A New Genus of Microteiid Lizard from the Atlantic Forests of State of Bahia, Brazil, with a New Generic Name for Colobosaura mentalis, and a Discussion of Relationships Among the Heterodactylini (Squamata, Gymnophthalmidae)

2007·1.6 MB·English

by  Miguel Trefaut Rodrigues

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3565, 27 pp., 7 figures, 3 tables May 16, 2007
A New Genus of Microteiid Lizard from the Atlantic
Forests of State of Bahia, Brazil, with a New Generic
Name for Colobosaura mentalis, and a Discussion of
Relationships Among the Heterodactylini
(Squamata, Gymnophthalmidae)
MIGUEL TREFAUT RODRIGUES1,2, KATIA CRISTINA MACHADO
PELLEGRINO3, MARIANNA DIXO4, VANESSA KRUTH VERDADE2, DANTE
PAVAN2, ANTOˆNIO JORGE SUZART ARGOLO5, AND JACK W. SITES, JR.6
ABSTRACT
Anewgenusandspeciesofmicroteiidlizardisdescribedfromaseriesofspecimensobtainedin
the leaf litter at Una (15u109S, 39u039W) in the Atlantic forest of southern Bahia, Brazil. It is
characterizedbythepresenceofprefrontals,frontoparietals,parietals,andinterparietal;parietals
longer than wide; distinct ear openings and eyelids; two pairs of genials, absence of collar and
occipitalscales;dorsalscalesanteriorlysmoothandbecominggraduallylanceolateandmucronate
posteriortotheforelimb;andfourregulartransverseseriesofsmoothventralsthatarelongerthan
wide, identical in size. A phylogenetic analysis based on external morphology, osteology, and
molecular data confirms this new lizard as a member of the Heterodactylini radiation of
Gymnophthalminae.Thetopologyrecoveredbymaximumparsimony(MP)analysesrevealsthat
its closest relatives are the sister taxa Colobosaura modesta and Iphisa elegans (BS 5 , 50%;
1DivisionofVertebrateZoology(Herpetology),AmericanMuseumofNaturalHistory.
2UniversidadedeSa˜oPaulo,InstitutodeBiocieˆncias,DepartamentodeZoologia,CaixaPostal11.461,CEP05422-970,
Sa˜oPaulo,Brazil([email protected];[email protected];[email protected]).
3UniversidadeFederaldeSa˜oPaulo,CampusDiadema,Sa˜oPaulo,Brazil([email protected]).
4UniversidadedeSa˜oPaulo,InstitutodeBiocieˆncias,DepartamentodeEcologia([email protected]).
5UniversidadeEstadualSantaCruz,Ilhe´us,Bahia,Brazil([email protected]).
6DepartmentofIntegrativeBiologyandM.L.BeanLifeScienceMuseum,BrighamYoungUniversity,Provo,Utah,US
84602([email protected]).
CopyrightEAmericanMuseumofNaturalHistory2007 ISSN0003-0082
2 AMERICAN MUSEUMNOVITATES NO. 3565
Bremer value 5 2) and the partitioned Bremer indexes indicated that the largest contribution to
thisrelationshipcomesfrommorphology;Colobosauramentalis,forwhichanewgenericnameis
here proposed, is basal to this radiation. Our analyses confirm a previous hypothesis suggesting
Stenolepis as a member of the Heterodactylini radiation and that the clade composed of
ColobodactylusandHeterodactylusisthesistergroupofthecladeformedbyColobosauramentalis-
Stenolepis (BS 5 100; Bremer value 5 18), Colobosaura modesta-Iphisa (BS 5 , 50%; Bremer
value51),andthenewgenusheredescribed.ThesupportforHeterodactylinimonophyly,onthe
basisofcombinedMPanalysesishigher(BS596,Bremervalue511)thanthatpreviouslyfound
in molecular-based studies only. Partitioned Bayesian methodology combining molecular and
morphologicaldatasetsrecoveredthenewgenusasthesistertaxon(PP50.94)oftheclade(PP5
0.94) formed by I. elegans-C. modesta (PP 5 0.51) and C. mentalis-S. ridleyi (PP 5 1.0). An
alternativetopologydemonstratingaparaphyleticHeterodactyliniisonlyweaklysupported(PP5
0.63). Based on the MP topology we discuss tentative scenarios for the evolution of
Heterodactylini.
INTRODUCTION have been made to infer relationships among
thegenera(MyersandDonnelly,2001;Hoyos,
The Gymnophthalmidae represent a large 1998), and one entirely new clade of micro-
South and Middle American radiation (41 teiids (characterized by scincoid scales) was
genera and about 180 species) of small to recognized (Rodrigues, 1991a, b, c). All of
medium-sized lizards occurring in terrestrial, these initiatives were based on morphology.
arboreal, fossorial, and semiaquatic habitats Only in this century, the first extensive
extending from sea level to the high Andes molecular-based phylogenetic hypothesis was
(Pellegrino et al., 2001; Doan, 2003; Doan proposedforGymnophthalmidae,whichwere
and Castoe, 2005; Rodrigues et al., 2005). recovered as monophyletic (Pellegrino et al.,
Taxonomic study of these so-called ‘‘micro- 2001).Basedonthestudyof26representatives
teiid’’ lizards (Ruibal, 1952) has been compli- of the 36 genera recognized at the time, four
catedbytherarityofspecimensincollections, subfamilies were recognized: Alopoglossinae,
which limits studies of geographical and Rhachisaurinae, Cercosaurinae (with two
individualvariation,andconvergence inchar- tribes, Cercosaurini and Ecpleopini), and
acter complexes (e.g., body elongation, limb Gymnophthalminae (also with two tribes:
reduction, earlessness, lack of eyelids, and the Gymnophthalmini and Heterodactylini).
fusion/fission of some major head scales), Genera not represented in the molecular
which has rendered higher-level taxonomy analysis of Pellegrino et al. (2001) were
of the Gymnophthalmidae problematic. Bou- tentatively allocated to the recognized clades
lenger (1885) first organized their chaotic on the basis of morphology. Pellegrino et al.
taxonomy by recognizing three groups of also demonstrated the paraphyly of several
microteiids and another group (macroteiids) microteiid genera, giving support to previous
in the Teiidae. After this, several attempts hypotheses suggesting extensive character
were made to validate or understand interge- convergence, and making understandable the
neric relationships between and within the unsuccessful attempts of previous classifica-
boulengeriangroups(Ruibal,1952;MacLean, tions.
1974; Presch, 1980;Harris, 1985; Sullivan and Castoe et al. (2004) using only four of the
Estes, 1997; Hoyos, 1998). The relationships five genes explored by Pellegrino et al. (2001)
between gymnophthalmids and teiids (macro- and a slightly improved sampling design
teiids) were generally recognized, with micro- (twelve additional species and one additional
teiids considered either as a distinct family genus), reanalyzed the Pellegrino et al. (2001)
(Presch, 1983, 1988; Estes, 1983; Estes et al., data. Their results, based on partitioned
1988) or as a subfamily (MacLean, 1974; Bayesian analyses, were highly consistent
Presch, 1978, 1980) of Teiidae. In the last two with those obtained by Pellegrino et al.
decades, several new taxa of the Gymno- (2001) with the following higher-level taxo-
phthalmidae have been described (Rodrigues, nomic changes: (1) Ptychoglossus was includ-
1991a, 1991b, 1991c, 1997; Myers and ed in Alopoglossinae; (2) Heterodactylini
Donnelly, 2001; Kok, 2005); some attempts and Gymnophthalmini were combined in
2007 RODRIGUES ETAL.: MICROTEIID LIZARD 3
aGymnophthalminaewithouttribaldivisions; biodiversity, phylogenetic relationships within
(3) Ecpleopini was raised to subfamily status, the family Gymnophthalmidae remain poorly
and (4) Bachia was allocated to the new tribe known. We therefore follow Rodrigues et al.
Bachiini within the Cercosaurinae. (2005) and use the classification proposed
Sincethen,bothschemeshavebeenreferred by Pellegrino et al. (2001) for the Gymno-
in the literature (Doan and Castoe, 2005; phthalminae until extensive character and
Rodrigues et al., 2005). In a paper focused on taxon sampling has been completed.
clarification of the phylogenetic taxonomy of Pellegrino et al. (2001) defined the tribe
the cercosaurines, Doan and Castoe (2005) Heterodactylini, on the basis of molecular
followed the arrangement of Castoe et al. evidence, to include the genera Colobo-
(2004). They described two new genera dactylus, Colobosaura, Heterodactylus, Iphisa,
(Potamites and Petracola) to accommodate and probably the genus Stenolepis (which was
a group of species formerly included in not sampled by Pellegrino et al.; or Castoe et
Neusticurus and Proctoporus, respectively, al. 2004) on the basis of morphology. In this
and resurrected Riama to allocate another paper we include the latter genus and add
group of Proctoporus. Rodrigues et al. (2005) morphological data (completely absent from
fully agreed with the reallocation of the studies of both Pellegrino et al., and
Ptychoglossus, but continued to follow the Castoeetal.)toextendourstudiesofrelation-
proposal by Pellegrino et al. (2001) in other ships within the Heterodactylini. We describe
respects because they considered the other an unknown genus and species within this
changes proposed by Castoe et al. (2004) clade,anddiscussitsrelationshipstotheother
premature; this decision was based on the heterodactyline genera. The first specimen of
evidence provided by morphological charac- the new taxon came to our attention in the
ters included in the study of Rodrigues et al. mid-1990s, when the first author received for
Although the study by Rodrigues etal. (2005) examination a lizard obtained at Ilhe´us, state
wasbasedonasmallerdataset,theyincluded ofBahia,Brazil.ItwassentbyP.E.Vanzolini
representatives of the major clades to test the with a note saying that it was possibly a new
relationships of the new genus described species of Colobosaura. That specimen was
(Dryadosaura) in that paper, and their com- discovered during the process of reorganiza-
bined morphological and molecular data re- tionoftheherpetologicalcollectionofthelate
covered monophyly for the Heterodactylini Werner Bokermann, now at the Museum of
and Gymnophthalmini, as well as a sister- Zoology,UniversityofSa˜oPaulo.Someyears
grouprelationshipbetweenthem,andallwith later, two additional specimens were obtained
strong support (their fig. 5B). They also near Itabuna, state of Bahia by one of us (A.
recovered the ecpleopines as a monophyletic Argolo). It was not until 2000 that a large
groupunrelatedtoCercosaurini,inagreement series of specimens was collected in pitfall
with Castoe et al. (2004). Nevertheless, based traps at Una by M. Dixo during a survey of
on the shape of the postorbital, its contact the herpetofauna of the Atlantic forests of
with the postfrontal, and the shape of the southern Bahia. Additional specimens were
interclavicle all suggesting a close relation- subsequently obtained atSerrado Teimoso in
ship between Ecpleopini and Cercosaurini, the same general region.
Rodrigues et al. (2005) retained the tribal Vanzolini’s tentative generic allocation was
assignment of Pellegrino et al. (2001) while accurate in the context of the taxonomy of
awaiting additional evidence. Additionally, that time, and this was also its provisional
based on the limited taxonomic and character allocationinthemolecularstudyofPellegrino
sampling of cercosaurines, Rodrigues et al. et al. (2001), where it was referred as Colo-
(2005) did not adopt the Bachiini proposal of bosaurasp.andrecovereddeeplynestedwithin
Castoe et al. (2004). the Heterodactylini. Nevertheless, as recom-
Although molecular studies have highlight- mended by that study and demonstrated
edandgreatlyimprovedourunderstandingof hereinafter, we allocate this taxon to its own
microteiid relationships, geographic distribu- genus to render classification compatible with
tions, character evolution, and Neotropical the recovered phylogenetic structure. Our
4 AMERICAN MUSEUMNOVITATES NO. 3565
study also shows that as currently recognized ing intraspecific polymorphism after the ex-
the genus Colobosaura is paraphyletic, and, amination of the MZUSP collections. Hemi-
for this reason, a new generic name is penes were examined when partly or totally
proposed to allocate the former Colobosaura everted in preserved specimens in order to
mentalis. improve descriptions and comparisons, but
The discovery of this new taxon in Atlantic due to their limited availability among the
forests of southern Bahia, one of the most generaincludedhere,theywerenotusedinthe
threatened habitats in the world (Myers et al., subsequent analyses.
2000; Rodrigues et al., 2002a, b), reveals The majority of DNA sequences used here
further undisclosed biological diversity and were those from Pellegrino et al. (2001), but
refines our understanding of heterodactyline new sequences for the same mitochondrial
relationships. This combined estimate of the (12S, 16S and ND4) and nuclear (c-mos and
evidence of phylogenetic relationships of the 18S) gene regions were included for the
Heterodactilinialsoraisesadditionalquestions previously unsampled ingroup taxon Ste-
about historical biogeography and speciation nolepis ridleyi (GenBank accession numbers
mechanismsofthisinterestinggroupoflizards. EF 405618-405622). The present molecular
partition is composed of 2,333 bp of aligned
sequences; some adjustments on the original
MATERIAL AND METHODS
Pellegrino et al. (2001) alignments for the
Snout-vent length was measured to the ribosomal 12S, 16S, and 18S regions were
nearest mm with a rule; scale counts and performed manually on the reduced matrix
osteological data on cleared specimens (ap- used in this study. This was necessary to
pendix 1) were taken with the aid of a stereo- accommodate the sequences of Stenolepis and
microscope. Scale counts and scale nomencla- exclude unnecessary gaps. Regions of ambig-
ture are according to Rodrigues and Borges uous alignment for the 12S and 16S were
(1997), and osteological nomenclature follows excluded from the final analyses on the
Presch (1980) and Estes et al. (1988). Sex was reduced matrix.
determinedbythepresence/absenceoffemoral Phylogenetic inference was first conducted
pores and confirmed by dissection in pre- on the separate morphological partition fol-
viouslyopenedspecimens.Alldataweretaken lowed by a combined analyses with the
from preserved specimens housed at MZUSP molecular partitions under equally weighted
(Museu de Zoologia, Universidade de Sa˜o parsimony (MP) in PAUP* v4.0b10
Paulo), MNRJ (Museu Nacional, Rio de (Swofford,2002).Foramatrixof42morpho-
Janeiro), MZUESC (Museu de Zoologia, logical characters (appendix 2), with all char-
Universidade Estadual de Santa Cruz), and acter states coded as unordered and all
RNHM (Nationaal Natuurhistorisch Mu- transformations uniformly weighted, a search
seum, Leiden). To compare abundance of with the branch-and-bound algorithm was
males and females and total abundance performed.Theanalysisofthecombineddata
among habitats of the new taxon sampled we setincludedanMPheuristicsearchwith1,000
used one-way analysis of variance (ANOVA; replicates of random stepwise addition and
Zar, 1996). When significant differences were TBR branch-swapping.
obtained,differencesbetweenpairsofhabitats Nodal support was assessed by bootstrap
were tested with a Tukey’s test (Zar, 1996). analysis (BS; Felsenstein, 1985) with a 1,000
Association among habitat categories and random stepwise additions per bootstrap
species or sex was detected by chi-square test. pseudoreplicate, and TBR branch-swapping,
All analyses were done using Statistica in both branch-and-bound (morphology) and
(StatSoft, 1998). Appendix 2 presents the 19 heuristic (combined data) searches; bootstrap
species, assigned to 17 genera of Gymno- valuesgreaterthan70%(HillisandBull,1993)
phthalmidae (including the new one herein wereinterpretedasstrong supportforanode.
described),usedinthepresentstudy.External Total and partitioned Bremer support (PBS)
morphological characters included in appen- values (Baker and DeSalle, 1997), the latter
dix 2 were selected among those not present- representingthecontributionofeachspecified
2007 RODRIGUES ETAL.: MICROTEIID LIZARD 5
data partition to each node, were calculated three supraoculars. Dorsal scales anteriorly
for all nodes of the combined data-partition smooth, isodiametric or subrectangular in
topology using the program TreeRot v. 2.0 occipital region, becoming progressively nar-
(Sorenson, 1999). rower, more elongate and rounded toward
Modeltest v.3.06 (Posada and Crandall, armlevelandthenlanceolate,stronglykeeled,
1998)wasusedtoselecttheappropriatemodel with lateral sides almost juxtaposed.
of evolution for each molecular data parti- Occipitals absent. Ventrals longer than wide,
tion: 12S (TrN+C), 16S (TrN+I+C), ND4 smooth, in four regular transverse series
(TVM+I+C), c-mos (K80+C) and 18S (JC). identical in size. Males with a continuous
Using the combined data set (molecular and series of very conspicuous pores with no gap
morphology) a partitioned Bayesian analysis between preanal and femoral; pores absent in
was implemented in Mr.Bayes 3.0 (Hue- females. Hemipenis without evident biloba-
lsenbeck and Ronquist, 2001), under the best tion, sulcus spermaticus single, marginated by
fit models of substitution for individual gene a naked area which is also present on the
regions and morphological data set as ‘‘stan- opposite side of sulcus. Between naked areas
dard’’, with 3,000,000 generations, four two symmetrical series of continuous trans-
chains, and trees sampled at intervals of verse rows of comb-like flounces with minute
1,000generations.Twoindependentrunswere spines or irregular tooth-like structures. A
conducted. Trees prior to stationarity were series of enlarged spines near the base of
discarded as ‘‘burn-in’’, and a 50% majority- hemipenis.
rule consensus tree was obtained from 2,900 CONTENT: Alexandresaurus camacan new
data points. Nodes on consensus trees from species; monotypic.
both runs with posterior probability (PP ETYMOLOGY: A homage to the Brazilian
values) $ 0.95 were considered as evidence naturalistAlexandreRodriguesFerreira,born
of significant support for a given clade in Salvador, state of Bahia, as a recognition
(Huelsenbeck and Ronquist, 2001). for his effort to document the natural history
Two species of Alopoglossus, members of of Brazil at the end of the 18th century. After
the subfamily Alopoglossinae formally recog- pursuing natural history studies under
nized by Pellegrino etal. (2001) and Castoeet Vandelli at the University of Coimbra,
al. (2004) and strongly supported as basal to Alexandre headed the famous viagem filoso´fi-
allothercladesofGymnophthalmidaeinboth ca (‘‘philosophical voyage’’) to Brazil at the
studies, were constrained to monophyly and end of 18th century. He remained in Brazil
used as the outgroup. Selected taxa from until 1789 where, among other places, he
Gymnophthalmini, Cercosaurini and Ecpl- traveled extensively along the Rio Negro,
eopini were also included to give Stenolepis Madeira and Guapore´ in the Brazilian
a chance to fail to be recovered within the Amazon. The travels of Alexandre were part
Heterodactylini (see appendix 2). of a larger project to publish what was meant
to be a monumental natural history of the
RESULTS Portuguese colonies (Farias, 2001). The plan
was abandoned by the end of 1807 when the
TAXON DESCRIPTION
royal family escaped to Brazil against the
Alexandresaurus, new genus backdrop of Lisbon’s imminent conquest by
Napoleonic forces. Most of the collections
DEFINITION: A large and slightly elongate assembledbyAlexandreduringthemorethan
gymnophthalmid (maximum SVL 70 mm) 35,000 km traveled in Brazil were published
with distinctive ear opening and eyelid, large by other naturalists following the request of
tail (1.63%–2.36% SVL), and slender penta- partofthematerialbyE´tienneGeoffroySaint
dactyl limbs; first toe lacking a claw. Hilaire,who was present inLisbonduring the
Frontonasal single; prefrontals, frontoparie- subsequent conquest.
tals, parietals, and interparietals present. COMPARISONS: Considering the now recog-
Parietals and interparietal longer than wide. nized multiple origins of character complexes
Collar fold absent. Two pairs of genials and associated with fossoriality in gymnophthal-
6 AMERICAN MUSEUMNOVITATES NO. 3565
mid lizards, and the chaotic state of their slightly wider than long in Colobosaura
taxonomy based on external attributes (until mentalis, whereas the reverse occurs in
recently), we compare Alexandresaurus with Colobosaura modesta. The latter species also
all other gymnophthalmids whileemphasizing differs from Alexandresaurus by having only
the external morphological differences with three pairs of genials. As in Alexandresaurus,
the genera Iphisa, Colobosaura, Coloboda- thehemipenisofColobosauramentalisalsohas
ctylus, Heterodactylus, and Stenolepis. Except a naked area on the side of organ opposite to
for the latter, this group of genera was thesulcus,as well as flouncesof right and left
recovered by Pellegrino et al. (2001) as a well- lobes medially interrupted by a naked area,
supported clade that included the closest a character absent in the former species. The
relatives of Alexandresaurus. Stenolepis, al- onlyotherspeciesreferredtoColobosaurawas
though not included in that study, was Perodactylus kraepelini. It is a young female
tentatively allocated to the same radiation. (SVL 5 40 mm) described by Werner in 1910
Most character states for the taxa com- havingPuertoMax:Paraguayastypelocality.
pared here are summarized in appendix 1. We have not examined the holotype, which is
Alexandresaurus differs from Iphisa (data for lost from the Hamburg Zoological Museum
the latter in parenthesis) by having: two pairs accordingtoDr.J.Hallermann(inlitt.);based
ofgenials(one);severalrowsoflanceolateand on the original description, this species is
keeled dorsal scales (smooth, in two longitu- identical to Colobosaura modesta, an opinion
dinalrows);ventralscalesregularlytransverse advanced long ago by Amaral (1933). The
in four longitudinal rows (not in transverse distributional data agree with this interpreta-
rows and in two longitudinal rows); and tion because the type locality is in the area of
regularly transverse rows of gulars (not occurrence of C. modesta. The only character
regular). Alexandresaurus also differs from that until now has precluded synonymy
Iphisa in having a naked area opposite to the between C. modesta and C. kraepelini was
sulcusspermaticusandlargespinesatthebase Werner’s reference to the tongue of his
of the hemipenis (naked area absent and only specimen as having ‘‘oblique plicae like
small spines at base of the organ). From Alopoglossus’’. Based on the agreement of all
Colobosaura Alexandresaurus can be distin- other characters mentioned in the description
guished from Colobosaura by its smooth neck we think that Werner was in error in de-
scales and small, smooth, and posteriorly scribing the tongue, evidently a strange char-
rounded scales in the first 7–8 dorsal rows, acter and restricted to Alopoglossinae.
rarely two enlarged nuchals are present at the Alexandresaurus differs from Stenolepis,
central part of the second dorsal row; occipi- Colobodactylus, and Heterodactylus in having
tals are never present. In Colobosaura neck apairofprefrontals(absent).Itfurtherdiffers
scales have keels; there are several enlarged from Stenolepis by having small dorsal neck
scales,muchwiderthanlong,inthefirsteight scales (large, in 3–4 longitudinal rows), larger
dorsal rows, the first two characteristically scalesonthesidesofneck(small,somealmost
smooth and enlarged, forming an occipital granular), ventral and gulars in regularly
pair separated or not by a smaller scale; the transverse rows (not so), and smooth scales
posterior 5–6 dorsal rows are striate or multi- near venter (keeled). In Colobodactylus and
carinate.VentralscalesinAlexandresaurusare Heterodactylus the parietals are wider than
longer than wide, laterally juxtaposed, round- long in contrast to the longer than wide
ed posteriorly, identical in size and shape, parietal scale observed in Alexandresaurus
regularly arranged transversally and in four and other Heterodactylini. This difference is
longitudinal series. In Colobosaura there are probably due to a possible scale rearrange-
also four longitudinal rows of quadrangular ment in Heterodactylus and Colobodactylus in
ventral scales, but they are not regularly which parietals (and interparietal in
transversalandareirregularinsizeandshape, Colobodactylus) were divided and their poste-
laterally imbricate, and in the first ventral rior regions incorporated into the highly
rows midline scales always distinctively wider conspicuous occipitals that characterize both
than long; the external ventral rows are genera. This hypothesis, further discussed
2007 RODRIGUES ETAL.: MICROTEIID LIZARD 7
under a phylogenetic framework below, re- Alexandresaurus camacan, new species
veals that even head scales traditionally used figures 1–4
in taxonomic descriptions may not be strictly
homologous. This is another example of the HOLOTYPE: MZUSP 93201, an adult male
taxonomic confusion that may result from from Una (15u109S, 39u039W): state of Bahia:
Brazil, collected by Marianna Dixo on 01
uncritical use of morphological characters in
November 1999, field number md-1106.
microteiid systematics. Alexandresaurus also
differs from Colobodactylus and Heterod- PARATYPES: All from the state of Bahia,
Brazil. MZUSP 93179–93200, Una, collected
actylus in having five toes and smooth flank
by M. Dixo between 24 January 1999 and 27
scales(onlyfourtoesandflankscaleskeeledin
February 2000; MZUSP 93331, Ilhe´us, col-
the latter genera). Heterodactylus further
lected by W. Bokerman on December 1971;
differs from Alexandresaurus by the absence
MZUSP 93462–93464 Ilhe´us (CEPLAC, sede
ofexternalear,absenceoffrontoparietals,and
regional),A.Argololeg.,collectedrespectively
absence or extreme reduction of interparietal.
onJuly1990,24April1991,and12May1999;
Finally, the hemipenis of Heterodactylus is
MZUSP 93227–93228, Jussari (Serra do
very different from that of Alexandresaurus.
Teimoso) collected by M. T. Rodrigues, D.
The organ is distinctively bilobate with a cen-
Pavan,M.Dixo,andV.K.VerdadeonMarch
tripetal sulcus spermaticus surrounded by
2001; MZUESC 2431, Ilhe´us (Campus
a naked area. The asulcate side of the organ
UESC), collected by C. N. Souza on 8
has a series of continuous and unornamented
March 2002; RMNH 29741, from Una,
naked flounces containing only one spine at
Ilhe´us.
each edge, very different from the interrupted
ETYMOLOGY: Thespecificepithetisahom-
flounces containing spines in a comb-like
age to the extinct Camacan Indians, a Boto-
arrangement observed in Alexandresaurus.
cudo group who lived in the exuberant forests
Alexandresaurus differs from members of
ofBahiafromnorthofRioPardotoIlhe´us.
Alopoglossinae by having imbricate, scalelike
DIAGNOSIS: A gymnophthalmid with ear
papillae covering the tongue (oblique plicae),
openings and eyelids, and slender pentadactyl
and from Rhachisauridae by having a distinc-
limbs lacking the claw on first toe.
tive ear opening and eyelid (absent), five toes
Frontonasal single; prefrontals, frontoparie-
(four), a normal body (clearly elongate),
tals, parietals, and interparietals present.
presence of frontoparietals (absent), and
Parietalslongerthanwide.Collarfoldabsent.
a nostril at the nasal border (in the center of
Two pairs of genials; three supraoculars.
the nasal scale). From all other Gymno-
Dorsal scales in 30–38 rows, anteriorly
phthalminae Alexandresaurus differs by hav-
smooth, isodiametric or subrectangular in
ing five toes (fifth toe reduced or absent),
occipital region, becoming progressively nar-
a distinctive eyelid (absent in all Gymno- rower,moreelongateandroundedtowardthe
phthalminiexceptforTretioscincus),nostrilat arm, and posteriortothe arm becoming more
the nasal border (in the center of the nasal lanceolate, strongly keeled, with lateral sides
scale), a wide and flattened clavicle enclosing almostjuxtaposed.Occipitalsabsent.Ventrals
a fenestra (simple, boomerang shaped and longer than wide, smooth, in four regular
without fenestrae), nasal bones in contact at longitudinal and 15–19 transverse rows, iden-
midline but broadly separated anteriorly tical in size. Scales around body 29–35; 11–15
(totally separated by contact between frontal and 16–20 infradigital lamellae under finger
and premaxilary), and, among other charac- IVandtoeIVrespectively.Maleswithaseries
ters,byhavingstraightlateralprocessesofthe of 20–25 very conspicuous pores without gap
interclavicle (posteriorly oriented). Alexan- between preanal and femoral; pores absent in
dresaurus also differs from all Cercosaurinae females.
by having a distinctive sternal fontanelle DESCRIPTION OF HOLOTYPE (fig. 2A–C):
process (absent), a glossohyal separated from Rostral broad, wider than high, contacting
basiyal (fused), nasal bones broadly separated first supralabial, nasal, and frontonasal.
anteriorly (in broad contact), and postorbital Frontonasal pentagonal, twice as wide as
covering postfrontal (covered by postfrontal). long, contacting rostral, nasal, loreal, and
8 AMERICAN MUSEUMNOVITATES NO. 3565
Fig.1. Alexandresauruscamacan(MZUSP93228),anadultmale(57mm)fromSerradoTeimoso,state
of Bahia,Brazil.
prefrontals. Prefrontals slightly wider than wider anteriorly and indenting suture between
long, in broad contact at midline. Frontal preocular and frenocular. Seven supralabials,
hexagonal, with almost parallel lateral mar- third and fourth under the eye, fifth the
gins, slightly longer than wide, anteriorly highest and contacting two smaller scales
indenting the prefrontal and posteriorly the following posteriorly the subocular, the smal-
frontoparietalsutures.Frontoparietalspentag- lest one clearly a postocular; seventh suprala-
onal, larger than prefrontals, in slight contact, bial the smallest, in contact with the granules
strongly indented by the interparietal. surrounding anterior margin of ear. Three
Interparietal longer than wide, longer and superciliaries, first largest, wider anteriorly,
narrower than frontal, as long as and longerthan first supraocular, contactingpreo-
narrower than parietals. Parietals heptagonal, cular, loreal, first and second supraoculars,
marginated laterally by two temporal scales second superciliary and upper eyelid. Central
with approximately the same size, anteriorly part of eyelid with a semitransparent un-
by the third supraocular and frontoparietal, divided disc surrounded by small and slightly
medially by the interparietal, and posteriorly pigmented granular smooth scales. Lower
by the first dorsals. Three supraoculars, first eyelid with eight strongly pigmented palpeb-
the smallest, second and third about the same rals. Temporal region with few smooth and
size, second slightly wider, its longest suture juxtaposed scales, irregular in size and shape,
with frontal, third slightly longer, in broad between parietals and supralabials; the larger
contact with frontoparietal. Nasal above first as large as the sixth supralabial. Ear opening
supralabial, large, longer than high, with the surrounded by a series of very small and
nostril in the center and lower part of scale, juxtaposed rounded granules; external audito-
indenting suture with labial. Loreal posterior rymeatuslarge,tympanumdistinct,subovoid,
to nasal, narrower and diagonally oriented; recessed. Lateral surface of neckwith smooth,
contacting posteriorly first superciliary, preo- imbricate and mostly longer than wide scales,
cular, and frenocular. Frenocular below preo- larger dorsally. All head scales smooth and
cular,followedposteriorlybyalongsubocular juxtaposed with scattered sensorial organs.
2007 RODRIGUES ETAL.: MICROTEIID LIZARD 9
Fig. 2. Ventral (A), lateral (B), and dorsal (C) views of the head of the holotype of Alexandresaurus
camacan(MZUSP93201). Scale51 mm.
Mental broad, wider than high. Postmental the level oflabial comissure. An enlarged pair
heptagonal, wider than long. Two pairs of of symmetric flat and chevron-like scutes
genials, both contacting infralabials; the first follows second pair of genials, preventing
smaller, in broad contact at midline; second their contact posteriorly. Six infralabials,
pair contacting only anteriorly and reaching second the largest. Gulars smooth, imbricate,
10 AMERICAN MUSEUMNOVITATES NO. 3565
Fig. 3. Dorsal (A) and ventral (B) views of the skull of Alexandresaurus camacan (MZUSP 94252-53).
Scale5 1mm.