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This paper aims to demonstrate how subfossil bone remains from Pleistocene and Holocene deposits can help to reconstruct the history of recently extinct taxa through the example of Pholidoscelis lizards from the Guadeloupe Islands in the French West Indies. To achieve this, we conducted a new anatomical and zooarchaeological study of fossil Pholidoscelis remains collected from 23 archaeological and paleontological deposits on the Guadeloupe Islands from which this genus is nowadays absent. Our results shed light on the past existence of large Pholidoscelis lizards on all the Guadeloupe islands but also on the difficulties of confident specific identification for these remains. Nevertheless, we suggest a possible past occurrence of the now extinct Pholidoscelis major on nearly all of the Guadeloupe islands. In addition, we identified a new Pholidoscelis species, Pholidoscelis turukaeraensis sp. nov., on Marie-Galante Island, where no Pholidoscelis lizards were previously reported. This new species underwent an increase in size after the end of the Pleistocene period, possibly due to reduced predation pressure. We also highlight the consumption of Pholidoscelis lizards by pre-Columbian Amerindians and the huge impact of European colonization, which led to the extinction of all these lizards in less than 300 years.
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Historical Biology
An International Journal of Paleobiology
ISSN: 0891-2963 (Print) 1029-2381 (Online) Journal homepage:
Evolution, diversity and interactions with
past human populations of recently extinct
Pholidoscelis lizards (Squamata: Teiidae) from the
Guadeloupe Islands (French West-Indies)
Corentin Bochaton, Renaud Boistel, Sandrine Grouard, Ivan Ineich, Anne
Tresset & Salvador Bailon
To cite this article: Corentin Bochaton, Renaud Boistel, Sandrine Grouard, Ivan Ineich, Anne
Tresset & Salvador Bailon (2017): Evolution, diversity and interactions with past human populations
of recently extinct Pholidoscelis lizards (Squamata: Teiidae) from the Guadeloupe Islands (French
West-Indies), Historical Biology, DOI: 10.1080/08912963.2017.1343824
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Evolution, diversity and interactions with past human populations of recently extinct
Pholidoscelis lizards (Squamata: Teiidae) from the Guadeloupe Islands (French
Corentin Bochatona,b, Renaud Boistelc, Sandrine Grouarda, Ivan Ineichb, Anne Tresseta and Salvador Bailona,d
aLaboratoire “Archéozoologie et Archéobotanique: Sociétés Pratiques et Environnements” UMR 7209 – CNRS, MNHN – Muséum national d’Histoire
naturelle, Sorbonne Universités 55 rue Buffon, CP 56, Paris, France; bInstitut de Systématique, Évolution, Biodiversité ISYEB – UMR 7205 – CNRS,
MNHN, UPMC, EPHE – Muséum national d’Histoire naturelle, Sorbonne Universités 57 rue Cuvier, CP 30, Paris, France; cInstitut International de
Paléoprimatologie et de Paléontologie Humaine – UMR 7262 – CNRS, Université de Poitiers, UFR SFA – Bât. B35 – 6 rue Michel Brunet – TSA 51106,
Poitiers, France; dLaboratoire “Histoire naturelle de l’Homme préhistorique” UMR 7194 – CNRS, MNHN, UPVD – Muséum national d’Histoire naturelle,
Sorbonne Universités, Paris, France
This paper aims to demonstrate how subfossil bone remains from Pleistocene and Holocene deposits can
help to reconstruct the history of recently extinct taxa through the example of Pholidoscelis lizards from
the Guadeloupe Islands in the French West Indies. To achieve this, we conducted a new anatomical and
zooarchaeological study of fossil Pholidoscelis remains collected from 23 archaeological and paleontological
deposits on the Guadeloupe Islands from which this genus is nowadays absent. Our results shed light on
the past existence of large Pholidoscelis lizards on all the Guadeloupe islands but also on the diculties of
condent specic identication for these remains. Nevertheless, we suggest a possible past occurrence of
the now extinct Pholidoscelis major on nearly all of the Guadeloupe islands. In addition, we identied a new
Pholidoscelis species, Pholidoscelis turukaeraensis sp. nov., on Marie-Galante Island, where no Pholidoscelis
lizards were previously reported. This new species underwent an increase in size after the end of the
Pleistocene period, possibly due to reduced predation pressure. We also highlight the consumption of
Pholidoscelis lizards by pre-Columbian Amerindians and the huge impact of European colonization, which
led to the extinction of all these lizards in less than 300years.
e West Indies are known to be a biodiversity hot-spot inhab-
ited by many endemic species of lizards and snakes from several
genera (Henderson & Powell 2009; Powell & Henderson 2012).
Yet most of these taxa remain enigmatic, since little attention has
been paid to the study of their subfossil remains. e lizards of the
genus Pholidoscelis Fitzinger, 1843 from the Guadeloupe Islands,
in the Lesser Antilles, where they are now extinct, represent a
good example of this lack of past documentation. Pholidoscelis
was traditionally included in Ameiva Meyer, 1795, a genus of
small to large-sized teiid lizards occurring in eastern South
America, Central America and the West Indies. However, this
group was recently demonstrated to be paraphyletic (Goicoechea
et al. 2016; Tucker et al. 2016) and the West Indian members
of Ameiva, which were already recognized as forming a mono-
phyletic group in both morphological (Harvey et al. 2012) and
molecular (Hower & Hedges 2003) phylogenetic analyses, were
regrouped in Pholidoscelis (Goicoechea et al. 2016; Tucker et al.,
2016). e ten Pholidoscelis species known to occur or to have
occurred in the Lesser Antilles from the northernmost island of
Anguilla to the southernmost island of Dominica (Henderson
& Powell 2009) form a monophyletic group in all proposed phy-
logenies (Hower & Hedges 2003; Harvey et al. 2012; Pyron et al.
2013; Goicoechea et al. 2016). e southernmost Lesser Antillean
islands lack Pholidoscelis lizards (La Martinique, Sainte-Lucia and
Barbados) and/or are occupied by members of the continental
Ameiva (Saint-Vincent, some of the Grenadines and Grenada)
(Henderson & Powell 2009; Powell & Henderson 2012). Among
West-Indian islands where Pholidoscelis living specimens have
been collected, the Guadeloupe Islands present the peculiarity of
now being Pholidoscelis-free because the two species described
in the archipelago, Pholidoscelis cineracea Barbour and Noble
1915 and Pholidoscelis major Duméril and Bibron 1839 are
now extinct. Yet, many questions regarding past Guadeloupe
Pholidoscelis persist as these species were described using a very
small set of living specimens collected on anisolated islet or of
uncertain origins (Baskin & Williams 1966; Breuil 2002).
e rst mentions of Pholidoscelis (originally called Ameiva)
in Guadeloupe come from the seventeenth century histor-
ical literature. Jean-Baptiste Du Tertre (1654) and Charles de
© 2017 Informa UK Limited, trading as Taylor & Francis Group
Extinction; herpetology;
osteology; paleontology;
Received 2 May 2017
Accepted15 June 2017
CONTACT Corentin Bochaton
Supplemental data for this article can be accessed at
exists in the form of osteological subfossil remains. Indeed, many
Pholidoscelis remains were collected, mostly in archaeological
but also in paleontological deposits, on Basse-Terre, Grande-
Terre, La Désirade, Petite-Terre, Les Saintes and Marie-Galante
Islands (Figure 1) (Pregill et al. 1994; Grouard 2001; Bochaton et
al. 2015; Bailon et al. 2015). However, these remains were never
fully investigated due to their minor role in terms of the subsist-
ence strategies of human archaeological populations and because
no data concerning the osteology of the two Guadeloupian
Pholidoscelis were available. Pregill et al. (1994) and Grouard
(2001) proposed the attribution of the remains collected on
Basse-Terre and Grande-Terre Islands to P. cineracea on the basis
of geographical arguments but no real specic identication was
ever attempted. e aim of the present study is to provide a broad
review of the Pholidoscelis remains collected from archaeological
and paleontological deposits on the Guadeloupe Islands in order
to further describe theirmorphology, discuss their taxonomic
attribution, and to provide new insight into the natural history
of this now-extinct genus in Guadeloupe.
To achieve this, we rst investigate the intraspecic and interspe-
cic osteological variability of Lesser Antillean Pholidoscelis in order
to assess the possibility of distinguishing Pholidoscelis species on the
basis of bone remains. Combined with this approach, we provide
the rst data concerning the osteology of the two Pholidoscelis from
Guadeloupe, P. cineracea and P. major, using X-ray microtomogra-
phies of museum specimens preserved in alcohol. We then provide a
morphological description and, when possible, taxonomic identi-
cations of the Pholidoscelis subfossil remains from 23 archaeological
Rochefort (1658) described a medium-sized terrestrial lizards
up to 50cm long called ‘anolis’ by the Amerindians (this name
originally referred to former Ameiva lizards and was later erro-
neously attributed to other small lizards now known as Anolis
Daudin, 1802). ese lizards were described as restricted to
the Grand Cul-de-Sac Marin on Basse-Terre and Grande-Terre
Islands but uniformly distributed on all the other Guadeloupe
islands (Du Tertre 1654). However, what Du Tertre called ‘the
other islands’ cannot be indisputably established and could
refer to the islands of the Guadeloupe archipelago or to the
West Indian islands he visited, including Martinique where no
Pholidoscelis were ever described. Subsequent data concerning
Guadeloupian Pholidoscelis come from the twentieth century and
from the specimen collections from which P. cineracea and P.
major were described. e three only known living specimens
of P. cineracea were collected on the Grand Isle of Petit-Bourg,
800m from the shore of Basse-Terre Island by Barbour & Noble
(1915), who believed that this islet was the last place where the
Guadeloupe Pholidoscelis occurred. However, no evidence of
the past occurrence of this species on other Guadeloupe islands
exists. P. major is only known from three museum specimens of
unclear origin, but on the basis of the archives of the Muséum
national d’Histoire naturelle of Paris, Breuil (2002) demonstrated
that these specimens are likely to have been collected on the
Petite-Terre islands in Guadeloupe. is hypothesis is also sup-
ported by the morphological proximity between P. major and
P. cineracea (Baskin & Williams 1966). Physical evidence of the
past occurrence of Pholidoscelis in the Guadeloupe Islands also
Figure 1.(A): Map of the Lesser Antilles indicating the position of Guadeloupe archipelago; (B): Map of Guadeloupe Islands with isobaths (−200m) from Münch et al. 2013
and location of the studied sites: 1: 24, rue Schœlcher; 2: Cathédrale de Basse-Terre; 3: Gare Maritime de Basse-Terre; 4: Embouchure de la Rivière Baillif; 5: Saint-Rose La
Ramée; 6: Grotte de l’Anse à l’Ecu; 7: Morel; 8: Grotte de l’Anse à la Gourde; 9: Anse à la Gourde; 10: Pointe du Helleux; 11: Anse Petite Rivière; 12: Pointe Gros Rempart 6; 13:
Caille à Bélasse; 14: Mouton de Bas; 15: Grotte de la ravine Jean-François; 16: Morne Rita; 17: Tourlourous – Stade José Bade; 18: Grotte Blanchard; 19: Grotte Blanchard 2;
20: Grotte Cadet 2; 21: Abri Cadet 3; 22: Folle Anse; 23: Grande-Anse de Terre de Bas.Source: Maps modified from NASA Worldview:
and paleontological deposits from the Guadeloupe Islands. We also
conduct a zooarchaeological analysis in order to assess the occur-
rence of Pholidoscelis remains in archaeological sites and the inter-
actions of these lizards with Amerindian populations. Finally, by
combining all the collected data, we try to reconstruct the recent
history of Pholidoscelis lizards in Guadeloupe.
Material and methods
Fossil specimens
e studied fossil osteological material originated from 23
archaeological open-air sites and paleontological cave deposits
for a total of 1148 Pholidoscelis bone remains (Table 1). ese
sites are located on dierent Guadeloupe islands: Grande-
Terre (5 sites), Basse-Terre (5 sites), Les Saintes (1 site), Petite-
Terre (2 sites), La Désirade (2 sites) and Marie-Galante (8
sites) (Table 1, Figure 1). Details concerning the deposits and
distribution of Pholidoscelis remains in these deposits can be
found in Supplementary Material 1 (S. 1) and 2 (S. 2). e ear-
liest archaeological deposit containing Pholidoscelis remains is
Morel I on Grande-Terre Island, dated to AD 80 and attributed
to the Huecan Saladoid culture (Neoindian or Early Ceramic
Age) (Clerc 1964; Hofman et al. 1999; Romon, Bertran, Fouéré,
Hildebrand, Serrand and Vallet, 2006; Bérard 2013; Fitzpatrick
2015). e other sites are attributed to Cedrosan Saladoid cul-
tures (300 – 900 AD) or the Marmoran Troumassoid culture (900
AD-1500 AD) (Late Ceramic Age) (Bonnissent 2008; Keegan
et al. 2013; Honoré 2014). e natural deposits (by opposition
to articial archaeological deposits) are dated from the few last
centuries apart from three deposits from Marie-Galante Island
with sediments dating from Late Pleistocene to historical periods
(Stouvenot et al. 2014; Bochaton et al. 2015; Bailon et al. 2015).
Most of the faunal remains collected from these deposits have
already undergone general zooarchaeological studies (Grouard
2001; Grouard 2003; de Waal 2006; Grouard 2007; Grouard 2010;
Boudadi-Maligne et al. 2016) or studies focusing on squamate
remains (Bochaton et al. 2015; Bailon et al. 2015; Bochaton,
Boistel, et al. 2016; Bochaton, Bailon, et al. 2016; Bochaton
et al. in press).
Comparative specimens
e set of modern specimens used to describe the morpholog-
ical variability of Pholidoscelis is composed of 64 dry skeletons
representing 11 species of Pholidoscelis from the West Indies and
5 specimens of Ameiva (A. ameiva and A. bifrontata), a genus
that included Pholidoscelis in previous classications (Appendix
1). ese specimens are from the collections of the Museum of
Comparative Zoology (Harvard, USA) (MCZ), Museum national
d’Histoire naturelle (Paris, France) (MNHN) and, Smithsonian
Institution National Museum of Natural History (Washington
In addition to this set of dry specimens, we also used microto-
mographies of a Pholidoscelis fuscata specimen (MNHN-RA
1985.0473) and of three now-extinct Guadeloupian Pholidoscelis
specimens preserved in alcohol: P. major (MNHN-RA1491 and
MNHN-RA 8357) and P. cineracea (MNHN-RA 9902). X-ray
microtomographies (XMTs) were performed according to the
protocols reported by Zanette et al. (2013). We used an EasyTom
XL duo RX solution scanner at the Centre for Microtomography
of the University of Poitiers (France). e acquisitions were per-
formed at 100kV and 34/32mA. For P. cineracea and P. major
specimens, the head and the whole specimen were scanned. Only
the head was scanned for the P. fu s cat a specimen. e geometry
was set to obtain a 39.0–18.0μm voxel size in the reconstructed
Table 1.Table presenting archaeological and paleontological deposits from which Pholidoscelis remains were studied.
Notes: The Number of Remains (NR) of each Pholidoscelis species collected in the different deposits is also indicated. Calendar ages corresponding to cultural periods are
the followings: Archaic=2000–500 BC, Saladoid =AD 80–900, Troumassoid=AD 900–1490, Modern=1493 – present. More details concerning sites and collected
remains can be found in supplementary materials S. 1 and S. 2.
Islands Deposits Age NR of Pholidoscelis cf. major
NR of Pholidoscelis turukaeraensis
sp. nov.
Basse-Terre 24, rue Schoener Early Saladoid 3 0
Cathédrale de Basse-Terre Early Saladoid 94 0
Gare Maritime de Basse-Terre Early Saladoid 193 0
Embouchure de la Rivière Baillif Late Saladoid 7 0
Saint-Rose La Ramée Early Troumassoid 57 0
Grande-Terre Anse à la Gourde Late Saladoid-Early Troumassoid 112 0
Grotte de l’Anse à l’Ecu Modern 16 0
Grotte de l’Anse à la Gourde Modern 8 0
Morel Early & Late Saladoid (Morel I, II) –
Early Troumassoid (Morel III, IV)
3 0
Pointe du Helleux Late Troumassoid 32 0
La Désirade Anse Petite Rivière Late Saladoid –Early Troumassoid 11 0
Pointe Gros Rempart 6 Amerindian, Modern 81 0
Petite-Terre Caille à Bélasse Early Troumassoid 12 0
Mouton de Bas Early Troumassoid 1 0
Marie-Galante Grotte Cadet 2 Pleistocene – Late Holocene 0 112
Abri Cadet 3 Pleistocene – Early Troumassoid 0 220
Grotte Blanchard Pleistocene – Late Troumassoid 0 88
Grotte Blanchard 2 Late Holocene -Modern 0 1
Grotte du Morne Rita Archaic, Saladoid 0 2
Grotte de la ravine Jean-Francois Modern 0 16
Folle Anse Early Saladoid-Troumassoid 0 1
Tourlourous- Stade José Bade Late Saladoid-Early Troumassoid 0 60
Les Saintes Grande Anse Terre de Bas Late Troumassoid 18 0
Total 648 500
of taxonomic identication. Details concerning the morphology
of these elements can be found in Supplementary Material (S.3).
Quantication and morphometry
e quantication units used in the study of subfossil remains
are: Number of Remains (NR), Minimum Number of Individuals
(MNI) using the most abundant anatomical part of a given lateral-
ity in each site (NMIf sensu Poplin 1976) and the ‘Proportional
Representation of each skeletal part’ (PR) (Dodson & Wexlar
1979). e PR was established on the basis of the number of
each bones contained in a single Pholidoscelis skeleton; this num-
ber was estimated using two modern Pholidoscelis specimens
(USNM 259529 and USNM 236388). Osteological measurements
are those dened by Bochaton (2016). SVL was estimated using
the equations dened for Pholidoscelis by Bochaton and Kemp
(2017). All statistical analyses were conducted using the R so-
ware v. 3.1.2 ( and the package stats v. 3.1.2.
Morphology of modern Pholidoscelis
Skeletal morphological variability of extant Pholidoscelis
Our observations show that the four primarily investigated bones
(parietal, frontal, maxilla and dentary) present morphological
ontogenetic changes that strongly impact intraspecic osteologi-
cal variability in the genus. In the smallest specimens, the parietal
(Figure 2(A)) is thin, wider than long in dorsal view and presents no
dermal ornamentation. In addition, the parietal table is not laterally
well delimited, presents no lateral constriction and the supratem-
poral processes are short. On larger specimens (Figure 2(A)), the
bone becomes increasingly thicker and longer than wide, the pari-
etal table extends posteriorly, presents an increasingly pronounced
lateral constriction and a tuberculous dermal ornamentation that
progressively approaches the anterior margin of the bone, and ulti-
mately covers nearly the entire parietal table. e supratemporal
processes also extend posteriorly and present well-developed dorsal
crests. All the studied species follow the same pattern but do not
attain the same ontogenetic stage in relation to their maximal size.
A similar ontogenetic phenomenon occurs on the frontal
(Figure 2(B)). On the smallest individuals the frontal is thin with
a well-marked interorbital constriction. On larger specimens, the
bone becomes thicker and the interorbital constriction weaker,
giving a more rectangular shape to the bone in dorsal view. A
tubercular and vermiculated dermal ornamentation also devel-
ops on the dorsal surface of the bone and becomes more and
more extensive in relation to the overall size of the bone.
On the dentary and maxilla (Figure 2(C)), the dentition pre-
sents considerable intraspecic variability and we observe the
general ontogenetic pattern for dentary and maxilla teeth pre-
viously described by Pregill et al. (1988). Following this pattern,
small specimens (Figure 2(C)-a) present short, monocuspid
anterior teeth with a pointed apex and slightly posteriorly curved;
wider, laterally compressed median teeth, oen bicuspid with a
small anterior cusp and a larger posterior cusp, and with a sharp
dorsal cutting edge; posterior teeth similar to the middle teeth
but wider with a tendency to be tricuspid with an additional small
cusp on the posterior edge of the tooth (Figure 2(C)-a). However,
during the growth of the specimen, teeth and especially posterior
3D images of the heads of the specimens and a voxel size of
74.86–38.22μm for the whole specimens. e reconstruction
was performed using the soware imageJ (http://imagej.nih.
gov) and the FDK algorithms of Xact ver. 1.1 (revision=6663M,
RXsolution, acceleration in GPU). e datasets consist of 992–
5760 projections taken over 360° on the head or body of the
specimens. Direct volume rendering was used to visualize the
sub-set of selected voxels of the skeleton, and the jaw in AVIZO
7.1 and 6.1 (FEI Visualization Sciences Group, http://www.fei.
com), aer having used ImageJ to mask the anatomical structures
we were not interested in.
Skeletal maturity
Observations made on our comparative sample of Pholidoscelis
and Ameiva indicate that fusions of skeletal elements follow a
strict succession. Trunk vertebrae, with the exception of the atlas
and axis, are the rst elements to fuse (centrum with neural arch).
ey are followed by the tarsus (astragalus with calcaneum),
scapulocoracoid (scapula with coracoid), pelvic girdle (ilium
with ischium and pubis), axis (centrum with neural arch) and
nally the occipital (bones of the occipital complex: supraoc-
cipital, exooccipital and basioccipital). e last elements to fuse
are the long bones of the limbs (diaphysis with epiphysis). ese
results dier slightly from those obtained by Bochaton (2016) on
Iguana, in which pelvic girdle fused later than in Pholidoscelis,
but otherwise skeletal maturation follows the same general pat-
tern. e succession we observed added more details to earlier
observations based on teiid lizards of the genus Cnemidophorus
Wagler, 1830 by Maisano (2002). Following this last author and
considering the simultaneous fusion of some elements, speci-
mens for which the pelvic girdle, axis or occipital are fully fused
reached at least half of their maximal size and are sexually mature
while specimens with fully fused long bones reached at least
80% of their maximal size. ese data are useful for the study of
subfossil Pholidoscelis remains combined with skeletal measure-
ments (see Bochaton 2016) and Snout-Vent Length reconstruc-
tion (referred to hereaer as SVL), obtained using size estimation
equations previously established for Pholidoscelis (Bochaton and
Kemp, 2017), to dene the Minimal eoretical Maximal Size
(MTMS) of a fossil population using the size of the largest imma-
ture bone (Bochaton 2016). is MTMS corresponds to the size
the specimen may have reached if it had not died beforehand
and display size information in the light of skeletal maturity
to avoid misinterpretation due to the occurrence of non-fully
mature specimens in fossil deposits.
Systematics and anatomical description
e taxonomy used is that of Goicoechea et al. (2016). We will
sometimes have to refer to the former paraphyletic genus Ameiva
(sensu lato), which include all members of the current genera
Pholidoscelis and Ameiva. Anatomical descriptions primarily
focus on the four most common Pholidoscelis bones found in
archaeological and paleontological deposits: the parietal, fron-
tal, maxilla and dentary. e occurrence of other subfossil bone
elements is taken into account in the quantitative study but
their scarcity in fossil deposits limits their usefulness in terms
stated that A. ameiva posterior teeth were strictly tricuspid. As
regards the number of teeth, we observed an intraspecic varia-
bility of a maximum of 4 teeth on the dentaries and maxillaries
of our modern Pholidoscelis sample; however, we lacked the full
size range for each taxon. e number of tooth positions varies
from 15 to 27 on the dentary and from 13 to 23 on the maxilla
of the investigated Pholidoscelis species. e number of teeth was
partially correlated with the Length of the Dental Row of both
dentaries (Linear regression; R2=0.68; p<0.05) and maxillaries
(Linear regression; R2=0.78; p<0.05) of the species for which
we had the largest number of specimens (P. griswoldi).
e main information provided by these observations is
that the considerable ontogenetic variability we observed in
Pholidoscelis prevents or at least impedes in most cases the estab-
lishment of specic diagnostic characters applicable to modern
Skeletal morphology of extinct Pholidoscelis museum
specimens from the Guadeloupe Islands
e large size of the two P. major specimens (MNHN-RA
8357; SVL=208mm and MNHN-RA 1491, SVL=190mm) is
reected in their osteological morphologies. Both possess orna-
mented parietal and frontals of ‘adult’ morphology and middle
teeth tend to lose cuspids to become bicuspid, then monocuspid
(Figure 2(C)-b). In the most extreme morphologies, the apex of
these monocuspid teeth tends to enlarge and to become blunt,
to form bulbous teeth, commonly called crushing teeth (Estes &
Williams 1984; Kosma 2004) (Figure 2(C)-c). is ontogenetic
dri occurs to a variable extent in all the observed species but we
still observed some signicant dierences between taxa. Indeed,
our observations combined with previous observations by Pregill
et al. (1988) show that most of the Lesser Antillean Pholidoscelis
present the full ontogenetic variability described above (P. plei, P.
griswoldi, P. f us ca ta and P. pluvianotata), but the crushing teeth
seem to be absent in P. corax. Most of the Greater Antillean
Pholidoscelis specimens present tri- or bicuspid posterior teeth
with the exception of our biggest P. chrysolaema parvoris speci-
men (USNM 259531) with monocuspid posterior teeth and a P.
auberi obsoleta specimen (MCZ 152525) with posterior crushing
teeth. In addition, Pregill (1981) reported that the occurrence of
crushing teeth in Greater Antillean fossil P. exsul. In this regard,
Ameiva ameiva diers from most Pholidoscelis as it presents pos-
terior teeth retaining a tricuspid morphology, even for the largest
specimens, with the exception of one large-sized specimen from
French Guyana (MNHN-RA 1991.4034) with bicuspid posterior
teeth. is observation contradicts those of Kosma (2004) who
Figure 2.Ontogenetic variability of Pholidoscelis bones. A: Parietal (dorsal view); B: Frontal (dorsal view); C: Dentary (medial view).
Abbreviations: b. t. : bicuspid teeth; c. t.: crushing teeth; d. o.: dermal ornamentation; i. c.: interorbital constriction; l. c.: lateral constriction; p. t .: parietal table; t. t.: tricuspid teeth; s. p. :
supratemporal process. (Drawings: C. Bochaton).
increase in tooth count (Estes et al. 1988); parietal lacking pineal
foramen (Gauthier et al. 2012); frontals fused together bearing
deep dorsolateral imprints of the nasal near their anterior mar-
gin, low ventral cranio-frontal crest and lacking antero-ventral
descending process (Estes et al. 1988; Evans 2008; Gauthier et
al. 2012); jugal lacking postero-ventral process and bearing an
ectopterygoid medial process (Tedesco et al. 1999; Evans 2008;
Gauthier et al. 2012); lack of well-dened structure for the
insertion of the basipterygoid on the pterygoid (Evans 2008);
postfrontal bearing a double process bordering the frontopa-
rietal suture (Evans 2008); occurrence of a ventromedial crest,
well-developed tympanic fossa and strong tympanic crest on
the quadrate (Evans 2008); coronoid with posterior branch and
well-developed antero-labial branch (Evans 2008); articular with
a ventrally oriented angular process (Conrad 2008; Evans 2008;
Gauthier et al. 2012); dentary with a weakly developed subdental
shelf (Gauthier et al. 2012), curved ventral margin and fully open
Meckel groove (Evans 2008); heterodont bicuspid or monocuspid
pleurodont teeth included in basal cement and basally pierced
by a subcircular replacement pit (Estes et al. 1988; Evans 2008;
Pujos et al. 2009); procelous vertebrae presenting well-developed
zygosphenes-zygantrum articulations that lack an anterior notch
(Hostetter & Gasc 1969; Gauthier et al. 2012); iguanid type IV
caudal vertebrae of Etheridge (1967) and the occurrence of a frac-
ture plane on nearly all caudal vertebrae except the most anterior
(Etheridge 1967); absence of scapular fenestra and occurrence
of all other morphological structures of the scapulo-coracoid
(Lécuru 1968); occurrence of an ilium tubercle (Gauthier et al.
2012), and humerus with a single distal fossa and a delto-pectoral
crest forming an angle of 90° with the proximal part of the sha
(Lécuru 1969). e following characters of Teiinae Presch, 1970
following Gauthier et al. (2012) also occur on the remains: pre-
maxilla nasal process widens across nares; facial process of the
maxilla covering more than 56% of the total length of the bone,
and occurrence of a sus-orbital bulge on the prefrontal. Finally,
the remains are attributed to Pholidoscelis on the basis of their
strong similarities with comparative specimens, geographical
occurrences and presence of the following characters reported
to occur in Ameiva (sensu lato): lack of pterygoid teeth (Evans
2008); opening of the Meckel groove of the dentary only visible
in lingual view (Evans 2008) and loss of lateral process ante-
rior to the fracture plane in the most posterior caudal vertebrae
(Etheridge 1967).
Maxilla (Figure 4(A) and (B)) – is bone displays a high and
long facial process (covering more than 70% of the total length
of the bone) with a vertical anterior margin. In medial view, the
facial process bears a more or less horizontal medial crest mod-
erately extended on the dorsal face of the supradental shelf (sensu
Rage & Augé 2010). is last structure is straight, thin, long and
slightly incurved in medial direction in its posterior part. e
facet of the jugal is visible near the posterior extremity of the
bone. e maxillary foramen (superior alveolar foramen sensu
Oelrich 1956) is large and occurs on the posterior quarter of the
supradental shelf. In dorsal view, the bone displays a short and
pointed external premaxillary process and a wider and longer
internal premaxillary process. e Length of the Dental Row
(or tooth row length) of this bone is between 13.8 and 28.5mm.
or posterior monocuspid teeth with an enlarged apex (previously
described as crushing teeth) on the dentary and maxilla (Figure
3(A) and (B)). ese specimens are the largest of our comparative
material, as reected by their osteological measurements: Parietal
(13.7 and 12.4mm of Greatest Length), Frontal (6 and 5.6mm
of Smallest Width – or interorbital length – and 19 and 15mm
of Greatest Length), Maxilla (Length of the Dental Row of 25.6
and 23.2mm), Dentary (Length of the Dental Row of 26.6 and
25.3mm). Skeletal maturity indicates that the largest specimen
where the long bone diaphyses are fully fused with their epiphy-
ses, was at least 80% of its maximal size (Maisano 2002), whereas
this is not the case for the smallest specimen. Apart from large
sizes and associated morphology, we did not discover any specic
osteological characteristics on these specimens.
e specimen of P. cineracea (MNHN-RA 9902) is smaller
(SVL=135mm) and presents a less ontogenetically advanced
skeletal morphology than P. major specimens (Figure 3(G) and
(H)). It also presents a combination of clearly bicuspid and
blunted posterior teeth, which also suggests that the specimen
did not reach the last ontogenetic stage. Measurements taken
on the main bones of this specimen are the following: Parietal
(9.7mm of Greatest Length), Frontal (14mm of Greatest Length
and 4mm of Smallest Width -or interorbital length-), Maxilla
(18.2mm of Length of the Dental Row), and Dentary (19.6mm
of Length of the Dental Row). is specimen, and the two other
specimens described by Barbour & Noble (1915), are indicated
as being adults by these authors. However, the fact that the long
bones are not fused with their epiphyses indicates that this spec-
imen did not reach its maximal size in spite of being mature.
e holotype specimen of this species (MCZ-R10577) is larger
(SVL=150mm) than the paratype from the Muséum national
d’Histoire naturelle, thus its skeletal morphology may dier from
our specimen. However, on the basis of size at similar maturity
stages (fusion stages of bones and occurrence of blunted teeth),
it seems unquestionable that P. cineracea is smaller than P. major.
Apart from size and associated morphological characteristics,
we were unable to detect any osteological dierences between
P. cineracea and P. major.
Systematic paleontology of fossil Pholidoscelis from
Squamata Oppel, 1811
Teiidae Gray, 1827
Pholidoscelis cf. major Duméril & Bibron, 1839
is material includes all the Pholidoscelis remains found in
Basse-Terre, Grande-Terre, La Désirade, Petite-Terre and Les
Saintes Islands for a total of 648 remains (Table 1, S. 2), rep-
resenting all skeletal parts. All these remains were collected in
pre-Columbian archaeological or natural Holocene deposits (see
details inTable 1, S. 1).
Diagnostic osteological characters
ese remains were attributed to Teiidae Gray, 1829 on the basis
of the combined occurrence of the following osteological char-
acters: single premaxilla (Evans 2008); maxillary presenting a
long and high facial process (Evans 2008); reduced ontogenetic
ridges and grooves running toward the nasal imprint. On larger
specimens, the ornamentation is more marked and occurs on the
whole dorsal surface of the bone. e tubercles become higher
but the longitudinal groove between the frontal scales is still well
visible. However, on the largest specimens, the tubercles are so
large that they fused together and cover all the dorsal side of the
bone in the same way as what we observed on parietals. In ventral
view, the cranio-frontal crests are low, prefrontal impressions
occur on the anterior half of the lateral margins of the bone,
and the impression of the frontal processes of the post-orbitof-
rontals are visible on the anterior margins of the postero-lateral
processes. e frontal dermal ornamentation pattern was unclear
on the scan of P. major (MNHN-RA 1491) (Figure 3(D)), in
which the suborbital area of the frontal seems free of dermal
ornamentation, but this could reect an early ontogenetic stage
where dermal ornamentation still does not occupy the whole
dorsal area of the bone, as shown by the weak development of the
tubercles occurring in the median area of the bone of this spec-
imen. Parietal – e parietals (Figure 4(D)) present the same
e dental row displays a tooth count between 18 and 23. As
regards the dental morphology, the full ontogenetic variability
observed on modern Pholidoscelis can be observed from juvenile
to adult morphology. e variability of the number of teeth (5)
slightly outranges the intraspecic variability observed on liv-
ing Pholidoscelis (4). Frontal –Frontals (Figure 4(C)) present the
same ontogenetic variability observed on modern Pholidoscelis.
eir Smallest Width ranges between 4.5 and 7.2mm. e frontal
table is slightly transversally convex and lacks dorsal longitudinal
ridges (sensu Harvey et al. 2012). In dorsal view, a longitudinal
groove marking the separation of the two posterior frontal scales
occurs on the posterior half of the bone. e anterior part of the
bone is trifurcated and displays two deep nasal impressions. Most
of the specimens bear dermal ornamentation dorsally. On the
smallest specimens, this dermal ornamentation is formed by a
lot of small and low tubercles of dierent size, randomly organ-
ized on the posterior half of the bone. ese tubercles tend to be
smaller in the most posterior part of the bone. On the anterior
part, the tubercles become vermiculated and form longitudinal
Figure 3.Direct volume rendering obtained from the X-ray microtomography of P. cineracea (MNHN- RA 9902; A-D) and P. major (MNHN-RA 1491; E-H): (A) & (E): Maxillary
(medial view); (B) & (F): Dentary (medial view); (C) & (G): Parietal (dorsal view); (D) & (H): frontal (dorsal view).
Abbreviations: b. t.: bicuspid teeth; c. t.: crushing teeth; d. o.: dermal ornamentation. (Scale bar=2mm).
Dentaries are mostly of large size, between 32.1 and 22.2mm of
the Length of the Dental Row. ey bear between 20 and 28 teeth
ranging in morphology from monocuspid teeth with pointed
or blunted apex for the most anterior to bicuspid, tricuspid or
broad monocuspid (crushing teeth) teeth for the more poste-
rior ones. Some bones also show slight dermal ornamentation
formed of small ridges and tubercles on the anterolabial surface.
e considerable variability observed regarding the number of
teeth exceeds the maximal intraspecic variability observed on
modern Pholidoscelis (4 teeth). is dierence could possibly be
linked to the occurrence of several species in the material but
on account of their morphological similarity, we cannot detect
them using only qualitative methodology.
Additional descriptions of other skeletal elements can be
found in Supplementary material S.3.
In regard to the size of these fossil Pholidoscelis (Figure 5(A)),
although we observed the morphological homogeneity of the
Pholidoscelis remains collected on Basse-Terre, Grande-Terre,
ontogenetic variability observed on all modern Pholidoscelis. e
Maximal Lengths of the only two complete remains are 11.45mm
and 16.6mm; these bones both present the morphology of a large
specimen, as described in our modern sample. ey bear dermal
ornamentation strictly limited to the anterior part of the parietal
table, even on the largest specimens with tubercles that tend to
randomly fuse together to form a bony plate on the anterior half
of the parietal table. Dentary (Figure 4(E) and (F)) – e bone is
elongated, relatively taller posteriorly in the largest specimens. In
lingual view, the Meckel groove is visible and open over its whole
length. is opening is wide in the posterior part of the bone
and very narrow in the anterior part. e subdental shelf (sensu
Rage & Augé 2010) is weakly developed, dorsally concave and
bears an imprint of the splenial in the posterior half of the bone.
e bone lacks a sulcus dentalis (sensu Rage & Augé 2010). e
mandibular symphysis is reduced and dorso-ventrally attened.
In labial view, an imprint of the antero-lateral process of the
coronoid is visible in the posterior part of the bone and reaches
the fourth to h tooth position from the rear. A row of ve to
eight labial foramina occurs on the anterior half of the bone.
Figure 4.Remains attributed to Pholidoscelis cf. major: (A): Left maxilla from La Désirade (Pointe-Gros Rempart 6 site- Décapage 10) (medial view); (B): Right maxilla from
Basse-Terre (Gare Maritime de Basse-Terre site, US 1009) (medial view); (C): Frontal from Basse-Terre (Cathédrale de Basse-Terre site – Sondage 7 US 1060) (dorsal view); (D):
Parietal from La Désirade (Pointe-Gros Rempart 6 site- Décapage 7) (dorsal view); (E): Right dentary from La Désirade (Pointe-Gros Rempart 6 site- Décapage 8) (medial
view); (F): Right dentary from Basse-Terre (Gare Maritime de Basse-Terre site, US 1008) (medial view).
Abbreviations: b. t.: bicuspid teeth; c. t.: crushing teeth; d. o.: dermal ornamentation; f. p.: facial process; l. g.: longitudinal groove; m. c.: medial crest; M. g.: Meckel groove; m. f.: maxillary
foramen; m. s.: mandibular symphysis; m. t.: monocuspid teeth; n. i.: nasal impressions; p. p.: postero-lateral process; p. t.: parietal table; sb. s.: subdental shelf; sp. s.: supradental shelf. (Scale
bar=2mm; Drawings: C. Bochaton).
sample of large Pholidoscelis species, which is necessary for
clearly understanding growth patterns (Brown et al. 2015) and
the intraspecic maximal variability of the dental rows. ese
observations were not repeated on the maxillaries as the vari-
ability in the number of teeth was too low. Most of the maxil-
laries fall into the range of the number of teeth observed on P.
major (19–21) (Figure 4(A)) and only one maxilla presents a
dental row bearing 23 teeth (Figure 4(B)). ese observations
do not represent clear arguments for a specic attribution of
fossil remains since no morphological character is characteris-
tic of a single Pholidoscelis species. However, size data revealed
that the size of fossil Pholidoscelis matches the Guadeloupian P.
major (around 190–200mm SVL) and diers from the smaller
P. cineracea (around 130–150mm SVL). e size of the fossils
is also comparable to the largest extant Pholidoscelis, P. f u sc at a
from Dominica but the dermal ornamentation of the frontal of
this species presents a dierent organization pattern (see above).
Fossil size data indicate possible sexual dimorphism, at least in
Pholidoscelis from Basse-Terre, since the two dierent MTMS
observed on this island are comparable to the sexual size dimor-
phism noted on modern P. f u s c a t a (males up to 200mm SVL
and females up to 154mm SVL (Schwartz & Henderson 1991)).
Observed sizes of Basse-Terre fossils could also match with the
two morphotypes of fossil dentaries bearing dierent numbers
of teeth described above but such a dierence related to sexual
dimorphism was never reported in Pholidoscelis or in Ameiva
(sensu lato). As a result of these observations, we suggest an
attribution of the fossil remains discussed above as Pholidoscelis
cf. major. In the event where P. major and P. cineracea might
enter into synonymy, as suggested by their strong morphological
similarity, the name P. major being the oldest would be kept and
our attribution would remain unchanged.
Pholidoscelis turukaeraensis sp. nov.
‘Cadet 2-O21-U5c-Phol-1’, nearly complete frontal recovered
from the Late Pleistocene layer U5c-O21 of Cadet 2 (Bochaton
et al. 2015) on Marie-Galante Island. is material, along with
other described specimens, is stored at the deposit of the
Service Régional d’Archéologie of Guadeloupe (Le Moule/
Paratype material
Two additional mostly complete frontals ‘Cadet 2-O21-U5a-
Phol-2’ and ‘Cadet 3-E4-C7-Phol-3’ were recovered from Cadet
2 and Cadet 3, Marie-Galante, deposits in U5a-O21 and E4-C7
layers dated respectively from the Late Pleistocene and Holocene.
Additional material
In addition of frontals, 497 non-diagnostic elements (MNI=30)
from all Pleistocene and Holocene Marie-Galante Island deposits
were attributed to Pholidoscelis turukaeraensis (Table 1, S. 2).
Dierential diagnosis
e frontals of this species dier from all frontals of the other
examined members of Pholidoscelis by the occurrence of
dorsal depressions posterior to the nasal impressions in dor-
sal view. ese bones also dier from other Guadeloupian
fossil Pholidoscelis in that they are slender and their dermal
Petite-Terre and Les Saintes Islands, clear size dierences
emerge between fossil specimens from the dierent islands.
e MTMS observed on the Pholidoscelis remains from Basse-
Terre Island was 242 mm, with a mean of all estimated SVL
of 143mm (n=85). On Grande-Terre Island, the MTMS was
194mm, with a mean of all estimated SVL of 144mm (n=14)
and on La Désirade Island the MTMS was 210mm, with a mean
of all estimated SVL of 176mm (n = 10). e only estimated
size obtained from Petite-Terre Island was 187 mm SVL and
no MTMS could be estimated due to the lack of maturity data
concerning this remain. A peculiarity of Basse-Terre Island, is
that the Minimal eoretical Maximal Size (MTMS) of the spec-
imens strongly diers if it is based on the smallest mature bone
(MTMS 1=208.7mm of SVL) and largest immature long bone
(MTMS 2=242mm of SVL). is seems to indicate the existence
of populations of two dierent sizes (Figure 5(A)) likely to be
related to sexual dimorphism. e low number of estimated size
data for the other Guadeloupe islands does not allow for similar
observations regarding the occurrence of dierent MTMS and
the global distribution of size data.
Identication and comments
Because of the considerable morphological intraspecic variabil-
ity and the morphological homogeneity of modern Pholidoscelis
species, it is dicult to identify these fossil remains to species
level. Nonetheless, some interesting morphological observations
can be made: 1) the fact that the dermal ornamentation of fossil
parietals (Figure 4(D)) is limited to the anterior part of the pari-
etal table of the large specimens is a character we only observed
on the Guadeloupian Pholidoscelis (P. major and P. cineracea;
see Figure 3(C) and (G)) and on P. f u s c a t a whereas ornamen-
tation covered the full parietal table on large specimens of P.
chrysolaema, P. exsul and P. plei. Specimens of allliving species
did not present well-developed dermal ornamentation so their
pattern of ornamentation wasin many cases impossible to asses;
2) the ornamentation pattern observed on fossil frontals (Figure
4(C)) is similar to that of many Lesser Antillean Pholidoscelis
(P. pluvianotata, P. plei, P. corax, P. griswoldi, P. erythrocephala),
including P. cineracea and P. major but is also similar to at least
one Greater Antillean Pholidoscelis (P. chrysolaema). However,
P. f u s c a t a from the neighboring island of Dominica seems to
present a dierent pattern with more regularly shaped tubercles
that do not extend to the lateral margins of the bone. e pattern
of P. f u s c a t a was also observed on P. auberi from Cuba and on
P. exsul from Puerto-Rico by Pregill (1981). is last pattern
also strongly diverges from the continental members of Ameiva
(sensu lato) since the A. ameiva frontal presents more regularly
shaped tubercles restricted to the posterior and medial parts of
the bone; 3) although some of the fossil frontals are larger than
in any extant Pholidoscelis, they bear weakly developed dermal
ornamentation suggesting that they were probably still far from
their maximal sizes; 4) the number of teeth observed on the
dentaries was highly variable. e fossil specimens presenting
the fewest teeth (20–23) (Figure 4(E)) display a number of teeth
similar to P. major specimens (22 and 24 tooth positions) and
their morphology cannot be distinguished from the latter. Yet
other specimens present more teeth (up to 28) (Figure 4(F)).
We are currently unable to understand the signicance of this
dierence in the number of teeth as we lack an appropriate
nearly complete frontals recovered on Marie-Galante Island are
smaller than the holotype and present smaller Smallest Width
(‘orbital constriction, SW), as low as 2.6mm. ey also present
weakly marked to absent dermal ornamentation, suggesting they
are from younger specimens than the holotype. Nasal impres-
sions on these frontals also seem weaker than in the holotype
but their morphology is otherwise similar and they all present
dorsal depressions. Other elements (Figure 7; S. 2): other skeletal
elements are similar to those described for P. cf. major, yet we
provide here some information regarding their size and minor
anatomical characteristics. Maxilla (Figure 7(A))- ree com-
plete adult maxillae presenting crushing teeth were collected
on Marie-Galante Island. eir LDR are 14.1, 20.2 and 22mm
and they bear respectively 16, 20 and 21 tooth positions. Some
smaller specimens presenting tricuspid posterior teeth were also
collected. Parietal (Figure 7(B)) – e two Pholidoscelis parietals
collected on Marie-Galante Island measure 10.5 and 13.7mm of
Greatest Length (GL) and 10.3 and 11.2mm of anterior Width
(Wa). ey present well-marked dermal ornamentation in the
anterior half of the parietal table and are formed of large well-in-
dividualized tubercles. Dentary (Figure 7(C) and (D))- Out of
ve juvenile dentaries bearing tricuspid posterior teeth, the mor-
phologies of Pholidoscelis dentary collected on Marie-Galante
Island is similar. Lengths of the Dental row (LDR) of the only
two complete dentaries are 19.1 and 22mm for 20 and 22 den-
tal positions. ese dentaries present the peculiarity of bearing
crushing teeth in spite of their small size compared to fossil ele-
ments from the other Guadeloupe islands and could thus be
associated with adult specimens of smaller size. e occurrence
of crushing teeth was also observed on fragmented specimens.
e only exception to this trend is a dentary collected from a
Holocene layer presenting a LDR of 20.6mm, bearing 20 dental
positions but lacking crushing teeth.
e MTMS indicates that size dierences exist between
Pholidoscelis remains collected in Pleistocene and Holocene layers.
Indeed, thisvalue ishigher in Holocene layers (MTMS=187mm,
n=33) than in Pleistocene layers (MTMS=142mm, n=19)
(Figure 5(B)).
ornamentation pattern is formed of strongly concentrated small
tubercles in the medial part of the frontal and homogeneous
groups of tubercles in the posterior part of the frontal.
e species name turukaeraensis (pronounced ‘touloukaeraen-
sis’) refers to the original name of Marie-Galante Island given
by Arawak Amerindians long before Europeans renamed the
island (Barbotin 1968). e meaning of this name is ‘the island of
touloulous’, and the touloulous are terrestrial crabs (Gecarcinus)
still very common on the island. We propose the English name of
‘Marie-Galante ameiva’ and the French name ‘améive de Marie-
Galante. We choose to retain the historically used terms ‘ameiva’
and ‘améive’ for these lizards.
Holotype specimen (Figure 6(A)) – is frontal measures
19.4mm maximal length (GL), 10.9mm posterior width (Wp)
and 5.7mm smallest width (‘orbital constriction, SW). e bone
is formed by the fusion of the two frontals. In dorsal view, the
bone is elongated, presents a moderately marked orbital con-
striction and a weaker constriction in its anterior half. e
anterior part of the bone is trifurcated but the anterior tips are
partly broken. Two very deep nasal impressions occur at the
anterior extremity of the bone. e bone is covered by dermal
ornamentation concentrated in the medial part of the bone and
nearly absent from the lateral margins in the posterior part of
the bone. is ornamentation forms homogeneous packs of
tubercles fused together in the posterior part of the bone and
large well-individualized tubercles at mid-length of the bone.
ese tubercles become long and vermiculated in the anterior
part of the bone. Posterior to the nasal impressions, two wide
and shallow depressions occur in which the dermal ornamen-
tation is less pronounced than in the other parts of the bone. In
ventral view, the cranio-frontal crests are low, and a very low
antero-ventral descending process occurs. In lateral view, the
impression of the prefrontal occurs on the whole anterior half
of the bone. Well marked impressions of the post-orbitofrontal
occur on the anterior margins of the postero-lateral processes
of the bone. Paratype specimens (Figure 6(B,C)) – Other two
Figure 5.Estimated SVL and MTMS of fossil Pholidoscelis from Basse-Terre (A) corresponding to the smallest mature bones (MTMS 1) and largest immature bones (MTMS
2) and from Marie-Galante (B) corresponding to the largest immature bones from Pleistocene and Holocene layers.
However, the size of Holocene specimens is roughly similar to
the size of fossil Pholidoscelis observed on other Guadeloupe
islands at the same period. We did not use these size dierences
as an argument to propose dierent taxonomic attributions for
Pleistocene and Holocene Pholidoscelis from Marie-Galante
as we could not test the occurrence of frontal depressions on
large Marie-Galante Holocene fossil Pholidoscelis based on the
available material. us, we postulated that P. turukaeraensis
increased in size between the Pleistocene and Holocene peri-
ods,but the possibility of the arrival of new Pholidoscelis species
from another islandcannot be unambigiously discarded. is
dierence in size cannot be explained by a sample bias between
natural and archaeological deposits as, by taking account of the
skeletal maturity of the specimens in size estimation, the MTMS
is robust to such bias (Bochaton 2016). In addition, the small
size of Marie-Galante Pholidoscelis is also visible on dentary and
maxillary bearing smaller crushing teeth than on other islands.
Indeed, the LDR of the smallest fossil maxillary presenting crush
ing teeth on Marie-Galante is only 14mm, as opposed to 22mm
on the other Guadeloupe islands. A similar trend occurs for the
dentary, with the smallest LDR of 19.1mm on Marie-Galante as
opposed to 26mm on the other islands.
Identication and comments
Following the observations made on our comparative sam-
ple, P. turukaeraensis diers from all extant and extinct Lesser
Antillean Pholidoscelis on account of the dorsal depressions
posterior to the nasal impressions of the frontal in dorsal view.
Such depressions were not observed in living Pholidoscelis but
however occur in members of the former genus Ameiva (sensu
lato): Ameiva ameiva (MCZ R-173127) and Ameiva bifrontata
(MNHN UMR7209–343) from Central America. e latter pre-
sents large and deep depressions in the same area as P. turukaer
aensis. In addition, the dermal ornamentation pattern observed
on P. turukaeraensis frontals diers from the other Guadeloupian
large fossil Pholidoscelis for which dermal ornamentation is
more disorganized on the whole surface of the bone. For these
reasons, we separated the Pholidoscelis remains collected on
Marie-Galante from other Guadeloupian Pholidoscelis on the
basis of its frontal morphology that is dierent from that of all
the other observed West Indian Pholidoscelis. Size dierences
between Marie-Galante Pholidoscelis and large Pholidoscelis (P.
major and P. fuscata) also exist but only concern Pleistocene
forms. Indeed, Pleistocene specimens (max. 142 mm SVL)
were clearly smaller than Holocene ones (max. 187mm SVL).
Figure 6.Frontals of Pholidoscelis turukaeraensis sp. nov.: (A): Holotype frontal from Cadet 2, Late Pleistocene layer U5c-O21 (Cadet 2-O21-U5c-Phol-1); (B): Additional
frontal from Cadet 2, Late Pleistocene layer U5a-O21 (Cadet 2-O21-U5a-Phol-2); (C): Additional frontal from Cadet 3, Holocene layer E4-C7 (Cadet 3-E4-C7-Phol-3).
Abbreviations: c. c.: cranio-frontal crest; d. o.: dermal ornamentation; d. d.: dorsolateral depression; l. g.: longitudinal groove; n. i.: nasal impression; o. c. : orbital constriction; p. p.: postero-
lateral process; p. tu.: pack of tubercles; pf. i.: prefrontal impression; po. i.: post-orbitofrontal impression. (Scale bar=2mm; Drawings: C. Bochaton).
Zooarchaeological analysis
As previously explained, the investigated Pholidoscelis remains
come from two dierent types of accumulations: archaeological
open-air sites (605 remains) and natural cave deposits (543
remains). Cave deposit accumulations were formed under the
inuence of many dierent biotic or abiotic processes but a
discussion of the accumulation processes of all the investi-
gated caves is beyond the scope of this study. is question
was tackled for Blanchard Cave (Bailon et al. 2015), where
we observed digestion traces on the fossil remains collected
from the site. However, humans are considered to be the main
Figure 7.Pholidoscelis turukaeraensis sp. nov.: (A): Left maxilla, Marie-Galante (Cadet 3-E3-C5-Phol-4) (medial view); (B): Parietal, Marie-Galante (Cadet 3-E3-C9-Phol-5)
(dorsal view); (C): Right dentary, Marie-Galante (Cadet 3-E3-C5-Phol-7) (medial view); (D): Right dentary, Marie-Galante (Cadet 2-O21-U5b-Phol-6) (medial view).
Abbreviations: c. t.: crushing teeth; d. o.: dermal ornamentation; p. t .: parietal table; t. t.: tricuspid teeth. (Scale bar=5mm; Drawings: C. Bochaton).
Figure 8.Anatomical distribution of Pholidoscelis remains from natural deposits (A) and archaeological deposits (B). The unit used is the ‘Proportional Representation of
each skeletal part’ (PR) (Dodson and Wexlar, 1979).
Figure 9.Pholidoscelis archaeological bones presenting fire traces: (A) Lateral view
of an anterior caudal vertebrae from Gare Maritime site (US1003), (B) Lateral view
of a coronoid from Tourlourous site (Z4 S3 P7). (Scale bar=5mm).
e vast majority of the specimens collected in the archaeo-
logical deposits present an adult size between 100 and 220mm
SVL, which is relatively similar to the estimated MTMS for each
island. However, the limited quantity of data does not allow for
inter-island comparisons of selection intensity, as for Iguana
(Bochaton, Bailon, et al. 2016). Collection methods in the eld
could also strongly impact this size distribution since smaller
specimens are less likely to be collected even if sieves are used
because the most commonly used mesh (2.7 mm or more)
is too large to collect parts of small specimen below 100mm
SVL. Consequently, small specimens are nearly absent from the
archaeological material and it is dicult to assess the possible
hunting practices on these lizards.
e observation of modern Pholidoscelis skeletal morphology
shows that species identication on the basis of qualitative char-
acters is nearly impossible. is is mainly due to the marked
intraspecic variability we observed in the genus and the strong
morphological similarity that made modern taxa nearly impos-
sible to dierentiate solely on the basis of their bones. Species
of Pholidoscelis extinct in Guadeloupe (i. e. P. cineracea and
P. major) were no exception to this general observation. us the
identication of subfossil remains of Pholidoscelis in Guadeloupe
was dicult and we had to rely on size and geographic argu-
ments to propose an attribution of the fossil remains from most
Guadeloupe islands to Pholidoscelis cf. major. However, the occur-
rence of a character we did not observe on any other Pholidoscelis,
the anterior dorsolateral depression on the frontal, wasused us
to describe a new species, Pholidoscelis turukaeraensis, restricted
to Marie-Galante Island. ese results are not surprising since
they are in accordance with the geological history of the archi-
pelago. Indeed, isobaths show that Basse-Terre, Grande-Terre,
La Désirade and Petite-Terre islands were connected during the
Pleistocene whereas Marie-Galante was never connected to any
other island throughout its geological history (Münch et al. 2013).
A similar phenomenon occurs for instance in the Anguilla Bank
Islands, which were connected during the Pleistocene and which
are now inhabited by the same species of Pholidoscelis (P. plei)
(Powell & Henderson 2012). e Marie-Galante Pholidoscelis
seems to share its frontal dermal ornamentation pattern with
P. fuscata, whereas it diers from other Guadeloupian Pholidoscelis.
e possibility of close phylogenetic proximity between P. t ur u -
kaeraensis and P. fu s ca ta would make the colonization scenario
of Marie-Galante Island by Pholidoscelis similar to that of Anolis,
since the Marie-Galante anole, Anolis ferreus, is also molecu-
larly closer to the anole from Dominica, Anolis oculatus, than
to any Guadeloupian anoles (Nicholson et al. 2012). is could
be explained by the fact that Marie-Galante is an isolated island,
which emerged later (800 ky ago) than the volcanic islands of the
Guadeloupe archipelago that started their formation between
4.7 and 2.7 My ago (Münch et al. 2013) and thus has a dierent
and more recent colonization history. Our work also demon-
strates that there is no fossil evidence showing that the so-called
Guadeloupe ameiva (P. cineracea) was occurring outside of the
Islet where living specimens were collected (Barbour & Noble
1915). However, the hypothesis that P. major occurred on all the
Guadeloupe islands also seems dubious because the collected
accumulator agent of open-air archaeological sites correspond-
ing to human settlements (see S. 1) which mainly contain rejec-
tion of culinary activities (Grouard 2001; Grouard 2003; de
Waal 2006; Grouard 2007; Bochaton, Bailon, et al. 2016). us,
investigating Pholidoscelis archaeological assemblages allows,
in addition to taxonomic and biogeographic indications, for
a better understanding of the relations between past human
populations (in our case Pre-ColumbianAmerindians) and
these teiid lizards.
e dierence between natural cave deposits and archaeolog-
ical deposits is primarily reected by the dierences in the ana-
tomical parts collected from both of these types of sites (χ2 test,
p<0.05) (Figure 8). In natural deposits, the mean Proportional
Representation of the dierent anatomical parts is low (Mean
PR= 12.7%), but anatomical distribution is relatively homo-
geneous with a better representation of bones from the skull
(PR=22%), sacral vertebrae (PR=29%) and femurs (PR=27%)
(Figure 8(A)). ese well-represented bones correspond to the
most robust, biggest or most easily identiable anatomical parts.
Only the smallest and most breakable remains which are dicult
to collect and to identify (radius, ulna, bula, scapulocoracoid,
ribs, posterior caudal vertebrae, metapodials and phalanx) are
absent or very scarce in the assemblages (Figure 8(A)). is ana-
tomical distribution does not reect a clear selection regarding
the anatomical parts occurring in the deposits despite a probable
collection, preservation and identication bias impacting some
elements. In archaeological deposits, the mean percentage of rep-
resentation is similar (Mean PR=14.3%), but the distribution is
more heterogeneous than in cave deposits (Figure 8(B)). As in
cave deposits, the largest and most robust elements like the femur
(PR=46%), tibia (PR=23%), and pelvic girdle (PR=24%) are
the best represented and the small and delicate elements are
nearly absent, probably reecting a strong collection bias. is
phenomenon could be even more marked than in natural depos-
its, as shown by the representation of skull elements (PR=10%)
which are scarcer in archaeological deposits. e anatomical dis-
tribution of the remains observed in archaeological sites can be
explained in terms of anthropogenic behavior. Indeed, the good
representation of trunk and limb proximal elements could reect
preparation of the carcasses whereby the elements with the least
meat (head, limb extremities and distal part of the tail) were
removed prior to cooking and consumption. However, many
phenomena probably impact the archaeological assemblages and
therefore no clear conclusions can be drawn.
As regards the surface alterations of bone remains, only three
localized charring traces (Stage 1 from Stiner & Kuhn 1995) were
observed on the archaeological remains, respectively on an ante-
rior caudal vertebra from Gare Maritime (US1003) (See S. 1, S. 2),
an ischium from Gare Maritime (US1004) and a coronoid from
Tourlourous (Z4 S3 P7) (Figure 9, S. 1, S. 2). ese marks indicate
that the bones were exposed to temperatures above 300° (Munro
et al. 2008). No cut marks were observed on the material. ese
observations indicate that at least some of our archaeological
specimens may have been cooked using re, as shown by the
small burnt areas indicating that the bones were still covered
by esh when they were exposed to re. No traces associated
with dismantling the carcasses were observed but this is hardly
surprising considering the small size of these lizards that can
probably be taken to pieces without strong eort.
in archaeological sites, as well as the anatomical distribution of
the remains, could also be used as arguments in favor of the
human consumption of Pholidoscelis lizards by pre-Columbian
populations, although the potential collection bias impacting
the archaeological assemblages prevents us from being fully
armative. As stated for archaeological iguanas from the same
contexts (Bochaton, Bailon, et al. 2016), consumption by dogs
may also have played a role in archaeological accumulations but
there is no clear evidence of this from the Pholidoscelis bones.
Finally, the best argument in favor of Pholidoscelis consumption
by archaeological Amerindian populations would be the occur-
rence of many Pholidoscelis remains in most of the investigated
archaeological sites. Yet the consumption of these lizards was
not mentioned by early European chroniclers who described
the daily life of Amerindian populations from the seventeenth
century (Anonyme de Carpentras 1618), and other chroniclers
mention that Pholidoscelis lizards were numerous near villages
searching for food scraps le by inhabitants (Rochefort 1658;
Du Tertre 1667).
Our observation conrms that a considerable diversity of
Pholidoscelis existed in the Guadeloupe Islands. We obtained
fossil evidence of the occurrence of large Pholidoscelis lizards
described as numerous in Guadeloupe by chroniclers from the
seventeenth century and which were only named and described
by naturalists more than 200years later. At that time, only a small
now-extinct population of Pholidoscelis subsisted on a small islet
(the ‘Grand îlet de Petit-Bourg’) (Honegger 1981). is evidence
shows that the numerous Pholidoscelis lizards that occurred in
Guadeloupe were all rapidly decimated within 200years of the
European colonization of the Guadeloupe Islands, between the
seventeenth century and their last observation in the rst half
of the twentieth century (Barbour & Noble 1915). is could be
partly explained by the biological characteristics of this taxon.
Indeed, extinction patterns of terrestrial vertebrates in the Lesser
Antilles have shown that terrestrial large lizards (Kemp & Hadly
2015; Slavenko et al. 2016) are the most likely to go extinct due to
anthropogenic phenomena (Case 1978). ese phenomena could
be explained by the fact that such lizards potentially represent
a signicant amount of meat and are easily catchable by alien
mammalian predators introduced by man (cat, dog and mon-
goose), which are acknowledged to be the predominant factor in
squamate extinction (Case 1978; Henderson 1992). is is prob-
ably especially true for terrestrial diurnal lizards, which are even
more exposed to predators than arboreal or nocturnal lizards.
To conclude, although the important morphological varia-
bility of Pholidoscelis precludes any denitive identication of
Pholidoscelis fossil remains at species level, we have shown that at
least two or three species occurred in Guadeloupe. Two species,
possibly synonymous, were already known through museum
specimens (P. cineracea and P. major), and the third fossil species
occurring on Marie-Galante Island (P. turukaeraensis sp. nov.)
was never described before. We also demonstrated that these
lizards attained a large size in Guadeloupe, especially on Basse-
Terre Island. In addition, we demonstrated a size increase of P.
turukaeraensis aer the Pleistocene period, possibly due to the
extinction of the larger predators (snakes) that occurred there
during this period. Aer the arrival of Amerindian populations
in Guadeloupe, Pholidoscelis probably started to be hunted and
consumed, as shown by archaeological evidence, but the real
museum specimens do not match the descriptions provided by
the rst naturalists who visited the Guadeloupe Islands during
the seventeenth century. Indeed, although size of Guadeloupe
ameivas described by historical sources seems to be the same
as P. major (50cm of total length), the chroniclers described a
lizard with an ashy-gray belly and a blue, green and gray striped
back (Rochefort 1658; Du Tertre 1667). is description does
not match the original descriptions of P. major which refer to
a yellowish belly and an olive back with tawny or yellow stripes
(Duméril & Bibron 1839), nor with P. cineracea which presents a
milky or straw-colored belly and an ashy-gray back striped with a
slightly darker tone of gray (Barbour & Noble 1915). Coloration
cannot be considered as denitive proof, but since the recovery
of new modern specimens is impossible, it is likely that only
paleogenetics can further advance the question of the specic
identity of the Guadeloupe fossil Pholidoscelis and the taxonomic
status of P. cineracea and P. major.
Size data reveal that the size of fossil P. cf. major was not
the same across the islands. Indeed, Pholidoscelis were larger on
Basse-Terre Island (MTMS=242mm SVL) than on Grande-Terre
(MTMS=194mm) and La Désirade (MTMS=210mm). e
large size of the Pholidoscelis from Basse-Terre could be related to
the geographic specicity of this island, which is a volcanic island
with wetter and more diversied habitats (Lasserre 1961; Lescure
1987). is size dierence between the Guadeloupe islands was
also recorded for fossil Iguana (Bochaton, Bailon, et al. 2016).
e only data concerning size evolution through time concerns
P. turukaeraensis from Marie-Galante, the only island on which
both Holocene and Pleistocene fossil records exist. Our results
seem to indicate an increase in the size of this lizard through
time, since the MTMS of Holocene Pholidoscelis (187mm SVL)
was unambiguously higher than the Pleistocene MTMS (142mm
SVL). is Holocene size increase of P. turukaeraensis is dicult
to explain. Several studies pointed out the importance of diet in
the evolution of lizard size and the limited impact of other factors
such as island size (Meiri 2007; Runemark et al. 2015). us, the
availability of prey may have impacted the size of Marie-Galante
fossil Pholidoscelis. Paleontological data show that Marie-Galante
vertebrate terrestrial fauna does not seem to have experienced
many modications during the Pleistocene/Holocene transition
(Bochaton et al. 2015; Bailon et al. 2015) and no data exist con-
cerning invertebrates. However, the extinction of a member of
Boa, which may have been the largest terrestrial predator on
the island at the end of the Pleistocene (Bochaton et al. 2015;
Bailon et al. 2015), could have diminished predation pressure on
Pholidoscelis, which was the largest non-snake terrestrial verte-
brate on the island, and led to an increase in their size, as shown
for other taxa (Herczeg et al. 2009).
About 0.5% of Pholidoscelis remains from archaeological
deposits were found to present re traces. e fact that these
traces were limited to some parts of the bone indicates that they
were not only thrown into the re but that they were probably
intentionally burnt, meaning that at least some specimens were
cooked. Yet, re traces are very rare in the material which pre-
cludes using them as denitive argument in favor of consumption
by Amerindians. A similar lack of re traces was documented on
Iguana remains from the same deposits (see Bochaton, Bailon,
et al. 2016) but other comparison points are still rare (Stoetzel et
al. 2012; Smith et al. 2013). e size of the specimens occurring
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is work clearly shows the importance of Pleistocene and
Holocene fossil evidence for a better understanding of past squa-
mate biodiversity and human impact on snake and lizard com-
munities in an area where such questions mostly rely on analysis
conducted on the few remaining survivors. More work is still
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e authors sincerely thank the Museum of Comparative Zoology (J.
Rosado), the Smithsonian Institution National Museum of Natural
History (K. de Queiroz) and the Muséum national d’Histoire naturelle
(N. Vidal) for the loan of the museum specimens. We also thank M. E.
Kemp for assistance during our stay at the MCZ and all the team of the
MCZ Herpetology collection. We are also grateful to the team of CeMIM
(USM 0504, Department RDDM, MNHN) for access to their three-dimen-
sional workstation. Finally, we thank R. Cornette and the USM 2700 of
the Muséum national d’Histoire naturelle (Paris) for providing us with the
material needed to analyze the scans. X-ray CT at the University of Poitiers
was supported by grants from the Poitou-Charentes Région (16-17-79-86).
We are also grateful to J.-C. Rage and to an anonymous reviewer who pro-
vided useful comments on a previous version of this manuscript.
We also wish to thank all the excavation directors of the studied sites: P.
Bodu, D. Bonnissent, M. Boudadi-Maligne, F. Casagrande, R. Chenorkian,
C. Colas, A. Delpuech, P. Courtaud, C. Etrich, P. Fouéré, J. Gagnepain, E.
Gassies, C. L. Hofman, M. l. P. Hoogland, A. Lenoble, T. Romon, N. Sellier-
Segard, N. Serrand, C. Stouvenot, and M. de Waal.
Disclosure statement
No potential conict of interest was reported by the authors.
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USNM 284475-76); Pholidoscelis chrysolaema (MCZ R-131791-92; USNM
259527-32); Pholidoscelis corax (USNM 236387-93); Pholidoscelis dor-
salis (MCZ R-131793); Pholidoscelis erythrocephala (MCZ R-131924);
Pholidoscelis fuscata (MCZ 28579; MCZ 28580; MCZ 28585); Pholidoscelis
exsul (MCZ R-131794-95; USNM 221746-48); Pholidoscelis griswoldi
(USNM 218347-55; USNM 236517-21); Pholidoscelis lineolata (USNM
259533); Pholidoscelis plei (MNHN-RA 1991.4275; MNHN-UMR 7209-
405; USNM 236375-86; USNM 236394); Pholidoscelis pluvianotata (USNM
Appendix 1
Comparative specimens used from the Museum of Comparative Zoology
(Harvard, USA) (MCZ), Museum national d’Histoire naturelle (Paris,
France) Reptile and Amphibians (MNHN-RA) and UMR 7209 (MNHN-
UMR 7209) collections and, Smithsonian Institution National Museum of
Natural History (Washington D. C., USA) (USNM).
Ameiva ameiva (MCZ R-173127; MCZ R-173128; MCZ R-173129;
MNHN-RA 1991.4034); Ameiva bifrontata (MNHN-UMR 7209-343);
Pholidoscelis auberi (MCZ R-32674; MCZ R-152525; MCZ R-131789;
... formerly Ameiva sp.). These lizards were identified to species in Trinidad and Tobago [52] and Guadeloupe [68]. The remaining lizard taxa include several small species that are more rarely identified in archaeological deposits. ...
... These medium-sized lizards (less than 55 cm in length) occur in 29 of the 85 reviewed sites containing squamate remains. The consumption of these lizards by Indigenous groups was not reported by European chroniclers, although their frequent occurrence in Indigenous faunal assemblages and associated zooarchaeological evidence from Guadeloupe suggest they were [68]. Ameiva lizards are still widespread in the Caribbean islands, where they are represented by numerous endemic species [8]. ...
... Open Sci. 9: 220256 9 species in the Caribbean is very difficult if not impossible [68], a problem reflected in the near absence of this information in currently available zooarchaeological data. ...
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Although the importance of the archaeological record for addressing questions of biodiversity is gaining ground, its relevance for describing past faunal communities is still under-exploited, particularly for the most under-documented areas and species. Among the most poorly documented taxa are reptiles and amphibians, which are rarely studied in detail in the archaeological record, even in tropical areas where most of these species occur today. Here I evaluate the archaeological and paleontological significance of reptiles and amphibians from the Indigenous archaeological record of the insular Caribbean. Quantitative (bone counts) and qualitative (taxonomic identification) analyses allow researchers to discuss the role of herpetofauna in the subsistence strategies of Indigenous populations as well as their interest for assessing past insular biodiversity. This overview sheds light on both the poor representation of herpetofaunal taxa in Caribbean archaeological deposits and trends in the potential exploitation of reptiles and amphibians by Indigenous populations. In terms of paleoecology, the presented results reveal strong regional differences in the quality and density of data as well as the inadequacy of available archaeofaunal data for addressing questions of past biodiversity.
... Extant cnemidophorines are widespread and common components of modern North American ecosystems [3]. Fossil cnemidophorines were recovered in large numbers from Quaternary deposits on mainland North America (north of Mexico) [13][14][15] and the Caribbean [16][17][18][19]. Some fossil cnemidophorines were reported from mainland Neogene localities [20][21][22][23][24][25][26][27][28][29][30] (figure 1). ...
... Open Sci. 7: 200317 the anterolabial surface of the bone was previously documented in Pholidoscelis [17]. I interpret labial sculpturing as an apomorphy of the least inclusive clade containing Aspidoscelis, Holcosus and Pholidoscelis. ...
... Aurivela, Contomastix and Ameivula were also reported to have tricuspid distal teeth [22,60,69], although the one specimen of Aurivela examined here has bicuspid distal teeth. At least one large and presumably adult specimen of Ameiva is known to have bicuspid distal teeth [17]. In Holcosus, Ameiva, Cnemidophorus, Kentropyx and Medopheos, the transition from unicuspid teeth to tricuspid teeth occurs in the anterior portion of the tooth row, often abruptly. ...
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Teiid lizards are well represented in the fossil record and are common components of modern ecosystems in North and South America. Many fossils were referred to the cnemidophorine teiid group (whiptails, racerunners and relatives), particularly from North America. However, systematic interpretations of morphological features in cnemidophorines were hampered by the historically problematic taxonomy of the clade, and the biogeography and chronology of cnemidophorine evolution in North America is poorly understood from the fossil record. Few fossil cnemidophorines were identified with an apomorphy- based diagnosis, and there are almost no fossil cnemidophorines that could be used to anchor node calibrations. Here, I describe a cnemidophorine from the Miocene Ogallala Group of Nebraska and diagnose the fossil using apomorphies. In that process, I clarify the systematic utility of several morphological features of cnemidophorine lizards. I refer the fossil to the least inclusive clade containing Aspidoscelis, Holcosus and Pholidoscelis. The most conservative minimum age of the locality of the fossil is a fission-track date of 6.3 Ma, but mammal biochronology provides a more refined age of 9.4 Ma, which can be used as a minimum age for the crown cnemidophorine clade in divergence time analyses. The fossil indicates that a cnemidophorine lineage that does not live in Nebraska today inhabited the area during the Miocene. I refrain from naming a new taxon pending discovery of additional fossil material of the lineage to which the fossil belonged.
... The genus includes 19 extant and three extinct species (Hedges et al. 2019), although P. alboguttatus and P. desechensis are considered by some authorities to be subspecies of P. exsul (Tucker et al. 2017;Uetz et al. 2020). Two wide-ranging taxa, P. auberi and P. chrysolaemus, likely represent species complexes (Harvey et al. 2012), and two of the extinct species, P. cineraceus and P. major, might be conspecific (Bochaton et al. 2019). ...
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Invasive species are a major threat and primary driver of vertebrate extinctions on islands, including native and endemic lizards such as the West Indian Groundlizard (Genus Pholidoscelis). The genus comprises 19 extant species that range collectively from the Bahamas through the Greater Antilles south to Dominica in the Lesser Antilles. Few studies have addressed aspects of groundlizard population ecology despite invasive mammals being implicated in groundlizard declines and three extinctions. From 2007 to 2011, we conducted a capture-recapture study of Pholidoscelis exsul on Guana Island. Guana Island is privately owned and hosts a small number of feral cats but is mongoose-free and has otherwise experienced only minor human modification since the mid-18th century. Thus, the island provides a unique opportunity to study lizard demography under conditions minimally impacted by human activities. Our objective was to establish benchmark demographic parameter estimates of P. exsul that may be used in future studies to detect ecological changes. We hypothesized that lizards would be encountered more frequently in disturbed habitat than less disturbed habitat because of available anthropogenic food subsidies. We also predicted that large lizards would have higher annual survival rates than small lizards, and lizards with higher indices of body condition would have corresponding higher annual survival rates than lizards of the same size class with lower indices of body condition. Our results showed that male growth rates were comparable to similar studies and individuals reached the presumed size of sexual maturity within one year, with growth rates declining thereafter. Annual survival increased with size class (small = 0.155, 95% CI = 0.069–0.312; large = 0.518, 95% CI = 0.374–0.660), but we found no evidence of a body condition effect on annual survival. The probability of mortality increased to 99.4% by age 6, but evidence of senescence via declining body condition was apparent sooner. Daily capture probabilities were higher in heavily modified habitat (0.081, 95% confidence interval = 0.049–0.132) than in relatively natural habitat (0.025, 95% confidence interval = 0.012–0.054). Mean maximum distance moved between recaptures was 24.4 m (2–350 m). Estimates of annual abundance ranged from 139 to 250 individuals. Density estimates among years based on the upper confidence limit ranged from 41.9 –81.5 individuals/ha within the core sampling area (6.01 ha) and 20.1–39.2 individuals/ha when a 12.5-m buffer strip was incorporated into the effective sampling area.
... The assemblage is dominated by iguana bones, accompanied by a few remains of two species of Alsophis snake, one of which (Alsophis sp. 2) is now extinct in Guadeloupe (Bochaton et al. 2019b). The other rare squamates are two lizard taxa that are also now extinct in Guadeloupe: an anguid lizard of the genus Diploglossus that has also been identified at other sites on the island (Bochaton et al. 2016b), and an unidentified species of ground lizard (Pholidoscelis sp.) that is frequently represented in Guadeloupian Amerindian assemblages (Bochaton et al. 2019a). The majority of the squamate remains represent at least 33 individual iguanas, represented by 444 elements. ...
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Evidence for chronological change in intra-site subsistence strategies is very rare in the Amerindian record of the Lesser Antilles. The study of the vertebrate assemblage from the archeological site of Roseau in the Guadeloupe Islands underlines the complexity and variability of Ceramic Age Amerindian subsistence behavior. This study establishes a more precise chronology of the previously identified strata of the site, and demonstrates that the Contact period was only represented by rare archeological artifacts dispersed in the stratigraphy. The results from this assemblage indicate that the earlier occupations of the site exhibit a more intensive exploitation of aquatic vertebrate resources compared to later occupations, which have a larger focus on terrestrial fauna, especially rodents and iguanas. This unusual pattern highlights how subsistence behaviors in the Lesser Antilles during the Late and Final Ceramic periods were highly variable. This new evidence of strong inter-site variability shows that the behaviors of Amerindians are not only dependent of large-scale environmental conditions but also influenced by more complex socio-cultural and local environmental parameters.
... For instance, some authors have explored the role of the fluvial action on the transportation of reptiles to paleontological deposits such as tanks, sinkholes, caves and fissures (Araújo-Júnior, 2016;Araújo-Júnior et al., 2013;Pacheco et al., 2011). Others have analyzed the preservation status of the fossil remains according with their fossorial conditions (Albino and Kligman, 2007;del Papa et al., 2017;Weissbrod and Zaidner, 2014); the accumulation of fossil remains caused by owl predation (Blain et al., 2010;Lev et al., 2020;Smith et al., 2013); the consumption of snakes, lizards and turtles by ancient humans (Biton et al., 2017;Blasco et al, 2011Blasco et al, , 2016Lev et al., 2020;Nabais and Zilhão, 2019); the use of reptile individuals in ritual activities of indigenous groups (Albino and Kligmann, 2009), the extinction of Quaternary lizards and snakes caused by human consumption and the Spanish colonization (Bochaton et al., 2019a(Bochaton et al., , 2019b; and the existence of predation interactions between crocodiles and turtles (Scheyer et al., 2018). ...
Records of Pleistocene reptiles are relatively well known in different countries. However, reptile fossil records of Pleistocene from Latin America are scarce and limited to presence data associated with records of megafauna remains. Taphonomic studies in Pleistocene reptiles are focused on finding out the origin of the paleontological remains; that is, if the remains found at a particular place correspond to organisms that lived and died there or if they were carried out and deposited there by water currents or predators. Pleistocene reptiles have been used as a proxy for qualitative paleoclimatic reconstructions of localities. In this study, we describe the fossil record of squamate reptiles from the Pleistocene of San José Buenavista (Puebla, Mexico), and the taphonomic relationship between them and the associated megafauna, as well as reconstruct the paleoclimate of the paleontological site in Central Mexico where the remains were found. We present the first Mexican Pleistocene record of Anolis and Storeria, the third fossil record of the alligator lizard Barisia sp. in Mexico, and the first North American Pleistocene record for the rattlesnake Crotalus triseriatus. The rattlesnake (C. triseriatus) was found inside the mandible of the American mastodon (Mammut americanum). This finding suggests that the mastodon remains were used as burrow by the rattlesnake during the Pleistocene. Paleoclimate was reconstructed using the Mutual Ecogeographic Range method, which uses the distribution and climate niche of the nearest extant representatives of the identified fossil taxa. A mean annual temperature of 13.91 ± 1.54 °C and an annual precipitation of 964.04 ± 316.82 mm were inferred for the locality. This suggests that the assemblage of fossil species found inhabited the locality during a glacial period.
... The frontal of adult teiids bears low, irregular and poorly defined dermal rugosities (e.g. Pholidoscelis, Fig. 2 in Bochaton et al. 2017: Ameiva according to the nomenclature followed here) and the subcircular swellings that appear on the frontal from La Bouffie may be interpreted in that sense. ...
Several teiid specimens (frontal, vertebra, maxillae) are described from the late Eocene of Europe (MP17, Phosphorites du Quercy). The results of phylogenetic analyses confirm that these European Eocene fossils belong to teiid lizards and more specifically to the subfamily Tupinambinae. So far, the Paleogene record of teiids is limited to South America and no occurrence of crown teiids is known in Europe. This disjunct distribution of teiids during the Eocene suggests transatlantic dispersal and this possibility is discussed. The presence of teiids in the European fossil record is brief (limited to standard level MP17). The circumstances that prevented the persistence of an invading clade in Europe are examined. Ecological (e.g. biotic interactions) and/or demographic (Allee effect) processes may have been involved.
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La liste taxinomique est actualisée pour les Amphibiens et les « Reptiles » de la Guadeloupe, département et région d' outre-mer dans les Petites Antilles. Elle tient compte des publications les plus récentes. À côté du nom scientifique zoologique international, nous attribuons un nom scientifique français à chaque taxon. Summary: Taxonomic list of the herpetofauna in the Overseas France: V Department of Guadeloupe. The taxonomic checklist is updated for the Amphibians and "Reptiles" of the French department of Guadeloupe, in the Lesser Antilles. It takes into account most recent publications. In addition to international zoological scientific names, we attributed a French scientific name to each taxon.
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Large-scale extinction is one of the defining challenges of our time, as human processes fundamentally and irreversibly reshape global ecosystems. While the extinction of large animals with popular appeal garners widespread public and research interest, the importance of smaller, less “charismatic” species to ecosystem health is increasingly recognized. Benefitting from systematically collected fossil and archaeological archives, we examined snake and lizard extinctions in the Guadeloupe Islands of the Caribbean. Study of 43,000 bone remains across six islands revealed a massive extinction of 50 to 70% of Guadeloupe’s snakes and lizards following European colonization. In contrast, earlier Indigenous populations coexisted with snakes and lizards for thousands of years without affecting their diversity. Study of archaeological remains provides insights into the causes of snake and lizard extinctions and shows that failure to consider fossil-derived data probably contributes to substantial underestimation of human impacts to global biodiversity.
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The past distribution of Boa snakes and their interactions with Pre-Columbian human populations in the Lesser Antilles (Caribbean) remain enigmatic. These snakes currently have a patchy distribution in the islands and are nearly absent from archaeological deposits. This raises questions about whether their absence from Pre-Columbian contexts should be interpreted from a biological or a cultural point of view. In this study, I provide three new references to Boa remains from archaeological and natural deposits on the islands of Guadeloupe and Martinique, most of which were manufactured into beads. These are the first objects in the Lesser Antilles known to be manufactured using snake bones and all appear to be made from Boa, despite a wider diversity of snakes occurring in this region. Using these new observations and combined pieces of evidence from archaeological, historical, and biological data sources, I propose that the extreme scarcity of Boa in zooarchaeological assemblages reflects their prominent status in Pre-Columbian Amerindian communities.
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Guadeloupe and nearby dependencies (la Désirade, Petite Terre, Marie-Galante, les Saintes, Marie-Galante) are situated in the middle of the Lesser Antilles Arc, whereas Saint-Martin and Saint-Barthélemy lie at the Northern end. Together with Anguilla and their satellites, these northern islands belong to the same bank, the Anguilla Bank. Thus they formed a unique piece of land 12000 years ago, before the ice melted. The same situation occurs on the Guadeloupian Bank where Basse-Terre, Grande-Terre, and nearby satellites (Kahouanne, Fajou, Goyaves ... ), la Désirade, and Petite Terre appear to belong to the same terrestrial unit. The different islands forming les Saintes are on another bank, but they form an isolated island before the last ice melting, as did Marie-Galante that stands alone. The herpetofauna of aIl these islands is richer than assumed by Schwartz and Henderson (1991) and following researchers. While prospecting, we found numerous herps yet undiscovered by other naturalists. With historical data drawn from earlier naturalists (L'Herminier, Plée ... ) in conjunction with the descriptions written by historical chroniclers, we propose a quite different vision of the Guadeloupian herpetofauna. Because of their small size and low altitude, and because they lie so far away from South America, Saint-Barthélemy and Saint-Martin are considered to have an impoverished fauna. This is partially true. Saint-Barthélemy has Chelonoidis carbonaria (introduced), Iguana delicatissima, Anolis gingivinus, Hemidactylus mabouia (introduced), Sphaerodactylus sputator, S. parvus, Thecadactylus rapicauda, Ameiva plei plei, Mabuya sloanii, Ramphotyphlops braminus (introduced), Typhlops annae, Alsophis rijgersmaei. Moreover, according to Plée's manusctipt, it is possible to asselt that an Eleutherodactylus species lived in Saint-Barthélemy in 1821. This presumably endemic species disappeared for unknown reasons and, at the beginning of 1980, E. martinicensis arrived on this island. In the Middle of the nineties, we found E. johnstonei in gardens and greenhouses, together with the Cuban Treefrog (Osteopilus septentrionalis), originating from Florida. This last species is invasive and poses a great threat to the whole ecosystem. In addition, Saint-Barthélemy is said to have Mabuya mabouya but it has not been found for decades, and possibly Anolis cf. pogus. In Saint-Martin, Eleutherodactylus johnstonei is everywhere, but it is possible that the same Eleutherodactylus species found in Saint-Barthélemy in the nineteenth century lived there as well. This endemic whistling frog may have been eliminated by the opportunistic E. johnstonei. E. martinicensisis known from collections dating from the eighties, and was later reintroduced with plants from Guadeloupe. Osteopilus septentrionalis is present in great numbers on the French side, and it is a predator of Eleutherodactylus. Chelonodis carbonaria is present on Tintamarre. There is a clear-cut ecological separation between S. sputator, which is xerophilic, and S. parvus that is mesophilic. Hemidactylus mabouia became rupicolous and lives under rocks, as does Anolis gingivinus in grasslands, whereas Thecadactylus rapicauda is truly arboricolous or edificarian. Anolis gingivinus that is found everywhel'e with the exception of grasslands lives in some places alongside with Anolis pogus. This last species was also present on Anguilla but it disappeared from there in the twentieth centul'y. The reason may have been the reduction of land surface with the ice cap melting and the competition with A. gingivinus. Iguana delicatissima is very rare and could be found in the highlands of Saint-Martin. Iguana iguana is also there as an escapee from pet shops. Ameiva plei analifera inhabits lowlands (mainly beach areas) and is very abundant despite the presence of mongooses. Same as on Saint-Barthélemy, it is Ameiva plei plei that is present on Tintamane, with a huge population. Alsophis rijgersmaei is still present on the Dutch side, but it was not found recently on the French side, and the same is true for Mabuya mabouya that seems to no longer exist. This snake inhabits Saint-Barthélemy but it is killed by people and by cars. The herpetofauna of SaintBarthélemy- Saint-Martin shows affinities with Greater Antilles (Sphaerodactylus parvus, Mabuya sloanii, Alsophis rijgersmaei). At the beginning of the volcanic activity in this region, about 6-4 M. years ago, there were only 3 islands: Proto-Guadeloupe (= the Northern part of Basse-Terre), Proto-Désirade and Proto-Saintes. Proto-Montserrat was there as weIl as ProtoDominica and Proto-Martinique. During the following period (between 1,6 and 0,6 M.y.) Tene-de-Bas, Grande-Terre (2 islands) and Marie-Galante emerged. Because, Les Saintes and Marie-Galante belong to "Guadeloupe", naturalists tend to consider them as natural parts of Guadeloupe, irrespective of their situation on two other independent banks. It is traditionaIly thought that these herps had not reached species level, although they would have been isolated for a very long time. Meanwhile, S. fantasticus, A. marmoratus and Alsophis antillensis have differentiated on these independent islands. With the fragmentation of the original island of Les Saintes when the sea level rose, they differentiated again for a second time. With S. fantasticus excepted, it is suggested here that Alsophis and Anolis from Les Saintes are two species (Alsophis sanctonum and Anolis terraealtae) with two subspecies each. Basse-Terre has two endemic Eleutherodactylus (E. barlagnei, E. pinchoni and perhaps two more) and E. martinicensis. E. johnstonei is also there and seems to have arrived recently. This whistling frog is an invasive species of great potential threat. Chelonoidis carbonaria and C. denticulata are present in Guadeloupe as pets, but also as feraI animaIs. Trachemys stejnegeri is abundant on Marie-Galante, but also on Terre-de-Haut and Terre-de-Bas des Saintes, and there are some individuais in ponds in Grande-Terre. Pelusios castaneus is abundant in Grande-Terre but is also found on Basse-Terre and one shell was found· on Grand-Ilet des Saintes. Anolis marmoratus is found as 4 endemic subspecies on Basse-Terre and 3 more on satellites (Kahouanne, Petite Terre, Désirade). The differentiation of the populations to the subspecies or species level on the satellites of the same bank is probably due to recent isolation by water. For example, Ameiva major -which was caught by Félix-Louis L' Herminier in 1825, on the islands of la Petite Terre (a nature reserve since 1998 for its huge Iguana delicatissima population) and therefore not on Martinique, as always stated, under the name "Lézard pilori" which was the French name of Ameiva erythrocephala from Saint-Christophe - is different from A. cineracea whereas they belonged to the same bank. The differentiation of the populations of Anolis marmoratus to subspecies level on Basse-Terre was traditionally correlated with climatic zonation. Here we suggest an allopatric subspeciation model for this species, with geological events as the main constraint. During the Pleistocene, Grande-Terre was in the state of two independent islands (Grands Fonds and plateau du Nord) where subspeciation occurred for Anolis marmoratus and Sphaerodactylus fantasticus. Monts Caraïbes and Morne Liquin (South of BasseTerre) where also two independent islands which began to be connected about 100000 years ago with Proto-Guadeloupe, when the Soufrière volcano began its formation. It is possible to find there the prints of subspeciation or speciation events as in Martinique in the same geological context with Montagne Pelée and Mont Conil. La Désirade seems to have expanded since the rniddle of the Pliocene, and Marie-Galante during the Pleistocene, before Grande-Terre. The uplift of the Guadeloupian bank appears to have been the consequence of the Soufrière's activity, which began about 100000 years ago, and with ice melting at the end of Würm, there was a decrease of lowland surfaces which led to the formation of small islands where isolation occurred for sorne thousands of years, which seems to have been sufficient for sorne species to give rise to at least subspecies (Anolis marmoratus) but not in other (S. macrolepis, S. sputator, A. gingivinus, Ameiva plei, Alsophis rijgersmaei) with the fragmentation of the Anguilla bank. Together with Dominica, Grande-Terre, BasseTerre and Marie-Galante are the only islands where a Liophis (coming from South-America) lives alongside an Alsophis (coming from Greater Antilles), but on the French islands these two snakes are on the verge of extinction and are probably extinct on Marie Galante. With climatic changes and cyclones, colonisation, invasive species and introduced predators, the Guadeloupian Archipelago lost a great part of its original biodiversity. Since the beginning of colonisation, the Guadeloupian bank lost Ameiva major, Ameiva cineracea, Ameiva spp. (la Désirade, Les Saintes), Leiocephalus cf. cuneus, Ceslestus or Diploglossus sp., Mabuya mabouya (7), Clelia sp. and perhaps Boa constrictor. Alsophis antillensis, Liophis juliae and Iguana delicatissima are nearly extinct on Basse-Terre and Grande-Terre. Iguana delicatissima is still abundant in Petite Terre and La Désirade. In Les Saintes, Basse-Terre and GrandeTerre I. delicatissima has been elirninated through competition and hybridization by Iguana iguana which was very rare (except in Les Saintes) in the Seventies-Eighties. This last species is found nearly everywhere in Basse-Terre and Grande-Terre.
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Despite advances within particular groups, systematics of the Teiidae has long been unsatisfactory, because fewmorphological characters have been described for this family. Consequently, most species have been assigned to the large,polyphyletic, and poorly defined genera Ameiva and Cnemidophorus. We describe 137 morphological characters andscore them for most species of Neotropical Teiidae. Important, but previously undescribed, character suites include pupilshape; the frontal ridge; longitudinal division of the interparietal; the rostral groove; patterns of supraciliary fusion; thepreauricular skin fold; the “toothy” first supralabial; modified apical granules; the pectoral sulcus; expansion of scales atthe heel; tibiotarsal shields; scales between the digital lamellae along the postaxial edges of the toes; scale surfacemicrostructure of macrohoneycomb, macroridges, or lamellae; distribution patterns and morphology of lenticular scaleorgans; types of epidermal generation glands; and several hemipenial structures. We propose a new taxonomy of theTeiidae based on recovered evolutionary history and numerous morphological characters surveyed in this study. Werecognize three subfamilies: Callopistinae new subfamily, Teiinae Estes et al., and Tupinambinae Estes et al. To resolvepolyphyly of Ameiva and Cnemidophorus, we erect four new genera for various groups of Neotropical Teiidae: Ameivulanew genus, Aurivela new genus, Contomastix new genus, and Medopheos new genus. We resurrect Holcosus Cope fromthe synonymy of Ameiva and Salvator Duméril and Bibron from the synonymy of Tupinambis. On the basis of sharedderived characters, we propose new species groups of our redefined Ameiva and Cnemidophorus. We incorporate our newcharacters into a key to the genera and species groups of Teiidae. A phylogenetic hypothesis of Teiidae based onmorphological characters differs substantially from hypotheses based on mitochondrial DNA. The phylogeny based onmorphology is consistent with well-established biogeographic patterns of Neotropical vertebrates and explains extreme morphological divergence in such genera as Kentropyx and Aurivela.
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Fossil remains provide useful insights into the long-term impact of anthropogenic phenomena on faunas and are often used to reveal the local (extirpations) or global (extinctions) losses of populations or species. However, other phenomena such as minor morphological changes can remain inconspicuous in the fossil record depending on the methodology used. In this study,we used the anole of Marie-Galante Island (Anolis ferreus) in Guadeloupe (French, West Indies) as a model to demonstrate how the morphological evolution of an insular lizard can be tracked through the Pleistocene/Holocene climatic transition and the recent anthropization of the island.We used a fossil assemblage of nearly 30 000 remains and a combination of anatomical description, traditional morphometry and geometric morphometrics. These fossils are attributed to a single taxon, most likely to be A. ferreus on the basis of morphological and morphometric arguments. Our results showthe disappearance of a distinct (sub)population of large specimens that were about 25% larger than the modern representatives of A. ferreus. We also demonstrate an apparent size stability of the main fossil population of this species since the Late Pleistocene but with the possible occurrence of a reduction in morphological diversity during the Late Holocene. These results highlight the impact of anthropic disturbances on a lizard whose morphology otherwise remained stable since the Late Pleistocene.
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Island ecosystems are, in general, more vulnerable to change than are those on continents, and the West Indies are no exception. In the past 155 years a minimum of 7-12 extinctions and 12-13 extirpations of amphibians and reptiles have occurred in the West Indies. In the Lesser Antilles (and other small islands), most extirpations can be attributed to the introduction of alien predators (primarily the mongoose, but also cats and dogs). Certain species appear more sensitive to predator introductions than others (e.g., teiid lizards of the genus Ameiva and colubrid snakes of the genera Alsophis and Liophis). On mongoose-infested islands, other factors were investigated (human population density, island area, physiographic complexity), but none absolved the mongoose as the primary agent of extirpation. In the Greater Antilles, although introduced predators have had a negative impact, extinctions due to habitat destruction appear more likely due to the potentially stenoecious adaptation of many taxa (e.g., tree crown-, buttress-, and bromeliad-dwelling species).
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Estimating the size of fossil organisms is essential for most paleontological studies, including studies of fossil squamates. Paradoxically, few works about the methods used to achieve this goal have been published. This study focuses on two genera of lizards found in the Lesser Antilles: Pholidoscelis and Anolis. We produced sets of size estimation equations based on skeletal measurements taken on modern specimens. We tested the effect of the anatomical part used and of the species composition of the comparative sample in order to produce reliable and statistically significant equations. We then compared our results with previously published estimates of fossil size for Anolis lizards. We found that past studies were inconsistent with each other in how body size was estimated even if the same value for a measurement was obtained independently. Further, we found that what others have described as Holocene dwarfism for Anolis bimaculatus is likely to reflect a methodological bias, and fossil representatives of this species likely had a similar size to modern individuals. Many conclusions made in previous studies could not be challenged, however, because even though body size estimates are given, the actual measurements made on skeletal elements remain largely unreported in the literature. These results show that more transparency and reproducibility are needed for studies concerning the body size of fossil squamates.
L'étude archéozoologique de près de 95000 vestiges squelettiques de vertébrés et de crustacés de sept sitesPrécolombiens de Guadeloupe révèle l'existence d'une très forte diversité taxinomique. Nous avons mis en placedes critères de détermination ostéologique des poissons et des crustacés, créé des référentiels permettantd'estimer la taille et la masse de trois familles de poissons et proposé une réflexion sur l'échantillormage. Lesrésultats d'analyse archéozoologiques indiquent que si les économies de subsistance des communautésdépendaient en partie de la localisation envirormementale des installations, des choix ont néanmoins été faits àchaque période chronologique. Une dualité des formes d'économies (généralisée et spécialisée) a coexisté entreles communautés d'un même complexe céramique. Cette complémentarité des stratégies de subsistancecorrespondrajt à des activités saisonnières spécifiques adaptées aux différents milieux géographiques. En milieucorallien, le bon équilibre des classes d'animaux au cours de la période Saladoïde (400 av. l-C. à 600 ap. l-C)témoigne d'une exploitation généralisée de tous les écosystèmes attenants au site. Celle-ci devient de plus en plusspécialisée sur les poissons du lagon et de son récif frangeant au cours du Troumassoïde ancien l et II (500/600 à1500 ap. J.-c.), puis se tourne au cours du Troumassoïde récent (800 à 1500 ap. l-C.) vers les zones d'estran,avec les crabes terrestres, même si l'économie de subsistance reste généraliste. Enfin, dès les premièresinstallations Saladoïdes, les communautés de Guadeloupe ont recherché les ressources marines de façon trèsratiormelle, avec une pêche et des techniques de locomotion et de capture sophistiquées, adaptées aux différentsmilieux marins exploités. D'après les analyses des tailles des poissons précolombiens capturés et d'après l'étudedes techniques traditiormelles (chroniqueurs et ethnographes) nous supposons que trois techniques de captureétaient employées.