -0.353 0.509 0.198 0.147 0.003 -0.334 -0.043 0.319 0.079
AG/SVL -0.220 -0.051 0.294 0.180 -0.408 -0.057 0.069 0.669 -0.195 0.238
SHL/SVL -0.081 0.395 0.227 -0.486 0.285 0.117 0.020 0.253 0.045 0.044
SHL/HL -0.341 0.135 0.039 -0.330 0.349 -0.171 -0.175 0.269 0.199 0.058
LEO/SVL 0.084 0.230 -0.071 -0.257 -0.672 -0.103 -0.157 -0.105 0.580 0.162
VEO/SVL 0.152 0.393 -0.142 0.268 0.120 -0.001 -0.540 -0.079 -0.259 0.587
LDPS/SVL 0.305 0.136 0.129 0.044 -0.032 -0.606 0.231 0.070 -0.222 0.029
TDPS/SVL 0.333 0.144 0.150 0.103 0.151 -0.511 -0.006 0.082 0.187 -0.219
SPW/SVL -0.098 -0.301 0.192 -0.456 -0.003 -0.274 0.150 -0.442 -2.170 0.458
NDSPW/SVL 0.187 -0.228 -0.032 -0.397 -0.215 -0.057 -0.603 0.121 -0.409 -0.397
2018 hofmann and TownsEnd—a cryPTic nEw anolE from honduras 111
aPPEndix 4
GenBank accession numbers for sequence data of Norops caceresae, sp. nov., generated by Hofmann and Townsend (2017).
Sequences with JHT eld numbers refer to specimens not accessioned.
Museum ID 16S ND2 COI PRLR BDNF PTPN12
UF 166186 KU688065 – KU687954 – – –
UF 166188 KU688066 – KU687955 – – –
UF 166190 KU688060 – KU687949 – – –
UF 166191 KU688061 – KU687950 – – –
[JHT 2891] KU688062 – KU687951 – – –
[JHT 2892] KU688063 – KU687952 – – –
[JHT 2894] MF094466 – MF094513 – – –
[JHT 2895] KU688067 – KU687956 – – –
MVZ:Herp:286123 MF094468 MF094560 MF094515 MF094598 MF094634 MF094661
CM 161307 MF094469 MF094561 MF094516 MF094599 MF094635 MF094662
CM 161309 MF094471 – MF094518 MF094601 MF094637 MF094664
CM 161311 MF094472 MF094562 MF094519 MF094602 MF094638 MF094665
