Underwood's classification of the geckos: a 21st century appreciation

9 Pages·2002·5.3 MB·

by  Anthony P Russell

XA
{'±> '"*>
~\J
Bull. not. Hist. Mux. Loud. (Zool.)68(2): 113-121 Issued28November2002
Underwood's classification of the geckos: a 21st
century appreciation
ANTHONY RUSSELL
P.
DepartmentofBiologicalSciences, UniversityofCalgary; 2500 UniversityDriveN.W., Calgary, Alberta,
Canada, T2N 1N4. email: [email protected]
AARON M. BAUER
BiologyDepartment, Villanova University, 800LancasterAvenue, Villanova, Pennsylvania 19085-1699, USA
SYNOPSIS. Thepublicationin 1954ofUnderwood's'Ontheclassificationandevolutionofgeckos'wasthefirstcomprehensive
attempttounderstandthesystematica,evolutionandbiogeographyofthisgroupoflizards.Combiningtheuseoftheexploration
ofnovelcharacterswithaglobaloverviewofgeckos.Underwooderectedhypothesesofrelationshipandpatternsofdistribution.
Inthe48yearssincethatlandmarkpublicationmuchhaschanged,butmuchhasstayedthesame.Underwood'sdivisionofgeckos
intofourmajorclustersisstillrecognisedtoday,althoughthesphaerodactylsarenowregardedasagroupderivedfrom within
the gekkonines, and the diplodactylines have been diminished by the removal ofseveral genera and their placement in the
gekkonines.TheframeworkthatUnderwoodestablishedhasresultedingenericand/orspecieslevelphylogeniesbeinggenerated
fortheeublepharids, somesphaerodactyls, thecarphodactylinediplodactylinesandsomeclusters within thegekkonines. The
latter group, because ofits size, has remained intractable to detailed systematic analysis at the generic level, although the
recognition of many discrete monophyletic clusters within the Gekkonidae (the Gekkoninae of Underwood) holds out the
possibilitythatgreaterlevelsofintergenericresolutionareclosetorealisation.
Underwood'sinitialapproachtothesystematicanalysisofgeckoswasdistinguishedbyitsuseofnovelcharactersofthevisual
systemthatledtonewinsights.Itispossiblethatthenextbreakthroughinhigherlevelsystematicanalysisofgeckosmayagain
comefromtheexploitationofnewcharactersources. Someexamplesofthesepossibilitiesarediscussed.
INTRODUCTION A comparison of the lizard families recognised by Boulenger
(1885),Camp(1923),andmostmodernworkers(e.g.,Maceyetal.,
1997; Harris et al, 1999) reveals almost no discrepancies. The
'[We]wouldliketomakeadistinctionbetweenhow[Dr.Underwood! gekkotan lizards,however,areanexception.Untilthemiddleofthe
thoughtabouttheclassificationof[gekkotans]andwhathethought, 20"' century, lizard systematists variously recognised the
sincein[our]viewhowamanthinksisfarmoreimportantthanwhat Eublepharidae and Uroplatidae as entities distinct from the
he thinks ... [we] suggest that how [Underwood] thought about Gekkonidae. Different classificatory schemes reflected not only
classificationsurvivesuntarnishedtothisday.Doeswhathethought different interpretations ofcharacters, but alternative views ofthe
about it bear critical scrutiny nearly fifty years later?' With these systematic meaning of novel morphologies. For most of the time
words (bracketed modifications aside) Garth Underwood (1971a) after the description ofthe first gecko genera by Laurenti (1768),
began hisA ModernAppreciation ofCamp's ''Classification ofthe gecko systematics was dominated by alpha systematic treatments
Lizards.'' Nearly 50 years after its publication we here consider and the allocation ofnewly discovered species to an evergrowing
Underwood'sclassificationoftheGekkota,itscentralpositioninthe numberofgenera, defined chiefly by externally discernible digital
studyoftheselizards, its influenceon subsequentwork inthefield features. This reliance on digital characters as almost the sole
and how its conclusions have been modified over the intervening determinant of affinity resulted in the widespread recognition of
decades.This work was Underwood'sfirst substantialcontribution compositegeneraconstitutedbydigitallyconvergenttaxa. Further,
to squamate evolution, preceding other major contributions to the thefocusonfootstructuredidlittletoresolvehigherorderrelation-
systematics of pygopods (Underwood, 1957) and snakes ships among gekkotans, as the digital characters then recognised
(Underwood, 1967) andestablishingthewayhewastothinkabout suggested many discrete clusterings of species, but provided few
andemploycharacteranalysisinapproachestowhathadpreviously putative links betweenthem.
been regarded as ratherintractable problems. Garth Underwood was led to the topic of gecko classification,
GarthUnderwood was interestedinboth the theory andpractice which he(1954:469)characterisedas 'farfrom stable', through his
of systematics and also in the evolutionary morphology of the researchonthereptilianeye.Hisearlierworkonretinalmorphology
organisms that he chose as the subjects ofhis systematic analyses. (Underwood 1951a) andpupil shape (Underwood 1951b) hadboth
He was bold in embracing novel sourcesofdata forhis systematic highlighted thedistinctivenessofthe geckoeyeand suggested that
analyses, put forward hypotheses ofrelationship in the hopes that ophthamological characters could be of use in the resolution of
they would be scrutinised and evaluated by others, and frequently higher order relationships among the many gecko genera.
returnedtosystematicproblemsthathehadalreadypublishedsome Underwood'soptimismthattheeyecouldprovideusefulcharacters
years earlier to bring fresh insights and approaches. Specimens was bolsteredby the then recent workofBellairs (1948), who had
always figured prominently as aprimary source ofinspiration and conclusively demonstrated that the true eyelids ofthe eublepharid
newdata. geckoswereprimitivetothederivedconditionofawell-developed
©TheNaturalHistoryMuseum.2002
114 A.P. RUSSELLANDA.M. BAUER
brille and lack of moveable lids typical of other geckos. Further. Stephenson and Stephenson (1956) regarded New Zealand geckos
Walls (1942) and Prince (1949) had examined the eyes of some (Hoplodactylus andNaultinus) as the most primitive forms on the
geckos in their broader ophthamological treatments, suggesting basisofUnderwood's(1955)revisedviewthatamphicoelousverte-
avenues forfurtherresearch. bral centra are primitive within lizards and within the Gekkota.
Walls' (1942)comprehensivetreatmentofthevertebrateeyeled Furthermore, Stephenson (1960) rejected Underwood's (1954)
Underwoodtohypothesisethatthisorgansystemcouldyielduseful ophthamalogicaldivisionoftheGekkonidaeintotwosubfamiliesas
andstableassociationsofcharacters.Thegeneralrecognitionofthe it was inconsistent with osteological characters, but neither
retinalcharacteristicsoftheeyesofgeckosasevidenceofsecondary UnderwoodnorStephenson'correctly'placedallAustraliangenera.
nocturnality, and a preliminary survey (Underwood 1951b) ofthe Also included in Underwood's Diplodactylinae were several
form ofthe pupil suggested that a more intensive survey ofpupil genera not now regarded as closely allied to the Australo-Pacific
formmayprovideameansbywhichgekkonidscouldbesubdivided diplodactylines: Aristelliger, Chondrodactylus, Colopus,
into more manageable and meaningful subsets reflective of their Gymnodactylus,Palmatogecko,Phelsuma,Ptenopus,Rhoptwpella,
evolutionary history. Underwood (1954) set himself the task of Rhoptropus, SaurodacTylus, and Teratoscincus. Four of these,
surveying a moderately comprehensive collection of preserved Chondrodactylus, Colopus, Rhoptropus, and Palmatogecko, share
geckos at the Museum ofComparative Zoology, Harvard Univer- many features in common with eachotherandwithPachydactylus
sity,andtoanalysetheresultingdata.Heusedthesedatatoerectthe (placed by Underwood [1954] in the Gekkoninae). Kluge (1967a)
first modern generic level analysis of gekkotan relationships. He movedthesetaxatotheGekkoninae.andRussell(1972)andHaacke
recognised potential problems with character state interpretation (1976)establishedtheaffinitiesoftheseformsaspartofthePachy-
caused by state of preservation and the limitations of a single- dactylusgroup(seebelow).
character classification, but nonetheless regarded pupil character Two other taxa, Rhoptwpella and Phelsuma, have also been
states to be sufficiently discrete for the purpose ofestablishing a regardedasbeingcloselyrelatedtooneanother(seebelow).Bothof
workableclassificationofgeckos,whichwouldbesubjecttomodifi- these genera, as well as all remaining ones, were moved to the
cation as additional data became available. Werner (1977) later Gekkoninaeby Kluge(1967a)andhaveremainedtheresince, with
demonstrated that pupil shape and dilation change with differing Teratoscincus as the sister group of all other gekkonines. The
lightlevels,andtheseobservationshavehelpedrefineUnderwood's affinities of Gymnodactylus have remained problematic (Abdala
(1954) initial conclusions (see below). 1988, 1996; Abdala and Moro 1996), as have those ofAristelliger
(Russell and Bauer 1993), and Ptenopus (Bauer 1990b), whereas
Saurodactylus has been considered allied to the sphaerodactyline
UNDERWOOD'S CLASSIFICATION OF THE lineage(Kluge 1995). Underwood'sPhyllurusalsoincludedwithin
GECKOS itaspecies now assignedtothe gekkoninegenusNactus.
The Gekkoninae were characterised by Gekko-lype pupils or
secondarily circular pupils. Underwood's (1954) Gekkoninae, al-
Underwood (1954) recognised three families ofgekkotan lizards. though lackingthe taxamentioned above (and withtheadditionof
The Eublepharidae was characterisedby trueeyelids, the lackofa Eurydactylodes, and Crenadactylus as Phyllodactylus ocellatus)
spectacleandverticalpupilsreflectiveofthenocturnaladaptations otherwise included all ofthe genera placed in the group by Kluge
ofthefamily. Thefivegenerahe includedwere thosesubsequently (1967a).Thisgroupingalsoincluded Uroplatus, whichbyvirtueof
placedbyKluge(1967a)inhisEublepharinae,andbyKluge(1987) alargesuiteofautapomorphicfeatureshadbeen accordedseparate
andGrismer(1988)intheEublepharidae. [Note:Thecurrentalloca- familial status by many previous workers (see Bauer and Russell
tion oftaxa employed in this article is based upon Kluge, 2001]. 1989forareview).Inthisregard,Underwood's(1954)resultswere
Underwood(1954)consideredAeluroscalabotesasthemostprimi- similartothoseofWellborn(1933),whohadbasedherconclusions
tivememberofthefamily. Ithassubsequentlybeenregardedasthe on osteological data. Underwood did not rely entirely on the pupil
sistergroupofallremainingeublepharids (Grismer, 1988). character,however,asLygodactylus,withroundpupils,wasplaced
Underwood's Sphaerodactylidae was supportedby the presence in the Gekkoninae on the basis ofother (digital) similarities with
ofaround,diurnal-typepupil(orelipticalorstraightverticalpupilin Hemidactylus.
somecases), theexistenceofafovea, andthepresenceofaspecta- Nine genera were not assigned to family or subfamily by
cle. He included five genera therein, corresponding to Kluge's Underwood. Five ofthese were unplaced due to lack ofmaterial.
(1967a)SphaerodactylinaeandlaterSphaerodactylini(Kluge 1987, Theremainingfourweretaxawithroundpupilsthatwereregarded
1995). as secondarily diurnal gekkonids, but which Underwood consid-
Allremaininggenera wereplaced in theGekkonidae, character- ered,onthebasisofexistingdata,couldnotbeallocatedtooneorthe
ised by a spectacle and lack ofa fovea. Pupil shape was variable. otherofhistwosubfamilies.Ofthelatter,onegenus,Ancylodactylus,
Within the Gekkonidae he recognised two subfamilies, the hasbeensynonymizedwithanother,Cnemaspis.Theothertwowere
DiplodactylinaeandtheGekkoninae.Theformerhadverticalpupils Quedenfeldtia and Pristurus. Of the genera not examined.
with straight margins, orcircularpupils. He included 22 genera in Ceramodactylus has since been subsumed in Stenodactylus, and
this group. Among them are all ofthe genera now assigned to the Dravidogeckohas been synonymized withHemidactylus.
Diplodactylinae by Kluge (1967a) except for Eurydactylodes, Underwoodalsorecognisedsomeinstancesofconvergenceamong
Pseudothecadactylus, and Crenadactylus. Underwood had doubts geckos. Specifically he addressed the allocation ofspecies ofleaf-
aboutthe placement ofthefirstofthesegenera(see below), speci- toed geckos (then chiefly distributed in Diplodactylus and
mens of which he had not examined himself, and changed its Phyllodactylus), and bent-toed geckos (then mostly placed in
allocation the following year (Underwood 1955). Crenadactylus Gymnodactylus). Among the leaf-toed geckos, pupil shape sug-
ocellatuswasexaminedbutwasincludedwithPhyllodactylusinthe gested the transfer of several species ofAfrican Diplodactylus to
GekkoninaebyUnderwood(1954).TheDiplodactylinaewassubse- Phyllodactylus. These geckos are nowregarded as members ofthe
quently retained by Kluge (1987) and Bauer (1990a), although its genusUrocotyledon(Kluge, 1983)andare,asUnderwoodindicated,
affinities with the Pygopodidae were uncertain (see below). correctlyassignedtotheGekkonidaeratherthantheDiplodactylidae.
GECKOCLASSIFICATION 15
Diplodactylyus, as recognised by Underwood, corresponds to two examined material ofonly Narudasia and Saurodactylus. Species
currently recognised genera, Diplodactylus and Strophurus. His level analyses within individual sphaerodactyl generaare ongoing
reconstituted Phyllodactylus included forms now placed in that and have been attempted for the largest genus, Sphaerodactylus
genus as well as Asaccus, Afrogecko, Euleptes, Christinus, (Hass 1991, 1996).
Crenadactylus, Paroedura and Urocotyledon (based on his list of
specimens examined). He also separated Narudasia from Diplodactylinae
Quedenfeldtia, and divided the then cosmopolitan Gymnodactylus
into four genera: Gymnodactylus (restricted to South America), The composition ofthe Diplodactylinae has changed most signifi-
Phyllurus (corresponding to the current Phyllurus and Saltuarius, cantly.Kluge(1967a,b)removedalargenumberofgenerafromthis
but also including the species vankampeni, now allocated to the group to the Gekkoninae, leaving only forms with parchment-
gekkoninegenusNadus), Cyrtodactylus(includingrepresentatives shelled eggs in his Diplodactylinae, and provided a generic level
ofCyrtodactylus,Geckoella,Tenuidactylus.Mediodactylus,Nactus), hypothesis of relationships among the remaining forms. Bauer
and Wallsaurus (the latter now synonymized with Homonota, a (1990a) erected a species level hypothesis ofrelationships among
genus listed as unexaminedby Underwood). theCarphodactylini,oneoftwotribal groupsestablishedby Kluge
(1967a). Additional hypotheses at the species level have been
presented by Good et al. (1997) and Vences et al. (2001). The
STEPS TOWARDS FURTHER SYSTEMATIC Diplodactylini, also established by Kluge (1967a), has yet to be
RESOLUTION investigated phylogenetically at the species level, although Kluge
(1967b)erectedagenericlevelhypothesisofrelationshipsandKing
(1987b)suggestedaspecieslevelphylogenyofDiplodactylusbased
Underwood's (1954)classification providedaspringboard forsub- onseveralkaryotypiccharacters.Underwood(1954)hadpurgedthe
sequent systematic work on geckos. The four large units he genusDiplodactylusoftwotaxawithGekko-typepupils,rendering
establishedwere 'corrected' by Kluge(1967a),butremainedasthe a cluster of taxa still accepted as monophyletic. However, he re-
chief elements in Kluge's higher order treatment of the group. tainedinPhyllodactylusthespeciesocellatus,whichhassincebeen
Moffat (1973) generally accepted Kluge's (1967a) allocation of recognisedasadiplodactylineandplacedinthegenusCrenadactylus.
genera to subfamilies but disagreed with his methodology and his AlthoughthecontentofUnderwood's(1954) Diplodactylinaeas
pattern ofsubfamilial relationships. a whole has changed little, argument persists over patterns of
internal relationship. In particular, the monophyly of the
Eublepharidae Carphodactylini has been called into question (Donnellan et al.
The most stable unit has been the Eublepharidae. This group was 1999) and the relationship ofNew Zealand taxa has also been re-
retained intact by Kluge (1967a), although reduced to subfamilial evaluated (Chambers et al. 2001). King (1987b) and King and
rank. All subsequent researchers have accepted the monophyly of Mengden (1990), based on chromosomal data, argued that Oedura
thisgroupandmorerecenttreatmentshavereflectedthephylogenetic was more closely allied to the Carphodactylini than to other
Diplodactylini. and that pygopods are also allied to the
positionoftheEublepharidaeasthesister-groupofallothergekkotans
bpayttaegranisnoafcrceolradtiinognshitipfawmiiltihailn rtahnekeu(eb.lge.phGarriisdmserhav19e88b)e.enFuerstthaerb,- c(a1r2pShoRdNaAc,tycl-imneoss.),DroengnaerldleadntehtealD.ip(l1o9d9a9c)t,ylbiansied, ionnclmuodliencguOleardudraat.a
lishedatthegenericandspecieslevels(Grismer, 1988, 1991, 1994; as monophyletic, but suggested that the Carphodactylini is
Grismeretal. 1999; Olaetal.. 1999). In this instance. Underwood paraphyletic. They found pygopods to be the sister group of all
Diplodactylines.
(1954)chieflyusedprimitivefeaturesindiagnosingthefamily(e.g.
Patterns ofrelationship within the Diplodactylinae have further
true eyelids present, etc.) but subsequent research has identified
numerous synapomorphies that support the reality of this mono- beencomplicatedbytherecognitionthatpygopodsaremoreclosely
related to this group (or some component thereof) than to other
phyletic unit(Grismer, 1988; Otaetai, 1999).
gekkotans (Kluge 1987). On this basis, Kluge (1987) recognised a
Sphaerodactylidae redefined Pygopodidae for the group that includes diplodactyline
geckos plus pygopods. Good et al. (1997), based in part on argu-
The Sphaerodactylidae ofUnderwood has remained unchanged in ments presented by Bauer (1990a), proposed an alternative higher
terms ofgeneric content. Kluge (1967a) recognised the group as a level scheme, recognising the Diplodactylidae as a family level
subfamily and considered it to be highly derived, in contrast to group. BasedonthepatternsofrelationshipretrievedbyDonnellan
Underwood(1954),whointerpreteditasaprimitivelydiurnalgroup etal. (1999), the Diplodactylidae and Pygopodidae are sistertaxa.
andarelativelyearlyoffshootofthegekkotanlineage.Subsequently As mentioned above, the genus Eurydactylodes proved particu-
Kluge (1987) demonstrated that sphaerodactyls are derived from larlyproblematictoUnderwood(1954)andheonlyincludeditinhis
within gekkonines, confirming their monophyly while obviating Diplodactylinae in the following year (Underwood, 1955). For a
theirrecognitionasahigherordergroup,assuchrecognitionwould varietyofreasons,thisgenushascontinuedtobeenigmatic,exhib-
render the Gekkoninae paraphyletic. This arrangement also re- iting an odd mosaic of characteristics. Although Eurydactylodes
ceivedsupportfromreproductivecharactersincludingtherestriction appears to be a member of a monophyletic New Caledonian
of the calcareous eggshell to gekkonines and sphaerodactylines carphodactyline radiation (Bauer 1990a), itpossesses a numberof
(Bustard 1968; Werner 1972). Kluge (1995) later conducted an features that are problematic and, at least superficially, link it to
explicitinvestigationofthephylogenyofthesphaerodactyls,yield- othergroups ofgeckos. One such feature isthe tail-squirtingappa-
ingafullyresolvedgenericlevelpatternforthegroup.Kluge(1995) ratus.Membersofthisgenushavecaudalglandsthatsecreteasticky
regarded the gekkonine Pristurus as the immediate sistergroup of substanceas adefensive mechanism. Such mechanisms havebeen
the sphaerodactyls and considered Quedenfeldtia, Cnemaspis, widely reported in arthropods (Deslippe et al. 1996), and amphi-
NarudasiaandSaurodactylusasotherappropriateoutgrouptaxafor bians(Arnold 1982),butamongamnioteshavebeennotedonlyfor
his analysis (see below). Of these outgroup genera, Underwood geckosoftheAustraliandiplodactylinegenusStrophurus(Rosenberg
116 A.P.RUSSELLANDA.M. BAUER
and Russell 1980) and Eurydactylodes (Bohme and Sering 1997). of smaller, but putatively monophyletic, genera. The problem re-
Althoughthesecretionhasnotbeencharacterised,itislikelysimilar mains, however, that relationships among these genera are poorly
tothatofStrophurusspp., whichisproteinaceous(Rosenbergetal. resolved. Whiletheidentificationofmonophyletic unitsisaneces-
1984)andiseffectiveindeteringatleastsomesmallpredators,such saryfirststepintheresolutionofgekkotanrelationships,theincrease
asspiders,whichbecomeentangledinthesecretion(Minton 1982). inthenumberofsuchunitsincreasesthesamplingrequiredinorder
However,boththeanatomyoftheglandandtheejectionmechanism toerectahypothesisofrelationshipacrossallmembersofthegroup.
ofsecretiondifferbetweenthetwogeckogenera,suggestingthatthe Thishasbeen the majorstumblingblockin the phylogenetic inter-
apparatusinconvergent(BohmeandSering 1997).Eurydactylodes pretation ofthe Gekkoninae: any attempt to resolve relationships
is also convergent with Strophurus in its bright yellow-orange among some subset ofgenera ofnecessity requires an analysis of
mouthcoloration. Most geckos haveunpigmentedbuccal linings. virtuallyallothergenera.Thesheerdiversityofthegrouphasbeen
Eurydactylodesalsosharessomefeatureswithgekkonidgeckos. an impedimenttoitsresolution.
Mostnotableis the presenceofextracranialendolymphatic sacs in Despite the difficulty of determining relationships among
the neck region, especially injuveniles and reproductive females. gekkonines,someclustersofgenerathatappeartobemonophyletic
Thesecalcium-storingstructuresfrequentlyformconspicuousbulges have been identified. These groups are chiefly those that share
onthenecksofgekkonids,butindiplodactylidsareintracranialand highlydistinctiveandgenerallyrestrictedderivedconditions.Thus,
contain little calcium. Eurydactylodes is an exception in that very suchgroupshavetypicallybeen identifiedonthebasisofinforma-
largesacsareoftenpresent,insomeindividualsartificiallyincreas- tion intrinsic to themselves rather than on the basis of outgroup
ing the apparent size ofthe head (Bauer 1989). Perhaps related to comparison. Indeed, when outgroup analysis has been attempted,
this, the eggshells of Eurydactylodes, although similar in most the choice of outgroup has been based on geography (e.g., Joger
regards to those of typical carphodactylines, are covered by a 1985; Bauer 1990b; Abdala 1996; Macey etal. 2000) or on some
calcified outer surface (Bauer and Sadlier 2000), which otherwise preconceivednotionofsimilarity,usuallybasedondigitalanatomy
typifiesgekkonids (Bustard, 1968; Werner, 1972). (e.g.,ArnoldandGardner 1994; Maceyetal. 2000).Chromosomal
characteristicsofgekkonidsarehighlyheterogeneous(King 1987c),
Gekkoninae butsuchvariationmayoccurwithingeneraandthushascontributed
little totheresolution ofhigherorderrelationships.
The Gekkoninae was the most heterogeneous and unwieldy of Oneofthemostsubstantiallysupportedsubgroupsofgekkonines
Underwood's higher order groups and it has remained largely is the Pachydacnlus group. This is aclusterofgenera sharing the
intractable to this day. Indeed, as a result ofthe resolution ofthe uniquefeatureofhyperphalangyofdigitIofboththemanusandpes.
content ofthe Diplodactyinae, the Gekkoninae has grown signifi- ThegroupincludesthechieflyMediterraneangeneraTarentolaand
cantly. Further, the vast majority ofall new or resurrected genera GeckoniaandthesouthernAfricaformsPachydactylus,Rhoptropus,
since 1954aregekkonines. Underwood(1954)initiatedtheprocess Chondrodactylus, Colopus, andPalmatogecko. Underwood(1954)
ofdismantling some ofthe larger gekkonid genera that he recog- recognised the relationship of all of these except Pachydactylus
nised as polyphyletic assemblages ofdigitally convergent taxa. In itself,placingthemintheDiplodactylinaeandidentifyingapeculiar
particular he addressed the composition of Phyllodactylus and pupilshape,theRhoptropus-type,thatallshared. Severalspeciesof
Gymnodactylus, twoofthe largestand mostcosmopolitantaxa. Pachydactylus(e.g.,P. austeni, P. kochi)arestrikinglysimilar,even
SubsequentreductionofPhyllodactylusoccurredwiththeremoval inexternal appearance, to Colopus and Palmatogecko. By chance,
of Crenadactylus and its shift to the Diplodactylinae (Dixon and however.Underwood's(1954)listoftaxaexaminedrevealsthathe
Kluge 1964),andtheplacementofseveralgeographicallycoherent did not examine any of these species. Hyperphalangy had previ-
gekkonineleaf-toedformsintoParoedura(DixonandKroll 1974). ously been identified in some members ofthe group by Wellborn
Asaccus(Dixon andAnderson 1973), Urocotyledon (Kluge 1983), (1933), but her sampling was inadequate to highlight the potential
and Christinus (Wells and Wellington 1983). All remaining Old phylogenetic valueofthefeature.Russell(1972, 1976)andHaacke
Worldleaf-toedgeckoswereremovedfromthenowstrictlyAmerican (1976) recognised the significance of hyperphalangy and argued
Phyllodactylusby Baueretal. (1997), whoerectedHaemodracon, convincinglythatthiswasevidenceoftherelatednessofthesetaxa.
Dixonius, Afrogecko, Cryptactites and Goggia, and resurrected Virtually all subsequent workers (Bauer 1990b, 2000; Kluge and
Euleptes. Nussbaum et al. (1998) further provided a new generic Nussbaum 1995; Lamb and Bauer 2002; but see Joger 1985) have
name for the elongate-bodied leaf-toed geckos of Madagascar, agreed that these seven genera (including collectively approxi-
Matoatoa.ArnoldandGardner(1994)alsoprovidedaspecieslevel mately80species)formamonophyleticgroup.Withcloselyrelated
phylogenyforAsaccus,usingavarietyofOldandNewWorldleaf- taxa thus identified, species level phylogenies have been possible
toedgeckosasoutgrouptaxa,butwithoutexplicitjustification.Both withinconstituentgenera(e.g.,Rhoptropus: BauerandGood 1996,
these authors and Nussbaum et al. (1998) suggested that at least Lamb and Bauer 2001; Pachydactylus: Lamb and Bauer 2000,
somephyllodactyl taxamightbecloselyrelated. 2002).
AsimilardismantlingofGymnodactyluswasbegunbyUnderwood Otherclusterings,althoughlesswellinvestigated,havealsobeen
(1954), who removed Phyllurus to the Diplodactylinae and recog- proposed, although not necessarily tested. The Gekko group, con-
nisedthegeneraGymnodactylus, Cyrtodactylusand Wallsaurusfor sisting of Gekko, Gehyra, Hemiphyllodactylus, Eepidodactylus,
a subset of the naked-toed geckos. Subsequently Golubev and Luperosaurus,Perochirus,Pseudogekko,andPtychozoon,allshare
Szczerbak (1981) and Szczerbak and Golubev (1984) divided the similarities ofdigital structure (Kluge 1968; Russell 1972, 1976)
Old World forms placed by Underwood in Cyrtodactylus, which andareprobablyamonophyleticgroup,althoughparticularpatterns
they regarded as polyphyletic, into several genera, including the ofintergenericrelationshipremainuntested.
Palearctic Tenuidactylus, Cyrtopodion, Mesodactylus, Carinato- The large and heterogeneous genus Hemidactylus seems to be
gecko,MediodactylusandAsiocolotes.Tropicalformsweredivided related to a number of much smaller genera that are also similar
into Cyrtodactylus, Geckoella andNactus (Kluge 1983). digitally,andareunitedbysynapomorphiesofsizeandshapeofthe
The effect ofthese actions has been to dismantle several larger, intermediatephalanges(Russell, 1977a).Dravidogecko,forexample,
clearlypolyphyleticgroupsandtoinsteadrecognisealargernumber has been synonymized with Hemidactylus on the basis of digital
GECKOCLASSIFICATION 117
morphology (Bauer and Russell 1995). In addition, Cosymbotus, unique crystallin ligand common to several of these taxa (plus
BribaandTeraiolepisarealsoverysimilarandarealmostcertainly Lygodactylus), that they interpreted as synapomorphic. This was
share a common ancestry with Hemidactylus, or are derived from subsequently found in Cnemaspis(Roll, in press), butwhetherthis
within it. indicates affinity orconvergence among secondarily diurnal forms
Bauer (1990b) found some evidence for the recognition of a remains tobedetermined.
Madagascanradiationincludingseveralgeneraofleafandfan-toed The naked-toed geckos have proved especially difficult to deal
geckos including Uroplatus, Ebenavia and Paroedura. Kluge and with. Szczerbak and Golubev (1984, 1986) provided evidence of
Nussbaum(1995)didnotretrieveidenticalpatternsofrelationship, relationship among some Palearctic forms, such as Tenuidactylus,
but these genera nonetheless grouped closely when only Afro- Mediodacrylus,Asiocolotes, and Cyrtopodion. Maceyetal. (2000)
Malagasygeckoswereincludedintheanalysis.AnexpandedIndian foundevidenceforthemonphylyofCyrtopodionandMediodacrylus
Oceanlineage,includingthesetaxaplusAiluronyx,Blaesodactylus, andhypothesizedrelationshipsamongasmall numberofspeciesin
Homopholis, and Geckolepis was retrieved by Bauer (1990b), al- each group. The generic allocation ofcertain Himalayan members
though not by KlugeandNussbaum (1995). ofthe group has provedespecially problematic (Khan 1993; Khan
AnotherputativelymonophyleticgroupistheLygodactyluscom- and Rosier 1999).
plex (Pasteur 1964), which includes twoadditional genera, at least Anothergroupofnaked-toedgeckosincludingAgamura,Bunopus,
one of which, Millotisaurus, is probably derived from within Alsophyla.x,Crossobamon,Microgecko,andTropiocoloteshasbeen
Lygodactylus (Pasteur, 1995; Kriiger, 2001). Lygodactylus itself evenlesswell investigated(LevitonandAnderson 1972;Szczerbak
clustered with Phelsuma in analyses constrained to include only and Golubev 1977; Golubev 1984; Golubev and Szczerbak 1985).
Afro-Malagasy genera(Bauer 1990b; Kluge andNussbaum 1995). The New World nakedtoed forms, GymnodactylusandHomonota,
Kriiger(2001) alsoclusteredLygodactylus andPhelsuma together. have been included in analyses by Abdala (1996) and Abdala and
Although some genera have been revised at the alpha level, and Moro(1996)buttheseinvestigationsincludedonlySouthAmerican
numerous new taxahavebeen erected, mostrevisions have merely gekkonines. Abdala 1988)alsoprovidedaspecieslevelphylogeny
(
proposed species groups, without providing explicit hypotheses of forHomonota (see alsoVanzolini 1968).
relationship(e.g. Pasteur 1964; BrownandParker 1977; Nussbaum While somedegreeofresolution forthegekkoninetaxaoutlined
and Raxworthy 2000). These, like many ofthe othergroups, share abovehasbeenreached,certainothergekkoninesremainenigmatic
digitalsimilaritiesandgeographiccohesiveness. Amongthosegen- and without any sound indication of affinities. Teratoscincus is
eraforwhich some ideaofrelationshipsexist, thereare several for highlyunusual initsmorphology,andappearstobethesistergroup
which species level phylogenies have been proposed, including ofallremaininggekkonines(Kluge 1987).Aspecieslevelphylogeny
Uroplatus (Bauer and Russell 1989) and Gchyra (selected species forthisgrouphasbeengenerated(Maceyetal. 1999).Stenodactylus
only; King 1979, 1983). hassometimesbeenconsideredtobealliedtoTeratoscincus(Kluge
Rhoptropellahasbeenassociatedwithseveraldifferentgeneraby 1967a;KlugeandNussbaum 1995).butitspositionremainsequivo-
differentauthors. Russell (1977b)useddigital morphologytoargue cal (Arnold 1980).
thatitwasinfactaPhelsuma,withnodirectaffinitiestoRhoptropus, AnotherperplexingpadlessgenusisPtenopus,asouthernAfrican
withwhichithadpreviouslybeenassociated(e.g. Boulenger 1885). endemic. Both Bauer (1990b) and Kluge and Nussbaum (1995)
Russell and Bauer (1990) found additional support for this from found little evidence for particular affinities, and constrained or
histological investigations and Good and Bauer (1995) presented retrieved a basal placement among African gekkonines. Ptenopus
allozyme evidence for Rhoptropella's links to Phelsuma. Both possesses a large number of autapomorphic traits (Haacke 1975;
Bauer (1990b) and Kluge and Nussbaum (1995) found the two Rittenhouse et al. 1998; Russell et al. 2000). This mirrors the
genera to be sister taxa when a generic analysis was conducted. situation that plagued analyses of Uroplatus in that many features
Rosier(2001),discussingpholidosis,alsoconcludedthatPhelsuma segregate these geckos from other taxa, but those traits that are
andRhoptropella are sistertaxa. Roll (1999), however, using oph- shared arechiefly primitive ones.
thalmological and digital surface data, interpreted it as displaying Fourpad-bearing genera, which appearunrelated toone another
features of both Rhoptropus and Phelsuma, which, if true, could and have no obvious affinities to previously discussed groups, are
suggestaffinitiesbetweenthechieflyAfricanPachydactylusgroup alsoproblematic.TheseareAfroedura,Aristelliger, Calodactylodes,
and the putatively monophyletic Indian Ocean complex. A variety and Paragehyra. Paragehyra was long known from asinglespeci-
ofcharactertypesalsosuggestthatBogertiaandThecadactylusmay menofasinglespecies,butasecondspecieswasrecentlydiscovered
beallied(Russell and Bauer 1988; Abdalaand Moro 1996). (Nussbaum and Raxworthy 1994). The availability of additional
Cnemaspis, Narudasia, Quedenfeldtia. Saurodactylus and materialallowedtherelationshipsofthegenustobeinvestigatedin
Pristurus have been proposed as gekkonine taxa basal to the moredetail,butthishasnotyieldedanydefinitivestatementsabout
sphaerodactyl lineage (Arnold 1993; Kluge 1995), demonstrating its position within the Gekkoninae (Kluge and Nussbaum 1995),
the paraphyly ofthe Gekkoninae. Although Kluge (1995) did not although Nussbaum and Raxworthy (1994) noted the similarity of
claim any specific relationships among these taxa, his analysis did thedigitsofthisformtothoseofanotherenigmatictaxon,theWest
yield patterns in which Pristurus was the sister group of the IndianAristelliger.
sphaerodactyls, and Narudasia, Saurodactylus and Cnemaspis Russell (1972) grouped Afroedura and Calodactylodes in the
formed aclade. Arnold (1993) advocated the pattern ({((Pristurus, samedigitallydefinedcluster.Loveridge(1944)hadinitiallysegre-
Quedenfeldtia)sphaerodactyls)Saurodactylus)Narudasia).Behav- gatedAfroedurafromtheAustralian Oedura,andthiswasreflected
ioural apomorphies unique to this cluster were documented by in Underwood's (1954) placement of the genera in different sub-
Rosierand Wranik (2001), who noted reproductive morphological families. Some question as to the distinctiveness of these taxa
apomorphiessharedbyQuedenfeldtiaandthesphaerodactylstothe remained, however, untilCogger(1964) conducted detailedosteo-
exclusion of Pristurus. Arnold (1993) provided a species level logical comparisons. Despite some similarities in digital design,
phylogeny forPristurus. The African members ofthis group were Russell and Bauer (1989) concluded that Calodactylodes and
alsoclusteredtogetherinananlysisofAfro-MalagasytaxabyKluge Afroeduraweremorelikelyconvergentthanrelated.BauerandDas
and Nussbaum (1995). Roll and Schwemer (1999) identified a (2000) noted some superficial similarity andgeographic proximity
118 A.P. RUSSELLANDA.M. BAUER
to Asaccus, but again concluded that the relationships of withdrawn in the face ofcompetition with gekkonine geckos. He
Calodactylodeswereobscure. viewedthegekkoninesasamoremodern,expandinggroupthatwas
Aristelliger was one of the taxa regarded as enigmatic by displacing diplodactylines from areas of previous occupancy. He
Underwood (1954). He placed it in the Diplodactylinae and re- regarded New Caledonia as marking the periphery ofthe range of
garded it as an archaic form, possibly unable to compete with the the gekkonines, with Lepidodactylus and Eurydactylodes being
gekkonines,whichheregardedasmorederived.Indeed,heregarded relatively recent invaders into diplodactyline (Rhacodactylus and
itasbeingabasal gekkonnid, retaining oil droplets in theeyes and Bavayia)territory.HebelievedthatPhelsuma,beingchieflyinsular,
displaying vertebral amphicoely. In part. Underwood's (1954) as- diurnal, and arboreal, was ecologically segregated from the
sessmentofthisgenusmayhavebeeninfluencedbythefactthathe gekkonines with which itco-occurs. He regarded its occurrence in
was,atthetime,basedintheWestIndiesandhadmoreinformation mainlandEast Africaasarecentevent. Its arrivalon islands ofthe
aboutitthanmostothergeckos,andcertainlymorethananythathe Indian Ocean was hypothesised to be as a nocturnal stock, an
alsoplacedintheDioplodactylinae.Aristelligerhasbeenemployed offshoot ofthe southern African cluster ofdiplodactylines, with a
inavarietyofevolutionary(Hecht 1952)andmorphological(Ruibal subsequentchangeinlifestyleenablingittocoexistwithgekkonines.
and Ernst 1965) studies, probably because ofease ofavailability. He regarded most continental diplodactylines as being terrestrial,
These studies, however, havehelpedlittletoclarifythepositionof witharboreal forms beingperipheral.
thetaxon. Althoughithasrathercomplexexternaldigital structure, TheforegoingrathertortuousscenariodevelopedbyUnderwood
anatomically it reveals a quite simple architecture. Thus more (1954) to account for diplodactyline biogeography and evolution
detailedstudiesofthedigits(Russell 1976, 1979;RussellandBauer wasthedirectresultoftherecognitionof,asitwasformulatedatthe
1990. 1993) have not assisted in placing it with other genera that time,apolyphyleticassemblage.RemovalofAristelliger,Phelsuma
typically show amore complex anatomy. and a variety ofother taxa (see above) from the Diplodactylinae
(Kluge, 1967)andinclusionofEurydactylodeswithinit(Underwood,
1955)renderedbiogeographicandevolutionaryconsiderationofthe
BIOGEOGRAPHIC AND EVOLUTIONARY remainingdiplodactylidsmoretractable(Bauer. 1990a),butleftthe
Gekkonidae (Underwood's Gekkoninae) yet more unwieldy. That
IMPLICATIONS OF UNDERWOOD'S
some gekkonine genera were present on multiple continents sug-
CLASSIFICATION OF THE GECKOS
gestedto Underwood (1954) that this was the dominant group. He
recognisedfourmajordigitalmorphologiesamonggekkonines,and
Underwood (1954) pioneered a comprehensive approach to gecko believedthateachhadreachedmostareasoftheworldandthatmost
systematics.Asaresultofthis,hewasfacedwithissuesofbiogeog- hadradiatedinsituineacharea,givingrisetonumerousregionally
raphy and evolution that begged an explanation. For geckos, this endemic genera. Thus, whileexpansion was an importanttheme in
was essentially uncharted territory and the recognition ofclusters, the evolution of gekkonines, there was significant within-region
especiallywithinhisGekkoninae,generatednewbiogeographicand evolutionaswell.Theseideaswereobviouslyheavilyinfluencedby
evolutionaryproblems. Chiefamongthesewasthe needtoexplain thoseofDarlington(1948)andbytheideaofcompetitiveexclusion
thebiogeographyandevolutionofhisDiplodactylinae.Thisproved (amoreecologicalthanhistoricalview).Henotedthewaifdispersal
especially challengingbecause, as noted above,thisclusteroftaxa capabilities of some geckos and opined that this complicated the
laterprovedtobethe leaststableofUnderwood's (1954)proposed picture ofdispersal via land bridges that served as his main para-
units. digm. The issue of waif dispersal, though recognised as being
Underwood(1954)interpretedeublepharids,withtheirscattered restrictedtocertaintaxa,remainstothisdayasaconfoundingfactor
distribution, as an ancient radiation with its own specialisations, in the interpretationofthe evolutionofgekkonidspatial patterns.
chiefly to arid conditions, rather than as a cluster of relicts. He Further systematic consideration (see above) has resulted in an
viewed the eublepharids as the primary, ancient Northern Hemi- increasedcomplementofgekkonidgenera, buthas alsoresultedin
sphereradiationoftheGekkota. some level of internal resolution, which, in turn, has influenced
The sphaerodactylids were biogeographically non-problematic some aspects of biogeographic interpretation. For many regions,
asalloccurintheNewWorld.Underwood(1954)viewedthemasan localradiationsofmonophyleticclustersofgenerahavebeenrecog-
early New World offshoot ofthe Gekkota, basedon his beliefthat nised, but resolution of pattern between these clusters remains
they were primitively diurnal, retaining certain plesiomorphic poorly understood.
lacertilian ophthalmological features. Kluge's (1967) demonstra-
tionthatthesphaerodactylsarederivedfromwithinthegekkonines,
and subsequent recognition of secondary diurnality in the
CONCLUSIONS
sphaerodactylines (Roll, in press) hasresultedin areinterpretation
ofsphaerodactylbiogeographyandevolutionaryhistory,withnorth
Africanaffinitiesbeingsupportedbymorerecentsystematicinves- Underwood's (1954) systematic, biogeographic and evolutionary
tigations (Arnold, 1993; Kluge, 1995). considerations of geckos marked the first attempt to comprehen-
Underwood (1954) undertook to explain the distribution ofthe sively assess this circumglobal and highly diverse cluster. His
Diplodactylinaewhich,inhisview,includedalargecoreofAustralo- analysesbrought somedegree ofordertoapreviously verypoorly
Pacifictaxa,butalsogenerafromAfricaandtheAmericas.Henoted understood set of problems, and his choice of ophthalmological
thatnogenusoccupiedmorethanonecontinentandthatmostgenera characters as those ofprimary consideration resulted in the estab-
hadratherlimitedorpatchydistributions. OnlyAristelligerandthe lishment of a basic pattern that has survived to the present in
New Zealand taxa did not co-occur with Gekkonines. He felt that modifiedform.AlthoughUnderwood(1968, 1970, 1971b, 1977a,b)
ovoviviparitymightexplaintheirabilitytosurviveinNewZealand. revisitedthe gekkotan eye repeatedly, thepromise ofphylogenetic
InthecaseofAristelliger,henotedthatitsoccurrencewasbasically utility originally held out by ophthalmolgical data has not. until
complementary to that of gekkonines, and suggested that it may recently, been pursued. Roll (1995, 1997, 1999) and Roll and
have formerly had a broader distribution but had subsequently Schwemer(1999)havedemonstratedthatmanydiurnalgeckosare
GECKOCLASSIFICATION 119
unable to modify pupil shape and instead regulate light through &Moro,S. 1996.CranialmusculatureofSouthAmericanGekkonidae.Journal
absorbancebycrystallinsinthe lens. AlthoughRollandSchwemer ofMorphology229: 59-70.
(1999) assumed that the use ofparticular crystallins was likely to ArnFoaludn,aEo.fN.Sa1u9d8i0.ArAabrieavi2e:w3o6f8t-h4e0l4i.zardgenusStenodactylus(Reptilia:Gekkonidae).
haveevolvedonlyonce,thereisnoevidencethatalldiurnalgekkonids 1993. Historical changes in the ecology and behaviourofsemaphore geckos
are allied (e.g. Phelsuma + Lygodactyhts and Sphaerodactylus + (Pristurus,Gekkonidae)andtheirrelatives.JournalofZoology.London229:353-
Quedenfeltia + Narudasia + Saurodactylus; Kluge and Nussbaum 384.
1995). This avenue of approach, however, suggests that at the & Gardner, A.S. 1994. A review of the Middle Eastern leaf-toed geckoes
(Gekkonidae:Asaccus)withdescriptionsoftwonewspeciesfromOman.Faunaof
anatomicalandmolecularlevel,datafromthevisualsystemmayyet SaudiArabia14:424-441.
be ofsignificance in assisting in the resolution ofpattern between Arnold,S.J. 1982. Aquantitativeapproachtoantipredatorperformance:salamander
nocturnal and secondarily diurnal clusters ofgekkonids (including defenseagainstsnakeattack.Copeia 1982: 247-253.
sphaerodactyls). Bauer, A.M. 1989. Extracranial endolymphatic sacs in Eurydactylodes (Reptilia:
Despite attempts to move away from digital architecture as a JGoeukrknoanlidoafe)H.erpweittohlocgoymm2e3:nt1s72o-n175e.ndolymphatic function in lizards in general.
primarymeansofidentifyingsupragenericclusters,thishascontin- 1990a.PhylogeneticsystematicsandbiogeographyoftheCarphodactylini(Rep-
uedtoplayaroleandhasbeeninstrumental,bywayofexamination tilia:Gekkonidae).BonnerzoologischeMonographien30: 1-217.
of internal architecture, in assisting in the circumscription of a 1990b. Phylogeny andbiogeographyofthegeckosofsouthernAfricaandthe
number ofapparently monophyletic assemblages (Russell, 1976). Gis.laannddsHouftttehreerw,esRt.e(rendsI).ndViearnteObcreaatne:sianptrheelTirmoipniacrsy.aZnoaollyosigsi,scphpe.s27F5o-r2sc8h4u.ngIsn:inPsettietrust.
Pedalanatomyremainsaprimarydeterminantofgeneric allocation undMuseumA. Koenig,Bonn.
and a major clue to potential higher order relationships (e.g., 2000. Evolutionary scenarios in the Pachydacrylus group geckos ofsouthern
NuCshsabnaguemsainndgRenaexrwiocrtalhiygn1m9e9n4)t.and more modern views ofplate ASfqruia&cma:aDtnaase),wfrIh.oymp2o0It0nh0de.isaeAsa.ndrAefSvrriiiecLwaannoJkfoau.trhJneoaulgrenokafkloHneorifpdSeotguoetlnhouAgssyiC4aa8nl:oN5da3at-cu6rr2ya.lloHdiessto(rRyep5t:il2i5a-:
tectonics have necessitated a rethinking of Underwood's (1954) 35.
biogeographic hypotheses. Essentially the eublepharids appear to &Good,D.A.1996.Phylogeneticsystematicsofthedaygeckos,genusRhoptropus
represent an ancient Laurasian radiation, in keeping with Under- (Reptilia:Gekkonidae).ofsouth-westernAfrica.JournalofZoology.London238:
wood's(1954)ideas.Theremaininggekkotansarenowregardedas 635-&66R3u.ssell, A.P. 1989. A systematic review ofthe genus Uroplatus(Reptilia:
being of Gondwanan origin and to consist of an essentially east Gekkonidae). withcommentson itsbiology.JournalofNaturalHistory23: 169-
GondwanandiplodactylidradiationandawestGondwanangekkonid 203.
radiation,withthelatterhavinggivenrise,inturn,totheNewWorld & 1995. The systematic relationships of Dravidogecko anamallensis
sphaerodactyls. (Gii&ntShaedrli1e8r7,5)R..AA.si2a0t0i0c.HTehrepeHteorlpoegtiocaflauRneusoefarNcehw6:Ca3l0e-d3o5n.ia.SocietyfortheStudy
Interpretation ofpatternsofrelationship must now deal with the ofAmphibiansandReptiles,Ithaca.NY.
recognition that the age of the Gekkota is much greater than was ,Good,D.A.&Branch,W.R. 1997.AtaxonomyoftheSouthernAfricanleaf-
believed in 1954 and that many genera might be quite ancient. toedgeckos(Squamata:Gekkonidae),withareviewofOldWorld'Phyllodacrylus'
Hence,genericbodyplansmayhavebeenestablishedforvery long andthedescriptionoffivenewgenera.ProceedingsoftheCaliforniaAcademyof
Sciences49:447^197.
periods, making them ratherdiscrete from one anotherand render- Bellairs, A.d'A. 1948. The eyelids and spectacle in geckos. Proceedings ofthe
ing it difficult toerect hypotheses ofrelationship. Even among the ZoologicalSocietyofLondon 118:420-425.
sphaerodacyls.genericdifferentiation isestimatedtohaveoccurred Bininda-Emonds,O.R.P.,Bryant,H.N.& Russell,A.R 1998.Supraspecifictaxaas
asmuchas40millionyearsago(Hass 1991). King(1987a, 1987b), ptehrymliongaelnsetiinccilnafdeirsetnicce.anBailyosliosg:iciamlplJiocuirtnaalssoufmtphteioLnisnnoefanmoSnocoipehtyyl6y4:an1d01e-r1r3o3r.s in
on the basis of chromosomal and immunological data correlated Bonnie.W.&Sering,M. 1997.TailsquirtinginEurydactylodes:independentevolu-
withtectonichistoryoftheAustralianregion,estimatedaminimum tionofcaudal defensiveglands in adiplodactylinegecko(Reptilia. Gekkonidae).
divergenceof66mybetweenthetwomajorcladesofdiplodactylines, ZoologischerAnzeiger235: 225-229.
and at least 120 my forthe origin ofthe gekkotans. BouHlisetnogreyr),.2Gn.dA.ed.18V8o5l.. C1.aTtraulsotgeueesoofftthheeBLriitziasrhdsMuisnetuhme,BrLiotnidsohnM.useum (Natural
Despitethemagnitudeoftheproblem,onlypatternsofrelationship Brown,W.C.&Parker,F.1977.LizardsofthegenusLepidodacty/us(Gekkonidae)from
withintheratheramorphousGekkonidae(Underwood'sGekkoninae) theIndo-AustralianArchipelagoandtheislandsofthePacific,withdescriptionsofnew
remainrelativelyunassailed.Evenhere,however,large,circumglobal species.ProceedingsoftheCaliforniaAcademyofSciences.4'"ser.61:253-265.
unwieldygenerahavebeenbrokenintosmaller,moregeographically Bustard,H.R. 1968.Theegg-shellofgekkonoidlizards:ataxonomicadjunct.Copeia
1968: 162-164.
circumscribedtaxaandthereisnowanopportunitytobegintomake Camp.C.L. 1923.Classificationofthelizards. BulletinoftheAmericanMuseumof
inroads intothedeterminationofthepatternsofinterrelationshipof NaturalHistory48: 89-481.
supragenericclustersofgekkonidtaxa.Thismaybestbebroachedby Chambers, G.K., Boon, W.M., Buckley,T.R., & Hitchmough, R.A. 2001. Using
taking exemplars, appropriately selected (Bininda-Emonds et al. bmioolteac:utlharreemceatsheodstsudtioes.unAduesrtsrtaalnidanthJeouGronnaldwoafnBaontanafyfi4n9i:tie3s77o-f38t7he.New Zealand
1998) from the putative clusters and the enigmatic genera, and Cogger,H.G. 1964.Thecomparativeosteologyandsystematicstatusofthegekkonid
investigating a combination ofmorphological and molecular data. generaAfroeduraLoveridgeandOeduraGray.ProceedingsoftheLinneanSociety
Giventhemagnitudeoftheproblem,thiswillbeaniterativeprocess ofNewSouthWales89: 364-372.
and will necessitate frequent cross-checking within and between Darlington,P.J.,Jr. 1948.Thegeographicaldistributionofcold-bloodedvertebrates.
QuarterlyReviewofBiology23: 1-26, 105-123.
clusters.TheboldnessofGarth Underwood'sapproachwillhaveto Deslippe,R.J.,Jelinski,L.,&Eisner,T.1996.Defensebyuseofaproteinaceousglue:
beadoptedinselectingnovelsourcesofdatatoallownewapproaches woodlicevs.ants.Zoology99(1995/96):205-210.
tobetakenand insights toberevealed. Dixon, J.R. & Anderson, S.C. 1973. A new genus and species ofgecko (Sauria:
Gekkonidae) from Iran and Iraq. Bulletin oftheSouthern CaliforniaAcademyof
Sciences72: 155-160.
&Kluge,A.G.1964.AnewgekkonidlizardgenusfromAustralia.Copeia1964:
REFERENCES 174-180.
& Kroll, J.C. 1974. Resurrection of the generic name Paroedura for the
phyllodactyline geckos ofMadagascar, and description ofanew species. Copeia
Abdala,V.1988.Analisiscladi'sticodelasespeciesdelgeneroHomonota(Gekkonidae). 1974: 24-30
RevistaEspanoladeHerpetologia 12: 55-62. Donnellan,S.C,Hutchinson,M.N.&Saint,K.M.1999.Molecularevidenceforthe
1996. Osteologia craneal y relaciones de los geconinos sudamericanos phylogeny of Australian gekkonoid lizards. Biological Journal ofthe Linnean
(Reptilia:Gekkonidae).RevistaEspanoladeHerpetologia 10:41-54. Society67:97-118.
120 A.P. RUSSELLANDA.M. BAUER
Golubev,M.L. 1984.StructureofthegenusTropiocolotes(Reptilia.Gekkonidae)[in ousPublications.MuseumofZoology.UniversityofMichigan(173):i-iv+ 1-54.
Russian]. VestnikZoologii1984(6): 12. 1995.Cladisticrelationshipsofsphaerodactyllizards.AmericanMuseumNovitales
&Szczerbak,N.N. 1981. Carinatogeckogen. n.(Reptilia,Gekkonidae)anew (3139): 1-23.
genusofgeckolizardsfromSouthwestAsia[inRussian]. VestnikZoologii1981(5): 2001.Gekkotanlizardtaxonomy.Hamadryad26:1-209.
34^U. & Nussbaum, R.A. 1995. A review of African-Madagascan gekkonid lizard
&Szczerbak,N.N. 1985.OntherelationshipsoftwoPalearcticgeckogenera: phylogenyandbiogeography(Squamata).MiscellaneousPublications,Museumof
TenuidactylusIAgamura (Reptila.Gekkonidae) [inRussian],pp.59-60. In: Prob- Zoology, UniversityofMichigan(183):i-iv+ 1-20.
lemsofHerpetology.SixthAll-UnionHerpetologicalConferenceAbstracts.Tashkent. Kriiger,J.2001. DiemadagassischenGekkoniden.Tiel II: DieGeckosderGattung
Nauka,Leningrad. LygodactylusGray, 1864(Reptilia:Sauria:Gekkonidae).Gekkota3:3-28.
Good, D.A. & Bauer, A.M. 1995. The Namaqua day gecko revisited: allozyme Lamb,T.&Bauer,A.M.2000.RelationshipsofthePachydactylusrugosusgroupof
evidence for the affinities of Phelsuma ocellata. Journal ofthe Herpetological geckos(Reptilia;Squamata:Gekkonidae).AfricanZoology35:55-67.
AssociationofAfrica44: 1-9. & 2001.MitochondrialphylogenyofNamibdaygeckos(Rhoptropus)based
, &Sadlier,R.A. 1997.AllozymeevidenceforthephylogenyofgiantNew oncytochromeband 16SrRNAsequences.Copeia2001:775-780.
Caledoniangeckos(Squamata:Diplodactylidae:Rhacodactylus),withcommentson & 2002. Phylogenetic relationships ofthe large-bodied members ofthe
thestatusofR.leachianushenkelii.AustralianJournalofZoology45:317-330. AfricanlizardgenusPachydactylus(Reptilia:Gekkonidae). Copeia.inpress.
Grismer, L.L. 1988. Phylogeny, taxonomy, classification, and biogeography of Laurenti, J.N. 1768. Specimen Medicum, Exhibens Synopsin [sic] Reptilium
eublepharidgeckos, pp.369^-69. In: Estes, R. andPregill.G. (eds). Phylogenetic Emendatam cum Experimentis circa Venena etAntidota ReptiliumAuslriacorum.
Relationships ofthe LizardFamilies: Essays Commemorating Charles L. Camp. Trattnern,Vienna.
StanfordUniversityPress,California. Leviton,A.E.&Anderson,S.C. 1972.DescriptionofanewspeciesofTropiocolotes
1991.CladisticrelationshipsofthelizardEublepharisturcmenicus(Squamata: (Reptilia: Gekkonidae) with arevisedkeytothegenus. OccasionalPapersofthe
Eublepharidae).JournalofHerpetology25: 251-253. CaliforniaAcademyofSciences96: 1-7.
1994. Phylogeny,classification,andbiogeographyofGoniurosauruskuroiwae Loveridge,A.1944.NewgeckosofthegeneraAfroedura.newgenus,andPachydactylus
(Squamata:Eublepharidae)fromtheRyukyuArchipelago,Japan,withdescriptionof fromAngola.AmericanMuseumNovitales 1254: 1—I.
anewsubspecies.ZoologicalScience11: 319-335. Macey, J.R., Ananjeva, N.B., Wang, Y. & Papenfuss, T.J. 2000. Phylogenetic
,Viets,B.E.&Boyle,L.J. 1999.TwonewcontinentalspeciesofGoniurosaurus relationships among Asian gekkonid lizards formerly ofthe genus Cyrtodactylus
(Squamata:Eublepharidae)withaphylogenyandevolutionaryclassificationofthe based on cladisitic analyses of allozymic data: monophyly of Cyrtopodion and
genus.JournalofHerpetology33: 382-393. Mediodactylus.JournalofHerpetology34: 258-265.
Haacke, W.D. 1975. The burrowing geckos of Southern Africa, 1. (Reptilia: ,Larson,A.,Ananjeva,N.B.,&Papenfuss,T.J.1997.Replicationslippagemay
Gekkonidae).AnnalsoftheTransvaalMuseum29: 197-243. causeparallelevolutioninthesecondarystructuresofmitochondrialtransferRNAs.
Haacke, W.D. 1976. The burrowing geckos of Southern Africa, 5. (Reptilia: MolecularBiologyandEvolution 14: 30-39.
Gekkonidae).AnnalsoftheTransvaalMuseum30: 71-89. , Wang, Y., Ananjeva, N.B., Larson, A. & Papenfuss, T.J. 1999. Vicariant
Harris,D.J.,Sinclair,E.A.,Mercader,N.L.,Marshall,J.C.&Crandall,K.A.1999. patterns of fragmentation among gekkonid lizards of the genus Teratoscincus
Squamate relationships based on c-mos nuclear DNA sequences. Herpetological produced by the Indian collision: a molecular phylogenetic perspective and an
Journal*. 147-151. area cladogram for Central Asia. Molecular Phylogenetics and Evolution 12:
Hass,C.A.1991.EvolutionandbiogeographyofWestIndianSphaerodactylus(Sauria: 320-332.
Gekkonidae):amolecularapproach.JournalofZoology,London225:525-561. Mutton,S.A. 1982. Highlightsinherpetology:it'snotallsnakes.Proceedingsofthe
1996.RelationshipsamongWestIndiangeckosofthegenusSphaerodactylus.a IndianaAcademyofSciences92:70-76.
preliminaryanalysisofmitochondrial 16sribosomalRNAsequences,pp.175-194. Moffat,L.A. 1973.Theconceptofprimitivenessanditsbearingonthephylogenetic
In:Powell,R.&Henderson.R.W.(eds).ContributionstoWestIndianHerpetology: classification ofthe Gekkota. Proceedings ofthe Linnean Society ofNew South
A TributetoAlbertSchwartz. Society forthe Study ofAmphibians and Reptiles, Wales97: 275-301.
Ithaca,NewYork. Nussbaum,R.A.&Raxworthy,C.J. 1994.ThegenusParagehyra(Reptilia:Sauria:
Hecht,M.K. 1952.NaturalselectioninthelizardgenusAristelliger.Evolution6: 112- Gekkonidae)insouthernMadagascar.JournalofZoology.London232: 37-59.
124. — & 2000. Systematic revision ofthe genusParoedura Giinther(Reptilia:
Joger,U. 1985.TheAfricangekkonineradiation preliminaryphylogeneticresults. Squamata: Gekkonidae). with the description offive new species. Miscellaneous
basedonquantitativeimmunologicalcomparisonsofserumalbumins,pp.479^1-94. Publications.MuseumofZoology, UniversityofMichigan(189):i-iv+ 1-26.
In: Schuchmann, K.-L. (ed). African Vertebrates: Systematics, Phylogeny and , &Pronk,O. 1998.TheghostgeckosofMadagascar:afurtherrevisionof
EvolutionaiyEcology. Zoologisches Forschungsinstitut und Museum A. Koenig. the Malagasy leaf-toed geckos (Reptilia, Squamata, Gekkonidae). Miscellaneous
Bonn. Publications,MuseumofZoology, UniversityofMichigan(186): i-iv+ 1-26.
Khan, M.S. 1993. A new angular-toed gecko from Pakistan, with remarks on the Ota,H.,Honda,M.,Kobayashi,M.,Sengoku,S.&Hikida,T. 1999.Phylogenetic
taxonomyandakeytothespeciesbelongingtothegenusCyrtodactylus(Reptilia: relationships ofeublepharid geckos (Reptilia: Squamata): a molecular approach.
Sauria:Gekkonidae).PakistanJournalofZoology25:67-73. ZoologicalScience 16:659-666.
& Rosier,H. 1999. Redescription andgeneric redesignationofthe Ladakhian Pasteur,G.1964.Recherchessurl'evolutiondeslygodactyles.lezardsAfro-Malgaches
gecko Gymnodactylus stoliczkai Steindachner.1969 [sic]. Asiatic Herpetological actuels. TravauxdeITnstitutScientifiqueCherifien.SerieZoologie29:1-132+ 12
Research8:60-68. plates.
King,M.1979.KaryotypicevolutioninGehyra(Gekkonidae:Reptilia).I.TheGehyra 1995. Biodiversite et reptiles: diagnoses de sept nouvelles especes fossiles et
variegata-punctatacomplex.AustralianJournalofZoology27: 373-393. actuellesdugenredelezardsLygodactylus(Sauria.Gekkonidae).Dumerilia2: 1-21.
1983. Karyotypicevolution in Gehyra(Gekkonidae: Reptilia). II. The Gehyra Prince,J.H. 1949. Visualdevelopment1.E.&S.Livingstone,Edinburgh.
australiscomplex.AustralianJournalofZoology31: 723-741. Rittenhouse,D.R.,Russell,A.P.&Bauer,A.M. 1998.Thelarynxandtracheaofthe
1987a.OriginoftheGekkonidae:chromosomalandalbuminevolutionsuggests barking gecko, Ptenopus garrulus maculatus (Reptilia: Gekkonidae) and their
Gondwanaland.Search 18:252-254. relationtovocalization.SouthAfricanJournalofZoology33:23-30.
1987b.ChromosomalevolutionintheDiplodactylinae(Gekkonidae:Reptilia).I. Roll, B. 1995. Crystallins in lenses of gekkonid lizards (Reptilia. Gekkonidae).
Evolutionary relationshipsandpatternsofchange.AustralianJournalofZoology HerpetologicalJournal5: 298-304.
35: 507-531. 1997.Photoreceptorsofdiurnalandnocturnalgeckos.In:Eisner,N.andWassle,
1987c.MonophyleticismandpolyphyleticismintheGekkonidae:achromosomal H.(eds),Proc.25GottingenNeurobiologyConferenceVol.II,502.Thieme,Stutt-
perspective.AustralianJournalofZoology35: 641-654. gart.
&Mengden,G. 1990.ChromosalevolutionintheDiplodactyinae(Gekkonidae: 1999.BiochemicalandmorphologicalaspectsoftherelationshipoftheNamaqua
Reptilia).II.ChromosomalvariabilitybetweenNewCaledonianspecies.Australian daygeckotoPhelsumaandRhoptropus(Reptilia,Gekkonidae).Zoology102:50-
JournalofZoology38: 219-226. 60.
Kluge, A.G. 1967a. Higher taxonomic categories of gekkonid lizards and their In press. Cnemaspis: tertiarily diumal (Reptilia. Gekkonidae). Herpetological
evolution.BulletinoftheAmericanMuseumofNaturalHistoty135: 1-60.pis. 1-5. Journal.
1967b. Systematics, phylogeny, and zoogeography of the lizard genus &Schwemer,J. 1999.L-CrystallinandvitaminA,isomersinlensesofdiurnal
DiplodactylusGray(Gekkonidae).AustralianJournalofZoology15: 1007-1108. geckos.JournalofComparativePhysiologyA 185:51-58.
1968. Phylogeneticrelationshipsofthegekkonid lizardgeneraLepidodactylus Rosenberg, H.I. & Russell, A.P. 1980. Structural and functional aspects of tail
Fitzinger, Hemiphyllodactylus Bleeker, and Pseudogekko Taylor. The Philippine squirting: a unique defense mechanism ofDiplodactylus (Reptilia: Gekkonidae).
JournalofScience95:331-352. CanadianJournalofZoology58: 865-881.
1983.Cladisticrelationshipsamonggekkonidlizards. Copeia 1983:465^75. , &Kapoor,M. 1984.Preliminarycharacterizationofthedefensivesecre-
1987.CladisticrelationshipsintheGekkonoidea(Squamata,Sauria).Miscellane- tionofDiplodactylus(Reptilia:Gekkonidae). Copeia 1984: 1025-1028.
GECKOCLASSIFICATION 121
Rosier, H. 2001. Zur Rumpfbeschuppung der Gattung Phelsuma Gray, 1825: & 1986.GeckoFaunaoftheUSSRandContiguousRegions[inRussian].
ZoogeographischeAspekte(Sauna:Gekkonidae).Gekkota3:47-73. — NaukovaDumka,Kiev.
2.U&ntWerrasnuickh,unWg.en20z0ur1.RBeepmreprokduukntgieonnzvuornFoPrrtipsftluarnuzsunobgsslbiioRloosgiieerv&onWGreacnkiko.ne1n999 Under1w9o5oldb.,PGu.pil19s5h1aa.peReinptcielritaaninregteicnakso.s.NaCtoupreeia1671:9511:83.211-212.
und Pristurus sokotrunus Parker. 1938 im Vergleich zu anderen Geckos (Sauria: 1954.Ontheclassificationandevolutionofgeckos.ProceedingsoftheZoologi-
Gekkonidae).Gekkota3: 125-182. calSociety ofLondon 124:469^192.
Ruibal,R.& Ernst,V. 1965.Thestructureofthedigitalsetaeoflizards.Journalof 1955.Classificationofgeckos.Nature175: 1089.
Morphology 117: 271-294. 1957.OnlizardsofthefamilyPygopodidae.Acontributiontothemorphology
Russell,A.P. 1972.Thefootofgekkonidlizards:astudyincomparativeandfunctional andphylogenyoftheSquamata.JournalofMorphology100:207-268.
anatomy. UnpublishedPh.D.thesis.UniversityofLondon.England. 1967.A ContributiontotheClassificationofSnakes. BritishMuseum(Natural
1976.Somecommentsconcerninginterrelationshipsamongstgekkoninegeckos. History) Publication No. 653. Trustees ofthe British Museum (Natural History),
pp.217-244. In: Bellairs. A.d'A. & Cox, C.B. (eds). MorphologyandBiologyof London. 179pp.
Reptiles.LinneanSocietySymposiumSeries3.AcademicPress.London. 1968. Some suggestionsconcerning vertebratevisualcells. VisionResearch8:
1977a. The phalangeal formula of Hemidactylus Oken, 1817 (Reptilia: 483-488.
Gekkonidae): a correction and a functional explanation. Anatomia Histologic! 1970.Theeye.pp.1-97.In:Gans.C.&Parsons,T.(eds).BiologyoftheReptilia,
Embryologia6: 332-338. vol.X.AcademicPress,London.
1977b.ThegeneraRkoptropusandPhelsuma(Reptilia:Gekkonidae)inSouthern 1971a.AmodernappreciationofCamp's'Classificationofthelizards.'Pp.vii-
Africa:acaseofconvergenceandareconsiderationofthebiogeographyofPhelsuma. xviiInCamp,C.L.,ClassificationoftheLizards.SocietyfortheStudyofAmphibians
ZoologicaAfricana 12: 393-408. andReptiles,Lawrence.Kansas.
1979. Parallelismand integrateddesign in the foot structureofgekkonineand 1971b. Vertebrate comparative anatomy at the cellular level: visual cells and
diplodactylinegeckos. Copeia 1979: 1-21. phylogeny. pp.64-69. In: Hecht. M. (ed). Vertebrate Evolution: Mechanism and
&Bauer,A.M. 1988.Paraphalangealelementsofgekkonidlizards:acompara- Process. ReportoftheNATOAdvancedStudy Institute, RobertCollege, Istanbul.
tivesurvey.JournalofMorphology197: 221-240. Turkey4-15August, 1969.NATO.
& 1989.Themorphologyofthedigitsofthegoldengecko.Calodactylodes 1977a. Simplificationanddegeneration in thecourseofevolutionofsquamate
aureus(Reptilia:Gekkonidae)anditsimplicationsfortheoccupationofrupicolous reptiles. Colloques internationaux C.N.R.S. N° 266 - Mecanismes de la
habitats.Amphibia-Reptilia 10: 125-140. RudimentationdesOrganeschezlesEmbryonsdeVertebres.pp.341-351.
& 1990.DigitIinpad-bearinggekkoninegeckos:alternatedesignsandthe 1977b. Comments on phyletic analysis ofgekkotan lizards. NATOAdvanced
potential constraints ofphalangeal number. Memoirs ofthe QueenslandMuseum StudiesInstituteSeries(LifeSciences) 14:53-55.
29:453^172. Vanzolini, P.E. l968.Geography of the South American Gekkonidae. Arquivos de
& 1993. Aristelliger Cope. Caribbean geckos. Catalogue ofAmerican Zoologia 17: 85-112.
AmphibiansandReptiles565.1^4. Vences, M„ Henkel, F.-W. & Seipp, R. 2001. Molekulare Untersuchungen zur
,Rittenhouse,D.R.&Bauer,A.M.2000.LaryngotrachealmorphologyofAfro- PhylogenieundTaxonomiederNeukalcdonischenGeckosderGattungRhacodactylus
Madagascangeckos-acomparativesurvey.JournalofMorphology245:241-268. (Reptilia: Gekkonidae).Sulamandra37: 73-82.
Stephenson, N.G. I960. The comparative osteology of Australian geckos and its Walls, G.L. 1942. The Vertebrate Eye. Cranhrook Institute Scientific Bulletin 19.
bearing on their morphological status. Transactions ofthe Royal Society ofNew Bloomfield Hills, Michigan.
Zealand84: 341-358. Wellborn, V. 1933. Vergleichende ostcologische Untersuchungen an Geckoniden,
& Stephenson,E.M. 1956.Theosteologyofthe NewZealandgeckosandits EublepharidenandUroplatiden.SitzungsberichtederGesellschaftNaturforschender
bearing on their morphological status. Transactions ofthe Royal Society ofNew FreundezuBerlin 1933: 126-199.
Zealand84: 341-358. Wells,R.W.& Wellington,C.R. 1983.AsynopsisoftheclassReptiliainAustralia.
Szczerbak,N.N.&Golubev,M.L.1977.SystematicaofthePalearcticgeckos(genera AustralianJournal<>/Herpetolog) 1:73-129.
Gymnodactylus,Bunopus,Alsophylax) [in Russian]. TrudyZoologiiskogoInstitute! Werner,Y.L. 1972.Observationsoneggsolcublepharidlizards,withcommentsonthe
AkudemiyaNaukUSSR74: 120-133. evolutionoftheGekkonoidea.ZoologischeMededelingen47: 211-224.pi. I.
& 1984. On generic assignement ofthe Palearctic Cyrtodactylui lizard 1977.EcologicalcommentsonsomegekkonidlizardsoftheNamibDesert,South
species(Reptilia,Gekkonidae)[inRussian]. VestnikZoologii 1984(2): 50-56. West Africa.Madoqua10: 157-169.