de Queiroz, K., Huie, J. M., Hammel, J. U., Müller, P., & Baranov, V. (n.d.). A New Fossil
Anolis Lizard in Hispaniolan Amber: Ecomorphology and Systematics. Journal of
Herpetology. https://doi.org/10.1670/23-058
A NEW FOSSIL ANOLIS LIZARD IN HISPANIOLAN AMBER: ECOMORPHOLOGY AND SYSTEMATICS 
K AEVIN DE QUEIROZ1 , JONATHAN M. HUIE2, JÖRG U. HAMMEL3, PATRICK MÜLLER4, VIKTOR BARANOV5 
1 VERTEBRATE ZOOLOGY, NATIONAL MUSEUM OF NATURAL HISTORY, 
2 BIOLOGICAL SCIENCES, GEORGE WASHINGTON UNIVERSITY, 
3 HELMHOLTZ-ZENTRUM GEESTHACHT, INSTITUTE OF MATERIALS PHYSICS, 
4 FRIEDHOFSTRASSE 9, 
5 ESTACIÓN BIOLÓGICA DOÑANA, CONSEJO SUPERIOR DE INVESTIGACIONES CIENTÍFICAS 
https://doi.org/10.1670/23-058 
ABSTRACT 
Pre-Pleistocene fossils of Anolis lizards are rare, although 20 Miocene fossils preserved in amber from the island of 
Hispaniola have been reported on previously. Using light-microscopy and computed-tomography imaging, we studied a 
new amber fossil Anolis lizard from Hispaniola. The fossil is likely a juvenile and preserves a largely intact left forelimb, 
including both scales and skeletal elements, as well as some additional skin, skin impressions, and fragmentary skeletal 
elements from other parts of the body. Using measurements and lamella counts from the forelimb of the fossil and 
other juvenile anoles, discriminant function analysis and three Euclidean-distance criteria derived from a principal 
components analysis consistently support classification of the fossil as a member of the trunk ecomorph category, and 
those results, in combination with two scale characters preserved in the fossil, suggest that it is a member of the Anolis 
distichus series within the Ctenonotus clade. These results represent only the third case of a well-supported assignment 
of an Anolis fossil to the trunk ecomorph category and the first to the A. distichus series. They also highlight that such 
assignments can sometimes be inferred from highly incomplete fossils. 
Anolis lizards provide a well-known example of the eco- bers of the trunk-crown ecomorph category; most speci-
morph phenomenon, in which organisms exhibit species- mens analyzed to date that can be assigned to an ecomorph 
specific quantifiable morphological adaptations related to with a high degree of confidence have been assigned to 
ecological specialization, in this case to use of structural that category (Sherratt et al., 2015). Only a few specimens 
habitats (e.g., Losos, 2009; Losos et al., 1998; Williams, have been assigned to three other of the ecomorph cate-
1972, 1983). Each anole ecomorph is named for the struc- gories—namely, trunk (2 specimens), trunk-ground (2), and 
tural habitat, or habitats, that are most commonly occupied twig (1) (Sherratt et al., 2015). Here, we report on a frag-
by the lizards assigned to that ecomorph, and six or seven mentary new specimen that nonetheless appears to be as-
are commonly recognized—crown giant, trunk-crown, signable to one of the rarer ecomorphs as well as to a small 
trunk, twig, trunk-ground, grass-bush, and, recently recog- subclade of anoles with reasonably high degrees of confi-
nized, ground (Huie et al., 2021). In the Greater Antilles dence. 
of the Caribbean Sea, where the ecomorph phenomenon in 
Anolis lizards is best documented, members of the various MATERIALS AND METHODS 
ecomorph categories have evolved largely independently 
through convergent evolution between lineages on differ- The specimen is cataloged in the paleontology collection 
ent islands (Losos, 2010; Losos et al., 1998; Mahler et al., of the Staatliches Naturhistorisches Museum Braunschweig 
2013). under the number 8198. It was studied with a combination 
The fossil record of Anolis lizards prior to the Pleistocene of optical microscopy, including photography, and x-ray 
is generally poor (Losos, 2009); however, one exception computed tomography. The specimen was examined and 
concerns amber deposits on the island of Hispaniola, from photographed using a Keyence VHX-6000 digital micro-
which some 20 fossil specimens of Anolis lizards have been scope with an overhead ring light. Stacks of images were 
studied (Sherratt et al., 2015). Amber fossils often preserve recorded at different focal lengths, then each stack was as-
not only skeletal elements but also skin, often, the pre- sembled into a single composite image using the micro-
served parts are intact, and sometimes, strongly supported scope’s built-in software. All photographs presented here 
inferences can be made about the systematic relationships are composite images. 
and ecological morphology of the fossils. The first three am- The specimen was scanned using micro-computed to-
ber fossil anoles reported on in the scientific literature (de mography (µCT) on the Imaging Beamline P05 (Lytaev et al., 
Queiroz et al., 1998; Polcyn et al., 2002; Rieppel, 1980) were 2014) that is operated by Helmholtz-Zentrum Hereon at the 
all inferred to be Hispaniolan green anoles (members of the PETRA III storage ring (Deutsches Elektronen-Synchrotron 
Anolis chlorocyanus species group of Williams, 1976 = Ano- DESY, Hamburg, Germany). Scans were performed using a 
lis chlorocyanus series of Burnell & Hedges, 1990) and mem- photon energy of 18 keV with a sample-to-detector distance 
of 100 mm. We recorded the projections with a custom 20 
A CORRESPONDING AUTHOR. E-MAIL: DEQUEIROZK@SI.EDU 
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A New Fossil Anolis Lizard in Hispaniolan Amber: Ecomorphology and Syst…
MP CMOS imaging system and an effective pixel size of 1.28 and both each ecomorph centroid and the members of each 
µm (Lytaev et al., 2014). For each tomographic scan, 3601 ecomorph. We then applied the following three distance cri-
projections were recorded at equal intervals between 0 and teria to determine whether the fossil could be assigned to 
π. We conducted a reconstruction by applying a transport- one or more of the ecomorphs: (1) Centroid distance (CD); 
of-intensity phase-retrieval approach and using the filtered the fossil was considered assignable to an ecomorph if the 
back-projection algorithm carried out with a custom recon- Euclidean distance from the fossil to the centroid of that 
struction pipeline (Moosmann et al., 2014) using MATLAB ecomorph was ≤ the distance of the furthest member of 
(Mathworks, 2022) with application of the ASTRA Toolbox that ecomorph to the centroid. (2) Mean pairwise distance 
(van Aarle et al., 2015, 2016). We binned raw projections (MPD); the fossil was considered assignable to an ecomorph 
twice for subsequent processing, which resulted in an effec- if the average Euclidean distance of the fossil to all mem-
tive pixel size of the reconstructed volume (voxel) of 2.56 bers of that ecomorph was ≤ the largest average pairwise 
µm. To decrease the burden on computer memory, we con- distance among the members of that ecomorph. (3) Near-
verted all the stacks into 8-bit tiffs, which we downscaled by est-neighbor distance (NND); the fossil was considered as-
50%; we also cropped the empty space around the specimen signable to an ecomorph if the Euclidean distance of the 
using the ‘scale’ and ‘crop’ functions in Fiji (Schindelin et fossil to its nearest member of that ecomorph was ≤ the 
al., 2012). We used Drishti version 2.6.6 to reconstruct the largest nearest-neighbor distance among the members of 
projections as volumes (Limaye, 2012). We took measure- that ecomorph. 
ments from the scan using the 3D Slicer software, version Comparisons of scale characters were also made with 
5.2.2 (Kikinis et al., 2013) equipped with the SlicerMorph specimens representing several species of Hispaniolan 
toolkit (Rolfe et al., 2021). A TIFF stack of the scan is de- anoles representing every ecomorph for which the fossil 
posited at www.morphdbase.de. had a nonzero DFA assignment probability or satisfied at 
Because the amber specimen is likely a juvenile, and least one of the Euclidean-distance criteria (Supplementary 
most existing Anolis ecomorphological data sets are for Data: Appendix 2). 
adults, (e.g., Armstead & Poe, 2015; Huie et al., 2021; Losos 
et al., 1998; Poe & Anderson, 2019) or were measured by RESULTS 
a different researcher (Sherratt et al., 2015), JMH collected 
new data for 43 juvenile anoles representing 14 Hispaniolan The amber piece containing the fossil (Fig. 1A) is irreg-
ecomorph species (Supplementary Data: Appendix 1). In ularly shaped and measures 24 mm in length by 21 mm in 
addition, because the best-preserved part of the fossil is the width and 10 mm in thickness. It was collected from an un-
left forelimb, our study focused on that limb. Thus, JMH known mine in the Dominican Republic and purchased by 
measured the length of the upper arm, lower arm, and the PM from Ambra Greco (Viale Ugo Foscolo, 9, 20900 Monza 
hand (from wrist to tip of the fourth finger), as well as the MB, Italy). Although the geological formation is unknown, 
width of the pad on the fourth finger. He also counted the all currently known Dominican amber is from two forma-
number of subdigital lamellae (laterally expanded subdigi- tions, La Toca and Yanigua (Penney, 2010). The age of Do-
tal scales) on the fourth finger from the base of the digit to minican amber has been debated (reviewed by Penney, 
the distal terminus of the adhesive pad (i.e., the lamellae 2010), but relatively recent estimates give the age as Early 
underlying the proximal three phalanges of digit IV, which to Middle Miocene or 15–20 Ma (Iturralde-Vinent, 2001; 
has five phalanges in total; the subdigital scales under the Iturralde-Vinent & Macphee, 1996). 
penultimate phalanx are not lamellar, while the terminal = The fossil lizard (Fig. 1B) is oriented diagonally within 
ungual phalanx is small and associated primarily with the the amber piece relative to both its width and its length. 
claw). We measured the soft tissues rather than the bones The fossil consists of a mostly intact (skin and skeleton) left 
of both the fossil and the modern specimens. forelimb, parts of the right forelimb, the posterior part of 
To assess the amber anole as a member of an ecomorph the head, skin from the left side of the body posterior to 
class, we first size corrected the continuous variables using the insertion of the forelimb, and skin (scale) impressions 
forelimb length (sum of the upper arm, lower arm, and hand from the ventral surface of the body, the posterior lateral 
measurements) as a proxy for size. The discrete variable, surface of the body on left side, the lateral surface of the 
lamella count, was not size corrected but was log trans- body on the right side, and the right temporal region of the 
formed. We then assessed the ecomorph assignment of the head. The main skeletal elements preserved (Fig. 1C) con-
fossil using the same approaches as Huie et al. (2021). First, sist of most of the bones of the left forelimb (humerus, ra-
we performed a discriminant function analysis (DFA) using dius, ulna, and some of the metacarpals and phalanges), 
the size-corrected, and log-transformed, variables and the ventral part of the left quadrate bone and posterior part 
trained with the ecomorph species using the MASS package of the lower jaw (articular and most of the surangular and 
(Venables & Ripley, 2002) in R Statistical Software version prearticular bones), and fragments of at least four ribs (both 
4.3.2 (R Core Team, 2021). We then performed a principal bony dorsal and cartilaginous ventral components) asso-
components analysis (PCA), also using R version 4.3.2, and, ciated with the skin on the left side of the body. The ra-
because of the small number of variables, used only the first dius and ulna are displaced distally relative to the skin, and 
two PC axes to calculate Euclidean distances between the some of the metacarpals are also displaced from their pre-
specimens and ecomorph centroids. Second, we calculated sumed natural positions. The proximal and distal ends of 
Euclidean distances between each member of an ecomorph most of the limb bones (diaphyzes) appear flat or concave 
and the ecomorph centroid, between all pairs of specimens and adjacent phalanges are separated by gaps, suggesting 
representing the same ecomorph, and between the fossil that the epiphyses were mostly or entirely cartilaginous 
de Queiroz et al. 2
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A New Fossil Anolis Lizard in Hispaniolan Amber: Ecomorphology and Syst…
TABLE 1. Component loadings (above) and standard deviation and proportion of variance explained by the components 
(below) from the PCA. Pad width and lamella count are for manual digit IV. 
PC1 PC2 PC3 PC4 PC5 
Upper arm 0.013 0.023 0.190 0.804 0.563 
Lower arm 0.013 0.132 0.650 −0.529 0.530 
Hand −0.019 −0.133 −0.711 −0.272 0.634 
Pad width −0.251 0.952 −0.176 0.017 0.002 
Lamella count −0.968 −0.242 0.071 0.004 0.002 
Standard deviation 0.22427 0.13542 0.06140 0.03405 0.00178 
Proportion of variance 0.68367 0.24928 0.05124 0.01576 0.00004 
Cumulative proportion 0.68367 0.93295 0.98420 0.99996 1.00000 
TABLE 2. Ecomorph assignment of the fossil anole. The table gives discriminant-function-analysis probabilities and 
Euclidean distances for the amber fossil Anolis specimen regarding its classification in the six traditional ecomorph 
categories (Losos, 2009; Williams, 1972). Abbreviations: CG = crown giant, GB = grass-bush, TC = trunk-crown, TG = 
trunk-ground, Tr = trunk, Tw = twig, DFA = discriminant-function-analysis-classification probability, CD = centroid 
distance, MPD = mean pairwise distance, NND = nearest-neighbor distance, ns = (criterion) not satisfied. Euclidean-
distance values are given only when the fossil satisfied the criterion based on the corresponding distance measure (see 
Methods). Best supported assignments according to each criterion (highest value for DFA, lowest for Euclidean distances) 
are in bold. 
Ecomorph 
Criterion CG GB TC TG Tr Tw 
DFA 0 0.034 0.002 0.213 0.751 0 
CD ns 0.201 ns 0.169 0.042 ns 
MPD ns 0.213 0.285 0.173 0.095 ns 
NND ns 0.089 0.117 0.073 0.007 ns 
(see Maisano, 2001). Some poorly preserved bones of the (Table 1). When the specimens of ecomorph species and the 
right forelimb are also present. The retroarticular process fossil are plotted in the morphological space defined by the 
of the lower jaw is highly fragmented but appears to have first two PCs (Fig. 2), the ecomorphs are reasonably well 
been well-developed; the presence and size of the angular separated. That separation reveals, for example, that crown 
process could not be determined. giant anoles have many lamellae while twig and grass-bush 
The dorsal and lateral scales are small and granular. The anoles have few lamellae, and that trunk-crown and twig 
ventral scales (impressions) are much larger (at least 4x anoles have wide pads while crown giant, trunk-ground, 
larger than the dorsals by area); they are smooth and sub- and grass-bush anoles have narrow ones (for their body 
imbricate (the posterior borders slightly overlap the scale sizes; see also Fig. S1). The fossil is positioned along the up-
or scales behind them) and are arranged in transverse rows per border of the space occupied by trunk anoles, very close 
(Fig. 1B). The supra-digital scales are smooth or perhaps to one of the trunk anole specimens (USNM 329155, a spec-
faintly keeled. The scales on the dorsal surface of the manus imen of A. distichus), which is an area of the space not over-
(metacarpal region) are large and overlapping. The subdigi- lapped by any of the other ecomorphs. 
tal scales associated with the antepenultimate phalanx and The DFA of the same ecologically relevant characters 
the phalanges proximal to it, when present, are wide and (Table 2) classified the fossil as a trunk anole with the high-
lamellar, forming the ventral surface of pads on digits II–V. est probability (P = 0.751) and a trunk-ground anole with 
There are 19 subdigital lamellae on the underside of manual the second highest probability (P = 0.213), with those two 
digit IV (counted using the method of Köhler, 2014). Indica- assignments accounting for > 0.96 of the probability. The 
tions of dark transverse markings are present on the upper remaining fraction was accounted for by the grass-bush (P = 
and lower forelimb. 0.034) and trunk-crown (P = 0.002) ecomorphs; assignment 
In the PCA of ecologically relevant morphological char- to the crown-giant and twig ecomorphs had zero probabil-
acters, the first two PCs explained 93.3 % of the variation ity. 
(Table 1). The loadings of both of those PCs primarily in- Comparing the fossil to the modern ecomorph species 
volved pad characters, with lamella number loading most under the three Euclidean-distance criteria (Table 2), the 
heavily on PC1 and pad width loading most heavily on PC2 fossil satisfied the three Euclidean-distance criteria for 
de Queiroz et al. 3
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A New Fossil Anolis Lizard in Hispaniolan Amber: Ecomorphology and Syst…
FIG. 2. Location of the amber fossil Anolis (SNHMB.G 8198) 
relative to representatives of modern ecomorph species in 
FIG. 1. SNHMB.G 8198, a fossil Anolis lizard preserved in the morphological space defined by the first two principal 
Miocene amber from the Dominican Republic. (A) components (PC) for the forelimb data. Modern specimens 
Photograph of the entire amber piece containing the fossil representing the different ecomorphs and the fossil are 
lizard. (B) Photograph of the fossil lizard in dorsal view represented by color-filled circles as indicated in the key on 
showing the impressions of the ventral scales (center), the the right, with the species assigned to the same ecomorph 
supinated left forelimb (lower right), and the lateral body united within a minimum convex polygon shaded with a 
skin (between the ventral scale impressions and the limb). similar (but lighter) color. The orange-filled circle 
(C) Radiographic image (composite z-projection) of the representing the fossil anole is positioned near the middle 
fossil lizard generated from the µCT data. Osseous of the upper edge of the polygon for the trunk ecomorph, 
elements include parts of the left quadrate (q) and lower partially overlapping the circle representing one of the 
jaw (lj), the left humerus (h), ulna (u) radius (ra), trunk-ecomorph members. 
metacarpals (m), and phalanges (p), bones of the right 
forelimb (rf; faint), and parts of several left ribs (ri). 
DFA assignment probability for that category is more than 
three times greater than for any other ecomorph, and its 
three to four of the six traditionally recognized ecomorphs, Euclidean distances to the next closest ecomorph are 1.8 
depending on the criterion. In all cases, the smallest Euclid- (MPD), 4.0 (CD), and 10.4 (NND) times greater than to the 
ean distances (indicating strongest support) were for the trunk ecomorph. Although there is some ambiguity in eco-
trunk ecomorph and the next smallest for the trunk-ground morph assignment resulting from the small number of rele-
ecomorph followed by grass-bush and trunk-crown, except vant traits that can be measured in the fossil, the only other 
that the fossil did not satisfy the centroid-distance criterion ecomorph assignment that deserves further consideration 
for the trunk-crown ecomorph. The fossil did not satisfy any is trunk-ground, as that is the only other assignment with 
of the Euclidean-distance criteria for the crown-giant and more than a negligible DFA assignment probability (i.e., P > 
twig ecomorphs. 0.05) and is the assignment that ranks second according to 
all three of the Euclidean-distance criteria. 
DISCUSSION Extant Hispaniolan trunk anoles are all members of the 
Anolis distichus series (Losos, 2009), which is part of the 
The amber fossil can be assigned unambiguously to the 
Ctenonotus subclade (sensu Poe et al., 2017), while the ex-
Anolis clade (sensu Poe et al., 2017) based on the morphol-
tant Hispaniolan trunk-ground anoles are all members of 
ogy of the toepads. Although toepads have evolved multi-
the Anolis cybotes series, which is equivalent to the Au-
ple times in squamatan reptiles (Hagey et al., 2017; Miller 
dantia subclade (sensu Poe et al., 2017). Members of these 
& Stroud, 2021), the pads of anoles are distinctive in having 
two clades differ in that those of the distichus series have 
the combination of a single row of lamellae and the claw 
smooth supra-digital scales while those of the cybotes series 
separated from the pad by a nonlamellar segment associ-
have keeled supradigitals (Cochran, 1941); moreover, based 
ated with the penultimate phalanx (see figures in Griffing et 
on our own observations, members of the distichus series 
al., 2022). In addition to the small size of the specimen and 
have the ventral scales arranged in transverse rows (as do 
the fact that nearly all previously known fossil anoles pre-
members of the chlorocyanus series of Hispaniolan trunk-
served in amber appear to be juveniles (see measured and 
crown anoles; see illustrations in Köhler & Hedges, 2016) 
estimated SVLs in Sherratt et al., 2015: Table S2), the lack 
while those of the cybotes series (= Audantia) have the ven-
(or very small size) of ossified epiphyses indicates that the 
trals arranged in oblique rows (see illustrations in Köhler et 
specimen is a juvenile (see Maisano, 2001). 
al., 2019). The fossil exhibits the conditions found in mem-
Regarding its ecological morphology, the amber fossil is 
bers of the distichus series for both characters. 
most likely a member of the trunk ecomorph category, as its 
de Queiroz et al. 4
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A New Fossil Anolis Lizard in Hispaniolan Amber: Ecomorphology and Syst…
Thus, assuming that the fossil is a member of one of ACKNOWLEDGMENTS 
the extant clades of Hispaniolan anoles, as most previously 
studied well-preserved Hispaniolan amber fossil anoles are M. Reich cataloged the specimen in the paleontology col-
inferred to be (Sherratt et al., 2015), its ecomorphological lection of the Staatliches Naturhistorisches Museum Braun-
and taxonomic characters in combination support the infer- schweig. E. Langan and T. Hsu provided information and 
ence that the fossil is a trunk anole of the A. distichus se- photos for specimens in the amphibian and reptile collec-
ries. The age of the fossil (15–20 Ma) is within that of the tion of the National Museum of Natural History, Smithson-
distichus series, whose crown age is estimated to be 20.78 ian Institution. Scanning of the specimen was supported 
Ma and was not estimated using fossil calibrations within by the DESY Block Allocation Group project “Scanning the 
that clade (Poe et al., 2017). Our findings represent only the past - Reconstructing the diversity in million years old fossil 
third case of a well-supported trunk ecomorph assignment amber specimens using SRµCT” at PETRA III. J. D. Daza and 
for a fossil Anolis lizard (the other two were by Sherratt et an anonymous reviewer provided helpful comments on the 
al., 2015) and the first case of assignment to the A. distichus manuscript. 
series (see Sherratt et al., 2015). They also highlight that 
such assignments can be inferred from only a small part of SUPPLEMENTARY DATA 
the body, in this case, primarily the forelimb, of a highly in-
complete fossil. Supplementary data associated with this article can be 
found online alongside the manuscript. 
Accepted: December 27, 2023 EDT 
de Queiroz et al. 5
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A New Fossil Anolis Lizard in Hispaniolan Amber: Ecomorphology and Syst…
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SUPPLEMENTARY MATERIALS 
Figure S1 
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