Bull. nat. Hist. Mus. Lond. (Zool.) 64(1): 63-89 Issued 25 June 1998 

Structural niche, limb morphology and 
locomotion in lacertid lizards (Squamata, 
Lacertidae); a preliminary survey 

E.N. ARNOLD 
Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK 
CONTENTS 
NVA CIC (OTe ease ns so cn core ste ihn cones ste scny! «cnoter eagomteseseatenbessts 
Phylogenetic relationships of the Lacertidae .................::cceeee 
SiMichialmmchesiah lacentic Unzand Ss f..c-scscesspseensecevsescreseseesareneshe= 
Overview of lacertid structural niche space ..........:..ssccseseeeeeeees 
Structural habitats occupied by groups within the Lacertidae 
Prinniitivea bal AC ANGI Ch OLAS ener saece exacesect ct aceese cucanesek caaset uncstugetschsevun Contonsdesaveni sven dens cectsseentectcavessduisdsistecesnesotavecnasastatcaress ans 65 
Wlemmbe rs On tne Amina te CLAM Cae axe anen ats aster <aacan ge exes sant deen aver agnnsartaseantatdargered Cin cvsoscecataiencivessecuesscansstersdusaueiatenesseaar 
I OUINCLOMeTayy CIALIS Cin STIG EVN all ON GING cate acesas conn aietasias cc accantaxcesasuveven: iste contac seecstdaterecerensecchcttncsunccceesdcasasvastasGusesuesvansnsvacnee 
INSU GOIPS EURO ON ee ees ar ese ese ante ae acne eee cece nen isa ian ans cavescs taractenan Mics s cota ete scope ence ev sean rac ence ee asnae ene seni woeaee Aer tateesor act 
ESO CLV APLOPOLMOUS AUG Vette OLel eM UNSC Te crear ease acec es cece ne ca axira-taseeae eaeteae Tatar oce Mace ec ons ans teana ce eaecceneswecot ca ceen neaetetee <cacvessee acer 
ING AUIV EM AUN SIO Cl wee ccvscracecctence sen stecac eves cose oat vee cares Sanson oaseccec er 68 tens Mote tees ees Ferenc PGE N eee ce coat ce nae So eeas Rat cece aeace eee. face ees 
FETT SLO POU OMS; ANIC SUNG CUA NLC UG reer tes ees ce-cencreccenas foc cevenc ces eure ar Seeee Ne dene ieestc eatevas ova ccoen devedet ces waves ty eetted save reetasvees ae stasoee 
De kal MM OnpkScm Mm lite Den Steen csccseeccecceseceoccsetsse@a-crensecseweesesecvscee 
Relative proportions of femur and tibia ...............00. 
PALLOLES(O lr MINDY OT OWL rastecedusacctoecvns csesoscestesteer¥csacseucsvscsescaveceesseatavasezss 
Evolutionary plasticity of limb proportions ........... 
Functional aspects of differences in limb proportions .............:::0c000 
DeTeNGe SMD EDEL CaUthe ml OLaltsM OME Se MMUSC] Grate gate, tenes cerns ce qchcen cus capsceetes acer sae asi cecseossscecsdatecenatsnhaccvacearaeadectacsdcersbeaetntee 
Ste ral pal ALO O LMC en NN CNL tere tareneee Beare: Seat wena eta evened nan cn acs tease nines acc vecedaresccaddrasecnnsetres aks stebencs ssvenencieestacdactvenesertat 
PNA CERES COUT © nN oe saad eect TNA Nr Reece nc Oper MON Me a onthe cig Seiny ceeecne means Favs treshe nasties teirwscennexseseishacesees idence Tieduareress 74 
URE ME Smin sroundicywellin lac ent cls TOM OPCW) SUA OUS 1202: cucex-cencencassnteoscenespssnsnacudncortoevivsndasesces+ esses caccennsntvarsissetsaneeatac 75 
IUD EMME SMMMACEKNASKe PT any Climbs, SLESMs [OMEN SUGLACES) sucanencssaxectausuderaccenecseceecesssesenesase cneeuteacenssrestvuscuxinceusstyccnsersscacs 75 
Walia VOnsiinithedicectOnlOn Kaka SMOiLOG ro! Of LACEKIS meecenccececscacnseesanesercndesec«tacetees-ccsessstensseacksteescusvasanerancasuschecsastoy an WL 
Zen Gals Ont Chieti l eval wtavex idayelhanoloveriss Oy old nls) eh tz Van tll hE) peat se sececentccace:Pucc togeorcoce-eoekencacbeee ber cir adeaceenck ecepcraccoeeseaee eecoacens THE 
SIUC nO LAU pI AIN US Peescata cee areeRec mee teeta ceo Ree esac wet nec aet ener trac ooROr cas scm tocereeeR ag tape tcsecccte rece ccasareacewsdecscasrenhesseseieotraccsteirieainee 78 
The manus in ground dwelling lacertids from open situations ............. 
The manus in lacertids regularly climbing on steep open surfaces ...... 
ChraractemSiics Ol the Lectin Otlen LACTIS i .cc.cesereteceessessteresecescsbensseaeerens 
PE CIAMS MIC HUINES OW UC CL GUUS recrenensetesecscencescessssenseascecectecnesscts-acesvessesers=x 
PX PANGE CISUDGLOttall ARICW Ae eeecmeeesenteat andecesss etcee ceeseee esice cecesseracenser? 
INES STOUNSUD GI EttalUSCal es memrrematee rs eeetet: caeataeeen rete deceecs ra cucccucs tec vartacceuasstisuctnecsceodeststesenteacconeascesctcgreestseneevacshissaseesasiccee 80 
WDD aaa ral Sie ey anes tetra ceseemcate ee ersten ics esate cece ree ee icc sese Sonseccs eacec dues savessesaeavactscastestistesettsssec?terios suteenensiesceressacascesssenie 
PE SOMMOUMOM AMG LL OMe aereeteeneneeceerera ete seen acca te en eeteeNaret ec acca nvens et nae Sov enecue sucsartotere success stacsssisncoeet taser sas suessessiscastssst veosteies ace 
ocompuonma sroundidwellersrotmiherArmiatinedicladene....-c.-eecccecceteeseeecr-csecaeeceadececsvscentsudescaesntoseeent'etucantreascetessioereececeres 
INMOWIGIMELAIS ONT (late. LnNiTAVGT IN0010) Senate sceneotceeosetonckk co adeee zens eosscor cone eeraannar cer pacar mee cockc Rea Fore Aac aE EERER coche Eno aAace eh eee cee coor 
Rotationlomtiketemunandisupposed mes thictlonsiOMitSMOVEMICIE cceee-cce- ceece cee sccerecee cece es co ceeeceesactace Geecascecsncceaessaseuces=neer 
MiheisuppoOsediproble mkomomalMmo Lat Olav ceeeencvestueerece-csscesnsvencevecenstecoaseteseesestecurerscarcurcustieeviristscsedertsecsnascentestrsaeatticcer 
PAGAN tia MOGE © tabUirrC Mn op OWE INCI en cee nes es ees cene Caster esas eeees seer nc reteeendoncreoroactecsctnaussnctendsdsescars oacrsgacecest <eestays setts eres 
Other hind limb gaits in ground dwelling lizards — continuous gearing 
Movements of the foreleg in ground dwellers .............:c:sesceeteeseeereeneeeees 
Functional aspects of the limbs and feet of ground dwelling lacertids ... 
GroundhOcomotonemMic tm SeSPOCLES tee veseeccterarecscseree-citees senecadseee cpr newtesee caneacis ioc react oancisescndtacenact snsv nus sodsccescassstarsece 
EOCOMOMOnMMIG IMB ERS OMIStCE PI OPEN SULLACES  wice scence asc ece sansa ste ceeuseeraeescete eon ae tea cnureuarnsceesme Rosser secu tances ssasen er scarecensccs cesses 
NTOVEIMOeN tS: Ofetinew nim QUI Dye cexcc- cess oan seess ave ceeceec acer ceea-tove.vcieoncueatacentl Susi sects on ceacsetenctsdutesc Fatvscucesrdcseattedsenaesncestovecraescess 
INZOVEIIEMTS 0 ile Ones NADY csc creccence see scvasnuteretect cae eet toes scevat Wace tte act nioren ste Zvertanain sveu se metrh ne dacaeaseadegeneae ee ov Reseevsncvernet che 
Other patterns of locomolonin specialised ic limmbimp AC Ertl Sireeeetecnsecneeteeses cesses -eteter scene ceney stn seseneccentsrse:sacesensestertee=es cs 
Broblemsrof upward: Venti calllOCOmMotlonerrsrs ctecccsstrecessux<-<cacsteetersosrercocnocastsereseesvestwortrrstcer: sascceetsususcerssretceverredranees ocests 
Functional aspects of the limbs and feet of specialised climbing lacertids 
Climbing in specialised ground dwelling species 
Concluding remarks 
Acknowledgements 
I ESSIEN See cach aedcnatcopechcecee Ace AeRE Ce RGAE Asch ce ba EAE LARC Hace Oar eR REE SSS pcr coe AES CEA CRANEE EE SoA RecA REO ROO eRe PEPE Ce REO eee 



























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