A New Genus of Microteiid Lizard from the Atlantic Forests of State of Bahia, Brazil, with a New...

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Number 3565, 27 pp., 7 figures, 3 tables May 16, 2007

A New Genus of Microteiid Lizard from the AtlanticForests of State of Bahia, Brazil, with a New GenericName 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, ANTONIO JORGE SUZART ARGOLO5, AND JACK W. SITES, JR.6

ABSTRACT

A new genus and species of microteiid lizard is described from a series of specimens obtained inthe leaf litter at Una (15u109S, 39u039W) in the Atlantic forest of southern Bahia, Brazil. It ischaracterized by the presence of prefrontals, frontoparietals, parietals, and interparietal; parietalslonger than wide; distinct ear openings and eyelids; two pairs of genials, absence of collar andoccipital scales; dorsal scales anteriorly smooth and becoming gradually lanceolate and mucronateposterior to the forelimb; and four regular transverse series of smooth ventrals that are longer thanwide, identical in size. A phylogenetic analysis based on external morphology, osteology, andmolecular data confirms this new lizard as a member of the Heterodactylini radiation ofGymnophthalminae. The topology recovered by maximum parsimony (MP) analyses reveals thatits closest relatives are the sister taxa Colobosaura modesta and Iphisa elegans (BS 5 , 50%;

Copyright E American Museum of Natural History 2007 ISSN 0003-0082

1Division of Vertebrate Zoology (Herpetology), American Museum of Natural History.2Universidade de Sao Paulo, Instituto de Biociencias, Departamento de Zoologia, Caixa Postal 11.461, CEP 05422-970,

Sao Paulo, Brazil ([email protected]; [email protected]; [email protected]).3Universidade Federal de Sao Paulo, Campus Diadema, Sao Paulo, Brazil ([email protected]).4Universidade de Sao Paulo, Instituto de Biociencias, Departamento de Ecologia ([email protected]).5Universidade Estadual Santa Cruz, Ilheus, Bahia, Brazil ([email protected]).6Department of Integrative Biology and M. L. Bean Life Science Museum, Brigham Young University, Provo, Utah, US

84602 ([email protected]).

Bremer value 5 2) and the partitioned Bremer indexes indicated that the largest contribution tothis relationship comes from morphology; Colobosaura mentalis, for which a new generic name ishere proposed, is basal to this radiation. Our analyses confirm a previous hypothesis suggestingStenolepis as a member of the Heterodactylini radiation and that the clade composed ofColobodactylus and Heterodactylus is the sister group of the clade formed by Colobosaura mentalis-Stenolepis (BS 5 100; Bremer value 5 18), Colobosaura modesta-Iphisa (BS 5 , 50%; Bremervalue 5 1), and the new genus here described. The support for Heterodactylini monophyly, on thebasis of combined MP analyses is higher (BS 5 96, Bremer value 5 11) than that previously foundin molecular-based studies only. Partitioned Bayesian methodology combining molecular andmorphological data sets recovered the new genus as the sister taxon (PP 5 0.94) of the clade (PP 50.94) formed by I. elegans-C. modesta (PP 5 0.51) and C. mentalis-S. ridleyi (PP 5 1.0). Analternative topology demonstrating a paraphyletic Heterodactylini is only weakly supported (PP 50.63). Based on the MP topology we discuss tentative scenarios for the evolution ofHeterodactylini.

INTRODUCTION

The Gymnophthalmidae represent a largeSouth and Middle American radiation (41genera and about 180 species) of small tomedium-sized lizards occurring in terrestrial,arboreal, fossorial, and semiaquatic habitatsextending from sea level to the high Andes(Pellegrino et al., 2001; Doan, 2003; Doanand Castoe, 2005; Rodrigues et al., 2005).Taxonomic study of these so-called ‘‘micro-teiid’’ lizards (Ruibal, 1952) has been compli-cated by the rarity of specimens in collections,which limits studies of geographical andindividual variation, and convergence in char-acter complexes (e.g., body elongation, limbreduction, earlessness, lack of eyelids, and thefusion/fission of some major head scales),which has rendered higher-level taxonomyof the Gymnophthalmidae problematic. Bou-lenger (1885) first organized their chaotictaxonomy by recognizing three groups ofmicroteiids and another group (macroteiids)in the Teiidae. After this, several attemptswere made to validate or understand interge-neric relationships between and within theboulengerian groups (Ruibal, 1952; MacLean,1974; Presch, 1980; Harris, 1985; Sullivan andEstes, 1997; Hoyos, 1998). The relationshipsbetween gymnophthalmids and teiids (macro-teiids) were generally recognized, with micro-teiids considered either as a distinct family(Presch, 1983, 1988; Estes, 1983; Estes et al.,1988) or as a subfamily (MacLean, 1974;Presch, 1978, 1980) of Teiidae. In the last twodecades, several new taxa of the Gymno-phthalmidae have been described (Rodrigues,1991a, 1991b, 1991c, 1997; Myers andDonnelly, 2001; Kok, 2005); some attempts

have been made to infer relationships amongthe genera (Myers and Donnelly, 2001; Hoyos,1998), and one entirely new clade of micro-teiids (characterized by scincoid scales) wasrecognized (Rodrigues, 1991a, b, c). All ofthese initiatives were based on morphology.

Only in this century, the first extensivemolecular-based phylogenetic hypothesis wasproposed for Gymnophthalmidae, which wererecovered as monophyletic (Pellegrino et al.,2001). Based on the study of 26 representativesof the 36 genera recognized at the time, foursubfamilies were recognized: Alopoglossinae,Rhachisaurinae, Cercosaurinae (with twotribes, Cercosaurini and Ecpleopini), andGymnophthalminae (also with two tribes:Gymnophthalmini and Heterodactylini).Genera not represented in the molecularanalysis of Pellegrino et al. (2001) weretentatively allocated to the recognized cladeson the basis of morphology. Pellegrino et al.also demonstrated the paraphyly of severalmicroteiid genera, giving support to previoushypotheses suggesting extensive characterconvergence, and making understandable theunsuccessful attempts of previous classifica-tions.

Castoe et al. (2004) using only four of thefive genes explored by Pellegrino et al. (2001)and a slightly improved sampling design(twelve additional species and one additionalgenus), reanalyzed the Pellegrino et al. (2001)data. Their results, based on partitionedBayesian analyses, were highly consistentwith those obtained by Pellegrino et al.(2001) with the following higher-level taxo-nomic changes: (1) Ptychoglossus was includ-ed in Alopoglossinae; (2) Heterodactyliniand Gymnophthalmini were combined in

2 AMERICAN MUSEUM NOVITATES NO. 3565

a Gymnophthalminae without tribal divisions;(3) Ecpleopini was raised to subfamily status,and (4) Bachia was allocated to the new tribeBachiini within the Cercosaurinae.

Since then, both schemes have been referredin the literature (Doan and Castoe, 2005;Rodrigues et al., 2005). In a paper focused onclarification of the phylogenetic taxonomy ofthe cercosaurines, Doan and Castoe (2005)followed the arrangement of Castoe et al.(2004). They described two new genera(Potamites and Petracola) to accommodatea group of species formerly included inNeusticurus and Proctoporus, respectively,and resurrected Riama to allocate anothergroup of Proctoporus. Rodrigues et al. (2005)fully agreed with the reallocation ofPtychoglossus, but continued to follow theproposal by Pellegrino et al. (2001) in otherrespects because they considered the otherchanges proposed by Castoe et al. (2004)premature; this decision was based on theevidence provided by morphological charac-ters included in the study of Rodrigues et al.Although the study by Rodrigues et al. (2005)was based on a smaller data set, they includedrepresentatives of the major clades to test therelationships of the new genus described(Dryadosaura) in that paper, and their com-bined morphological and molecular data re-covered monophyly for the Heterodactyliniand Gymnophthalmini, as well as a sister-group relationship between them, and all withstrong support (their fig. 5B). They alsorecovered the ecpleopines as a monophyleticgroup unrelated to Cercosaurini, in agreementwith Castoe et al. (2004). Nevertheless, basedon the shape of the postorbital, its contactwith the postfrontal, and the shape of theinterclavicle all suggesting a close relation-ship between Ecpleopini and Cercosaurini,Rodrigues et al. (2005) retained the tribalassignment of Pellegrino et al. (2001) whileawaiting additional evidence. Additionally,based on the limited taxonomic and charactersampling of cercosaurines, Rodrigues et al.(2005) did not adopt the Bachiini proposal ofCastoe et al. (2004).

Although molecular studies have highlight-ed and greatly improved our understanding ofmicroteiid relationships, geographic distribu-tions, character evolution, and Neotropical

biodiversity, phylogenetic relationships withinthe family Gymnophthalmidae remain poorlyknown. We therefore follow Rodrigues et al.(2005) and use the classification proposedby Pellegrino et al. (2001) for the Gymno-phthalminae until extensive character andtaxon sampling has been completed.

Pellegrino et al. (2001) defined the tribeHeterodactylini, on the basis of molecularevidence, to include the genera Colobo-dactylus, Colobosaura, Heterodactylus, Iphisa,and probably the genus Stenolepis (which wasnot sampled by Pellegrino et al.; or Castoe etal. 2004) on the basis of morphology. In thispaper we include the latter genus and addmorphological data (completely absent fromthe studies of both Pellegrino et al., andCastoe et al.) to extend our studies of relation-ships within the Heterodactylini. We describean unknown genus and species within thisclade, and discuss its relationships to the otherheterodactyline genera. The first specimen ofthe new taxon came to our attention in themid-1990s, when the first author received forexamination a lizard obtained at Ilheus, stateof Bahia, Brazil. It was sent by P.E. Vanzoliniwith a note saying that it was possibly a newspecies of Colobosaura. That specimen wasdiscovered during the process of reorganiza-tion of the herpetological collection of the lateWerner Bokermann, now at the Museum ofZoology, University of Sao Paulo. Some yearslater, two additional specimens were obtainednear Itabuna, state of Bahia by one of us (A.Argolo). It was not until 2000 that a largeseries of specimens was collected in pitfalltraps at Una by M. Dixo during a survey ofthe herpetofauna of the Atlantic forests ofsouthern Bahia. Additional specimens weresubsequently obtained at Serra do Teimoso inthe same general region.

Vanzolini’s tentative generic allocation wasaccurate in the context of the taxonomy ofthat time, and this was also its provisionalallocation in the molecular study of Pellegrinoet al. (2001), where it was referred as Colo-bosaura sp. and recovered deeply nested withinthe Heterodactylini. Nevertheless, as recom-mended by that study and demonstratedhereinafter, we allocate this taxon to its owngenus to render classification compatible withthe recovered phylogenetic structure. Our

2007 RODRIGUES ET AL.: MICROTEIID LIZARD 3

study also shows that as currently recognizedthe genus Colobosaura is paraphyletic, and,for this reason, a new generic name isproposed to allocate the former Colobosauramentalis.

The discovery of this new taxon in Atlanticforests of southern Bahia, one of the mostthreatened habitats in the world (Myers et al.,2000; Rodrigues et al., 2002a, b), revealsfurther undisclosed biological diversity andrefines our understanding of heterodactylinerelationships. This combined estimate of theevidence of phylogenetic relationships of theHeterodactilini also raises additional questionsabout historical biogeography and speciationmechanisms of this interesting group of lizards.

MATERIAL AND METHODS

Snout-vent length was measured to thenearest mm with a rule; scale counts andosteological data on cleared specimens (ap-pendix 1) were taken with the aid of a stereo-microscope. Scale counts and scale nomencla-ture are according to Rodrigues and Borges(1997), and osteological nomenclature followsPresch (1980) and Estes et al. (1988). Sex wasdetermined by the presence/absence of femoralpores and confirmed by dissection in pre-viously opened specimens. All data were takenfrom preserved specimens housed at MZUSP(Museu de Zoologia, Universidade de SaoPaulo), MNRJ (Museu Nacional, Rio deJaneiro), MZUESC (Museu de Zoologia,Universidade Estadual de Santa Cruz), andRNHM (Nationaal Natuurhistorisch Mu-seum, Leiden). To compare abundance ofmales and females and total abundanceamong habitats of the new taxon sampled weused one-way analysis of variance (ANOVA;Zar, 1996). When significant differences wereobtained, differences between pairs of habitatswere tested with a Tukey’s test (Zar, 1996).Association among habitat categories andspecies or sex was detected by chi-square test.All analyses were done using Statistica(StatSoft, 1998). Appendix 2 presents the 19species, assigned to 17 genera of Gymno-phthalmidae (including the new one hereindescribed), used in the present study. Externalmorphological characters included in appen-dix 2 were selected among those not present-

ing intraspecific polymorphism after the ex-amination of the MZUSP collections. Hemi-penes were examined when partly or totallyeverted in preserved specimens in order toimprove descriptions and comparisons, butdue to their limited availability among thegenera included here, they were not used in thesubsequent analyses.

The majority of DNA sequences used herewere those from Pellegrino et al. (2001), butnew sequences for the same mitochondrial(12S, 16S and ND4) and nuclear (c-mos and18S) gene regions were included for thepreviously unsampled ingroup taxon Ste-nolepis ridleyi (GenBank accession numbersEF 405618-405622). The present molecularpartition is composed of 2,333 bp of alignedsequences; some adjustments on the originalPellegrino et al. (2001) alignments for theribosomal 12S, 16S, and 18S regions wereperformed manually on the reduced matrixused in this study. This was necessary toaccommodate the sequences of Stenolepis andexclude unnecessary gaps. Regions of ambig-uous alignment for the 12S and 16S wereexcluded from the final analyses on thereduced matrix.

Phylogenetic inference was first conductedon the separate morphological partition fol-lowed by a combined analyses with themolecular partitions under equally weightedparsimony (MP) in PAUP* v4.0b10(Swofford, 2002). For a matrix of 42 morpho-logical characters (appendix 2), with all char-acter states coded as unordered and alltransformations uniformly weighted, a searchwith the branch-and-bound algorithm wasperformed. The analysis of the combined dataset included an MP heuristic search with 1,000replicates of random stepwise addition andTBR branch-swapping.

Nodal support was assessed by bootstrapanalysis (BS; Felsenstein, 1985) with a 1,000random stepwise additions per bootstrappseudoreplicate, and TBR branch-swapping,in both branch-and-bound (morphology) andheuristic (combined data) searches; bootstrapvalues greater than 70% (Hillis and Bull, 1993)were interpreted as strong support for a node.Total and partitioned Bremer support (PBS)values (Baker and DeSalle, 1997), the latterrepresenting the contribution of each specified


data partition to each node, were calculatedfor all nodes of the combined data-partitiontopology using the program TreeRot v. 2.0(Sorenson, 1999).

Modeltest v.3.06 (Posada and Crandall,1998) was used to select the appropriate modelof evolution for each molecular data parti-tion: 12S (TrN+C), 16S (TrN+I+C), ND4(TVM+I+C), c-mos (K80+C) and 18S (JC).Using the combined data set (molecular andmorphology) a partitioned Bayesian analysiswas implemented in Mr.Bayes 3.0 (Hue-lsenbeck and Ronquist, 2001), under the bestfit models of substitution for individual generegions and morphological data set as ‘‘stan-dard’’, with 3,000,000 generations, fourchains, and trees sampled at intervals of1,000 generations. Two independent runs wereconducted. Trees prior to stationarity werediscarded as ‘‘burn-in’’, and a 50% majority-rule consensus tree was obtained from 2,900data points. Nodes on consensus trees fromboth runs with posterior probability (PPvalues) $ 0.95 were considered as evidenceof significant support for a given clade(Huelsenbeck and Ronquist, 2001).

Two species of Alopoglossus, members ofthe subfamily Alopoglossinae formally recog-nized by Pellegrino et al. (2001) and Castoe etal. (2004) and strongly supported as basal toall other clades of Gymnophthalmidae in bothstudies, were constrained to monophyly andused as the outgroup. Selected taxa fromGymnophthalmini, Cercosaurini and Ecpl-eopini were also included to give Stenolepisa chance to fail to be recovered within theHeterodactylini (see appendix 2).

RESULTS

TAXON DESCRIPTION

Alexandresaurus, new genus

DEFINITION: A large and slightly elongategymnophthalmid (maximum SVL 70 mm)with distinctive ear opening and eyelid, largetail (1.63%–2.36% SVL), and slender penta-dactyl limbs; first toe lacking a claw.Frontonasal single; prefrontals, frontoparie-tals, parietals, and interparietals present.Parietals and interparietal longer than wide.Collar fold absent. Two pairs of genials and

three supraoculars. Dorsal scales anteriorlysmooth, isodiametric or subrectangular inoccipital region, becoming progressively nar-rower, more elongate and rounded towardarm level and then lanceolate, strongly keeled,with lateral sides almost juxtaposed.Occipitals absent. Ventrals longer than wide,smooth, in four regular transverse seriesidentical in size. Males with a continuousseries of very conspicuous pores with no gapbetween preanal and femoral; pores absent infemales. Hemipenis without evident biloba-tion, sulcus spermaticus single, marginated bya naked area which is also present on theopposite side of sulcus. Between naked areastwo symmetrical series of continuous trans-verse rows of comb-like flounces with minutespines or irregular tooth-like structures. Aseries of enlarged spines near the base ofhemipenis.

CONTENT: Alexandresaurus camacan newspecies; monotypic.

ETYMOLOGY: A homage to the Braziliannaturalist Alexandre Rodrigues Ferreira, bornin Salvador, state of Bahia, as a recognitionfor his effort to document the natural historyof Brazil at the end of the 18th century. Afterpursuing natural history studies underVandelli at the University of Coimbra,Alexandre headed the famous viagem filosofi-ca (‘‘philosophical voyage’’) to Brazil at theend of 18th century. He remained in Braziluntil 1789 where, among other places, hetraveled extensively along the Rio Negro,Madeira and Guapore in the BrazilianAmazon. The travels of Alexandre were partof a larger project to publish what was meantto be a monumental natural history of thePortuguese colonies (Farias, 2001). The planwas abandoned by the end of 1807 when theroyal family escaped to Brazil against thebackdrop of Lisbon’s imminent conquest byNapoleonic forces. Most of the collectionsassembled by Alexandre during the more than35,000 km traveled in Brazil were publishedby other naturalists following the request ofpart of the material by Etienne Geoffroy SaintHilaire, who was present in Lisbon during thesubsequent conquest.

COMPARISONS: Considering the now recog-nized multiple origins of character complexesassociated with fossoriality in gymnophthal-


mid lizards, and the chaotic state of theirtaxonomy based on external attributes (untilrecently), we compare Alexandresaurus withall other gymnophthalmids while emphasizingthe external morphological differences withthe genera Iphisa, Colobosaura, Coloboda-ctylus, Heterodactylus, and Stenolepis. Exceptfor the latter, this group of genera wasrecovered by Pellegrino et al. (2001) as a well-supported clade that included the closestrelatives of Alexandresaurus. Stenolepis, al-though not included in that study, wastentatively allocated to the same radiation.

Most character states for the taxa com-pared here are summarized in appendix 1.Alexandresaurus differs from Iphisa (data forthe latter in parenthesis) by having: two pairsof genials (one); several rows of lanceolate andkeeled dorsal scales (smooth, in two longitu-dinal rows); ventral scales regularly transversein four longitudinal rows (not in transverserows and in two longitudinal rows); andregularly transverse rows of gulars (notregular). Alexandresaurus also differs fromIphisa in having a naked area opposite to thesulcus spermaticus and large spines at the baseof the hemipenis (naked area absent and onlysmall spines at base of the organ). FromColobosaura Alexandresaurus can be distin-guished from Colobosaura by its smooth neckscales and small, smooth, and posteriorlyrounded scales in the first 7–8 dorsal rows,rarely two enlarged nuchals are present at thecentral part of the second dorsal row; occipi-tals are never present. In Colobosaura neckscales have keels; there are several enlargedscales, much wider than long, in the first eightdorsal rows, the first two characteristicallysmooth and enlarged, forming an occipitalpair separated or not by a smaller scale; theposterior 5–6 dorsal rows are striate or multi-carinate. Ventral scales in Alexandresaurus arelonger than wide, laterally juxtaposed, round-ed posteriorly, identical in size and shape,regularly arranged transversally and in fourlongitudinal series. In Colobosaura there arealso four longitudinal rows of quadrangularventral scales, but they are not regularlytransversal and are irregular in size and shape,laterally imbricate, and in the first ventralrows midline scales always distinctively widerthan long; the external ventral rows are

slightly wider than long in Colobosauramentalis, whereas the reverse occurs inColobosaura modesta. The latter species alsodiffers from Alexandresaurus by having onlythree pairs of genials. As in Alexandresaurus,the hemipenis of Colobosaura mentalis also hasa naked area on the side of organ opposite tothe sulcus, as well as flounces of right and leftlobes medially interrupted by a naked area,a character absent in the former species. Theonly other species referred to Colobosaura wasPerodactylus kraepelini. It is a young female(SVL 5 40 mm) described by Werner in 1910having Puerto Max: Paraguay as type locality.We have not examined the holotype, which islost from the Hamburg Zoological Museumaccording to Dr. J. Hallermann (in litt.); basedon the original description, this species isidentical to Colobosaura modesta, an opinionadvanced long ago by Amaral (1933). Thedistributional data agree with this interpreta-tion because the type locality is in the area ofoccurrence of C. modesta. The only characterthat until now has precluded synonymybetween C. modesta and C. kraepelini wasWerner’s reference to the tongue of hisspecimen as having ‘‘oblique plicae likeAlopoglossus’’. Based on the agreement of allother characters mentioned in the descriptionwe think that Werner was in error in de-scribing the tongue, evidently a strange char-acter and restricted to Alopoglossinae.Alexandresaurus differs from Stenolepis,Colobodactylus, and Heterodactylus in havinga pair of prefrontals (absent). It further differsfrom Stenolepis by having small dorsal neckscales (large, in 3–4 longitudinal rows), largerscales on the sides of neck (small, some almostgranular), ventral and gulars in regularlytransverse rows (not so), and smooth scalesnear venter (keeled). In Colobodactylus andHeterodactylus the parietals are wider thanlong in contrast to the longer than wideparietal scale observed in Alexandresaurusand other Heterodactylini. This difference isprobably due to a possible scale rearrange-ment in Heterodactylus and Colobodactylus inwhich parietals (and interparietal inColobodactylus) were divided and their poste-rior regions incorporated into the highlyconspicuous occipitals that characterize bothgenera. This hypothesis, further discussed


under a phylogenetic framework below, re-veals that even head scales traditionally usedin taxonomic descriptions may not be strictlyhomologous. This is another example of thetaxonomic confusion that may result fromuncritical use of morphological characters inmicroteiid systematics. Alexandresaurus alsodiffers from Colobodactylus and Heterod-actylus in having five toes and smooth flankscales (only four toes and flank scales keeled inthe latter genera). Heterodactylus furtherdiffers from Alexandresaurus by the absenceof external ear, absence of frontoparietals, andabsence or extreme reduction of interparietal.Finally, the hemipenis of Heterodactylus isvery different from that of Alexandresaurus.The organ is distinctively bilobate with a cen-tripetal sulcus spermaticus surrounded bya naked area. The asulcate side of the organhas a series of continuous and unornamentednaked flounces containing only one spine ateach edge, very different from the interruptedflounces containing spines in a comb-likearrangement observed in Alexandresaurus.

Alexandresaurus differs from members ofAlopoglossinae by having imbricate, scalelikepapillae covering the tongue (oblique plicae),and from Rhachisauridae by having a distinc-tive ear opening and eyelid (absent), five toes(four), a normal body (clearly elongate),presence of frontoparietals (absent), anda nostril at the nasal border (in the center ofthe nasal scale). From all other Gymno-phthalminae Alexandresaurus differs by hav-ing five toes (fifth toe reduced or absent),a distinctive eyelid (absent in all Gymno-phthalmini except for Tretioscincus), nostril atthe nasal border (in the center of the nasalscale), a wide and flattened clavicle enclosinga fenestra (simple, boomerang shaped andwithout fenestrae), nasal bones in contact atmidline but broadly separated anteriorly(totally separated by contact between frontaland premaxilary), and, among other charac-ters, by having straight lateral processes of theinterclavicle (posteriorly oriented). Alexan-dresaurus also differs from all Cercosaurinaeby having a distinctive sternal fontanelleprocess (absent), a glossohyal separated frombasiyal (fused), nasal bones broadly separatedanteriorly (in broad contact), and postorbitalcovering postfrontal (covered by postfrontal).

Alexandresaurus camacan, new speciesfigures 1–4

HOLOTYPE: MZUSP 93201, an adult malefrom Una (15u109S, 39u039W): state of Bahia:Brazil, collected by Marianna Dixo on 01November 1999, field number md-1106.

PARATYPES: All from the state of Bahia,Brazil. MZUSP 93179–93200, Una, collectedby M. Dixo between 24 January 1999 and 27February 2000; MZUSP 93331, Ilheus, col-lected by W. Bokerman on December 1971;MZUSP 93462–93464 Ilheus (CEPLAC, sederegional), A. Argolo leg., collected respectivelyon July 1990, 24 April 1991, and 12 May 1999;MZUSP 93227–93228, Jussari (Serra doTeimoso) collected by M. T. Rodrigues, D.Pavan, M. Dixo, and V. K. Verdade on March2001; MZUESC 2431, Ilheus (CampusUESC), collected by C. N. Souza on 8March 2002; RMNH 29741, from Una,Ilheus.

ETYMOLOGY: The specific epithet is a hom-age to the extinct Camacan Indians, a Boto-cudo group who lived in the exuberant forestsof Bahia from north of Rio Pardo to Ilheus.

DIAGNOSIS: A gymnophthalmid with earopenings and eyelids, and slender pentadactyllimbs lacking the claw on first toe.Frontonasal single; prefrontals, frontoparie-tals, parietals, and interparietals present.Parietals longer than wide. Collar fold absent.Two pairs of genials; three supraoculars.Dorsal scales in 30–38 rows, anteriorlysmooth, isodiametric or subrectangular inoccipital region, becoming progressively nar-rower, more elongate and rounded toward thearm, and posterior to the arm becoming morelanceolate, strongly keeled, with lateral sidesalmost juxtaposed. Occipitals absent. Ventralslonger than wide, smooth, in four regularlongitudinal and 15–19 transverse rows, iden-tical in size. Scales around body 29–35; 11–15and 16–20 infradigital lamellae under fingerIV and toe IV respectively. Males with a seriesof 20–25 very conspicuous pores without gapbetween preanal and femoral; pores absent infemales.

DESCRIPTION OF HOLOTYPE (fig. 2A–C):Rostral broad, wider than high, contactingfirst supralabial, nasal, and frontonasal.Frontonasal pentagonal, twice as wide aslong, contacting rostral, nasal, loreal, and


prefrontals. Prefrontals slightly wider thanlong, in broad contact at midline. Frontalhexagonal, with almost parallel lateral mar-gins, slightly longer than wide, anteriorlyindenting the prefrontal and posteriorly thefrontoparietal sutures. Frontoparietals pentag-onal, larger than prefrontals, in slight contact,strongly indented by the interparietal.Interparietal longer than wide, longer andnarrower than frontal, as long as andnarrower than parietals. Parietals heptagonal,marginated laterally by two temporal scaleswith approximately the same size, anteriorlyby the third supraocular and frontoparietal,medially by the interparietal, and posteriorlyby the first dorsals. Three supraoculars, firstthe smallest, second and third about the samesize, second slightly wider, its longest suturewith frontal, third slightly longer, in broadcontact with frontoparietal. Nasal above firstsupralabial, large, longer than high, with thenostril in the center and lower part of scale,indenting suture with labial. Loreal posteriorto nasal, narrower and diagonally oriented;contacting posteriorly first superciliary, preo-cular, and frenocular. Frenocular below preo-cular, followed posteriorly by a long subocular

wider anteriorly and indenting suture betweenpreocular and frenocular. Seven supralabials,third and fourth under the eye, fifth thehighest and contacting two smaller scalesfollowing posteriorly the subocular, the smal-lest one clearly a postocular; seventh suprala-bial the smallest, in contact with the granulessurrounding anterior margin of ear. Threesuperciliaries, first largest, wider anteriorly,longer than first supraocular, contacting preo-cular, loreal, first and second supraoculars,second superciliary and upper eyelid. Centralpart of eyelid with a semitransparent un-divided disc surrounded by small and slightlypigmented granular smooth scales. Lowereyelid with eight strongly pigmented palpeb-rals. Temporal region with few smooth andjuxtaposed scales, irregular in size and shape,between parietals and supralabials; the largeras large as the sixth supralabial. Ear openingsurrounded by a series of very small andjuxtaposed rounded granules; external audito-ry meatus large, tympanum distinct, subovoid,recessed. Lateral surface of neck with smooth,imbricate and mostly longer than wide scales,larger dorsally. All head scales smooth andjuxtaposed with scattered sensorial organs.

Fig. 1. Alexandresaurus camacan (MZUSP 93228), an adult male (57 mm) from Serra do Teimoso, stateof Bahia, Brazil.


Mental broad, wider than high. Postmentalheptagonal, wider than long. Two pairs ofgenials, both contacting infralabials; the firstsmaller, in broad contact at midline; secondpair contacting only anteriorly and reaching

the level of labial comissure. An enlarged pairof symmetric flat and chevron-like scutesfollows second pair of genials, preventingtheir contact posteriorly. Six infralabials,second the largest. Gulars smooth, imbricate,

Fig. 2. Ventral (A), lateral (B), and dorsal (C) views of the head of the holotype of Alexandresauruscamacan (MZUSP 93201). Scale 5 1 mm.


Fig. 3. Dorsal (A) and ventral (B) views of the skull of Alexandresaurus camacan (MZUSP 94252-53).Scale 5 1 mm.


rounded posteriorly, in seven transversal rows;at midline similar to ventrals, much smallertoward side of neck. Gulars followed bya distinct interbrachial region with seven much

larger and elongate scales, about twice as longas gulars. Collar fold absent.

Dorsal scales imbricate and arranged inregular transversal rows; smooth, isodiametric

Fig. 4. Right hand (A), right foot (B), hyoid (C), shoulder girdle (D), and pelvic girdle (E) ofAlexandresaurus camacan (MZUSP 94252-53). Scale 5 1 mm.


or subrectangular in occipital region, becom-ing progressively narrower, more elongatedand rounded at arm level and then posteriorlylanceolate, strongly keeled, with lateral sidesalmost juxtaposed. Thirty transverse rowsbetween interparietal and the posterior levelof hind limbs. Lateral scales about the samesize as dorsals but smooth, strongly imbricatelaterally, less acuminate and more diagonallyarranged than dorsals; those closer to ventralslarger. A distinctive area with granular scalessurrounds the area of arm insertion. Thirty-one scales around midbody. Ventral scalessmooth, longitudinally imbricate, laterallyjuxtaposed, longer than wide, rounded poster-iorly; 17 transverse rows from interbrachials(excluded) to preanals. Six scales in precloacalregion, central and paramedials the largest.Total pores 23, continuous, with no gapbetween femoral and preanal ones.

Scales of tail smaller than midbody dorsals,but otherwise identical; keeled, lanceolate,strongly imbricate longitudinally; those frombase of tail slightly larger ventrally, becominggradually identical around tail toward theextremity.

Forelimbs with large, smooth, and imbri-cate scales; those on ventral part of brachiummuch smaller. Anterior and ventral parts ofhind limbs with irregularly large, smooth, andimbricate scales, identical to the correspon-dent parts of fore-limbs. Posterior part of hindlimbs with granular, juxtaposed scales, grad-ing progressively to larger, imbricate, andkeeled scales in dorsal part of tibia. Carpaland tarsal scales large, imbricate; supradigitallamellae, smooth imbricate. Palmar and plan-tar surfaces with smooth, small granules;infradigital lamellae single, 12 on Finger IVand 19 on Toe IV. Toes and fingers, except forFinger I, clawed, and respectively in thefollowing relative sizes: 1 , 2 5 5 , 3 5 4and, 1 , 2 , 5 , 3 , 4.

Dorsal surfaces of body and tail and lateralpart of tail dark brown with an irregularlydistributed darker punctuation smaller thanone fourth of the scale size. Flanks pre-dominantly yellowish to cream, as the ventralparts of body and tail, but strongly mottled byan irregularly black pattern. Lateral parts ofhead with an identical pattern, black blotchesconcentrated in central parts of supra- and

infralabials, yellow at their sutures. Ventralparts of body and tail creamy yellow, immac-ulate, except in the external rows of gulars andventrals, which present irregular black spots.Ventral part of tail becomes gradually darkertoward the extremity. Limbs dark browndorsally, irregularly mottled with yellow;creamy yellow and immaculate ventrally.

Hemipenis partly everted at preservationexcept for the apex; bilobation not evident;sulcus spermaticus single, marginated by anenlarged naked area that is also present, butnarrower, on the opposite side of sulcus.Between naked areas of the organ, twosymmetrical series of 29 (or more) continuoustransverse rows of comb-like flounces withminute spines or irregular tooth-like struc-tures. Flounces continuous even near basalpart of the organ where their ornamentation ismedially interrupted by a small area withoutspines. A basal series of very enlarged spines ispresent near the base of hemipenis in itsasulcate face.

Measurements of the holotype: snout-ventlength: 66 mm; tail length: 145 mm.

VARIATION: Males are slightly larger thanfemales but have much longer tails: maximumSVL for males and females was 70 mm and66 mm. Tail length varied respectively inmales and females from 1.76 to 2.36 and 1.8to 2.03 times SVL (fig. 5: r2 males 0.93; r2

females 0.29). No sexual differences werefound in squamation. Variation in meristiccharacters (N 5 28) was the following (meanand standard error, respectively, in the paren-thesis): dorsal rows, 28–30 (29 and 0.72);ventrals 15–19 (16.8 and 1.06); scales aroundmidbody 29–35 (31.5 and 1.42); infradigitallamellae under finger IV 11–15 (12.9 and0.19); and infradigital lamellae under Toe IV16– 20 (18.1 and 1.0). All specimens showthree supraoculars and three superciliaries. Allthe specimens have seven supralabials, and allhave six infralabials except RM 29791, whichhas seven. One specimen (MZUSP 93187) hasfour ventral rows fused midventraly resultingin two longitudinal rows of ventrals. We notethat this arrangement is similar to the patternfound in Iphisa, but differs from it because therows of scales are regularly arranged transver-sally in Alexandresaurus and not so in Iphisa.Only males have pores, they are absent in


females. Pores are highly conspicuous, placedin distinctively elevate scales, aligned on eachside without gaps between preanal and femo-ral. Total number of pores varied from 20–25(N 5 16). Sexual color dichromatism isevident; males are characterized by darkcoloration in the flanks mottled with yellowishor whitish blotches and a reddish venter;females and juveniles are more dull coloredhaving unpatterned light brown flanks anda creamy white venter.

OSTEOLOGY (figs. 3–4): The following de-scription is based on two alizarin-preparedskeletons (MZUSP 94252–53). Premaxillary aslong as wide, contacting but not articulatinglaterally with the maxillary. Its dorsal laminatriangular posteriorly, long, just coveringanteriorly the nasals and deeply indentingtheir suture, preventing their anterior contact.Twelve premaxillary conical teeth. Nasalslarge, longer than wide, wider anteriorly,diagonally arranged, widely separated anteri-orly, posterior third in midline contact,covering frontal anteriorly. Frontal longerthan wide, wider posteriorly, covering parietaland articulating with it by a pair of frontopar-ietal tabs. Parietal longer than wide, wider andconcave posteriorly and covering laterally theoccipital region. Epipterygoid contacting ex-ternally a descending epipterygoid process ofparietal. Maxillary dorsally contacting nasal,and lateral parts of frontal and lacrimal,

without overlapping, and extensively coveringprefrontal and jugal; 21 maxillary teeth.Prefrontal large, its posterior process longbut not reaching the level of middle of orbit; inbroad contact with frontal. Lacrimal large,very conspicuous, contacting prefrontal andmaxillary and indenting the orbit. Postfrontaland postorbital single. Postfrontal triangular,contacting jugal, frontal, postorbital andjugal, closing posteriorly the orbita. Postor-bital long and wide, contacting squamosalposteriorly, almost closing the supraorbitalfenestra. Squamosal long, posteriorly curvedand abutting the top of quadrate. Supra-temporal fenestra almost closed and stronglyconstrained laterally by parietal and post-orbital. Supratemporal present, in straightcontact with posterior part of parietal andsquamosal. Fifteen scleral ossicles in the eye.Vomer, palatine, pterygoid, ectopterygoid,and pterygoid present. Vomer, palatine, pre-maxillary, and maxillary in contact restrictingthe fenestra exochoanalis. Infraorbital fenes-tra large, bordered posteriorly by ectoptery-goid. Stapes rodlike, wider at the base. Suturesbetween supraoccipital, exoocipital, basiocci-pital and the otic area of skull are not asvisible as those between basioccipital andbasisphenoid. In the lower jaw dentary,articular, splenial, angular, and supraangularare distinct; there are 14–15 dentary teeth,conic anteriorly, bicuspid or tricuspid poster-iorly.

Glossohyal long, fused to basihyal. Firstceratobranchial curved posteriorly; hypohyaland ceratohyal present. A second short pair ofceratobranchials is present and positionedparallel to the anterior part of trachea.

Anterior part of clavicle very enlarged,flattened, enclosing a fenestra. Interclaviculelong, cruciform, with very long lateral pro-cesses reaching the sternum but not sternalfenestra. Scapulocoracoid with coracoid, scap-ular, and scapulocoracoid fenestrae; suprasca-pula present. Sternum with a large fenestra,receiving three sternal ribs and a xiphisternumwith two inscriptional ribs. Ilium, ischium, andpubis present, the latter with a conspicuouspectinate apophysis. Hypoischium long, largerat the base, almost reaching the preanal border;a small elongate preischium and a small qua-drangular ossificate prepubis are present.

Fig. 5. Regression lines between body lengthand tail length for males and females ofAlexandresaurus camacan from Una, State ofBahia, Brazil.


Twenty-seven procoelous presacral verte-brae, neural spines low, higher anteriorlyhypopophyses present in the first six verte-brae; zigantrum-zygosphene present. Lastpresacral vertebra lacking ribs. Two sa-cral vertebrae. First caudal vertebrae withlong transverse processes, lacking autotomicsepta.

Humerus and femur are slightly larger thanradius and ulna and tibia and fibula.Remaining elements of the forelimbs and hindlimbs as in figure 4.

DISTRIBUTION AND NATURAL HISTORY

Alexandresaurus camacan is currentlyknown only from four localities in thesouthern Atlantic rainforest region of the stateof Bahia: Ilheus, Una, Jussari (Serra doTeimoso), and Urucuca. The maximumstraight-line distance between the most distantof these localities is less than 60 km. Pristinetropical forest with high levels of humidity andrainfall (about 2,000 mm) regularly distribut-ed throughout the year, with a dense leaf litter,large trees reaching up to 30 m high and morethan 50 cm diameter at breast height, and anabundance of bromeliads and other epiphytes,was the typical habitat in the Atlantic rain-forest at this latitude. Most of this area isdominated by lowlands or low hills dissectedby small- to medium-sized streams, inter-rupted by isolated higher mountains. TheSerra do Teimoso is one these higher moun-tains reaching about 1,000 m altitude andreceiving the highest levels of humidity at thetop as a result of the orographic effect andcondensation of air masses coming from thesea. Currently most of the forest cover isrestricted to scattered patches of remnantforest, the Una reserve (Reserva Biologica deUna—ca. 11,500 ha) being the largest one.The largest area of original forest wasdestroyed by agriculture (for cacao planta-tions), cattle ranching, and timber harvestactivities. In cacao groves (‘‘cabrucas’’ locally)where large and emergent trees have beenpreserved to shade the understory cacaoplants, leaf litter is abundant and humidity ishigher than in other disturbed areas.

The localities where Alexandresaurus cama-can was collected show slight differences

related to soil type, levels of humidity, anddistance from the coast. Urucuca (14u339S,39u189W) is located on fertile soils originatingfrom volcanic deposition and is intenselycultivated for cacao production. Ilheus andthe Reserva Biologica do Una (15u109S,39u039W) are located near the coast on thesandy soils of Barreiras Formation and receivemore rainfall. The RPPN (Reserva Particulardo Patrimonio Natural; Serra do Teimoso;15u199S, 39u319W) is an isolated peak towardthe interior of the state and is covered bya semideciduous forest, along with cacaogroves at the lower areas and tropical rain-forest with high humidity levels near the top.

Pitfall traps were installed at Una, Serra doTeimoso, and Urucuca to sample areas ofprimary and secondary forest and cacaogroves. At Serra do Teimoso a series of pitfalltraps also sampled semideciduous forest.Alexandresaurus camacan was always ob-tained in sheltered areas in primary forest orcacao groves, never in secondary forests. AtSerra do Teimoso the only individual collectedwas found in semideciduous forest at highelevation (about 900 m) in the humid forest.No association between habitat categories andsex was detected (p . 0.05, chi-square test).Alexandresaurus camacan was always found inshady areas in primary forest or cacao groves,and never in secondary forests. Total abun-dance (F(2, 27) 5 1,06, p 5 0.36) and femaleabundance (F(2, 27) 5 0.32, p 5 0.73) did notvary significantly among habitats (forest in-terior, forest edge, and cacao groves), whilemale abundance did (F(2, 27) 5 4.70, p 50.018). Male abundance was significantlyhigher in forest edges than in cacao groves (p5 0.03) and marginally higher than in forestinterior (p 5 0.06).

PHYLOGENETIC ANALYSES

Parsimony analyses to determine thephylogenetic placement of Alexandresauruscamacan and its relationships within theHeterodactylini, were first conducted on thepartition composed of 42 morphologicalcharacters (38 of which are parsimony in-formative; appendix 2) with all states coded asunordered. This analysis recovered 211 mostparsimonious trees; the poorly resolved strict


consensus tree (L 5 106; CI 5 0.55 and RI 50.72; fig. 6A) left the position ofAlexandresaurus unresolved within theHeterodactylini, but, although weakly sup-ported, recovered the tribe as monophyletic(BS 5 58, Bremer value 5 4). Except fora well-supported sister relationship betweenColobodactylus taunayi-Heterodactylus imbri-catus (BS 5 95, Bremer value 5 4) this tree leftall other ingroup taxa as an unresolvedpolytomy. Two other well-supported sister-group relationships recovered by the morpho-logical partition were those of Procellosaur-inus tetradactylus-Micrablepharus maximiliani(BS 5 99, Bremer value 5 4) and Colo-bosauroides cearensis-Anotosaura vanzolinia(BS 5 88, Bremer value 5 2).

Combined analyses of the morphologicaland molecular partitions resulted in a singlemost parsimonious tree of 2,740 steps, 660parsimony-informative characters, and CI 50.47 and RI 5 0.43 (fig. 6B). The recoveredrelationships are very similar to those sug-gested by the previous study of Pellegrino etal. (2001) based on molecular data only. TheHeterodactylini was recovered with strongsupport (node 8: BS 5 92, Bremer value 511; see appendix 4). Within the Hetero-dactylini, A. camacan is weakly resolved asthe sister group of I. elegans-C. modesta (node5: BS , 50; Bremer value 5 2; see appen-dix 4); (C. mentalis + S. ridleyi) is recovered asa strongly supported clade (node 6: BS 5100, Bremer value 5 18, see appendix 4); andboth the monophyly of the clade (Colobo-dactylus taunayi + H. imbricatus) and that ofits sister group (A. camacan, C. modesta, I.elegans + C. mentali -S. ridleyi) are stronglyand moderately well supported (respectively,node 3: BS 5 99, Bremer value 5 13, and node7: BS 5 75, Bremer value 5 5; appendix 4).The Gymnophthalmini represented by theProcelosaurinus-Micrablepharus clade (node9: BS 5 100, Bremer value 5 28) is resolvedas the sister clade of the Heterodactylini withhigh support (node 10: BS 5 96, Bremer value5 11). Slightly different from the Pellegrino etal. (2001) hypothesis are the weakly supportedsister relationship Rhachisaurus-Bachia (node1: BS , 50, Bremer value 5 1) and theplacement of Arthrosaura reticulata as thesister taxon of the C. cearensis-A. vanzolinia

clade (node 13: BS 5 63, Bremer value 5 8).The monophyly of the ecpleopines was alsostrongly supported (node 15: BS 5 92, Bremervalue 5 11).

Externally, the (C. ocellata (Rhachisaurusbrachylepis-B. bresslaui)) clade was weaklysupported (node 2: BS , 50, Bremer value 56), and the Heterodactylini-Gymnophthalminiclade (node 10) and the Ecpleopini clade(node 15) were recovered with high sup-port indexes (BS 5 96, Bremer value 5 11;BS 5 92, Bremer value 5 11; fig. 6B, re-spectively).

Figure 7 shows the majority-rule consensustree estimated using partitioned Bayesianmethod, which differs from that recoveredfrom parsimony analysis (fig. 6B). Themajor difference refers to the paraphyly ofthe Heterodactylini with respect to theGymnophthalmini, but this alternative topol-ogy is only weakly supported by posteriorprobabilities (PP 5 0.63). Bayesian tree isunique in recovering a quite well-supportedsister relationship (PP 5 0.94) betweenI. elegans–C. modesta (PP 5 0.51) andC. mentalis–S. ridleyi (PP 5 1.0), withAlexandresaurus basal to this grouping (PP5 0.94). Also, the sister-group relationshipbetween Bachia and Cercosaura is highlysupported (PP 5 0.99). Similar to the MPtree (fig. 6B), the support for Ecpleopinimonophyly is high (PP 5 1.0), although itsposition is unresolved (fig. 7).

TAXONOMIC IMPLICATIONS: Our resultsshow that the position of Colobosaura mentalisin the tree is incompatible with a monophyleticColobosaura. This was also indicated in theanalysis of Pellegrino et al. (2001), whichalthough based exclusively on molecular dataincorporated a much larger dataset. The sister-group relationship between morphologicallyhighly divergent species like Iphisa elegans andColobosaura modesta, suggests the recognitionof another new genus to allocate Colobosauramentalis. We propose the following name toaccommodate it:

Acratosaura, new genus

DEFINITION: A microteiid characterized bya distinctive ear opening and eyelid, large tail,and slender pentadactyl limbs. Frontonasal


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single; prefrontals, frontoparietals, parietals,and interparietals present. Parietals longerthan wide. Collar fold absent. Three pairs ofgenials and three supraoculars. Dorsal scalesanteriorly smooth, large in the occipital re-gion, becoming gradually elongated, lanceo-late, and strongly keeled posteriorly. Occi-pitals present. Ventral scales quadrangular, inlongitudinal but not in regularly transverse

rows; laterally imbricate, irregular in size andshape, scales of external rows slightly wider.Males with continuous series of very conspic-uous pores without gap between preanal andfemoral; pores absent in females. Frontal boneanteriorly divided.

CONTENT: Acratosaura mentalis, new ge-neric name for Colobosaura mentalis Amaral,1933.

Fig. 7. Phylogenetic tree estimated under Bayesian method of the combined morphology and moleculardata set and posterior probabilities (above nodes) for clades recovered at the 50% majority ruleconsensus topology.


ETYMOLOGY: From the Greek akratos,meaning ‘‘unmixed, pure’’; here intended toreflect its unmixed condition after beingdismembered from the polyphyletic Colo-bosaura.

DISCUSSION

Our study provides evidence for the mono-phyly of the Heterodactylini and its sisterrelationship with Gymnophthalmini (nodes 8and 10 respectively; fig. 6B). Castoe et al.(2004) admitted monophyly of Gymno-phthalminae as proposed by Pellegrino et al.(2001), but removed their tribal arrangement.In contrast to Castoe et al. (2004), ourparsimony analyses (MP) recover both themonophyly of the Heterodactylini andGymnophthalmini (nodes 8 and 9; fig. 6) andtheir sister-group relationship (node 10;fig. 6B). The support for Heterodactylinimonophyly, on the basis of combined analysesof molecular and morphological characters, ishigher (BS 5 96, Bremer value 5 11) than thatpreviously obtained using molecular data only(BS 5 71, Bremer value 5 2 in Pellegrino etal., 2001). The partitioned Bayesian analyseswe performed here differently from Castoe etal. (2004), included 42 morphological char-acters. Agreeing with them, our analyses alsorecovered a paraphyletic Heterodactylini withrespect to the Gymnophthalmini, but there islow support for such alternative topology (PP5 0.65). More extensive sampling of morpho-logical and molecular characters is essential toconfirm the result of our Bayesian analyses;these studies are in progress, but until they arecomplete we accept the tree topology re-covered by MP analyses as the best hypoth-esis.

Monophyly of the Heterodactylini is re-inforced by the unique condition of the shapeof their interclavicle and the condition of theirnasals (characters 25 and 34, appendix 1). Thebetter support now obtained is due not only tothe inclusion of morphological characters inthis analysis but also to the addition of thegenus Stenolepis, absent in the Pellegrino et al.(2001) study. Our results also indicate that theHeterodactylini consists of two strongly sup-ported clades, and that morphological char-acters are the major contribution to each

(appendix 4). One is the Heterodactylus-Colobodactylus assemblage; the other assem-bles all other Heterodactylini (respectivelynodes 3 and 7, fig. 6B). This conclusion issupported by the condition of the supratem-poral fenestra, the lateral expansion of theparietals, the postorbital width, shape ofpostfrontal, and the shape and size of parietalscale (characters 23, 31, 33, 35, 37 and 39;appendix 1). The assemblage Colobodactylus-Heterodactylus also differs from the remainingHeterodactylini by their more elongated bodyand degree of limb reduction (characters 17and 21). The genera Alexandresaurus, Colo-bosaura, Iphisa, Acratosaura, and Stenolepisare also characterized by a strong sexualdichromatism absent in Colobodactylus andHeterodactylus. Reproductive males of allgenera in the first clade exhibit a reddishventer and have a lateral color patternconsisting of cream, white, or yellowishblotches against a dark brown or blackground color, which is absent in females.

Alexandresaurus camacan was recovered asthe sister group of the (Iphisa elegans-Colobosaura modesta) clade with weak sup-port, and the partitioned Bremer index showsthat the largest contribution to this relation-ship comes from morphology (node 5, fig. 6B,appendix 4). The present study also confirmsthe previously hypothesized inclusion of thegenus Stenolepis within the Heterodactylini, incontrast to its affinities with Tretioscincus assuggested by Presch (1980). Acratosaura(Colobosaura mentalis in fig. 6B) is recoveredas the sister group of Stenolepis, and thisrelationship received high support (BS 5 100,Bremer value 5 18).

Although our taxonomic sampling is limitedwith respect to total microteiid diversity and isfocused primarily on the Heterodactylini, themonophyly of the Ecpleopini was also con-firmed. Relationships among the genera aresimilar to those obtained by Pellegrino et al.(2001), except for the position of Arthrosaura,which was basal and here is recovered as thesister group to the (Colobosauroides-Anotosaura) clade (node 13; fig. 6B), withmorphology making the second largest con-tribution to the partitioned Bremer index(node 13, appendix 4). In contrast toPellegrino et al. (2001), Castoe et al. (2004)


recovered the Ecpleopini assemblage distantlyrelated to Cercosaurini, which led them toraise the status of the former to subfamiliallevel. Our results confirm monophyly of theEcpleopini with higher support than previous-ly obtained (BS 5 92, BS 5 11 vs. BS 5 75, BS5 6 in Pellegrino et al., 2001) and not relatedto Cercosaurini, a result consistent with theposition suggested by Castoe et al. (2004).

In a different part of the tree, we recoveredthe Rhachisaurinae Rhachisaurus brachylepis,as nested with Bachia and Cercosaura, a resultwe regard as highly suspect. Pellegrino et al.(2001) recovered the Rhachisaurinae as thesister group of Gymnophthalminae, and ap-pendix 4 shows that the strongest contributionfor this relationship (node 1) comes frommorphology, specifically from characters clas-sically subject to convergence (see appen-dix 2). Further, because our taxonomic sam-pling of Cercosaurini is limited (of the 13genera now accepted [Doan, 2003] only Bachiaand Cercosaura are represented here), weprefer to keep the scheme adopted byPellegrino et al. (2001) until more completetaxonomic sampling and additional data setsprovide better resolved phylogenetic hypothe-sis upon which sound taxonomic decisions canbe made.

Most of the Heterodactylini are restrictedto eastern Brazil. The two species ofColobodactylus (C. dalcyanus and C. taunayi)are restricted to the Atlantic forest of south-eastern Brazil (Vanzolini and Ramos, 1977).The genus Heterodactylus includes one species(H. imbricatus) occurring in the Atlanticforests of southern Brazil but extending intoelevated areas in the mountains of the state ofMinas Gerais, where the only other species (H.lundi) seems to be restricted. Apart from somerecords near the coast, the distribution ofColobodactylus-Heterodactylus appears to berestricted to mountainous terrain once cov-ered by or influenced by the Atlantic ForestDomain (see Vanzolini and Ramos, 1977).Except for Iphisa elegans and Colobosauramodesta, the other genera of the Hetero-dactylini are also associated with Atlanticforest or contiguous elevated areas. Alex-andresaurus camacan is restricted to theAtlantic forests of southern Bahia. Stenolepisridleyi occurs in the Atlantic forest north of

the Rio Sao Francisco, but its association withmountainous areas is evident because it alsooccurs in isolated remnant forests of thesemiarid northeastern Brazil (for example,Serra de Baturite [1115 m altitude], state ofCeara, and Pico do Jabre [1044 m], state ofParaiba). Acratosaura mentalis has a largerbut seemingly insularized distribution; it isknown from some isolated areas in theCaatingas of northeastern Brazil, and else-where in the Serra do Espinhaco, a largemountain range reaching to 2,400 m inaltitude that extends from the state of MinasGerais to Bahia. Colobosaura modesta andIphisa elegans are the only Heterodactylini notdirectly associated with mountains or Atlanticforests; the first is a typical Cerrado lizardoccurring both in open areas and in galleryforests, and the second is restricted toAmazonian forests.

We are aware that the ideal approach tounderstand the historic biogeography of theheterodactyline should be based on a well-supported phylogenetic hypothesis derivedfrom more extensive molecular and morpho-logical character sampling than that nowavailable. In our hypothesis, for example, itis evident that nodes 4, 5, and 7 needadditional support. However, some considera-tions based on the present topology might beappropriate at this time and advanced asa preliminary working hypothesis to be testedby additional data.

Assuming the topology of figure 6B, animportant point to note is the ancestralposition of Atlantic forest forms comparedto those from central Brazilian Cerrados(Colobosaura modesta) or Amazonia (Iphisaelegans). Another interesting issue suggestedby the resulting topology is the insularizeddistribution of the older genera as originallyhypothesized by Vanzolini and Ramos (1977).They hypothesized specifically that Colo-bodactylus and Heterodactylus were morewidely distributed during colder episodes ofthe Quaternary, and are presently isolated inmountains or especially favorable areas. Theoccurrence of Stenolepis ridleyi in a forestrefuge in northeastern Brazil also suggestsa range contraction from times when theAtlantic forest extended more to the west thanthe present day distribution indicates. The


sister relationship recovered betweenStenolepis ridleyi and Acratosaura mentalisconfirms this pattern. Although we did nothave any previous idea of the relationshipsbetween entities grouped at node 7 (fig. 6B),in hindsight the recovery of Stenolepis andAcratosaura as sister species is not unexpected.This hypothesis is consistent with an emergingbiogeographic pattern based on sister grouprelationships between Atlantic forest speciesand those occurring along the Serra doEspinhaco in other taxa (see for exampleParrini et al. 1999). Third, why are Iphisaelegans and Colobosaura modesta so wide-spread compared to their relatives? Is theirbroad distribution across Amazonia and theCerrados due, respectively, to recent originand dispersal under favorable conditions?

To answer this last question we should havemore data on divergence times for the speciesinvolved, particularly for those with currentlyisolated populations. Calibration of diver-gence times could also help us to under-stand another intriguing feature of theHeterodactylini radiation. The monophyleticassemblage formed by Alexandresaurus,Iphisa, Colobosaura, Acratosaura, and Steno-lepis consists of monotypic genera, which issurprising when compared with other generaof Gymnophthalmidae occurring in the sameareas. Leposoma, for example, is a much morespeciose genus, and morphological differencesbetween its species are slight, sometimesinconspicuous, contrasting with the strongmorphological gaps we observe among thegenera under study. In the latter, anteropos-terior axes of differentiation affecting size,shape, and ornamentation of body scales areclearly observable. This suggests that devel-opmental processes might be differentiallyassociated with or contributing to speciationamong these lineages.

Since the submission of this paper twohighly distinctive new Heterodactylini werediscovered from widely isolated mountains ofeastern Brazil. Our initial observations of theirtentative phylogenetic positions suggest thatthey support the primary split of this in-teresting radiation, and the biogeographicscenario outlined above. Considering the rateof habitat loss in Brazil and the highsensitivity of these lizards to deforestation,

accelerated study is needed in order to un-derstand better the interesting biologicalproblems they present.

ACKNOWLEDGMENTS

We thank Fundacao de Amparo a Pesquisado Estado de Sao Paulo (FAPESP), ConselhoNacional de Desenvolvimento Cientıfico eTecnologico (CNPq), Comissao Executiva doPlano da Lavoura Cacaueira (CEPLAC),Universidade Estadual Santa Cruz (UESC),the staff of Restauna project for support andthe following persons for their valuable helpin the lab or field, collecting specimens andsending tissues: G. Skuk, V.X. Silva, M.J.Jesus Silva, D. Borges, M.N. Stevaux, A.Langguth, C. Alonso, F. Curcio. Thanks arealso due to Luciana Lobo for the drawingsand to Hussam Zaher, Carolina Castro-Mello,and Gabriel Skuk for access to specimens. J.Sites acknowledges support of NSF awardDEB 9815881 for some of the lab work.

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APPENDIX 1

MORPHOLOGICAL CHARACTERS AND CHARACTER STATES USED FOR PHYLOGENETIC ANALYSES

External morphology and scalation:

1. External ear opening: (0) present; (1) absent2. Ornamentation of head scales: (0) smooth; (1)

rugose3. Posterior margin of head scutes: (0) curved; (1)

straight4. Prefrontal scales: (0) present; (1) absent5. Frontoparietal scales: (0) present; (1) absent6. Superciliary scales: (0) four or more; (1) three or

less7. Number of temporal scales: (0) six or more; (1)

four8. Pairs of enlarged genials: (0) three; (1) two; (2)

one9. Size of interparietal scale: (0) reaching anteriorly

the level of the parietal; (1) shorter10. Anterior dorsal scales: (0) keeled; (1) smooth11. Posterior dorsal scales: (0) keeled; (1) smooth12. Shape of dorsal scales at midbody: (0) lanceo-

late; (1) quadrangular or slightly mucronate; (2)cycloid

13. Ventral scales: (0) strongly imbricate; (1)juxtaposed

14. Flank scales: (0) lanceolate, imbricate; (1)rectangular, smooth, imbricate; (2) quadrangu-lar, juxtaposed; (3) cycloid

15. Collar: (0) absent; (1) present16. Limbs: (0) normal, slender; (1) stout, compact;

(2) reduced17. Body form: (0) normal, 29 presacral vertebrae

(pv.) or less; (1) elongate, more than 29 pv.18. Eyelid: (0) present; (1) absent19. Disc of lower eyelid: (0) divided; (1) single20. Position of nostril: (0) in nasal border; (1) in

center of nasal21. Number of toes: (0) five, without reduction; (1)

less than five22. Interparietal scale: (0) present, large; (1) vesti-

gial or absent23. Parietal scale: (0) longer than wide; (1) short, as

wide as long24. Enlarged occipitals: (0) absent, (1) present

Osteology:

25. Interclavicle: (0) cross-shaped, with central areaextremely enlarged, lateral processes large butpointed; (1) same, but lateral processes short,ending abruptly, not pointed; (2) cruciform,central area reduced and lateral processes

extremely long and straight; (3) cruciform,central area reduced, lateral processes poster-iorly oriented; (4) a longitudinal rod-shapedelement, lateral processes absent; (5) trans-versal, clavicular, and sternal processes absent.

26. Sternal process of interclavicle: (0) long, reach-ing sternal fontanelle; (1) small, not reachingfontanelle.

27. Sternal fontanelle process: (0) absent; (1)present

28. Number of sternal ribs: (0) three; (1) two; (2)one

29. Glossohyal: (0) separated from basihyal; (1)fused

30. Second pair of ceratobranchials: (0) present; (1)absent

31. Supratemporal fenestra: (0) almost closed; (1)opened; (2) opened only posteriorly

32. Postorbital: (0) distinct; (1) fused to postfrontal33. Postorbital width: (0) narrow; (1) wide34. Nasals: (0) wide, almost parallel, in broad

contact under and with premaxillary; (1) wide,divergent and in contact at midline but broadlyseparated anteriorly by the subtriangular lam-ina of premaxillary, in slight contact withpremaxila; (2) separated by contact betweenfrontal and premaxillary.

35. Supratemporal fenestra: (0) closed by parietaland postorbital; (1) opened

36. Clavicle: (0) wide, flattened anteriorly andenclosing a single fenestra; (1) simple, boomer-ang shaped, fenestra absent; (2) axe shapedanteriorly, fenestra absent

37. Postfrontal: (0) irregular, posteriorly widerand longer leaving opened the supratemporalfenestra; (1) triangular; (2) boomerang-shaped

38. Supratemporal: (0) angulose, extremely curvedat the end; (1) slightly curved

39. Lateral expansions of parietal: (0) present; (1)absent.

40. Postorbital: (0) covers postfrontal; (1) coveredby postfrontal; (2) contacts postfrontal withoutoverlap

41. Premaxillary dorsal lamina: (0) wide, poster-iorly triangular; (1) wide, posteriorly straight;(2) sub-triangular with end pointed toward butnot reaching frontal; (3) large, subrectangular,contacting frontal.

42. Frontal: (0) normal; (1) anteriorly divided.


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APPENDIX 3

ADDITIONAL SPECIMENS EXAMINED

Alcohol. Alopoglossus atriventris: MZUSP 13922–24.Estiron, Rio Ampiyacu, Peru. Alopoglossus carini-caudatus: MZUSP 53683–84. Cachoeira do Limao,Rio Tapajos, Para. Anotosaura vanzolinia: MZUSP47754, Serra Negra, Inaja, Pernambuco; MZUSP60773, 60775–77, 60780–85, Cabaceiras, Paraiba;MZUSP 80151, Xingo, Alagoas. Arthrosaura reticu-lata: MZUSP 82644–46 Aripuana, Mato Grosso.Bachia bresslaui: MZUSP 10300. Utiaritı, MatoGrosso. Cercosaura ocellata: MZUSP 82425–27Juruena, Mato Grosso. Colobodactylus taunayi:MZUSP 91446–48 Pinhalao, Parana. Colobosauramentalis: MZUSP 56933, Grao Mogol, MinasGerais. Colobosaura modesta: MZUSP 81346–62,Caldas Novas, Goias. Colobosauroides carvalhoi:MZUSP 89453, Parque Nacional da Serra daCapivara, Piauı. Colobosauroides cearensis: 87580–83, Mulungu, Ceara. Ecpleopus gaudichaudii:MZUSP 78975, 79559, Caraguatatuba, andMZUSP 81401, Ariri, Sao Paulo. Heterodactylusimbricatus: MZUSP 89185–86, Parque Estadual daCantareira (Nucleo Pedra Grande), Sao Paulo;MZUSP 3309 Santa Barbara, Serra do Caraca,Minas Gerais. Iphisa elegans: MZUSP 82654–76,Aripuana, Mato Grosso. Leposoma osvaldoi:MZUSP 8270–37, Aripuana, Mato Grosso.Micrablepharus maximiliani: MZUSP 50172–74,Exu, Pernambuco. Procellosaurinus tetradactylus:MZUSP 71598–604, Alagoado, Bahia. Rha-chisaurus brachylepis: MZUSP 54897, 54898, Serrado Cipo, Minas Gerais. Stenolepis ridleyi: MZUSP78724, Mulungu (Sitio Lorena), Ceara.

Cleared and stained. Alopoglossus atriventris:MZUSP 13922, Peru, Loreto, Rio Ampiyacu.Alopoglossus carinicaudatus: MZUSP 67488,79544, no further data. Anotosaura vanzolinia:MZUSP 60772, 93417, 93418, Cabaceiras,Paraıba. Arthrosaura kocki: MZUSP 82951 VilaRica, Mato Grosso. Arthrosaura reticulata:MZUSP 52504, no further data. Bachia bresslaui:MZUSP 93419, Brasılia, Distrito Federal.Cercosaura ocellata: MZUSP 79556, Chapada dosGuimaraes, Mato Grosso. Colobodactylus taunayi:MZUSP 94254, Salesopolis, Sao Paulo.Colobosaura mentalis: MZUSP 79557, MaracasBahia; MZUSP 66231, Cabaceiras, Paraıba;MNRJ 9968, no further data. Colobosaura modesta:MZUSP 93420, Serra da Mesa, Goias.Colobosauroides cearensis: MZUSP 79595, 93421,Serra de Baturite, Ceara. Ecpleopus gaudichaudii:MZUSP 79558–59, 93424, 93425, no further data.Heterodactylus imbricatus: MZUSP 79562, nofurther data; MZUSP 94255, Caucaia, Ibiuna, SaoPaulo. Iphisa elegans: MZUSP 82654, 82658,Aripuana, Mato Grosso. Leposoma osvaldoi:MZUSP 93426, UHE Samuel, Rondonia.Leposoma scincoides: MZUSP 79564-65, no furtherdata. Micrablepharus maximiliani: MZUSP 66233,Joao Pessoa, Paraıba; MZUSP 79589-91, no furtherdata. Procellosaurinus erythrocercus: MZUSP74935, 74942, Ibiraba, Bahia. Rhachisaurus brachy-lepis: MZUSP 55533, Serra do Cipo, Minas Gerais.Stenolepis ridleyi: MZUSP 66049, Serra dosCavalos, Pernambuco.


APPENDIX 4

MEASURES OF SUPPORT FOR ALL INTERNAL NODES OF MOST PARSIMONIOUS TREE RECOVERED FROM

COMBINED ANALYSIS OF MORPHOLOGY (MORPH) AND MOLECULAR PARTITIONS (fig. 6)

Columns present the bootstraps proportions, totaland partitioned Bremer values; positive and negativepartitioned values indicate support for a given re-lationship in the combined analysis over the alterna-tive relationship in separate analyses, and contradic-tory evidence for a particular relationship in the

combined analysis, respectively. Zero score indicatesthe indifference of a given data set at a specific node(Baker and DeSalle, 1997; Gatesy and Arctander,2000). Node # 5, highlighted in bold, present thesupport for the relationship between Alexandresauruscamacan and its closest relatives (see fig. 6).

Node #

Bootstrap

support Bremer support

Partitioned Bremer

12S 16S ND4 18S c–mos morph

1 , 50 1 1.5 0.5 25.0 22.0 1.0 5.0

2 , 50 6 20.8 4.2 2.2 21.5 1.8 0.0

3 99 13 1.3 4.0 20.3 0.0 3.0 5.0

4 , 50 1 3.0 20.5 23.5 21.0 0.5 2.5

5 , 50 2 2.0 0.5 22.5 21.0 0.5 2.5

6 100 18 7.0 1.0 6.0 1.0 3.0 0.0

7 75 5 23.0 21.0 3.0 0.0 3.0 3.0

8 92 11 1.0 0.5 5.0 21.0 21.0 6.5

9 100 28 2.5 3.0 6.5 22.0 12 6.0

10 96 11 1.0 5.3 25.3 21.7 7.7 4.0

11 50 6 0.0 2.5 1.0 20.5 1.5 1.5

12 100 32 7.0 4.0 4.0 22.0 1.0 18

13 62 8 28.0 21.0 14.8 21.5 0.8 3.0

14 64 8 28.0 21.0 14.8 21.5 0.8 3.0

15 92 11 26.0 0.5 13 21.0 2.0 2.5

16 100 73 0.0 16 20 2.0 28 7.0


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A New Genus of Microteiid Lizard from the Atlantic Forests of State of Bahia, Brazil, with a New...

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