DE QUEIROZ, K., MÜLLER, P., HAMMEL, J. U., & BARANOV, V. (n.d.). A Fossil Anolis Lizard
Tail in Mexican Amber: Phylogenetic Relationships and Biogeographic Significance. Journal
of Herpetology. https://doi.org/10.1670/23-070
A FOSSIL ANOLIS LIZARD TAIL IN MEXICAN AMBER: PHYLOGENETIC RELATIONSHIPS AND 
BIOGEOGRAPHIC SIGNIFICANCE 
KEVIN DE QUEIROZ1A, PATRICK MÜLLER2, JÖRG U. HAMMEL3, VIKTOR BARANOV4 
1 DEPARTMENT OF VERTEBRATE ZOOLOGY, NATIONAL MUSEUM OF NATURAL HISTORY, 
2 ZWEIBRÜCKEN, GERMANY, 
3 INSTITUTE OF MATERIALS PHYSICS, HELMHOLTZ-ZENTRUM HEREON, 
4 ESTACIÓN BIOLÓGICA DOÑANA, CONSEJO SUPERIOR DE INVESTIGACIONES CIENTÍFICAS 
https://doi.org/10.1670/23-070 
ABSTRACT 
Pre-Pleistocene fossils of Anolis lizards from the mainland of the Americas are exceedingly rare: only two specimens 
referred to a single species have been described previously. Here we report on a third specimen, preserved (as are the 
other two) in Miocene amber from Chiapas, Mexico, and consisting primarily of the anterior vertebrae of the caudal 
sequence. Despite the fragmentary nature of the fossil, it preserves key osteological characters that permit confident 
referral to the Anolis clade and further suggest placement within the Dactyloa subclade in a clade of three extant species 
within the Anolis aequatorialis series. The Chiapan provenance of the fossil indicates that the geographic distribution 
of the Dactyloa clade (and possibly that of the A. aequatorialis series) extended considerably farther north during the 
Miocene. Although the new fossil represents a different part of the body than the two fossils representing the fossil 
species Anolis electrum, its inferred phylogenetic relationships are the same as one of the several possible phylogenetic 
relationships of that species and thus allow for the possibility that all three specimens belong to the same species. 
Anolis lizards (anoles) form a clade of approximately 439 as the remains of an Anolis lizard and suggest a more pre-
currently recognized extant species (Uetz et al., 2023) that cise phylogenetic placement than for the two previous Mex-
have been subjects of a wide variety of ecological and evo- ican amber specimens. The inferred phylogenetic relation-
lutionary studies (e.g., Alföldi et al., 2011; Gamble et al., ships of the new fossil have implications for the 
2014; Muñoz et al., 2023; Velasco et al., 2020; Winchell biogeographic history of anoles in Middle America and bear 
et al., 2016). Anoles are distinctive among iguanian lizards on the relationships of the previously named Anolis elec-
(Reptilia/Lepidosauria/Squamata/Iguania) in that the males trum (Lazell, 1965). 
of most species and the females of some possess a dewlap, 
an extensible gular appendage that is used for signaling MATERIALS AND METHODS 
(Harrison & Poe, 2012), and both males and females of all 
but one species possess adhesive toe and finger pads that The amber piece containing the specimen was collected 
facilitate climbing (Losos, 2009). Although the evolution- in the mine in the Simojovel Municipality in the state of 
ary history of anoles is becoming increasingly better under- Chiapas, Mexico, and is estimated to be 15–20 Ma = Early to 
stood based on data from extant species (e.g., Jackman et Middle Miocene (Solórzano-Kraemer, 2010). PM purchased 
al., 1999; Nicholson et al., 2005; Poe et al., 2017), the fossil the piece in 2019 and donated it to the paleontology col-
record of anoles is generally poor, and most known fossils lection of the Staatliches Naturhistorisches Museum Braun-
are of Pleistocene age (e.g., Bochaton et al., 2015; Holman, schweig (SNHMB.G), Germany, where it is catalogued under 
1995; Pregill et al., 1994; but see Pregill, 1999). The main the number 8197. 
exception is a set of Anolis fossils preserved in Miocene am- We studied the specimen (SNHMB.G 8197) using both 
ber from the island of Hispaniola in the West Indies, of microscopy (including photographs) and microcomputed 
which some 21 specimens have been reported on in the tomography (μCT). We examined and photographed the 
scientific literature (de Queiroz et al., 1998, 2024; Polcyn specimen using a Keyence VHX-6000 digital microscope 
et al., 2002; Rieppel, 1980; Sherratt et al., 2015). Among with an overhead ring light. We recorded stacks of images 
the few pre-Pleistocene Anolis fossils from the mainland of at different focal lengths, then assembled each stack into 
the Americas are two specimens preserved in amber from a single composite image using the microscope’s built-in 
Chiapas, Mexico, assigned to a single species (Lazell, 1965) software. 
whose phylogenetic position within the Anolis clade is un- We scanned the piece of amber containing the specimen 
certain (Castañeda et al., 2014). on Imaging Beamline P05 (Lytaev et al., 2014) run by 
Here we report on a new fossil in amber from Chiapas, Helmholtz-Zentrum Hereon at the PETRA III storage ring 
Mexico. Although this specimen is highly incomplete, con- (Deutsches Elektronen-Synchrotron DESY, Hamburg, Ger-
sisting primarily of the anterior caudal vertebrae, it many). We performed the scans using a photon energy of 18 
nonetheless preserves characters that identify the specimen keV with a sample-to-detector distance of 100 mm. We cap-
A CORRESPONDING AUTHOR. EMAIL: DEQUEIROZK@SI.EDU 
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A Fossil Anolis Lizard Tail in Mexican Amber: Phylogenetic Relationships a…
tured the projections with a customized 20 MP CMOS imag- with a transverse break in the vertebra that is posterior to 
ing system and an effective pixel size of 1.28 μm (Lytaev the transverse processes, which is inferred to be an artifact 
et al., 2014). For each tomographic scan, we recorded 3,601 because it is asymmetrical relative to the midsagittal plane. 
projections at equal intervals between 0 and π. We then In addition, the inferred partial septum is in approximately 
performed a reconstruction, applying a transport-of-inten- the same location as the full and unambiguous septum of 
sity phase-retrieval approach with application of the fil- CV 8, that is, roughly one-third of the distance from the an-
tered back-projection (FBP) algorithm with a custom recon- terior articulation with CV 6 to the posterior articulation 
struction pipeline (Moosmann et al., 2014) using MATLAB with CV 8; by contrast, the inferred break is located at more 
(MathWorks, 2022) and applying the Astra Toolbox (van than one-half that distance). The neural spines of the pre-
Aarle et al., 2015, 2016). We binned raw projections to- sacral and sacral vertebrae are low and short, each form-
gether twice for subsequent processing, which resulted in ing a middorsal ridge that extends along the entire dorsal 
an effective pixel size of the reconstructed volume (voxel) of surface of the vertebra and does not project over the ver-
2.56 μm. To decrease the strain on the PC’s RAM, we con- tebra posterior to it. Those of the first three caudal verte-
verted all stacks into 8-bit TIFFs, which we then downscaled brae also form middorsal crests that decrease in size poste-
by 50%. To further reduce the burden on computer mem- riorly and are absent from CV 4 on. In addition, the neural 
ory we cropped the empty space around the specimen us- spines of all caudal vertebrae form narrow processes that 
ing the “'scale” and “crop” functions in Fiji (ImageJ) 2.3.0/ project posterodorsally from the posterior end of each ver-
1.53q (Schindelin et al., 2012). We used Drishti v2.6.6 to tebra over the anterior end of the next vertebra posteriorly, 
reconstruct the projections as volumes (Limaye, 2012). We although the spine of CV 8 is missing. The neural spines 
examined a 3D model using Meshlab 2022.02 (Cignoni et decrease in size posteriorly. Haemal arches are attached to 
al., 2008) and 2D models using QuickTime Player 10.5 (Ap- the posteroventral termini (immediately anteriorly to the 
ple, Incorp., 2022). The scan is deposited as a TIFF stack at condyles) of CV 3–CV 7, with remnants present on CV 8 and 
www.morphdbase.de (see Supplementary Data). CV 9. 
RESULTS DISCUSSION 
The amber piece containing remains of the lizard is ap- The Mexican amber fossil (SNHMB.G 8197) exhibits pro-
proximately 26 mm long and 13 mm wide. In addition to the coelous vertebrae, a condition in which the anterior end of 
lizard, it contains at least five ant workers (Hymenoptera/ each centrum is concave and articulates with a convex sur-
Formicidae), a pselaphine beetle (Coleoptera/Staphylin- face at the posterior end of the centrum of the vertebra 
idae/Pselaphinae), and a gall midge (Diptera/Cecidomyi- anterior to it. The procoelous condition is considered di-
idae/Cecidomyiinae) (Fig. S1). Remains of the lizard (Figs. 1, agnostic of squamatan reptiles (lizards, including snakes), 
2) extend through most of the length of the amber piece and although it has reversed to the amphicoelous (biconcave) 
consist of parts of the vertebral column and pelvic girdle as condition in some gekkotans (Estes et al., 1988). 
well as various scattered scales, at least some of which bear The fossil also exhibits the condition of the caudal verte-
a single keel. Because skeletal elements are more informa- brae referred to as the first iguanid type by Etheridge (1959), 
tive on the systematics of the fossil, we focus on them. the second (iguanid) type by Etheridge (1967), and the alpha 
The skeletal remains consist of part of the second-to-last condition by Guyer and Savage (1986), in which there is a 
presacral vertebra, the last presacral vertebra, both sacral short anterior series of nonautotomic vertebrae each bear-
vertebrae, and all or parts of 11 caudal vertebrae, as well ing a single pair of transverse processes followed by a long 
as most of the left side and highly fragmented parts of the series of vertebrae that may or may not be autotomic with-
right side of the pelvic girdle (although only the left ilium out transverse processes (Etheridge, 1967). Autotomic ver-
is not severely damaged). The pelvic girdle is displaced (de- tebrae, when present, include all those in the series that 
pressed anteriorly and rotated clockwise around a longitu- lack transverse processes and occasionally the last of those 
dinal axis when viewed from anterior) so that the ilia (of that bear transverse processes (Etheridge, 1967). Although 
which only the left is present) are not articulated to the this type of sequence occurs in several clades of iguanid 
sacral diapophyses. The posterior part of caudal vertebra lizards (Iguanidae sensu Etheridge, 1967; Torres-Carvajal 
(CV) 7 and the caudal vertebrae posterior to it are crushed et al., 2020; Pleurodonta of Frost et al., 2001), in most of 
inwards on the left side. The vertebrae are procoelous. The them the caudal vertebrae are either nonautotomic (Coryto-
first seven caudal vertebrae bear transverse processes, some phanes, Laemanctus, Enyalius, Polychrus, Urostrophus) or, if 
of which are broken and/or partly missing. The transverse an autotomy septum is present in the last vertebra bearing 
processes increase in size from CV 1 to CV 2, then decrease transverse processes, the septum is located posterior to the 
in size successively to CV 7. The remaining caudal vertebrae transverse processes (Basiliscus, Enyalioides, Uranoscodon) 
lack transverse processes. CV 8 and CV 9 are mostly intact (Etheridge, 1967). Among iguanid lizards exhibiting this 
(except for the neural spines and haemal arches); CV 10 is type of caudal sequence with autotomy septa, only in some 
fragmented but most of it is present, and only the anterior species of Anolis is the septum located anterior to the trans-
part of CV 11 is preserved. Autotomy septa are clearly pre- verse processes (Etheridge, 1967). 
sent in CV 8–CV 11. A partial septum, most evident in the Moreover, other anoles exhibit a caudal sequence termed 
ventral one-half to two-thirds of the vertebra, also appears the second iguanid type by Etheridge (1959), the third 
to be present in CV 7 (Fig. 2B) and is located anterior to (iguanid) type by Etheridge (1967) and the beta condition 
the transverse processes (the septum is not to be confused by Guyer and Savage (1986), in which the autotomic caudal 
DE QUEIROZ et al. 2
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A Fossil Anolis Lizard Tail in Mexican Amber: Phylogenetic Relationships a…
FIG. 1. Photographs of the fossil remains of an Anolis lizard in amber from Chiapas, Mexico (SNHMB.G 8197). (A) Right 
ventrolateral view. (B) Approximately dorsal view. The remains consist mostly of the sacral and anterior caudal vertebrae. 
Anterior is to the right; scale bars (1 mm) are included. 
vertebrae bear anterolaterally directed and terminally bi- age, 1986; beta anoles of Etheridge, 1959). The morphol-
furcated transverse processes and the fracture planes are ogy of the fossil (alpha condition) suggests that it is outside 
located anterior to the transverse processes. Because the of the Norops clade. The Norops clade is inferred to have 
anterior location of the autotomy septa relative to the originated in the West Indies (Nicholson et al., 2005; Poe et 
transverse processes is unique to anoles among iguanids al., 2017), so the Mexican provenance of the fossil suggests 
(Etheridge, 1967), it appears to be a derived condition that that it is unlikely to belong to the Norops stem group, early 
is diagnostic for that clade. The condition also occurs in members of which might have lacked the derived beta con-
dibamids and a few scincids (Etheridge, 1967), distantly re- dition of the caudal vertebrae. 
lated squamatans whose vertebrae differ from those of Among anoles with the alpha type of caudal sequence, 
anoles in other respects (Etheridge, 1967) and which are un- (non-Norops) members of the Digilimbus clade are almost 
likely to be preserved in amber. The Mexican amber fossil entirely West Indian, while members of the Dactyloa clade 
has the (partial) autotomy septum of CV 7 located anterior occur primarily on the mainland (Poe et al., 2017). Among 
to the transverse processes, and this constitutes strong evi- members of the Dactyloa clade, only a few species exhibit 
dence that the fossil is an Anolis lizard. modes of seven non-autotomic anterior caudal vertebrae 
Following the phylogenetic taxonomy of Poe et al. (2017) (Etheridge, 1959; Poe, 2004), corresponding approximately 
and relevant to the present discussion, Anolis is divided into to those that bear transverse processes (Etheridge, 1959, 
two primary crown clades, Dactyloa and Digilimbus, the sec- 1967), and mostly the same species exhibit modes of seven 
ond of which contains the subclade Norops, which in turn caudal vertebrae that bear transverse processes in a more 
contains the subclade Draconura. The alpha type of caudal extensive data set for the Dactyloa clade (Castañeda, un-
sequence found in the Mexican amber fossil appears to be published data for Castañeda & de Queiroz, 2013). The low 
ancestral for Anolis (Guyer & Savage, 1986) and is retained number of such vertebrae appears to be a derived condition 
in anoles of the Dactyloa clade as well as some members within Dactyloa, and those species exhibiting it belong to 
of the Digilimbus clade. The ancestral alpha type of caudal the aequatorialis and roquet series, which occur in western 
sequence contrasts with the derived beta condition, which South America extending (barely) into Central America and 
characterizes members of the Norops clade (Guyer & Sav- in the southern Lesser Antilles, respectively (Castañeda & 
DE QUEIROZ et al. 3
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A Fossil Anolis Lizard Tail in Mexican Amber: Phylogenetic Relationships a…
FIG. 2. CT reconstructions of the fossil remains of an Anolis lizard in amber from Chiapas, Mexico (SNHMB.G 8197). (A) 
Dorsal view of the caudal vertebrae. (B) Detail of caudal vertebrae 6 (part), 7, and 8 in lateral view. Anterior is to the right; 
scale bars (1 mm) are included. Abbreviations: tps, transverse processes; d, diapophysis of second sacral vertebra; fp, 
fracture plane (autotomy septum); ns, neural spine; pfp, partial fracture plane; ha, haemal arch. 
de Queiroz, 2011, 2013). The Mexican amber fossil would Chiapas in the early Miocene. The Simojovel mine is ca. 
thus appear to be a member of the Dactyloa clade, and pos- 6.29 degrees of latitude and ca. 696 km farther north than 
sibly a member of the aequatorialis series. the northernmost locality for any extant Dactyloa species 
The fossil could possibly be a stem anole, but crown that we found (Anolis insignis; Haroutiounian, 2017), as well 
anoles are thought to have originated much earlier than as ca. 1054 km to the northwest of that locality, all on 
the age of the Mexican amber fossil, by the early Eocene the current continental configuration. Global average tem-
(Poe et al., 2017; Sherratt et al., 2015). Therefore, if the peratures during the interval of the Miocene estimated for 
fossil is a stem anole, it must represent a late surviving the age of the fossil (15–20 Ma) are thought to have been 
but now extinct lineage that evolved the derived low num- warmer than at the present time (Scotese et al., 2021), 
ber of anterior nonautotomic caudal vertebrae convergently which presumably would have allowed species of warm-
with those in the aequatorialis and roquet series. Although adapted clades to extend their geographic distributions far-
this possibility cannot be ruled out by the currently avail- ther north. Although a continuous land connection between 
able evidence, there is no evidence that favors it over a North and South America appears not to have formed until 
simpler alternative. Similarly, the fossil could possibly be a ca. 3 Ma (Pliocene), the earlier existence of an island arc 
non-Norops member of the Digilimbus clade representing a permitted dispersals of terrestrial vertebrates between the 
colonization of the mainland by that clade separate from continents as early as ca. 20 Ma (reviewed by O’Dea et al., 
the one involving the Draconura subclade of Norops. How- 2016). In addition, a phylogenetic analysis of anole bio-
ever, that would require not only an otherwise undocu- geography found evidence for at least two dispersals from 
mented separate colonization of the mainland from the Central America to South America that predate the contin-
West Indies but also an otherwise undocumented extinction uous land connection (Poe et al., 2017). 
of that clade or lineage on the mainland. Again, the simpler Patton et al. (2021) found that when the Draconura clade 
hypothesis that the fossil is a member of the Dactyloa clade came into contact with the Dactyloa clade the ensuing in-
seems preferable. teractions favored the Draconura clade in both area colo-
Within crown anoles, only members of the Draconura nized and diversification rates. It seems unlikely, however, 
subclade of the Norops clade currently occur in Chiapas that the Mexican amber fossil, inferred to be a member 
(Etheridge, 1959; compare the list of Anolis species for Chi- of the Dactyloa clade, provides evidence of displacement 
apas in Reynoso et al., 2011 with the composition of the by the Draconura clade. At the time of the Mexican amber 
Norops and Draconura clades in Nicholson et al., 2012 and fossil, Draconura-clade anoles had already been in Central 
Poe et al., 2017). Members of the Dactyloa clade extend only America 15–20 Ma and diversified into at least 42 lineages 
as far north as northern Costa Rica in Central America (Cas- based on the time-calibrated phylogeny in Poe et al. (2017). 
tañeda & de Queiroz, 2011, 2013; Etheridge, 1959). The Instead, the fossil appears to represent a greater incursion 
likely assignment of the Mexican amber fossil to the Dacty- of the Dactyloa clade into the area previously occupied by 
loa clade indicates that this clade (and perhaps specifically the Draconura clade than at present (or previously known) 
the aequatorialis series) extended at least as far north as as well as an earlier (15–20 Ma) instance of northward dis-
DE QUEIROZ et al. 4
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A Fossil Anolis Lizard Tail in Mexican Amber: Phylogenetic Relationships a…
persal by the Dactyloa clade than the 14 Ma estimated by Queiroz et al. (1998) noted that the specimens are diffi-
Patton et al. (2021). The Mexican fossil may also represent cult to distinguish from extant species of Hispaniolan green 
an exception to the inference by Patton et al. (2021) that anoles (chlorocyanus species group of Williams, 1976; 
the expansion of Dactyloa into areas occupied by Draconura chlorocyanus series of Burnell & Hedges, 1990). Further-
was only accomplished by anole species of large body size, more, Sherratt et al. (2015) inferred that eight specimens of 
which are mostly members of the latifrons series (Castañeda Miocene Dominican amber anoles (including the two stud-
& de Queiroz, 2011). If the fossil anole is a member of ied by de Queiroz et al., 1998) are members of the crown 
the aequatorialis series (see above), species in that series clade of Hispaniolan green anoles. As noted above, 
are composed of small- to medium-sized anoles, generally SNHMB.G 8197 may be closely related to A. chloris. In the 
less than 100 mm maximum snout-vent length (SVL), while results of Poe et al. (2017), A. chloris is inferred to form 
those in the clade composed of Anolis chloris, Anolis festae, a clade with A. gorgonae and A. festae, neither of which 
and Anolis gorgonae (see below) are less than 70 mm maxi- were mentioned by Lazell (1965) or included in the phylo-
mum SVL (data from Velasco et al., 2020). genetic analysis of Castañeda et al. (2014). All three species 
Two partial specimens of Anolis lizards in amber from exhibit a mode of seven “aseptate” anterior caudal verte-
Simojovel, Chiapas have been reported previously and as- brae (Etheridge, 1959) or anterior caudal vertebrae bearing 
signed to a single extinct species, A. electrum (Lazell, 1965). transverse processes (Castañeda, unpubl. data; along with 
The two specimens may represent parts of a single individ- data for an additional specimen of A. festae from a radi-
ual, as they constitute nonoverlapping parts of the body: ograph of QCAZ R9160 in Torres-Carvajal et al., 2022). The 
the paratype consisting of the head, forelimbs, and anterior crown and stem ages of the chloris-gorgonae-festae clade 
part of the trunk and the holotype consisting of the right have been estimated at 16.91 Ma and 23.63 Ma, respectively, 
hindlimb, left pedal digit IV, and the posterior part of the overlapping the age range of the fossil (15–20 Ma), which 
trunk (Castañeda et al., 2014; Lazell, 1965). Lazell (1965) lies entirely within the 95% credible interval for the crown 
considered A. electrum to be most similar to three anole age and partially within that interval for the stem age (see 
species now considered to be members of two distantly re- Poe et al., 2017, Fig. S7). Ecologically, all three species are 
lated groups: A. chloris, a member of the aequatorialis series similar to trunk-crown, trunk, or trunk-ground anoles (Miy-
in the Dactyloa clade (see Castañeda & de Queiroz, 2013) ata, 2013; Phillips et al., 2019; Rengifo M. et al., 2015; Ve-
and A. fuscoauratus and A. maculiventris, both members of lasco & Herrel, 2007), which are also the anole ecomorphs 
the Draconura subclade of the Norops clade (see Poe et al., most commonly preserved as amber fossils (Sherratt et al., 
2017). Castañeda et al. (2014) were able to score A. electrum 2015). Thus, the limited available data suggest that the fos-
for only 10 characters commonly used in phylogenetic sil SNHMB.G 8197 (as well as A. electrum) could be part of 
analyses of anoles, and they found the relationships of that the chloris-gorgonae-festae clade and possibly even a mem-
species to be highly ambiguous. Specifically, they inferred ber of its ancestral species. 
14 alternative placements in six distantly related anole sub-
clades (clade names sensu Poe et al., 2017): (1) sister to A. 
chloris within the aequatorialis series of Dactyloa; (2) five al-
ternative positions within the latifrons series of Dactyloa; ACKNOWLEDGMENTS 
(3) sister to A. darlingtoni within Deiroptyx; (4) five alterna-
tive positions within the cristatellus series of Ctenonotus; (5) KdQ dedicates this article to the memories of Richard 
sister to A. opalinus within the grahami series (= Placopsis Etheridge (1929–2019) and James D. “Skip” Lazell 
subclade) of Norops, and (6) sister to A. ortonii within the (1939–2023). Etheridge was KdQ’s mentor and MS advisor 
Draconura subclade of Norops. at San Diego State University; his work on anole skeletal 
Because there is no overlap in the parts of the body pre- morphology (Etheridge, 1959) and lizard caudal vertebrae 
served in SNHMB.G 8197 and the holotype and paratype (Etheridge, 1967) provided important foundations for this 
of A. electrum, it is currently unknown whether they repre- contribution (see Espinoza & de Queiroz, 2008, for a biog-
sent the same or different species. However, it is notewor- raphy). Lazell facilitated anole field work by KdQ on Guana 
thy that the new specimen appears to be a member of the Island, British West Indies, and published the original de-
Dactyloa clade and exhibits a character state, seven anterior scriptions of the two previously known amber fossil anole 
“aseptate” (CV 7 of our specimen has a partial septum) or specimens from Chiapas, Mexico (Lazell, 1965). M. Reich 
transverse-process-bearing caudal vertebrae, that is known catalogued the specimen reported on here in the paleon-
in only a few species within Dactyloa, including A. chloris. tology collection of the Staatliches Naturhistorisches Mu-
Anolis chloris is one of three species (and the only one in seum Braunschweig, M. del R. Castañeda provided data on 
the Dactyloa clade) that Lazell (1965) considered most sim- numbers of caudal vertebrae bearing transverse processes 
ilar to A. electrum as well as one of the species inferred to in members of the Dactyloa clade, E. Sherratt and J. M. Huie 
be a potential sister species of A. electrum in the analysis helped with the processing of CT data, and J. B. Losos pro-
by Castañeda et al. (2014). Anolis chloris is also one of the vided helpful comments on the manuscript. CT scanning 
species in the Dactyloa clade that occurs in Central America, was supported by the DESY Block Allocation Group project 
although only in southeastern Panama (Batista et al., 2015). “Scanning the past - Reconstructing the diversity in million 
The Miocene age of the amber specimen, approximately years old fossil amber specimens using SRμCT” at PETRA 
15–20 Ma, does not preclude a close relationship to extant III. 
anole species. In the case of two fossil anoles in Dominican 
amber, which are also considered to be of Miocene age, de 
DE QUEIROZ et al. 5
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A Fossil Anolis Lizard Tail in Mexican Amber: Phylogenetic Relationships a…
SUPPLEMENTARY DATA https://www.morphdbase.de/?V_Bara-
nov_20230818-M-60.1. 
Supplementary data associated with this article can be 
found online alongside this manuscript. See also Accepted: September 04, 2024 EST 
DE QUEIROZ et al. 6
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A Fossil Anolis Lizard Tail in Mexican Amber: Phylogenetic Relationships a…
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