pachydactylus labialis , commonly known as the calvinia thick-toed gecko , western cape gecko , or western cape thick-toed gecko , is a gecko species endemic to the western and northern cape in south africa , often found taking shelter under stones .

pachydactylus labialis

pachydactylus waterbergensis bauer & lamb 2003 pachydactylus weberi acuminatus — mertens 1955 : 49 ( part . ) pachydactylus weberi werneri — griffin 2003 : 33 ( part . ) pachydactylus waterbergensis — bauer et al . 2006 pachydactylus waterbergensis — mashinini & mahlangu 2013
pachydactylus capensis ( a . smith , 1846 ) – cape thick - toed gecko
pachydactylus gaiasensis steyn & j . mitchell , 1967 – brandberg thick - toed gecko
pachydactylus
. the reptile database . www . reptile - database . org .
pachydactylus is a genus of insectivorous geckos , endemic to africa , and commonly known as thick - toed geckos .
pachydactylus is a genus of arboreal insectivorous geckos , endemic to africa , and commonly known as thick - toed geckos .
neue / new pachydactylus - art / species p . boehmei online ( 4 : 43 pm , 11 / 25 / 2010 )
the geographic range of the genus pachydactylus is centred on southern africa , some reaching east africa , the northernmost limit of their distribution .
pachydactylus kochii v . fitzsimons , 1959 = colopus kochi ( v . fitzsimons , 1959 ) – koch ' s thick - toed gecko
pachydactylus bibronii ( a . smith , 1846 ) = chondrodactylus bibronii ( a . smith , 1846 ) – bibron ' s thick - toed gecko
the genus pachydactylus is characterised by dilated toe tips , usually with undivided scansors . body scales are small , granular and non - overlapping , with scattered , large keeled tubercles .
a lineage accumulation in the pachydactylus group . the plot depicts ltt curves for 1000 trees randomly sampled from the beast posterior distribution . b histogram of γ statistic estimates for the 1000 ltt curves depicted in ( a )
bauer , a . m . & lamb , t . 2003 . a new species of the pachydactylus weberi - group ( reptilia : squamata : gekkonidae ) from the waterberg plateau , namibia . cimbebasia 19 : 1 - 12
beolens b , watkins m , grayson m . 2011 . the eponym dictionary of reptiles . baltimore : johns hopkins university press . xiii + 296 pp . isbn 978 - 1 - 4214 - 0135 - 5 . ( pachydactylus barnardi , p . 17 ) .
beolens b , watkins m , grayson m . 2011 . the eponym dictionary of reptiles . baltimore : johns hopkins university press . xiii + 296 pp . isbn 978 - 1 - 4214 - 0135 - 5 . ( pachydactylus barnardi , p . 17 ) .
maximum snout–vent length for species in the genera pachydactylus , chondrodactylus , colopus , and elasmodactylus . values are based on literature sources along with observations of field - collected and museum - preserved specimens . the small - bodied species p . geitje and large - bodied p . namaquensis are illustrated
branch , william r . ; aaron m . bauer , todd r . jackman , and matthew heinicke 2011 . a new species of the pachydactylus weberi complex ( reptilia : squamata : gekkonidae ) from the namibrand reserve , southern namibia . breviora ( 524 ) : 1 - 15 . - get paper here
bauer am , lamb t , branch wr . 2006 .
a revision of the pachydactylus serval and p . weberi groups ( reptilia : gekkota : gekkonidae ) of southern africa , with the description of eight new species
. proc . california acad . sci . 57 ( 23 ) : 595 - 709 .
bauer , aaron m . lamb , trip . branch , william r . 2006 . a revision of the pachydactylus serval and p . weberi groups ( reptilia : gekkota : gekkonidae ) of southern africa , with the description of eight new species . proc . cal . acad . sci . 57 ( 12 - 24 ) : 595 - 709 . - get paper here
the results are consistent with a model in which lineages more likely to become geographically isolated diversify to a greater extent , although some patterns also resemble those expected of an adaptive radiation in which ecological divergence acts as a driver of speciation . therefore , the pachydactylus group may represent an intermediate between clades in which radiation is adaptive versus those in which it is non - adaptive .
we test whether body size or habitat preference is associated with the formation of geographic isolation in the pachydactylus group in a phylogenetic context . we have generated a comprehensive time - calibrated multi - locus phylogeny of the group , and obtained body size and habitat preference trait data for all ingroup species . geographic range size estimates are produced for all species , and the association between trait data and range size is quantified . we also estimate patterns of lineage accumulation through time and trait - associated estimates of diversification . our data show that both body size and habitat preference affect range size , and that variation in these traits is also correlated with variation in diversification rate , suggesting that allopatric divergence following isolation has played a key role in speciation in the pachydactylus group .
time - calibrated phylogeny of pachydactylus and related genera . the topology is the maximum clade credibility tree estimated in beast with non - gekkotan outgroups cropped for clarity . support values ( bayesian posterior probabilities / ml bootstrap ) are given at nodes ; asterisks indicate nodes with bayesian support values = 1 . 0 and ml bootstrap values > 95 . named species groups and genera are given to the right . geologic epochs and eras are indicated on the timescale ; post - miocene epochs ( pliocene , pleistocene , holocene ) are not labeled
both body size and habitat use are inferred to have shifted multiple times across the phylogeny of the pachydactylus group , with large size and generalist habitat use being ancestral for the group . geographic range size is correlated with both of these traits . small - bodied species have more restricted ranges than large - bodied species , and rock - dwelling species have more restricted ranges than either terrestrial or generalist species . rock - dwelling and small body size are also associated with higher rates of diversification , and subclades retaining ancestral conditions for these traits are less species rich than subclades in which shifts to small body size and rocky habitat use have occurred . the phylogeny also illustrates inadequacies of the current taxonomy of the group .
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
maritz , b . , de silva , r . , milligan , h . t . , wearn , o . r . , wren , s . , zamin , t . , sears , j . , wilson , p . , lewis , s . , lintott , p . & powney , g .
justification : widespread and common with no major threats ; hence this species is assessed as least concern .
endemic to south africa . it occurs in western portions of the western and northern cape provinces , from fonteinskop in the ceres karoo northwards to gelykwerf in the richtersveld national park ( bauer and branch 2003 [ 2001 ] ) .
the population is stable . an estimate of the number of existing subpopulations is not possible .
found in moderately mesic situations in a diversity of habitat types that provide suitable rocky or vegetative ground cover . prefers coastal habitats and river valleys with sandy substrates . occurs from sea level to at least 800 m ( branch 1998 , bauer and branch 2003 [ 2001 ] ) .
to make use of this information , please check the < terms of use > .
republic of south africa ( cape province ) type locality : steinkopf , little namaqualand .
bates , m . f . ; branch , w . r . , bauer , a . m . ; burger , m . , marais , j . ; alexander , g . j . & de villliers , m . s . ( eds . ) 2014 . atlas and red list of the reptiles of south africa , lesotho , and swaziland . suricata 1 . south african national biodiversity institute , pretoria , 512 pp .
bauer , a . m . , and branch , w . r . 2003 . the herpetofauna of the richtersveld national park , northern cape province , republic of south africa . herpetological natural history 8 : 111 - 160 [ 2001 ]
fitzsimons , v . f . m . 1938 . transvaal museum expedition to south - west africa and little namaqualand , may to august 1937 - reptiles and amphibians . ann . transvaal mus . ( pretoria ) 19 ( 2 ) : 153 - 209 - get paper here
loveridge , a . 1947 . revision of the african lizards of the family gekkondiae . bull . mus . comp . zool . harvard 98 : 1 - 469 - get paper here
mashinini , p . l . and mahlangu , l . m . 2013 . an annotated catalogue of the types of gekkonid lizards ( reptilia : squamata : gekkonidae ) in the herpetology collection of the ditsong national museum of natural history , south africa . annals of the ditsong national museum of natural history 3 : 165 - 181
rösler , h . 2000 . kommentierte liste der rezent , subrezent und fossil bekannten geckotaxa ( reptilia : gekkonomorpha ) . gekkota 2 : 28 - 153
this database is maintained by peter uetz ( database content ) and jakob hallermann , zoological museum hamburg ( new species and updates ) .
western cape . [ photo price w . © , from sarca virtual museum ]
western cape . [ photo dorse c . & van rooyen s . © , from sarca virtual museum ]
confused by a class within a class or an order within an order ? please see our brief essay .
to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
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fitzsimons , v . f . m . ( 1938 ) : transvaal museum expedition to south - west africa and little namaqualand , may to august 1937 . reptiles and amphibians . — ann . transvaal mus . , 19 ( 2 ) : 168 , fig . 7 — terra typica : steinkopf , little namaqualand , 23 august 1937
four specimens were collected : t . m . 18052 - 18055 , at steinkopf , little namaqualand , 23 august 1937 . type . t . m . 18055 .
colour . above , grey to greyish brown with dark markings arranged irregularly , in transverse series or in thin longitudinal stripes over back ; a dark band on side of head from second upper labial through eye and over temporal region above ear and then curving inwards round occiput , but not quite meeting its fellow behind ; a dark streak from nostril to just above eye ; labials largely infused with dark brown ; below , creamy white to greyish . tail more or less distinctly barred with dark brown to blackish ; reproduced tails spotted .
dimensions . type , t . m . 18055 , h . and b . 44 , tail ( partly reproduced ) 38 , length head 10 , 5 , breadth head 7 , 5 , forelimb 11 , hindlimb 15 mm .
field notes . taken under loose stones lying on the ground on the slopes of and along the bottom of a valley among the hills .
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disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
discrete geographic regions , both continentally and on islands , often have biotas dominated by a relatively small number of species - rich lineages . the most obvious of these dominant groups are adaptive radiations in which a single ancestral species has given rise to descendants filling numerous niches , examples of which include galapagos finches ( 14 sp . , 58 % of breeding songbird species in the archipelago ) , lake victoria cichlids ( 169 sp . , 67 % of ray - finned fishes in the lake ) , and west indies
] . however , there are many other less - known examples of regionally prominent radiations . among lizards , one of the most striking are geckos of the genus
( 2 species ) . by species number , these geckos are the most successful radiation of lizards in southern africa . sixty four of 66 species occur in the southern african subcontinent , defined as that part of africa south of the zambezi and kunene rivers , and most are endemic to this region . these species occupy all major habitat types in southern africa , and many species in the group display morphological novelties such as loss of adhesive toe pads or the evolution of interdigital webbing [
numerous possible causative factors have been posited or shown to explain the relative success of species - rich organismal groups . in classic adaptive radiations , the rapid evolution of morphological disparity may be the key process in spurring lineage accumulation [
] . in other cases , the evolution of a novel trait may allow organisms possessing that trait to access underutilized resources , with utilization promoting ecological diversification and lineage accumulation . examples include the evolution of antifreeze proteins in antarctic icefishes and evolution of the pharyngeal jaw structures in parrotfishes [
and its relatives , none of these potential explanations are likely to fully account for the observed species diversity .
] , each of which is believed to be monophyletic , and each of which is morphologically conservative . the degree of morphological disparity between these groups is not of a magnitude expected in a classic adaptive radiation [
] . this is not to say that there is no morphological variation within the genus . some morphological novelties have evolved , including the previously mentioned foot characteristics as well as variation in scalation . however , such morphological novelties do not appear to be strong drivers of speciation in the group . for example , the most species - rich radiation of
many geckos do have visual display systems or other traits which could theoretically promote diversification via divergent sexual selection . examples include the semaphore geckos (
] . in these genera , males are boldly patterned and use signaling behaviors to defend territories or attract mates .
and its relatives are strictly nocturnal , however , and typically have a drab pattern . nor are any other prezygotic isolating mechanisms evident that could plausibly be hypothesized to be under sexual selection . finally , the relative age of
] , which also occurs mainly in southern africa but includes only nine species .
and its relatives , then traits promoting the formation of geographic isolation should affect both species’ range sizes and diversification rate . the heritability of range size has been a matter of debate , but increasing numbers of studies demonstrate its heritability [
and its relatives , two variable traits of interest that may promote geographic isolation are body size and habitat preference . there is substantial body size variation in
varies , with species showing preferences ranging from sand dunes to rocky cliffs to houses . in southern africa , the periodic advance and retreat of kalahari and namib sands over geological time is linked to climatic variation [
] ; this process has likely allowed intermittent connections to form between adjacent rocky habitats , but the prevailing pattern is that terrestrial habitats are relatively continuous while rocky habitats are more discontinuous . as a result , a preference for rocky habitats may be expected to be associated with geographic isolation and smaller range sizes . such substrate specialization has been suggested to facilitate speciation in the
species to serve as a near outgroup . an additional 18 gekkotan and 4 non - gekkotan taxa (
) were included as more distant outgroups , with outgroup species choice partially determined based on utility for molecular clock calibration . nearly all ingroup sequences are associated with vouchered museum specimens . sequences for four species (
) are exceptions , with sequences derived from genetic material obtained from captive - bred individuals ; in these cases the live specimens were viewed by the authors to confirm identification and associated genetic material has been deposited in the cryogenic collection at the museum of comparative zoology , harvard university .
) . all newly generated sequences were deposited in genbank ( accession numbers ky224166–ky224347 ) .
for new sequences generated in this study , dna was obtained from frozen or ethanol - preserved tissue samples using qiagen dneasy tissue kits under the manufacturer’s protocol . pcr ( polymerase chain reaction ) amplification of fragments was performed in 25 μl reactions , under standard reaction conditions [
] . pcr purification was performed using ampure magnetic beads , followed by cycle sequencing and purification using cleanseq magnetic beads . capillary electrophoresis was performed on an applied biosystems 3730xl sequencer . sequence assembly was performed using bioedit [
phylogenetic analyses were performed using maximum likelihood ( ml ) and bayesian ( bi ) optimality criteria . for each analysis , model and partition choices were separately identified under the bayesian information criterion using partitionfinder [
] . in each case considered models of evolution were limited to those models that can be implemented by the programs used for phylogeny estimation . greedy search schemes were employed and thirteen potential data blocks were considered : twelve data blocks corresponding to the three codon positions for each of the four protein - coding genes and the trna data comprising the thirteenth data block .
] . one hundred independent searches were implemented on the original data set to identify the best tree , followed by 1 , 000 non - parametric bootstrap replicates to assess branch support . based on the partitionfinder results , the data were divided into eight partitions , each using one of two models : nd2 codon position 1 , nd2 codon position 2 , trnas , and ( pdc position 1 + 2 + rag1 position 1 + 2 ) used the gtr ( general time reversible ) + i + γ model , while nd2 position 3 , ( pdc position 3 + rag1 position 3 ) , ( kif24 position 1 + 2 ) and kif24 position 3 used the gtr + γ model .
] , using a yule tree prior and uncorrelated lognormal relaxed clock . based on the partitionfinder results , the data were divided into ten partitions employing six distinct models : nd2 position 1 , nd2 position 2 , and trnas used the gtr + i + γ model . nd2 position 3 used the gtr + γ model . rag1 position 1 + 2 used the trn ( tamura - nei ) + i + γ model . rag1 position 3 and kif24 position 1 + 2 used the hky ( hasegawa - kishino - yano ) + γ model . pdc position 3 and kif24 position 3 used the k80 ( kishino 1980 ) + γ model . pdc position 1 + 2 used the trnef + i + γ model . four replicate analyses were run for 50 million generations , sampled every 1000 generations . the first 5 million generations were discarded as burn - in . effective sample sizes were estimated in tracer 1 . 5 ( > 300 for all parameters in each run ) to confirm the chain length was adequate .
beast 1 . 8 . 2 was also used to estimate divergence times simultaneously with phylogenetic relationships . the root prior ( lepidosauria - archosauria divergence ) was given a normal distribution ( mean = 275 ma [ million years ago ] , sd = 15 ) encompassing the range of estimates for this divergence [
] . maximum body size was treated in two ways depending on analysis . when possible , svl ( snout - vent length ) was treated as a continuous character and the log - transformed maximum svl was used . however , when treatment of size as a continuous character was not computationally feasible we instead treated size as a binary character . in
) . those species with a maximum snout–vent length ( svl ) < 70 mm comprised the “small” category , and those with a maximum svl > 75 mm comprised the “large” category . habitat preference was divided into three categories . those species that primarily shelter in burrows or under surface debris ( logs , loose stones , aloe leaves , etc . ) , and forage actively on the ground , were classified as “terrestrial . ” species that primarily shelter in rock cracks and forage on cliff faces or boulders were classified as “rupicolous . ” finally , unspecialized species that both shelter and forage on a variety of surfaces ( rock faces , tree trunks , buildings , etc . ) were classified as “generalist climbers . ”
] . eoo was calculated as the area of the minimum convex polygon enclosing distribution records for each respective taxon . aoo was initially calculated as the sum of the total area of the quarter degree grid squares within which at least one record occurs . the final aoo was adjusted to an estimate of the actual suitable habitat within the occupied quarter degree squares based on the literature and the authors’ field knowledge of each species . for all endemic south african species and for most species with a portion of their distribution occurring in south africa , eoo and aoo values were previously estimated as part of the red list evaluation carried out in association with the atlas and red list of the reptiles of south africa , lesotho and swaziland [
] . calculated eoo usually provides the broadest possible interpretation of the space used by a species , whereas the aoo represents a quite conservative estimate . however , for taxa known from single localities or several localities that are very close to one another , aoo as calculated above may yield a greater area than eoo . we used eoo or aoo , which ever was the greater , as our estimates of species’ ranges . these values were log transformed when used in analyses .
we performed a variety of comparative analyses to investigate the relationship among phylogeny , divergence times , trait data , and range sizes . all comparative analyses were completed in replicate on both the beast maximum clade - credibility tree and on 1000 post - burnin trees randomly sampled from the beast posterior distribution . these trees were pruned to remove outgroups ( for which we have incomplete taxon sampling and no trait data ) . the package phytools [
] , which allows for tests of correlation between a multistate vs . continuous trait in a phylogenetic context . because body size evolution and habitat choice may be coupled [
] , we also tested for auto - correlation between these two traits , for a total of three analyses : ( 1 ) svl vs . range size , ( 2 ) habitat vs . range size , and ( 3 ) svl vs . habitat . svl was treated as a multistate ( binary ) trait in test ( 1 ) , but was treated as a continuous trait in test ( 3 ) to facilitate analysis . in all three cases , we tested the hypothesis that the optimum continuous trait value ,
, differed depending on the identity of the multistate trait value , i . e . whether large - and small - bodied species differ in range size ( test 1 ) , whether species differing in habitat use differ in range size ( test 2 ) , or whether species differing in habitat use differ in body size ( test 3 ) . we performed these tests by estimating ancestral states of the multistate character on the phylogeny , and then fitting values of
( rate of change of trait ) for the continuous trait under two model regimes . the null brownian motion ( bm ) model regime estimated single values of
that did not depend on the state of the multistate character . this was tested against a more complex ornstein - uhlenbeck ( ou ) model in which there were multiple
parameters , one per multistate character state . we used δaicc ( corrected akaike information criterion ) values to identify which model provided a better fit for the data . all three tests were performed on 1000 trees randomly sampled from the post - burnin beast posterior distribution , and on each of these 1000 trees we performed 100 ancestral reconstructions of the multistate trait using stochastic character mapping [
] implemented in phytools , resulting in a total of 100 , 000 model fits per test , each with a unique combination of phylogeny and ancestral state estimate .
] model for habitat data implemented in diversitree . because hypothesis testing in a bisse or musse framework can have a high type i error rate [
] , we refrain from explicitly testing the statistical significance of character - associated variation in model parameter estimates . instead , we fit models in which each trait was given individual
parameters strictly to determine estimates of these model parameters . model fitting was performed in an markov chain monte carlo ( mcmc ) framework with runs lasting 1100 generations and the first 100 discarded as burn - in . these estimates were obtained for each of 1000 trees randomly sampled from the beast posterior distribution , resulting in each parameter estimate being obtained from 1 , 000 , 000 observations .
the phylogenies estimated in both the ml and bi analyses are very similar ( fig .
) , and most branches receive strong support . as expected , the grouping of
is monophyletic ( bi / ml support values 1 . 0 / 97 ) and these are in turn most closely related to
] , which included 26 fewer ingroup taxa and was estimated from ~ 1 , 600 fewer nucleotide sites , but there are some notable differences . most notably , both the genera
group ( support values 1 . 0 / 93 ) , a set of four species represented by a single taxon in [
species being its closest relatives ( support values 0 . 76 / 54 ) , although there is strong support for its association with
being more closely related , but with poor support ( 0 . 37 / 46 ) . within
] are recovered as monophyletic with strong support as are many of the species - level relationships within these groups . however , within the speciose
and northwestern groups , in which many new taxa have been added , species relationships are more highly modified . in the first of these , the basal division into reciprocally monophyletic
groups is not supported , and the former makes the latter paraphyletic . relationships among species groups in
remain unresolved , with most groups connected by exceptionally short internodes . there are two exceptions . the
occurred in the early cenozoic ( 66–43 ma ) . this is a similar pattern as observed in other gekkonids , in which relatively species - rich regional radiations undergo initial diversification in the early cenozoic ( e . g . [
species groups are indicative of a relatively high diversification rate in the mid - cenozoic ~ 30–35 ma . the lineage through time ( ltt ) plot shows that the rate of lineage accumulation remains steady or slowly increases to this point after which there is a noticeable decline ( fig .
) . the overall trend is of significantly decreasing lineage accumulation through time ( mean γ value = −5 . 8 , p < 1 x 10
group suggests that being large - bodied is ancestral for the group ( fig .
) . a shift to small body size occurred once early in the evolutionary history of the group , and there have been only two reversals . reconstruction of habitat preference is more equivocal , but the common ancestor of the group is most commonly reconstructed as a generalized climber ( in 80 % of reconstructions ) . what is clear is that more shifts in habitat preference have occurred than shifts in body size , with approximately 26 transitions indicated in total , most commonly between rock - dwelling and terrestrial habitat preferences . although both habitat and body size are estimated to have shifted multiple times , including reversals , correlation between the two traits is not particularly strong based on fits of bm and ou models — out of 100 , 000 model fits , the bm model incorporating only a single global svl optimum was favored according to the aic 31 % of the time ( fig .
ancestral states for body size and habitat preference , based on 100 stochastic character maps for each trait on the maximum clade credibility tree . range size values for each species are given to the right of each terminal branch
histogram of aicc values for model fits of brownian motion ( bm ) and ornstein - uhlenbeck ( ou ) models of trait diversification estimated in ouwie . a habitat preference vs . svl . b habitat preference vs . range size . c svl vs . range size
in contrast , both body size and habitat preference are strongly correlated with range size ( fig .
) . range size displays significant phylogenetic signal based on pagel’s λ ( λ = 0 . 46 , p = 0 . 13 ) , but the estimate of
) . trait - associated estimates of speciation and extinction rates are less variable ( fig .
) . small - bodied species are estimated to have slightly higher speciation ( mean λ [ small - bodied ] = 0 . 055 ; mean λ [ large - bodied ] = 0 . 040 ) and lower extinction rates , but there is extensive overlap . habitat - associated estimates of diversification rate also overlap , especially between terrestrial species and generalized climbers , although rock - dwelling species are estimated to have speciated at somewhat higher rates ( mean λ [ generalized climber ] = 0 . 032 ; mean λ [ terrestrial ] = 0 . 012 ; mean λ [ rock - dwelling ] = 0 . 065 ) .
trait - associated estimates of speciation and extinction generated using diversitree . intermediate colors indicate overlap . a habitat - associated speciation rate . b habitat - associated extinction rate . c body size - associated speciation rate . d body - size associated extinction rate
] . in many cases , these factors may be interlinked . in this study , we focus on two traits , body size and habitat requirements , that were expected to affect dispersal ability either directly because smaller organisms , including some lizards , may disperse shorter distances [
group the smaller - bodied species occupying more patchily distributed habitats are the species with the smallest geographic ranges . other studies that have measured dispersal ability directly have shown that reduced dispersal ability does not always lead to reduced range size [
and its relatives our data suggest that dispersal ability and range size are correlated . traits affecting dispersal ability are likely not the only factors affecting range size , however . minimally , it is likely that geographic barriers , including major river systems and mountain ranges , also play a significant role in restricting the ranges occupied by individual species . for example , the species
group is similar to patterns documented in many lineages that are often attributed to reduced ecological opportunity through time as niches are filled ( e . g . [
group , a general pattern of morphological conservatism within species groups , exemplified by the small number of shifts in body size ( fig .
] through time is in line with expectations if ecological opportunity has decreased through time . however , shifts in habitat use are more frequent , and the number of co - occurring
group species varies from 1 to 13 , suggesting that ecological niche space has not been exhausted . an alternative explanation that may also partly explain the observed rate slowdown is a geographic model as described above . in clades dominated by allopatric speciation , diversification rates may decline as vicariance events affect fewer species as species’ geographic ranges decline through time [
, which includes mostly large - bodied habitat generalists ( i . e . , species with large geographic ranges ) , compared to
as indicated above , a jump in lineage accumulation coincident with the appearance of habitat - specialist clades in the mid - cenozoic ~ 30–35 ma is contrary to the general pattern of declining diversification rate through time . it is possible that climatic or geomorphic processes active at this time were especially favorable for isolating lineages , resulting in increased speciation . major periods of tectonic uplift in eastern and southern africa did not commence until approximately the oligocene - miocene boundary ( 23 ma ) [
] , making large - scale geomorphological change incompatible with the observed rate increase . however , a major climatic regime shift did occur at the eocene - oligocene boundary : a global cooling associated with antarctic glaciation [
group geckos , potentially facilitating rapid radiation . a similar pattern occurs in forest - adapted chameleons , where rapid radiation is coincident with wide availability of suitable habitat , in the case of chameleons during the eocene [
] , which is why we refrain from ascribing statistical significance to these results . alternate methods of trait - dependant diversification ( e . g . [
] ) likewise are best suited to larger data sets . one possible way to increase data set size is to incorporate taxa from across the afro - malagasy clade of geckos , but the interrelationships of genera within this large radiation are still poorly resolved and relevant trait data are missing for many species . finally , range size estimates are based on known collection localities and a correct interpretation of species - level taxonomy in the group . collecting effort varies greatly by country , with , for example , less than 10 , 000 amphibian and reptile collection records in angola , 38 , 000 in namibia , and > 100 , 000 in south africa [
] . some species also vary phenotypically and have named subspecies that further study may reveal to warrant specific status ( e . g . ,
] ) . however , given that trait - associations with range size varied by orders of magnitude , we do not expect refinement of species’ range limits or taxonomy to strongly influence our results .
beyond interpretation of evolutionary patterns , the results of this study also have significant implications for taxonomy and conservation . the phylogenetic results indicate that
. performing a shimodaira - hasegawa ( sh ) test also shows that the likelihood of our best - scoring tree ( lnl −97035 . 168404 ) is not significantly higher than the likelihood of a tree in which
] . thus , we suggest that taxonomic decisions regarding these species be delayed until each species’ phylogenetic position is better established . we refer both
group inheriting traits promoting smaller ranges also inherit traits promoting greater rarity . however , our analyses show these same traits to be associated with higher rates of diversification . given the difficulty in estimating extinction rates from phylogenies [
group . notwithstanding this difficulty , these results stress the importance of defining a frame of reference when measuring evolutionary “success . ” in the case of the
group , more widespread , common species tend to belong to relatively species - poor subclades .
group points to a history of geographic isolation contributing significantly to the group’s species richness compared to other african geckos . even so , some aspects of diversification in the
group , including early evolution of divergent traits within the group , are consistent with patterns observed in classic adaptive radiations . in this sense , the process of diversification of
] ) on the other . it is likely that many other species - rich groups share this same intermediate pattern .
we thank the late donald g . broadley , mirko barts , jon boone , william r . branch , marius burger , and krystal tolley for providing access to some specimens used in the study . johan marais , bill branch , werner conradie , trip lamb , don broadley , randy babb , paul moler , ross sadlier , glenn shea , jens vindum , and many villanova university masters students participated in collecting .
this project was funded by the university of michigan ( mph ) , nsf deb - 0515909 ( amb , trj ) , deb - 0844523 ( amb , trj ) , deb - 1019443 ( amb ) , deb - 1556255 ( amb ) , deb - 1556585 ( mph ) , and ef - 1241885 ( subaward 13–0632 ) ( amb ) . funding bodies played no role in study design or implementation .
mph and amb designed the project . data were collected by mph , trj , and amb . data analysis was performed by mph . mph wrote the paper with input from trj and amb . all authors read and approved the final manuscript .
use of animals to obtain tissue samples for genetic analysis was reviewed and approved by the villanova university institutional animal care and use committee . field work in southern africa that generated the tissues used in this work was made possible by permits ( 1987–2014 ) to amb from the relevant wildlife authorities of south africa , namibia and zimbabwe .
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) applies to the data made available in this article , unless otherwise stated .
froese r , pauly d . fishbase , version ( 10 / 2015 ) .
lamb t , bauer am . footprints in the sand : independent reduction of subdigital lamellae in the namib–kalahari burrowing geckos . proc r soc lond b biol sci . 2006 ; 273 : 855–64 .
gamble t , greenbaum e , jackman tr , russell ap , bauer am . repeated origin and loss of adhesive toepads in geckos . plos one . 2012 ; 7 : e39429 .
bauer am , branch wr . the herpetofauna of the richtersveld national park and the adjacent northern richtersveld , northern cape province , republic of south africa . herpetological nat hist . 2001 ; 8 : 111–60 .
schluter d . the ecology of adaptive radiation . oxford : oxford university press ; 2000 .
losos jb , mahler dl . adaptive radiation : the interaction of ecological opportunity , adaptation , and speciation . in : bell ma , futuyma dj , eanes wf , levinton js , editors . evolution since darwin : the first 150 years . sunderland : sinauer ; 2010 . p . 381–420 .
alfaro me , brock cd , banbury bl , wainwright pc . does evolutionary innovation in pharyngeal jaws lead to rapid lineage diversification in labrid fishes ? bmc evol biol . 2009 ; 9 : 1 .
rutschmann s , matschiner m , damerau m , muschick m , lehmann mf , hanel r , salzburger w . parallel ecological diversification in antarctic notothenioid fishes as evidence for adaptive radiation . mol ecol . 2011 ; 20 : 4707–21 .
deutsch jc . colour diversification in malawi cichlids : evidence for adaptation , reinforcement or sexual selection ? biol j linn soc . 1997 ; 62 : 1–4 .
masta se , maddison wp . sexual selection driving diversification in jumping spiders . proc natl acad sci . 2002 ; 99 : 4442–7 .
wiens jj , graham ch , moen ds , smith sa , reeder tw . evolutionary and ecological causes of the latitudinal diversity gradient in hylid frogs : treefrog trees unearth the roots of high tropical diversity . am nat . 2006 ; 168 : 579–96 .
group ( squamata : gekkonidae ) . afr j herpetol . 2005 ; 54 : 105–29 .
bauer am , heinicke mp , jackman tr , branch wr . systematics of the
, gekkonidae ) and their relatives . j zool . 1993 ; 229 : 353–84 .
from a dry forest in madagascar . amphibia - reptilia . 2005 ; 26 : 475–83 .
regalado r . does dichromatism variation affect sex recognition in dwarf geckos ? ethol ecol evol . 2015 ; 27 : 56–73 .
coyne ja . genetics and speciation . nature . 1992 ; 355 : 511–5 .
coyne ja , orr ha . speciation . sunderland : sinauer associates ; 2004 .
jablonski d . heritability at the species level : analysis of geographic ranges of cretaceous mollusks . science . 1987 ; 238 : 360–3 .
gaston kj . species - range size distributions : products of speciation , extinction and transformation . philos transact a math phys eng sci . 1998 ; 353 : 219–30 .
webb tj , gaston kj . on the heritability of geographic range sizes . am nat . 2003 ; 161 : 553–66 .
waldron a . null models of geographic range size evolution reaffirm its heritability . am nat . 2007 ; 170 : 221–31 .
vamosi sm , vamosi jc . perspective : causes and consequences of range size variation : the influence of traits , speciation , and extinction . front biogeography . 2012 ; 4 : 168–77 .
lee ms , skinner a , camacho a . the relationship between limb reduction , body elongation and geographical range in lizards (
slatyer ra , hirst m , sexton jp . niche breadth predicts geographical range size : a general ecological pattern . ecol lett . 2013 ; 16 : 1104–14 .
maddison wp , midford pe , otto sp . estimating a binary character’s effect on speciation and extinction . syst biol . 2007 ; 56 : 701–10 .
branch wr . field guide to snakes and other reptiles of southern africa . 3rd ed . struik : cape town ; 1998 .
gaston kj , blackburn tm . range size - body size relationships : evidence of scale dependence . oikos . 1996 ; 479–85 .
lancaster n . late quaternary paleoenvironments in the southwestern kalahari . palaeogeogr palaeoclimatol palaeoecol . 1989 ; 70 : 367–76 .
partridge tc . the evidence for cainozoic aridification in southern africa . quat int . 1993 ; 17 : 105–10 .
thomas ds , shaw pa . the evolution and characteristics of the kalahari , southern africa . j arid environ . 1993 ; 25 : 97–108 .
group geckos of southern africa : new hypotheses . afr j herpetol . 1999 ; 48 : 53–62 .
lamb t , bauer am . phylogenetic relationships of the large - bodied members of the african lizard genus
bauer am . phylogeny and biogeography of the geckos of southern africa and the islands of the western indian ocean : a preliminary analysis . in : peters g , hutterer r , editors . vertebrates in the tropics . bonn : zoologisches forschungsinstitut und museum a . koenig ; 1990 . p . 274–84 .
heinicke mp , greenbaum e , jackman tr , bauer am . phylogeny of a trans‐wallacean radiation ( squamata , gekkonidae ,
) supports a single early colonization of australia . zool scr . 2011 ; 40 : 584–602 .
heinicke mp , greenbaum e , jackman tr , bauer am . evolution of gliding in southeast asian geckos and other vertebrates is temporally congruent with dipterocarp forest development . biol lett . 2012 ; 8 : 994–7 .
heinicke mp , daza jd , greenbaum e , jackman tr , bauer am . phylogeny , taxonomy and biogeography of a circum - indian ocean clade of leaf - toed geckos ( reptilia : gekkota ) , with a description of two new genera . syst biodivers . 2014 ; 12 : 23–42 .
and the intraspecific evolution of coastal melanism in the western rock skink . afr zool . 2010 ; 45 : 147–59 .
hall ta . bioedit : a user - friendly biological sequence alignment editor and analysis program for windows 95 / 98 / nt . nucleic acids symp ser . 1999 ; 41 : 95–8 .
kearse m , moir r , wilson a , stones - havas s , cheung m , sturrock s , buxton s , cooper a , markowitz s , duran c , thierer t . geneious basic : an integrated and extendable desktop software platform for the organization and analysis of sequence data . bioinformatics . 2012 ; 28 : 1647–9 .
larkin ma , blackshields g , brown np , chenna r , mcgettigan pa , mcwilliam h , valentin f , wallace im , wilm a , lopez r , thompson jd . clustal w and clustal x version 2 . 0 . bioinformatics . 2007 ; 23 : 2947–8 .
laslett d , canbäck b . arwen , a program to detect trna genes in metazoan mitochondrial nucleotide sequences . bioinformatics . 2008 ; 24 : 172–5 .
lanfear r , calcott b , ho sy , guindon s . partitionfinder : combined selection of partitioning schemes and substitution models for phylogenetic analyses . mol biol evol . 2012 ; 29 : 1695–701 .
stamatakis a . raxml version 8 : a tool for phylogenetic analysis and post - analysis of large phylogenies . bioinformatics . 2014 ; 30 : 1312–3 .
drummond aj , suchard ma , xie d , rambaut a . bayesian phylogenetics with beauti and the beast 1 . 7 . mol biol evol . 2012 ; 29 : 1969–73 .
kumar s , hedges sb . a molecular timescale for vertebrate evolution . nature . 1998 ; 392 : 917–20 .
reisz rr , müller j . molecular timescales and the fossil record : a paleontological perspective . trends genet . 2004 ; 20 : 237–41 .
kluge ag . cladistic relationships of sphaerodactyl lizards . am mus novit . 1995 ; 3139 : 1–23 .
iturralde - vinent ma , macphee rd . age and paleogeographical origin of dominican amber . science . 1996 ; 273 : 1850 .
lee ms , hutchinson mn , worthy th , archer m , tennyson aj , worthy jp , scofield rp . miocene skinks and geckos reveal long - term conservatism of new zealand’s lizard fauna . biol lett . 2009 ; 5 : 833–7 .
hutchinson mn . the first fossil pygopodid ( squamata , gekkota ) and a review of mandibular variation in living species . memoirs queensland museum . 1997 ; 41 : 355–66 .
lee ms , oliver pm , hutchinson mn . phylogenetic uncertainty and molecular clock calibrations : a case study of legless lizards ( pygopodidae , gekkota ) . mol phylogenet evol . 2009 ; 50 : 661–6 .
macey jr , wang y , ananjeva nb , larson a , papenfuss tj . vicariant patterns of fragmentation among gekkonid lizards of the genus
produced by the indian collision : a molecular phylogenetic perspective and an area cladogram for central asia . mol phylogenet evol . 1999 ; 12 : 320–32 .
loveridge a . revision of the african lizards of the family gekkonidae . bull mus comp zool . 1947 ; 98 : 1–469 .
alexander gj , marais j . a guide to the reptiles of southern africa . cape town : struik ; 2007 .
bates mf , branch wr , bauer am , burger m , marais j , alexander gj , de villiers ms ( eds ) . atlas and red list of the reptiles of south africa , lesotho and swaziland . pretoria : south african national biodiversity institute ; 2014 .
revell lj . phytools : an r package for phylogenetic comparative biology ( and other things ) . methods ecol evol . 2012 ; 3 : 217–23 .
r core team . r : a language and environment for statistical computing . vienna , austria : r foundation for statistical computing ; 2015 .
blomberg sp , garland t , ives ar . testing for phylogenetic signal in comparative data : behavioral traits are more labile . evolution . 2003 ; 57 : 717–45 .
pagel m . inferring the historical patterns of biological evolution . nature . 1999 ; 401 : 877–84 .
pybus og , harvey ph . testing macro–evolutionary models using incomplete molecular phylogenies . proc r soc lond b biol sci . 2000 ; 267 : 2267–72 .
beaulieu jm , jhwueng dc , boettiger c , o’meara bc . modeling stabilizing selection : expanding the ornstein–uhlenbeck model of adaptive evolution . evolution . 2012 ; 66 : 2369–83 .
collar dc , schulte ii , james a , losos jb . evolution of extreme body size disparity in monitor lizards (
huelsenbeck jp , nielsen r , bollback jp . stochastic mapping of morphological characters . syst biol . 2003 ; 52 : 131–58 .
fitzjohn rg . quantitative traits and diversification . syst biol . 2010 ; 59 : 619–33 .
fitzjohn rg . diversitree : comparative phylogenetic analyses of diversification in r . methods ecol evol . 2012 ; 3 : 1084–92 .
rabosky dl , goldberg ee . model inadequacy and mistaken inferences of trait - dependent speciation . syst biol . 2015 ; 64 : 340–55 .
rabosky dl , huang h . a robust semi - parametric test for detecting trait - dependent diversification . syst biol . 2016 ; 65 : 181–93 .
davis mp , midford pe , maddison w . exploring power and parameter estimation of the bisse method for analyzing species diversification . bmc evol biol . 2013 ; 13 : 1 .
group of southern african geckos . afr zool . 2002 ; 37 : 209–20 .
groups ( reptilia : gekkota : gekkonidae ) of southern africa , and with the description of eight new species . proc calif acad sci . 2006 ; 57 : 595–709 .
group of geckos ( reptilia : squamata : gekkonidae ) . afr zool . 2000 ; 35 : 55–67 .
wood pl , heinicke mp , jackman tr , bauer am . phylogeny of bent - toed geckos (
) reveals a west to east pattern of diversification . mol phylogenet evol . 2012 ; 65 : 992–1003 .
böhning‐gaese k , caprano t , van ewijk k , veith m . range size : disentangling current traits and phylogenetic and biogeographic factors . am nat . 2006 ; 167 : 555–67 .
lester se , ruttenberg bi , gaines sd , kinlan bp . the relationship between dispersal ability and geographic range size . ecol lett . 2007 ; 10 : 745–58 .
olifiers n , vieira mv , grelle ce . geographic range and body size in neotropical marsupials . glob ecol biogeogr . 2004 ; 13 : 439–44 .
machac a , zrzavý j , storch d . range size heritability in carnivora is driven by geographic constraints . am nat . 2011 ; 177 : 767–79 .
sinervo b , calsbeek r , comendant t , both c , adamopoulou c , clobert j . genetic and maternal determinants of effective dispersal : the effect of sire genotype and size at birth in side‐blotched lizards . am nat . 2006 ; 168 : 88–99 .
beier p , noss rf . do habitat corridors provide connectivity ? conserv biol . 1998 ; 12 : 1241–52 .
rabosky dl , lovette ij . density - dependent diversification in north american wood warblers . proc r soc lond b biol sci . 2008 ; 275 : 2363–71 .
gavrilets s , losos jb . adaptive radiation : contrasting theory with data . science . 2009 ; 323 : 732–7 .
burbrink ft , pyron ra . how does ecological opportunity influence rates of speciation , extinction , and morphological diversification in new world ratsnakes ( tribe lampropeltini ) ? evolution . 2010 ; 64 : 934–43 .
mahler dl , revell lj , glor re , losos jb . ecological opportunity and the rate of morphological evolution in the diversification of greater antillean anoles . evolution . 2010 ; 64 : 2731–45 .
pigot al , phillimore ab , owens ip , orme cd . the shape and temporal dynamics of phylogenetic trees arising from geographic speciation . syst biol . 2010 ; 59 : 660–73 .
moen d , morlon h . why does diversification slow down ? trends ecol evol . 2014 ; 29 : 190–7 .
partridge tc . of diamonds , dinosaurs and diastrophism : 150 million years of landscape evolution in southern africa . s afr j geol . 1998 ; 101 : 167–84 .
sepulchre p , ramstein g , fluteau f , schuster m , tiercelin j , brunet m . tectonic uplift and eastern africa aridification . science . 2006 ; 313 : 1419–23 .
cowling rm , proches s , partridge tc . explaining the uniqueness of the cape flora : incorporating geomorphic evolution as a factor for explaining its diversification . mol phylogenet evol . 2009 ; 51 : 64–74 .


acryptolechia torophanes is a moth in the depressariidae family .

it was described by meyrick in 1935 .

it is found in china ( henan , shaanxi , zhejiang , hubei ) and korea .

acryptolechia torophanes

this is the place for torophanes definition . you find here torophanes meaning , synonyms of torophanes and images for torophanes copyright 2017 © urltoken
acryptolechia torophanes ; lvovsky , 2010 , zool . zh . 89 ( 3 ) ent . review 90 ( 2 ) : 255
here you will find one or more explanations in english for the word torophanes . also in the bottom left of the page several parts of wikipedia pages related to the word torophanes and , of course , torophanes synonyms and on the right images related to the word torophanes .
cryptolechia torophanes meyrick , 1935 ; mat . microlep . fauna chin . prov . : 81
acryptolechia facunda ; lvovsky , 2010 , zool . zh . 89 ( 3 ) ent . review 90 ( 2 ) : 255
acryptolechia malacobyrsa ; lvovsky , 2010 , zool . zh . 89 ( 3 ) ent . review 90 ( 2 ) : 255
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= meleonoma ; park & park , 2016 , j . asia - pacif . biodiv . 9 ( 4 ) : 485
assam , japan , n . china , e . china . see [ maps ]
leptosaces facunda meyrick , 1910 ; j . bombay nat . hist . soc . 20 ( 1 ) : 155 ; tl : khasis
japan , korea , china ( fujian , henan , jiangxi , shaanxi , sichuan ) , taiwan . see [ maps ]
= ; ridout , 1981 , ins . matsumurana 24 : 35 ; wang , 2003 , ent . sinica 9 ( 3 ) : 196
china ( henan , shaanxi , zhejiang , shanghai ) , korea . see [ maps ]
[ maps ] warning ! the maps are automatically generated from the textual information , and the process does not always produce acceptable result ; see about maps for more info .
wang , 2003 a study of cryptolechia zeller ( lepidoptera : oecophoridae ) in china ( i ) , with descriptions of fifteen new species ent . sinica 9 ( 3 ) : 195 - 213
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